RHODORA Journal of the New England Botanical Club CONTENTS Effect of achene morphology and mass on germination and seedling growth of Boltonia decurrens (Asteraceae), a threatened floodplain species. Marian Smith and John Cawly | Crossing effects on seed viability and experimental germination of the Federal Threatened Platanthera leucophaea (Orchidaceae). Marlin L. Bowles, Karel A. Jacobs, Lawrence W. Zettler, and Tonya Wilson Delaney 14 The identity and history of Myrica caroliniensis (Myricaceac). Robert L. i 31 A Honus inventory of Manatee Springs State Park, Levy County, Florida. berely J. Gulledge and Walter S. Judd 42 NEW ENGLAND NOTE Aneura maxima (Hepaticae: Aneuraceae) in Maine, U.S.A. Norton 77 NOTES Sci dpe ancistrochaetus Pe First record in Canada. Stuart G. and Gordon C. Thc 83 Schizaea Ente in North Carolina. Richard J LeBlond and Alan S. Weakley 86 IN MEMORIAM Rolla Milton Tryon, Jr. 1916-2001: Scientist, Teacher, and Mentor. David S. Conant, Gerald J. Gastony, and David S. Barrington 92 BOOK REVIEW Lichens of North America 96 NEW BOOKS 100 NEBC MEETING NEWS 101 ANNOUNCEMENT Invasive Plant Survey of New England: A call for volunteers ............ 106 Checklist for Contributors to Rhodora 107 NEBC Membership Form 114 Order Form for Index to Volumes 76—100 115 Statement of Ownership 116 BC Officers and Council Members inside back cover Vol. 104 Winter, 2002 No. 917 Issued: April 15, 2002 The New England Botanical Club, Inc. 22 Divinity Avenue, Cambridge, Massachusetts 02138 RHODORA JANET R. SULLIVAN, Editor-in-Chief Department of Plant Biology, University of New Hampshire, Durham 24 e-mail: janets@cisunix.unh.edu ANTOINETTE P. HARTGERINK, Managing Editor Department of Plant Biology, University of New Hampshire, 24 e-mail: Ahartgrink@aol.com Associate Editors ROBERT I, BERTIN STEVEN R. HILL DAVID S. CONANT THOMAS D, LEE GARRETT E. CROW THOMAS MIONE KANCHI GANDHI—Latin diagnoses and nomenclature RHODORA (ISSN 0035-4902). 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RHODORA, Vol. 104, No. 917, pp. 1-13, 2002 EFFECT OF ACHENE MORPHOLOGY AND MASS ON GERMINATION AND SEEDLING GROWTH OF BOLTONIA DECURRENS (ASTERACEAB), A THREATENED FLOODPLAIN SPECIES MARIAN SMITH Department of Biological aaa Southern [linois University, Edwardsville, IL 62026-1651 e-mail: Bao eri JOHN CAWLY Department of Plant Microbiology and secure ‘sai of Missouri, Columbia, MO 65211 ( e-mail: jdc3d9@ missou. on ABSTRACT. Boltonia decurrens (Asteraceae), a plant species endemic to the Illinois River Valley, is threatened with extinction. Alterations of the hydrologic regime of the river have resulted in habitat loss and population decline. Lack of information about the complex life cycle of this species frustrates efforts to develop an effective recovery ch . An essential part of ee recovery plan is an understanding of seed germination, seedling recruit- m ind early growth, and how each contributes to the maintenance of a Shpdtien: In B. decurrens, the dimorphic achenes have different masses and may provide different dispersal mechanisms. This study examined the effect of achene morphology and mass on seed germination and early seedling growth of B. decurrens under controlled environmental conditions. There was no difference in timing of germination of disk and ray achenes in B. decur- rens, however, there was a distinct difference in early growth of seedlings de aved from disk versus ray ae es and from larger a achenes compared to smaller ones. Disk achenes, which ae seedlings with more leaf area during the first 1O—15 days, may provide a competitive advantage over those produced by ray achenes. Since leaf area and photosynthetic rates are closely correlated, seedlings with more leaf area early in their development may also be more competitive with seedlings of other species. Key Words: dimorphic seeds, germination, seedling growth, seed _ size, threatened species, floodplain, Boltonic Boltonia decurrens (Torr. & A. Gray) A. Wood (Asteraceae) is an herbaceous perennial whose distribution is restricted to the Hlinois River floodplain (Torrey and Gray 1841; U.S. Fish and Wildlife Service 1990). The species is on the Federal List of threatened species (U.S. Fish and Wildlife Service 1988), and is currently listed as a Species of Concern in Missouri (Missouri | i) Rhodora [Vol. 104 Department of Conservation 1999) and as Threatened in Illinois (Herkert 1991). Inflorescences are borne on panicles and produce prolific numbers of dimorphic achenes (Smith and Keevin 1998). Additionally, vegetative ramets that overwinter and reproduce sexually the following year are produced at the base of the se- nescing flowering plants each fall (Redmond 1993; Smith 1991; Smith et al. 1998). Despite these reproductive strategies, the con- struction of levees and navigation dams along the Illinois River have resulted in habitat loss and a decline in population size and number (Schwegman and Nyboer 1985; Smith et al. 1998; U.S. Fish and Wildlife Service 1990). Dissimilarities in seed morphology within plant taxa may con- tribute to different dispersal and germination patterns as well as different growth and fitness of the resulting plants (Banovetz and Scheiner 1992; Zhang 1993). Disk flowers of Boltonia decurrens produce flattened, heart-shaped achenes that are characterized by a pappus of two bristles. Disk achenes average 1.8 mm in length and 1.3 mm in width with an average mass of 0.1 mg (Smith and Keevin 1998). Achenes produced from ray flowers are smaller and wedge-shaped, possessing a distinct third side, and they have greatly reduced bristles. Ray achenes average |.3 mm in length and 0.9 mm in width, and they average 0.05 mg in mass (Smith and Keevin 1998). The increased surface area/mass ratio of the disk achenes provides greater buoyancy and allows them to float for extended periods of time (> 30 days; Smith and Keevin 998), and this may facilitate the establishment of remote popu- lations after flood waters recede. The wedge-shaped ray achenes do not float and may contribute to the maintenance of the species at or near an extant population (Smith and Keevin 1998; Smith etal. 1998); Germination and early seedling development are the most crit- ical stages in the life cycle of a plant (Harper et al. 1970), and any variation in achene morphology or mass that could affect germination or early growth could also influence the establish- ment of new populations or the maintenance of extant popula- tions. Although germination studies have been conducted on Bol- tonia decurrens (Baskin and Baskin 1988; Smith and Keevin 1998; Smith et al. 1995), little is known of germination patterns or seedling growth specific to achene morphology or mass. The present study examines the effect of achene morphology and mass on germination and early seedling growth. — 2002]. Smith and Cawly—Boltonia decurrens Seedlings 3 MATERIALS AND METHODS Achenes were collected from West Alton, St. Charles County, Missouri (lat. 38°52'06"N, long. 90°12'22”W) and maintained at 5°C for two years before the initiation of this study. Size classes of each achene morphology were differentiated by sorting through a series of screens with mesh sizes of 0.841 mm, 0.595 mm, and 0.420 mm, followed by manual segregation of each morphological type using a stereoscope. The largest size class (> 0.841 mm) contained predominantly disk achenes (D1); the smallest (> 0.420 mm but < 0.595 mm) contained mature ray achenes (R3) and a very small proportion of immature or non- viable disk achenes (lacking a visible embryo). Both mature disk and ray achenes (D2 and R2) were represented in the intermediate size class (> 0.595 mm but < 0.841 mm). Seeds with evidence of herbivory, or apparently nonviable (no visible embryo), were discarded. Otherwise, seeds were selected randomly to minimize possible sampling bias. Each of the ten replicates consisted of 25 disk and 25 ray achenes in each of two size classes, for a total of 1000 achenes. Dry mass was recorded for each sample of 25 seeds, and a mass: size correlation was calculated. Since the achenes require light for germination (Baskin and Baskin 1988; Smith and Keevin 1998), they were germinated in 10 cm square pots on the surface of a commercial, peat-based growing medium in a Sherer CEL-25 7HL environmental cham- ber at 20°C and 200 pmol m ?s'!' PPF (photosynthetic photon flux), which was measured using an LI-185B quantum meter and LI-190SB quantum sensor (LI-COR, Lincoln, NE). Germination was determined by radicle emergence, recorded daily, and seed- lings were identified with a color-coded pin. Length and width measures of the cotyledons were taken after five days of growth. Measures were obtained for both cotyledons and true leaves at 10 and 15 days. Cotyledons and primary leaves are not lobed, do not have serrated edges, and are approximately rectangular; there- fore, leaf area was calculated using the algorithm for area of a rectangle (L X W). After 15 days the seedlings were transferred to a greenhouse, where they were initially placed under 50% shade cloth to minimize photodamage and were exposed to higher ambient light levels over a two-week period. Light levels were measured in the greenhouse at 12 noon daily during the study (June and July) and averaged 1500 + 350 wmol m~’ s~! PPE Pots 4 Rhodora (Vol. 104 were watered daily and rotated to minimize environmental vari- ation due to position. Seedling height was measured at 30 days and at 60 days. Because Boltonia decurrens is endangered, plants were not harvested for biomass measurements, but were trans- planted into the population from which the seeds had been col- lected. All analyses were performed using SYSTAT 5.2 (SPSS, Chi- cago, Illinois). The achene size class and mass relationship was determined using linear regression, and germination and seedling survival data were analyzed using a chi square statistic of a con- tingency table (Steel and Torrie 1980). For the x’ analyses of germination and survival, data from replicates were pooled within each achene class. The unequally represented size classes, which resulted from differences in the numbers of seeds that germinated within each achene class, produced a non-orthogonal design that would affect the relationship of the other classes in calculating the F-ratio (Steel and Torrie 1980); therefore, we used Multivariable General Linear Hypothesis (MGLH; type III sum of squares) for all anal- yses of variance (ANOVAs) of leaf area and height. Log trans- formation was used to normalize leaf area and height data (Steel and Torrie 1980). Linear contrast analyses were used for pairwise comparisons of seedling leaf area and seedling height for each size class and morphology combination in accordance with Steel and Torrie (1980). RESULTS Germination of achenes and seedling survival. There was no significant difference in germination among disk and _ ray achenes of any class (y? = 2.751; P = 0.432; df = 3; Figure 1). Of the 1000 seeds used in this experiment, 667 germinated: 339 disk achenes and 328 ray achenes. Although there was no statis- tical difference in survival among achene classes (x? = 2.570; P = (0.497: df = 3), there was a trend for a decrease in survival with a decrease in achene size (Figure |). Disk and ray achenes of all classes demonstrated the same germination pattern (Figure 2): germination peaked on day 4, with no germination occurring before day 3 or after day I1. There was a positive linear rela- tionship (r? = 0.8165; P < 0.05; df = 3) between achene size (area) and mass (1.e., the larger the achene, the greater the mass). 2002]. Smith and Cawly—Boltonia decurrens Seedlings 5 80 - earea Gg =3Germination iL 50 [= Survival ae e Oo 604 40 _ = o w > £ Ss i E -30 3 “”) 2 40 4 ~ oO c ~ o c 7 20-2 oO o . oO Oo. 20 | 10 0 , + 0 D1 D2 R2 R3 Achene Class gure |. Percent germination and percent seedling oo or achenes from all size classes. = disk achene size class | (> 41 mm). D2 = disk achene size class 2 (> 0.420 mm, but < 0.841 es = ray achene size class 2 (> 0.420 mm, but < 0.841 mm). R3 = ray achene size class 3 (< 0.420 mm). Seedling growth. One-way analysis of variance (ANOVA) of leaf area measurements taken after 10 days of growth indicated a statistical significance (Table |). Linear contrast analysis com- paring leaf areas of seedlings from both seed types in all size classes indicated statistical significance when contrasting all disk achenes versus all ray achenes (Table 2; Figure 3), the two size classes of ray achenes (R2 and R3), and the largest size class (D1) versus the smallest class (R3; Table 2; Figure 4). Similarly, ANOVA of 15-day measurements showed statistical difference (Table 1), and linear contrast analysis exhibited statis- tical significance in comparing all disk achenes versus all ray achenes (Table 2; Figure 3). Additionally, at 15 days, the larger class of disk achenes (DI) proved to have significantly greater leaf area when compared with all other size classes (Table 2; Figure 4). No statistical significance was found by ANOVA of plant height measurements taken after 30 days or after 60 days of growth; furthermore, linear contrast analysis revealed no sta- tistically significant differences in any size class or morphology comparison at 30-day or 60-day measurements. There is, how- 6 Rhodora [Vol. 104 Number of Seeds Germinated Figure 2. Number of achenes of Boltonia decurrens germinating at 22°C and 200 wmol m * s | PPF over a 12-day period. DI = disk achene size class | (> 0.841 mm). D2 = disk achene size class 2 (> 0.420 mm, but < 0.841 mm). R2 = ray achene size class 2 (> 0.420 mm, but < 0.841 mm). R3 = ray achene size class 3 (< 0.420 mm). Table 1. Results of the one-way ANOVAs for seedling leaf area at 10 and I5 days, and seedling height at 30 and 60 days among all four achene classes. Source SSE df MSE F P 10 days Type 1.923 3 0.641 4.498 0.004 Error 28.517 200 0.143 15 days Type 1.499 3 0.500 4.420 0.005 Error 16.501 146 O.113 30 days Type 0.313 3 0.104 1.358 0.256 Error 19,273 25] 0.077 60 days Type 0.196 3 0.128 1.253 0.29] Error 25.709 251 0.102 Table 2. Linear contrast analyses for seedling leaf area and seedling height. DI = disk achene size class | (> 0.841 mm). D2 = disk achene size class 2 (> 0.420 mm, < 0.841 mm). R2 = ray achene size class 2 (> 0.420 mm, < 0.841 mm), R3 = ray achene size class 3 (< 0.420 mm). Disk vs. Ray D1 vs. D2 DI vs. R2 DI vs. R3 D2 vs. R2 D2 vs. R3 R2 vs. R3 Contrasts Ee P F P F P Fi P F P F P iA P 10 days 8.461 0.004 0.067 0.796 1.308 0.254 10.842 0.001 0.768 0.382 9.079 0.003 4.689 0.038 15 days 4.870 0.029 7.684 0.006 6.618 0.011 11.173) 0.001 0.039 0.843 0.301 0.584 0.559 0.456 30 days 2.277 0.133 1.770 O.185 3.023 0.083 2.821 0.094 0.174 0.677 0.194 0.660 0.002 0.961 60 days 1.910 0.168 1.816 0.179 2.400 0.123 2.887 0.091 0.045 0.833 0.197 6 0.060 0.807 Qa [ZOOT SSUI[P9IG SUatNIap DIUO]JOG—A|[MLD pue YS 8 Rhodora [Vol. 104 Leaf Area (mm’) Age (days) Figure 3. Mean leaf area (+ SE) produced by disk and ray achenes, regardless of size class, after 10 and 15 days of growth. Bars with different letters are significantly different between achene types (see Table 2 for P values). ever, an indication that height in the plants produced from the smallest size class, R3, began to lag behind the others in growth at 60 days (Figure 5). DISCUSSION Our results suggest that seed morphology influences seedling establishment during the early stages of development; however, any differential competitive advantage due to achene type or mass is less obvious after 10—I5 days of growth. Grime (2001) pro- posed that differences in seed morphology may influence seedling establishment and growth, and our results with Boltonia decur- rens appear to corroborate his findings. Total leaf area after 10— 15 days of growth differed significantly between disk and ray achenes. Additionally, comparison of 10-day growth was signif- icant by size class (.e., larger achenes produced seedlings with greater leaf area). Since the photosynthetic area of the cotyledon, rather than its mass or stored energy, is the primary consideration in early seedling development (Harper et al. 1970), achenes pro- 2002] Smith and Cawly—Boltonia decurrens Seedlings 9 40 4 a Mmmm 10 day b (fem) «15 day | _ b —~ 30 tf N £ 7 D b = a © 7 a T @ 204 0.841 mm). D2 = disk achene size class 2 (> 0.420 mm, but < 0.841 mm). R2 = ray achene size class 2 (> 0.420 mm, but < 0.841 mm). R3 = ray achene size class 3 (< 0.420 mm). Bars with different letters are significantly different for each comparison between achene classes (see Table 2 for P values). pane ducing small cotyledons would have relatively less potential for growth than those possessing large cotyledons. After 15 days of growth, the advantage of greater cotyledon area is reinforced by the greater total leaf area (cotyledons plus true leaves). Boltonia decurrens requires light for germination (Baskin and Baskin 1988; Smith and Keevin 1998) and high light for growth and seed production (Smith et al. 1993), and seedlings have high mortality when germinated under plant litter (Smith et al. 1995). In natural populations, seedling establishment is extremely low after one year of succession (< 0.01%; Moss 1997; Smith et al. 1998), and B. decurrens is often completely replaced by com- peting vegetation 3—5 years after population establishment (Schwegman and Nyboer 1985; U.S. Fish and Wildlife Service 1990). If achene morphology or mass determines cotyledon and leaf size for the first 1O—-15 days, greater photosynthetic surface area during this period may enable these seedlings to be more competitive with rapidly growing seedlings of other species. Data 10 Rhodora [Vol. 104 | 40 - ——— | mm 30 day | [== 60 day | | L si ua | Height (cm) D1 D2 R2 R3 Achene Class e 5. Mean seedling height (+ SE) in each achene class after 30 and 60 ae of growth. DI = disk achene size class | (> 0.841 mm). D2 = disk achene size class 2 (> 0.420 mm, but < 0.841 mm). R2 = ray achene size class 2 (> err mm, but < 0.841 mm). R3 = ray achene size class 3 (< 0.420 mm). ee from our study indicate that seedlings from disk achenes, regard- less of size, would have the highest probability of surviving, and that seedlings from larger ray achenes would fare better than those from smaller ones. Although there may have been height or mortality differences among seed morphologies and sizes during the earliest stages of growth, at 30 and 60 days there were no statistically significant differences in either. In the present study, the widely spaced dis- tribution of seedlings across the soil surface minimized interac- tions between individuals. Similar conditions do not exist in the field, however, where thousands of seeds germinate simultaneous- ly. This is particularly so in the case of Bolftonia decurrens, be- cause its achenes and the seeds of other species are often depos- ited in densely packed rows by receding floodwaters (Smith and Keevin 1998; Smith et al. 1995). In this environment, where re- sources are limited and competition increases seedling mortality, a 10-15 day advantage provided by greater photosynthetic sur- face area may be critical for seedling growth and survival. 2002] Smith and Cawly—Boltonia decurrens Seedlings i In other Asteraceae, Fenner (1983) found that performance of seedlings in the period immediately after germination is critical for establishment. Although Fenner’s study was not designed to be a competition experiment, or to represent a field situation, some important inferences can be drawn from his data. Clearly, chronological differences in seedling emergence affect competi- tive interactions among seedlings: those emerging earlier poten- tially shade later-germinating seedlings and inhibit their growth. That this effect is largely due to differential seedling size is sup- ported by the results of Gross (1984), who found that within- species differences in seed size had a significant effect on early seedling growth and survival. Although both types of achenes of Boltonia decurrens have similar temporal patterns of germination, size differences due to achene morphology or mass produce the same result—larger seedlings that are less likely to be overtopped by other seedlings. In B. decurrens, this is particularly important due to its requirement for high light during all stages of growth. A competitive advantage during early seedling development may be essential to the survival of Bolftonia decurrens in its cur- rent habitat. The construction of a series of levees and navigation dams on the Illinois River over the past 70 years has altered the natural flood regime of the [Illinois River Valley (Sparks 1995; Sparks et al. 1998). Areas that once provided the open moist shorelines suitable for the establishment and regeneration of pop- ulations of B. decurrens are now seldom flooded, resulting in the invasion of the sites by a number of aggressive species that are less flood tolerant than B. decurrens (Schwegman and Nyboer 1985; Smith 1991; Smith et al. 1998). In these areas, individuals of B. decurrens become smaller and produce fewer and smaller achenes each year following population establishment (Smith 1993). Seedling survival declines rapidly (Moss 1997) as the number and density of aggressive competitors increase. Recent work by Mettler et al. (2001) and Smith (unpubl. data) indicates that the loss of an annual nutrient pulse in years without floods may contribute to the decline of B. decurrens by reducing plant size and achene number and mass. Information provided in the current study indicates that a re- duction in achene size would affect seedling growth, and adds to the accumulating evidence that alterations in the natural flood regime in the Hlinois River Valley are implicated in the decline of Boltonia decurrens (Smith and Mettler 2002; Smith et al. — LD, Rhodora [Vol. 104 1998), as has been reported for other native species (Sparks 1995; Sparks et al. 1998). This information may help stimulate efforts to restore connections between the river and the floodplain and to re-establish native plant communities in the Illinois River Val- ley. ACKNOWLEDGMENTS. This project was supported by a grant (National Science Foundation DEB95-09763) to M. Smith. LITERATURE CITED BANOVETZ, S. J. AND S. M. SCHEINER. 1992. The effects of seed mass on the seed ecology of Coreopsis lanceolata. Amer. Midl. Naturalist 131: 65— 4, BASKIN, C. C. AND J. M. BASKIN. 1988. Germination ecophysiology of her- baceous plant species in a temperate region. Amer. J. Bot. 75: 286-305. FENNER, M. 1983. Relationships between seed weight, ash content, and seed- ling growth in twenty-four species of ee New Phytol. 95: 694— 706. Grime, J. P. 2001. Plant Strategies and Vegetation Processes, 2nd ed. John Wiley and a Ltd., Chichester, UK. Gross, K. L. 1984. Effects of seed size and growth form on seedling estab- lishment we SIX spe aaaiee perennial plants. J. Ecol. 72: 369-387. Harper, J. L., P. H. LOVELL, K. ae MOORE: — The shapes and sizes f seeds. Annual Rev. Eco on : 327-3 HERKERT, J. R. 1991. Padansered and oe Species of Illinois: Status and Distribution. Illinois Endangered Species Protection Board, Spring- field, IL. METTLER, P. A., M. SMITH, AND K. Vicrory. 2001. The effects of nutrient pulsing on the threatened floodplain species, Boltonia decurrens. J. Plant Scol. 155: 89-96. MISSOURL DEPARTMENT OF CONSERVATION. 1999, Missouri species of conser- vation concern checklist. Jefferson J 997. X< 1 City, MO. Moss, J. K. Stage-based demography of Boltonia decurrens, a threat- ened ooo species. M.S. thesis, Southern [linois Univ., Edwards- ville, REDMOND, i‘ 1993. Population study of Boltonia decurrens, a pce threatened plant species. M.S. thesis, Southern Illinois Univ., Edwards- ille ree SCHWEGMAN, J. E. AN . W. Nyporr. 1985. The taxonomic and a status of ae urrens (Torrey and Gray) Wood. Castanea 50: 115. SmitH, M. 1991. Life history research for the Decurrent false aster. Report, Illinois Dept. C onservation, Springfield, IL. 1993. Effects of the Flood of 1993 on the aera false aster Balin decurrens). U.S. Army Corps of Engineers, St. uis, MO — RANDT, AND J. STONE. 1995. Effect of soil texture wee microto- 2002} Smith and Cawly—Boltonia decurrens Seedlings i pography on germination and seedling growth in Boltonia decurrens (Asteraceae), a threatened floodplain species. Wetlands (Wilmington) 15: age ND T. KEEvIN. 1998. Achene morphology, production and germi- nation, and potential for water dispersal in Boltonia decurrens Ahan false aster), a threatened floodplain species. Rhodora 100: I. , —,. P. MeTrLer-McC ure, AND R. BARKAU. 1998. es of the flood of 1993 on Bolin decurrens, a rare floodplain plant. Regul. Rivers: Res. Memt 191-202 AND P. METTLER. pe The aie of the flood a in maintaining Boltonia decurrens, a fugitive plant species of the [linois River floc plain: A case history of a threatened species, pp. 109-144. In: B. A Middleton, ed., Flood Pulsing and Wetlands: ieee the Natural Hy- eee Balance. John Wiley and Sons, Inc Y. Wu, AND O. GREEN. 1993. Effect of light mt water-stress on photosynthesis and biomass production in Boltonia i urrens (Astera- ie), a threatened species. Amer. J. Bot. 80: 859—8€ Sparks, R. E. 1995. Need for ecosystem management of ae rivers and their floodplains. BioScience 45: 168-182 NELSON, AND Y. YIN. 1998. ‘Naturalization of the flood regime in regulated rivers. BioScience 48: 706—720. STEEL a G. D. AND J. H. Torrit. 1980. a and Procedures of Sta- 7A Reese Approach. McGraw-Hill, New York. oe L AN . GRAY. 1841. The Flora of North America, Vol. 2. Wiley and eee New York. U.S. FISH AND WILDLIFE SERVICE. 1988. Endangered and threatened wildlife and plants, determination of threatened status for Boltonia decurrens ae false aster). Fed. Reg. ° 585 586 . 1990. Decurrent False Aster recovery plan. U.S. Fish and Wildlife Sc Twin Cities, ZHANG, J. 1993. Seed dimorphism in relation to germination and growth of Cakile edentula. Canad. J. Bot. 71: 1231-1235 RHODORA, Vol. 104, No. 917, pp. 14-30, 2002 CROSSING EFFECTS ON SEED VIABILITY AND EXPERIMENTAL GERMINATION OF THE FEDERAL THREATENED PLATANTHERA LEUCOPHAEA (ORCHIDACEAE) MARLIN L. BOWLES! AND KAREL A. JACOBS The Morton Arboretum, 4100 Hlinois Route 53, Lisle, IL 60532 e-mail: mbowles@ mo1 tonarb.org LAWRENCE W. ZETTLER AND TONYA WILSON DELANEY Department of Biology, The [linois a a West College Ave., acksonville, 5265( ABSTRACT. We conducted experimental pollination and seed germination trials with Platanthera leucophaea, a threatened north temperate grassland orchid species for which propagation and restoration are ee sae conser- vation needs. Our objectives were to determine how the species’ breeding ieee. and modes of pollination interact to affect production of viable seed and seed germination, and how seed germination responds to stratification and to inoculation by mycorrhizal fungi. Outcrossing by hand pollination produced a higher percentage of viable seed than did natural pollination, < did outcrossing between populations in comparison to outcrossing or selfing within populations, indicating a facultative outcrossing breeding system. Out- ssing also enhanced percent germination, which was positively correlated with the percentage, but not number, of viable seeds. A 16 wk. stratification period resulted in significantly higher percent germination than 8 wk. strati- fication or no stratification. Germination was further enhanced by inoculation with a mycorrhizal fungus (Ceratorhiza sp.) isolated from P. leucophaea. These results indicate that the breeding system of P. lewcophaea allows for greater numbers of viable seeds with greater germination rates when plants are outcrossed. Thus, hand pollination: and outcrossing can enhance propa- gation and restoration efforts, especially when coupled with scarification and stratification treatments that maximize germination rates. = S Key Words: Orchidaceae, Platanthera, seed germination, breeding system, pollination, conservation Gaining a better understanding of the propagation and resto- ration requirements of terrestrial orchids has become crucial to their conservation (Johansen and Rasmussen 1992; Zettle1 1996a), and is an important conservation objective for the Federal Threatened eastern prairie fringed orchid Platanthera leucophaea (Nutt.) Lindl. (Bowles and Bell 1999; U.S. Fish and Wildlife Service 1999). This orchid occurs in tallgrass prairie remnants 14 2002] Bowles et al.—Platanthera germination [5 and wetlands in eastern Jowa, southern Wisconsin, and northern Illinois and Indiana, and in shoreline prairies, sedge meadows, bogs, and fens eastward to Maine (Bowles 1983; Sheviak 1974: Sheviak and Bowles 1986). It has declined by more than 70% from original county records due to habitat conversion to agri- culture. Remaining populations are often small and continue to be threatened by succession to woody vegetation, competition from exotic species, illicit collecting, and drainage (Bowles 1983; U.S. Fish and Wildlife Service 1999), In this study, we report on effects of experimental pollination on its seed production, and effects of seed treatment and fungal inoculation on seed germi- nation. Showy Platanthera are thought to have facultative outcrossing breeding systems (Gregg 1990), a well known strategy for avoid- ance of deleterious effects of inbreeding (e.g., Willson 1983). Platanthera leucophaea has the largest flowers and nectar spurs of eastern North American Platanthera, and its large floral dis- play and lack of vegetative reproduction indicate a strong in- vestment toward pollination and seed production (Bowles 1983, 1985; Sheviak and Bowles 1986). Pollination is by hawkmoths. The orchid pollinarium, which comprises a pollinium (pollen mass), caudicle, and viscidium, adheres to a hawkmoth’s probos- cis by the viscidium. Caudicle movement (taxis) positions the pollinium for contact with the stigmatic surface after about 40 seconds, thereby promoting outcrossing (Bowles 1985). Pollen grains are then removed from the pollinium as they adhere to the plant’s stigma. Selfing or geitonogamy may occur when moths revisit flowers or inflorescences, especially in small orchid pop- ulations, and could potentially influence production of viable seeds if this species is affected by inbreeding depression. As in P. praeclara Sheviak & M. L. Bowles (Sieg and King 1995), most plants flower once and the median number of flowering seasons is less than three. Seedling establishment is therefore an important life history stage for this species, and pollination and seed production are critical factors in population viability (Bowles and Bell 1999), Terrestrial orchids are difficult to propagate due to physiolog- ical seed dormancy and the need for mycorrhizal fungi for suc- cessful seed germination and seedling development (Rasmussen 1995: Stoutamire 1974; Zettler 1996b:; Zettler and McInnis 1992), Experimental propagation with mycorrhizal fungi has been re- —_ 16 Rhodora [Vol. 104 ported for a few species of the widespread north temperate orchid genus Platanthera, including P. integrilabia (Correll) Luer (Zet- tler and McInnis 1992), P. clavellata (Michx.) Luer (Zettler and Hofer 1998), and P. leucophaea (Zettler et al. 2001). Seed ger- mination was highly variable among seed sources in these studies, and was facilitated by, but not dependent upon, the presence of mycorrhizal fungi. Such variation may be influenced by many factors, including sensitivity to inbreeding and levels of genetic diversity within populations of different sizes (e.g., Fenster and Dudash 1994; Weller 1994), and differing germination require- ments or different experimental methods used by researchers (Rasmussen 1995: Zettler 1996b). Seed germination and mycorrhizal fungi of Platanthera leu- cophaea were first investigated by Curtis (1939), who isolated species of the soil fungus Rhizoctonia (now Ceratorhiza, Ander- sen and Rasmussen 1996) from P. leucophaea roots in different habitats. Curtis was unable to germinate seeds inoculated with these fungi, perhaps due to failure to properly scarify or stratify seeds. Stoutamire (1996) increased asymbiotic seed germination rates by increasing scarification time in diluted NaOCl, and rec- ommended two or more months of cold stratification. Stoutamire (1996) also germinated P. leucophaea seeds in 35 mp Nitex bolt cloth (following Rasmussen and Whigham 1993) buried in prairie sod that contained soil fungi, but neither mycorrhizae nor further seedling development occurred. Zettler et al. (2001) achieved my- cotrophic germination of P. /eucophaea seeds using a Ceratorhiza species isolated from roots of this orchid, with development of leaf-bearing seedlings occurring after a second cold treatment. More information is needed about optimum pollination and germination requirements of Platanthera leucophaea, factors that will lead to a better understanding of its reproductive biology, population demographic processes, and restoration requirements. Our study had two related objectives. First, we assessed how different modes of pollination (i.e., natural versus hand pollina- tion, selfing, outcrossing within, and outcrossing between popu- lations) affect the percentage of viable seeds and the germinabil- ity of seeds. Because of the breeding system of this species, we expected that outcrossing would enhance seed viability by reduc- ing inbreeding depression. Second, using scarified seed, we tested how stratification periods and symbiotic versus asymbiotic con- 2002] Bowles et al.—Platanthera germination ey Table |. Seed sources, sample sizes, and seed collection dates for ex- perimental crossing and germination of Platanthera leucophaea, n = number | of plants sampled for seed. Sampling Date and Sample Size y Experiment Germination Crossing Site Location Experiment Experiment Abbot Lake Co., Il. Aug 1996 Sep 1997 Park (n = 3) (n = 5) Sep 1998 (n = 1) Wadsworth Lake Co., III. Aug 1995 Sep 1997 Prairie (n = I11) (n = 3) Sep 1996 Sep 1998 (n= 30) (n = 18) yons Lake Co., II. Sep 1996 Woods (n = 82) Pickerel Sandusky Co., Oct 1995 Creek Ohio (n > 10) ditions affected germination. We expected that longer stratifica- tion and symbiotic conditions would enhance germination. MATERIALS AND METHODS Seed and fungus sources. Platanthera leucophaea seeds were obtained from one site in Ohio and three sites in Illinois (Table 1). The Pickerel Creek, Ohio, site contains one of the largest known P. leucophaea populations, where plants occupy successional wetland habitat of the Lake Erie lake plain (U.S. Fish and Wildlife Service 1999). The Illinois sites are in Lake County, in the Chicago region of northeastern Illinois, and are < 15 km from one other. The Wadsworth Prairie, Lyons Woods, and Abbott Park populations are among the largest in Illinois, with plants occurring in successional prairie and sedge meadow (U.S. Fish and Wildlife Service 1999). All seeds were collected from mature capsules prior to dehiscence in late August or early October, and were stored at 5°C in a desiccator (containing 18 Rhodora [Vol. 104 CaSO,) prior to sowing. The mycorrhizal fungus used in germi- nation treatments has been tentatively identified as a species of Ceratorhiza (L. W. Zettler, pers. obs.), and was isolated from the roots of a mature P. /eucophaea specimen obtained from Abbott Park in 1995, and cultured at the Morton Arboretum. Crossing effects on seed viability. This study investigated Whether manually placing entire pollinia on stigmatic surfaces (= hand pollination) yielded a greater percentage of viable seeds than natural pollination. Hand pollination consisted of crosses between plants within populations, while natural pollination could also have included selfing through geitonogamy. Pollinated plants were not bagged to exclude subsequent natural pollination be- cause placement of the entire pollinium on the stigma excludes additional pollen deposition. Pollinations were conducted in 1998, with one or two mature capsules sampled per plant from ten nat- urally pollinated plants and from nine hand-pollinated plants (Ta- ble 1). Seeds were pooled from capsules within plants, and ap- proximately 200 seeds were sampled per plant. Seeds were briefly surface disinfested in dilute NaOCl, moist stratified for 11 mo. by suspending in sterile deionized water (SDW) at 6°C, and sown onal xX 4 cm filter paper strip in a 9 cm diameter petri dish containing 20 ml of modified oats medium (Dixon 1987). The dishes were then examined with a dissecting microscope to count numbers of apparently viable and non-viable seeds based on the presence or absence of distinct, rounded and hyaline embryos (Zettler et al. 2001). Viable seed numbers were expressed as a percent of the total seeds in each sample. These percentages were arcsine-transformed (Zar 1974), and tested against the null hy- pothesis that hand-pollinated capsules did not contain a greater percentage of viable seeds. We used a one-tailed t-test based on our expectation of more viable seeds with hand pollination be- cause 1t maximizes pollen availability. A second study examined crossing effects on production of viable seed using the Wadsworth Prairie and Abbott Park popu- lations. We compared self-pollination (1 Wadsworth plant and | Abbott plant), outcrossing within populations (1 Abbott plant and 3 Wadsworth plants), and reciprocal outcrossing between popu- lations (2 plants). These pollinations included > 5 flowers per inflorescence, and were conducted in 1997 (Table 1). Seeds col- lected from mature capsules were pooled within each plant, dis- 2002] Bowles et al.—Platanthera germination 19 infested, scarified by shaking in 0.5% NaOCl for one hr., and stratified for 8 wk. at 5°C in SDW. Seed suspension samples were removed from stratification with an eyedropper. Each sample con- tained about 100 seeds, with = 10 samples per cross. As de- scribed above, the numbers of seeds containing round distinct embryos were counted with a dissecting microscope and ex- pressed as a percent of the total seeds. One of the selfed plants did not produce mature capsules with seeds, resulting in 0% vi- able seeds for this cross. Differences between crossing treatments were analyzed by inspection because the non-normal distribution of data and unbalanced design prevented appropriate statistical testing of the hierarchical nesting of seed sources within treat- ments. Stratification and symbiotic culture effects on germina- tion. In these experiments, we tested effects of duration of moist stratification, seed age (storage time), and presence or ab- sence of fungal inoculant on seed germination. Seeds were dis- infested and scarified in 0.5% NaOCl for | hr. We used seedling development stages as defined by Hadley (1983), where Stage | germination is achieved by rupturing of the testa (seed coat) by the enlarging embryo, and Stage 2 germination coincides with enlargement of the protocorm beyond the original seed size and development of rhizoids. Our observations suggest that scarifi- cation may promote Stage | germination by facilitating water imbibition and rupture of the testa by the enlarging embryo. In contrast, other studies (e.g., Zettler and Hofer 1998; Zettler et al. 2001) using unscarified seed describe germination to Stage | as production of rhizoids, and Stage 2 as rupture of the testa. In this situation, unscarified seeds may initiate germination by producing rhizoids that help imbibe water and then cause rupture of the testa. The effect of stratification period on germination to Stage | was tested on seeds collected in 1996 (Table 1). After scarifica- tion, seeds were plated on sterile filter paper moistened with SDW and given treatments of either no stratification (Abbott popula- tion), or moist stratification in darkness at 4°C for 8 wk. (all sites) or 16 wk. (Wadsworth and Lyons populations). Seed numbers ranged from 450 to 1140 per population. Seeds were plated onto modified oats medium in petri dishes, with six to nine replicates per treatment and 30—160 seeds per dish. Petri dishes were 20 Rhodora [Vol. 104 wrapped in foil and incubated at 25°C in darkness, and germi- nation was monitored biweekly for 11 mo. with a dissecting mi- croscope. Additional germination that might have occurred with a second cold treatment was not considered in this experiment. As described above, seed viability counts were based on the pres- ence of round distinct embryos. Counts of Stage | germinated seeds were pooled among replicate plates within each treatment and tested by Chi-square analysis for differences in numbers of germinated and ungerminated seeds among stratification periods, and between the Wadsworth and Lyons seed sources. Effects of inoculant, stratification, and seed storage time on germination to Stage 2 were tested between the 1995 seed batch 3621 seeds), which was stored for 18 mo., and the 1996 seed batch (n = 1315 seeds), which was stored for 6 mo. (Table 1). In this study, only scarified seeds were used, and seeds were pooled among seed sources. To test whether a fungal inoculant and stratification resulted in greater germination than either treat- ment separately, replicate plates for the 1995 and 1996 seed batches were given treatments of 16 wk. stratification, 16 wk. stratification plus inoculant, or inoculant with Ceratorhiza sp. As above, seeds were plated onto modified oats medium and seed viability counts were based on presence of distinct embryos. Ger- mination was monitored biweekly for 11 mo. The number of Stage 2 seedlings on each plate was expressed as a percent of the number of viable seeds originally present on the plate. A two-factorial ANOVA was used to compare germination treatment and seed storage time effects on percent germination. Exclusion of contaminated plates resulted in an unbalanced ex- perimental design (replicates ranged from 4—16 plates), which we tested using a General Linear Model. Prior to analysis, all per- centages were arcsine transformed (Zar 1974). We also tested for a correlation between the percentage of viable seeds and the per- centage of those seeds reaching Stage 2. To determine whether seedling development (and seed viability) was independent of seed density, we tested for a correlation between the percentage of viable seeds reaching Stage 2 and the total number of seeds (both viable and non-viable) in each plate. — RESULTS Crossing effects on percent viable seed. Seed viability var- ied among pollination crossing treatments made in 1997 and in 2002] Bowles et al.—Platanthera germination 21 Crossing Effect on Percent Seed Viability 1.0 Population Cross Between GF) Within @ Self 0.8 Ww) CG oO oO ep) & 0.5 a a = 4 = g 0.3 = 0.0 Seed Source Figure |. Differences in mean percent viable seed produced by selfing, crossing within, or crossing between populations of Platanthera leucophaea. Seed source replicates: | = Wadsworth x Abbott, 2 = Wadsworth * Wads- worth, 3 = Abbott x Abbott, 4 = Abbott self. One Wadsworth selfed plant produced no viable seed. Lines represent standard errors. 1998. About 50% of the seeds in capsules obtained from hand cross-pollinations made in 1998 contained viable embryos, almost twice the percentage from naturally pollinated plants (t = —1.785, P = 0.046). Within-population outcrosses made in 1997 also ay- eraged about 50% viable seed, but between-population crosses averaged almost 70% with wide variation among means (Figure 1). One self-pollinated plant produced no capsules with viable seeds, while the second averaged 15% viable seeds. Effects of stratification period on Stage 1 germina- tion. Stratification period, but not seed source, significantly af- fected germination to Stage |, with percent germination increas- ing with increasing stratification period across all seed sources (Figure 2). Overall, germination was < 5% for unstratified seeds, a2 Rhodora [Vol. 104 Stratification Period and Seed Source Effects on Seed Germination [1 No stratification 50 - 45 4 8 weeks stratification 40 5 . w 16 weeks stratification Stage 1 Germination (%) Abbott Wadsw orth Lyons Pooled Seed Source Figure 2. Longer stratification period increases percent seed germination of Platanthera leucophaea, with similar effects among seed sources. Chi- aap canes ation period (xy? = 275.76, P < 0.001), Seed source (x? = 0.945 = 0.332) 10-20% after 8 wk. stratification and > 30% after the 16 wk. stratification. Effects of seed storage time, germination treatment, and seed viability on germination to Stage 2. No significant ef- fects of seed storage time or germination treatment were found for Stage | germination. However, Stage 2 germination (rhizoid production) was significantly higher for seeds from the 1996 seed batch than for seeds collected in 1995 and stored for an additional 12 mo. (Figure 3). Moreover, Stage 2 germination in both seed batches was higher for stratified seeds that were also germinated symbiotically with Ceratorhiza than for either treatment alone (Figure 3). Among the 1996 seeds, percent germination to Stage 2 was also significantly correlated with percentage of viable seeds (Figure 4). Percent germination was not, however, significantly correlated with total seed number per plate (r?7 = 0.003, P = 0.85) NM eS) 2002] Bowles et al.—Platanthera germination Treatment and Seed Age Effect on Germination 800 —- Treatment 7 ©) Inoculated 4 @ __'\noculated/stratified J @ Stratified - 600 4 2 7 = a & | E 4 tb) bas Oo 40.0 - N 4 o 4 D 8 4 (op) + xe 4 c 200°. = oO 4 © 4 = 4 O06: 4 O. Seed Age Figure 3. The mean percentage of viable Platanthera leucophaea seeds germinating to Stage 2 is lower for older seeds and greater for stratified seeds inoculated with Ceratorhiza sp. isolated from P. leucophaea. ANOVA: Treat- ment (F = 6.97, P = 0.003), Age (F = 20.74, P < 0.0001), Treatment x Age (F = 1.78, P = 0.1846). Lines represent standard errors. nor with total number of viable seeds per plate (r? = 0.008, P = O:759). DISCUSSION Crossing effects on seed viability. As suggested for showy Platanthera (Gregg 1990) our pollination experiments indicate that P. leucophaea has a facultative outcrossing breeding system. Because this system allows mixed mating, it is apparently vul- nerable to inbreeding depression, which can be expressed at dif- ferent plant life-history stages (e.g., Carr and Dudash 1996; Du- dash 1990; Fenster and Dudash 1994), In P. leucophaea, inbreed- ing depression appears to have cascading effects by decreasing the percentage of capsules formed, the percentage of viable seeds within capsules, and the percent germination of those seeds. For outcrossing species, this process may be alleviated in larger pop- ulations that maintain high levels of genetic diversity (Schaal et 24 Rhodora [Vol. 104 Relationship Between Viable Seeds and Germination Stage 2 Germina tion (%) o = ios. tee pe ee ot ' 40.0 50.0 60.0 75.0 85.0 Viable (%) Figure 4. Within plates, the percentage of viable Platanthera leucophaea seeds cemetery oe to Stage 2 is positively correlated with the total percentage f viable seeds = 0.374, P = 0.015 O al. 1991; Weller 1994). The amount of inbreeding in small P. leucophaea populations could therefore be greater than in large populations because opportunities for outcrossing may be less in small populations. In our study, the lower percentage of viable seeds in naturally pollinated plants than in hand-pollinated plants may have resulted from inbreeding due to geitonogamy and cross- ing among closely related individuals, as well as from low rates of pollen deposition by hawkmoths. The wide variation we de- tected in seed viability among outcrosses within populations could reflect different levels of inbreeding based on chance. Stratification and fungal symbiont effects on germina- tion. Platanthera leucophaea seed germination is highly re- sponsive to both stratification time and the presence of a fungal symbiont. As indicated by Stoutamire (1996), optimum germi- nation requires a sequential combination of scarification and i) ‘Nn 2002] Bowles et al.—Platanthera germination moist stratification. Although all of our seeds were scarified, we found extremely low (< 5%) germination without stratification. Further, increasing stratification time from 8 to 16 wk. more than doubled germination from < 15% to > 30%. This suggests that for north temperate orchids, which may have evolved under se- lective pressure of cold dormant season conditions, both scarifi- cation and long-term stratification are necessary to attain high rates of seed germination. Seed germination experiments that do not include sufficiently long moist stratification periods coupled with scarification and a fungal inoculant could lead to improper conclusions about seed germinability (Rasmussen 1995). For example, although north temperate species seem to depend on mycorrhizal fungi for seed- ling development (Johansen and Rasmussen 1992), variable re- sults are reported. Zettler and McInnis (1992) found higher ger- mination for Platanthera integrilabia with inoculated seeds, but Stoutamire (1996) reported that initial seedling germination for P. leucophaea did not require a fungal symbiont if cultured on artificial media containing a carbon source. Our results, along with those of others (e.g., Zettler and Hofer 1998; Zettler and McInnis 1992) underscore the importance of a fungal inoculant for successful germination in Platanthera species. The effect was especially apparent for seedling development to Stage 2 germi- nation, which was maximized by the combination of 16 wk. strat- ification and presence of a fungal inoculant. Seed age and storage techniques are also important factors in orchid seed germinability (Seaton and Hailes 1989). Stoutamire (1996) reported complete loss of Platanthera leucophaea seed viability after 6 mo. of storage at 5°C. Our results were less dras- tic, but similar, in that about 60% of the seeds reached Stage 2 germination after 6 mo. of storage at 5°C, but only 10% reached Stage 2 after 18 mo. of storage. However, Zettler (1996b) reported viable P. integrilabia seeds after 6 yr. of storage at —7°C and 6°C. Loss of viability may be related to failure to adequately dry seeds prior to and during storage (L. W. Zettler, pers. obs.), com- plex dormancy mechanisms (Johansen and Rasmussen 1992), and different species characteristics. For north temperate Platanthera, including P. leucophaea, propagation beyond Stage 2 may be accompanied by high rates of mortality, especially if using an aggressive fungal symbiont (Zettler and Hofer 1998; Zettler and McInnis 1992; Zettler et al. 26 Rhodora [Vol. 104 2001). As a result, symbiotic orchid propagation without host- specific fungi may be problematic. Screening of P. leucophaea fungal inoculants from naturally germinating seeds and seedlings could help alleviate this problem. Research is also needed to de- termine whether the combination of proper scarification, stratifi- cation, inoculation, and secondary cold treatments can enhance the transition from Stage 2 germination to further leaf and tuber development (Johansen and Rasmussen 1992), and how pollina- tion outcrossing rates affect this transition. Conservation applications and concerns. Although Pla- tanthera leucophaea is perennial, most individuals flower once, and seedling establishment appears to be a critically important stage in its life cycle (Bowles and Bell 1999). Thus, factors that increase production of viable seeds should enhance population viability. In that regard, hand pollination may be an important tool because it can increase viable seed numbers by maximizing pollen deposition and avoiding inbreeding. Hand crossing among fragmented populations appears most likely to enhance viability, but it is controversial because of concerns that outbreeding de- pression may result from the disruption of locally adapted gene complexes (Bowles and Whelan 1994). For example, genetic al- lozyme (Cowden 1993) and random polymorphic DNA (Havens and Buerkle 1999) studies of P. leucophaea have found compar- atively high levels of genetic differentiation among populations, which indicates potential for outbreeding depression. However, human-caused population fragmentation and reduced gene flow could have contributed to such differences, and it is unknown whether outbreeding depression would actually occur. For ex- ample, Fenster and Galloway (2000) found outbreeding depres- sion to be important in Chamaecrista fasciculata Michx. only for crosses of = 1000 km, a distance far greater than among our study sites. Hawkmoths are well known for long-distance move- ment, which may have facilitated landscape-scale gene flow in P. leucophaea that would have tended to minimize population dif- ferentiation. Human-mediated crosses can alleviate potential in- breeding within fragmented populations of outcrossing species (Richards 2000). For example, heterosis from long-distance cross- es has been observed in the orchid Liparis lilifolia (L.) A. Rich. ex Lindl. (Whigham and O’Neill 1991), and such crosses have been used to obtain viable seed of the orchid Cypripedium cal- 2002] Bowles et al.—Platanthera germination aa ceolus L. var. pubescens (Willd.) Correll in Britain (Light and MacConaill 1998). Our results indicate this may be possible for P. leucophaea, as the greatest percentage of viable seeds resulted from inter-population crosses. Another concern is that high levels of seed production from hand pollination could impose a significant cost on terrestrial or- chids, as found for 7ipularia discolor (Pursh) Nutt. (Snow and Whigham 1989) and Cypripedium acaule Aiton (Primack and Hall 1990). Calvo (1993) argued that the naturally low rates of orchid seedling recruitment would not select for increased polli- nation and seed production. Kull (1998) also found that microsite factors, rather than pollinators, limited population growth in the long-lived perennial C. ca/ceolus. However, such effects may be less important in short-lived orchid species. The short life span of Platanthera leucophaea, its lack of vegetative spread, and its showy, and apparently costly, inflorescence structure suggest that high rates of seed production, and more importantly, high levels of seed viability, are important for population maintenance in this species. Ultimately, successful seedling establishment will depend upon chance coupling of germinating seeds with hyphae of fa- vorable soil fungi, and rates of this demographic process remain essentially unknown for terrestrial orchids. These concerns, and our crossing experiments, indicate that further work is needed to assess the impacts of translocating ge- netic material among populations, and whether there are negative demographic consequences of increased rates of pollination. Fully replicated crossing experiments are also needed to test for plant and site effects on seed viability. ACKNOWLEDGMENTS. We thank Jenny McBride, Janette Ja- cobs, Scott Stewart, Jennifer Sunley, Lisa Berg, and Kristin Lud- wig for assistance with seed germination experiments; June Kei- bler, Louise Clemancy, Warren Stoutamire, Russ Dombeck, and many volunteers for providing orchid seeds; and Lee Thomas for fungal isolations. We also thank Tim Bell, Kay Havens, Sue Wie- grefe, and Warren Stoutamire for assistance and valuable com- ments; the U.S. Fish and Wildlife Service, []linois Department of Natural Resources, and Illinois Nature Preserves Commission for funding and permits; and Glen Kruse and Mike Sweet for coor- dinating the project funds. Rhodora [Vol. 104 LITERATURE CITED ANDERSEN, T. EF AND H. N. RASMUSSEN. 1996. The mycorrhizal species of Rhizoctonia, pp. 379-390. In: B. Sneh, S. Jabaji-Hare, S. Neate, and G. Dist, eds., Rhizoctonia Species: Taxonomy, Molecular Biology, Ecolo- gy, Pathology and Disease Control. Kluwer Academic Publ., Netherlands. Bow Les, M. L. 1983. 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Naturalist 145: 168-175 RHODORA, Vol. 104, No. 917, pp. 31-41, 2002 THE IDENTITY AND HISTORY OF MYRICA CAROLINIENSIS (MY RICACEAE) ROBERT L. WILBUR Department of Biology, Duke University, Durham, NC 27708 e-mail: rwilbur@duke.edu ABSTRACT. The protologue of Myrica caroliniensis 1s more than adequate to identify it as depicting the bayberry ae at ene 2 southern New Jersey to Florida and westward into eastern species has been mostly known for the past half century as M. Aes The alleged dif- ferences between the commonly recognized and more northern populations known most recently as M. pensylvanica (presumably ranging from New- foundland at least into northeastern North Carolina) are that the southern elements have more persistent to even evergreen leaves and lack the minute trichomes on both the hardened fruit wall and the young glandular projections or papillae that completely cover the young to just maturing fruit of the northern representatives. The pubescence on the fruit cannot be readily de- tected on mature fruit due to its heavy deposit of wax. The alleged differ- ences, which seem to be more like tendencies than sharply delineated differ- ences, are not of specific significance any more than those suggested between the northern and southern populations of Magnolia virginiana. The name for the bayberry that ranges from Newfoundland on into Florida and ace into eastern Texas a therefore be Myrica caroliniensis, the binomial by hich it was known throughout most of ‘the nineteenth centut ry. It has been rather eet demonstrated that the waxy fruited, papillate species ought to be placed in the genus Morella, clearly . from the genus Myrica with the latter’s smooth, non-papillate, non-waxy nutlet. Key Words: Myrica caroliniensis, M. cerifera, M. heterophylla, M. pensyl- vanica, Myricaceae, Morella caroliniensis, Morella cerifera Phillip Miller (1768) published the binomial Myrica caroli- niensis with the following protologue: 3. Myrica (caroliniensis) foliis lanceolatis serratis, caule suf- fruticosa. Myrica with spear-shaped sawed \eaves, and shrubby stalk. Myrtus Brabanticae similis caroliniensis hu- milior; foliis latioribus & magis serratis. Catesb. Car. vol. I. p. 13. Lower Carolina Myrtle, or Candleberry-tree resem- bling that of Brabant, having broader leaves which are more sawed. The third sort grows naturally in Carolina; this doth not rise al 32 Rhodora [Vol. 104 so high as the former, the branches are not so strong, and they have a grayish bark; the leaves are shorter, broader, and are sawed on their edges, but in other respects is like the second sort [M. cerifera L.|; the berries of this are also col- lected for the same purpose |i.e., for a “‘sort of green wax from the berries, which they make into candles.” The above scanty account does not describe unequivocally any one species but it does contrast Myrica caroliniensis in a manner adequate to distinguish it from M. cerifera for those familiar with the plants in the field. Miller cited Catesby’s account (1730, I: 13, t. 13.), which was accompanied by a convincing illustration. Catesby is quoted tn full below: *“Myrtus Brabanticae similis Caroliniensis humilior; foliis la- tioribus et magis serratis. The broad leaved Candle-berry Myrtle. This grows usually not above 3 feet high; in which, and its having a broader leaf than the tall Candleberry Myrtle, it principally differs from it.” Linnaeus (1753, 2: 1024) described Myrica cerifera [var.] B, based solely upon the same Catesby polynomial and illustration (cited as “‘Catesb. car. I: p. 13, t. 13”) noting its presence in “Carolina, Virginia, Pennsylvania.” Anyone familiar with both Myrica cerifera and M. carolinien- sis in the field in the Carolinas would readily assign the above descriptions of Miller and Catesby to the bayberry (M. caroli- niensts) with its broader leaves and shorter stature and not to the more commonly encountered wax myrtle, M. cerifera. As is to be expected when a protologue is so lacking in details as is that of Myrica caroliniensis, there has been much disagree- ment for almost two and a half centuries as to the identity of the binomial, especially by those with little or no familiarity with both species in the field. At various times the binomial has been attributed to what has been passing as M cerifera, M. pensylvan- ica Mirb., and M. heterophylla Raf. or M. curtissii A. Chev. Not surprisingly, our knowledge of the morphological distinctions be- tween these species, as well as their distributional ranges, has greatly increased with the passage of time. Hence we are now better able to determine what the various authors were describing. oS) ios) 2002] Wilbur—lIdentity of Myrica caroliniensis Although apparently there is no extant original material of Phillip Miller’s M. caroliniensis, it seems that the protologue’s emphasis on the low stature and the shorter and broader leaves would strongly suggest that a bayberry was being described, and not the wax myrtle (M. cerifera). This view is strengthened especially when one considers that M. cerifera sensu stricto was already included in a reasonably definitive manner elsewhere in each of the respective publications of Catesby, Linnaeus, and Miller. Most recent authorities, at least since Fernald (1938), have rec- ognized two bayberries in eastern North America, collectively ranging from southern Newfoundland south into northern Florida and westward into Arkansas and Texas. Myrica pensylvanica re- portedly is found southward as far as northeastern North Carolina while what has been most recently called M. heterophylla re- portedly ranges northward from Florida along the coastal plain at least as far as southern New Jersey and perhaps southeastern Pennsylvania as well as westward into Texas. Approximately half the plants are staminate and everyone agrees that staminate plants are exceedingly difficult, if not impossible, for one to distinguish between the two supposed eastern species of bayberry. Bornstein (1997, p. 434) reports that M. pensylvanica hybridizes quite read- ily with both M. cerifera and M. heterophylla which, if proven to be true, would surely make for an even more bafflingly com- plex problem in identification. My field experience with these two species in the southeast is considerable and I have not noted ev- idence of hybridization. I am unconvinced that there are two species of bayberry in eastern North America. Nothing suggests to me rampant hybrid- ization between the wax myrtles and the bayberries. I cannot re- call ever encountering a plant in field or herbarium that could not be identified immediately to species in the southeast. Miller, in publishing Myrica caroliniensis failed to distinguish it sharply from even the sympatric M. cerifera, not to mention the suppos- edly largely allopatric M. pensylvanica. Only those familiar with the pronounced tendencies exhibited by the plants in the field could expect to recognize the distinction between the species. If there is one bayberry in eastern North America, as my exami- nation of thousands of specimens has convinced me, we can safe- ly conclude that it is M. caroliniensis which, besides being the first of the bayberries to be described, is the only bayberry known from South Carolina, the area of Catesby’s intensive observations 34 Rhodora [Vol. 104 while preparing his Natural History of the Carolinas, Florida and the Bahama Islands (1730-1747). In this case, M. pensylvanica, M. heterophylla, and M. curtissii are all ater synonyms of Miller’s M. caroliniensis. Hf, contrary to my conclusion, after ex- amining over two thousand specimens from throughout their col- lective range, there actually are two species of bayberry in eastern North America, the southernmost of them is M. caroliniensis (Miller 1768) with M. heterophylla (Rafinesque 1838) and M. curtissti (Chevalier 1901) as later synonyms, the northern bay- berry would then be M. pensylvanica (Mirbel 1804). The principal distinction previously employed to separate Myr- ica pensylvanica trom M. heterophylla has been the presence of rather abundant but short, stiff trichomes on the densely com- pacted glandular papillae covering the usually pubescent, hard- ened wall of the fruit prior to the deposition of the heavy waxy layer. Only a minority of the thousands of specimens examined were of the gender and stage in which this feature could be em- ployed. I have found specimens whose papillae were hirsutulous among collections from the Gulf Coast and the frequency of such puberulently fruited specimens was even much higher in eastern North Carolina than further south. In my experience, species are separable by more and stronger characters than those differenti- ating these alleged species (1.e., M. caroliniensis and M. pensyl- vanica). Leaves of M. caroliniensis sensu lato are retained throughout most of the winter in the more southern parts of its extensive range; plants from the more northern portion of the range of M. caroliniensis sensu stricto retain their leaves for a shorter period of time. The reverse is true for those plants pre- viously called M. pensylvanica, which lose their leaves rather promptly at the approach of winter in the more northern part of the species’ collective range. The only other distinctions claimed to differentiate the two generally accepted species is the color of the young twigs, but I have found color to be so highly variable as to be of no help in distinguishing the alleged northern and southern taxa. My understanding is that we are dealing with one not particularly variable species. Those who persist in cleaving the bayberries into two species should at least accept the fact that M. caroliniensis has priority over either M. heterophylla or M. curtissiti and that Philip Miller’s name applies to the southern representatives of this somewhat variable, widespread species. Below are keys extracted from two leading treatments (Born- — 2002] Wilbur—lIdentity of Myrica caroliniensis 50 stein 1997; Fernald 1950) purportedly distinguishing the two east- ern North American bayberries from one another. In a majority of cases these keys do not separate the taxa, since much fewer than half the specimens bear fruit in a state in which the keys can be applied. Since the vegetative features of the twigs are even less applicable due to the great variability of twig color seemingly dependent upon exposure to light and other environmental vari- ables, I question whether we would be able to distinguish sta- minate plants, or pistillate plants, in most stages of their annual growth unless we first knew their provenance. An unpublished Master’s thesis from the University of Georgia (Houghton 1988) analyzed the morphological characteristics as well as the flavo- noid profiles of the eastern North American bayberries and wax myrtles concluding that the two bayberries (1.e., Myrica caroli- niensis and M. pensylvanica) were only varietally differentiated. To date the suggested varietal combinations have not been validly published. Fernald (1950, p. 524) differentiated the two eastern bayberries in his key as shown below: Bark of mature branches whitish-gray or drab; leaves dull above, membranaceous, deciduous (subpersistent south); inflorescenc- es all borne below the leafy tips; young fruit densely pubescent, Pe Ui o. 0 L ete waereawaeaeraenace Myrica Pensy lvanica Bark of mature branches blackish; leaves lustrous above; coria- ceous, evergreen; inflorescences below or in the axils of the old leaves; young fruit glabrous, ripe fruit 3—3.5 mm in di- OWNER “ovo ee eee eee eee es Myrica heterophylla Bornstein (1997, 3: 431) distinguished the eastern bayberries in his key as follows: Fruit wall and warty protuberances densely hirsute when young; branches whitish gray in age; leaves deciduous, membranous; Ihe 39-9 TO ee ec aeees ee eens 64 Myrica pensylvanica Fruit wall glabrous or sparsely glandular, warty protuberances + glandular; branches black; leaves persistent or tardily de- ciduous, leathery; fruits 3-4.5 mm..... Myrica heterophylla 36 Rhodora [Vol. 104 Type material or original specimens of Philip Miller voucher- ing his Myrica caroliniensis has not been found although dili- gently sought by several investigators. Rendle (1903) first re- ported his failure to find original material of Catesby’s two myr- icas depicted on his Plates 13 and 69. Reveal (in litt.) has also searched without success for original material of Miller vouch- ering his publication of M caroliniensis. Catesby’s Plate 69 clear- ly represents M. cerifera and Plate 13 depicts a bayberry. Al- though specimens of Catesby vouchering his “‘Myrtus Brabanti- cae similis Caroliniensis humilior” have not been located, this has not prevented three recent publications from confidently iden- tifying to species, the rather crude drawing and meager descrip- tion provided by Catesby. Ewan (1974) and Howard and Staples (1983) identified it as the more northern M. pensylvanica, while Wilbur (1990) concluded that it was M. heterophylla, a deter- mination clearly based more on the largely allopatric distribution of the two alleged species than upon the detail presented in the drawing and description provided by Catesby. Although previ- ously Catesby had lived and observed nature for several years in southeastern Virginia, he was not then focused on the goal of producing a sumptuously illustrated Natural History. A later ex- tended trip by Catesby was mostly spent in South Carolina and the Bahamas as well as allegedly in Florida, a claim questioned by Reveal (in litt.), for the intensive observation and painting that preceeded his long-protracted presentation of The Natural History of the Carolinas, Florida and the Bahama Islands. Fernald (1935, p. 423) made a major effort to straighten out the nomenclature of the eastern wax myrtles and bayberries of eastern North America without complete success. Fernald stated that “the wrong interpretation of Myrica caroliniensis is clearly discussed by Chevalier who correctly takes up for the deciduous- leaved and northern species the name of M. pensilvanica Loise- leur.”’ [Later, Fernald (1938, p. 410), upon the urgings of Rehder, adopted the spelling pensylvanica since Loiseleur (actually the author/editor was Mirbel 1804) employed both spellings and Che- valier (1901) had adopted the more usual form.] Chevalier’s clar- ification of M. caroliniensis, which earned Fernald’s approval, was that Chevalier refused to take up the earlier M. caroliniensis since that binomial had been frequently applied to a more south- ern species which Chevalier described on the next page as ““M. another name for the more southern bayberry. If Che- o° CUFLISSI, 2002] Wilbur—lIdentity of Myrica caroliniensis a) valier ever explained why he felt that those who employed ™. caroliniensis as the binomial for the southern bayberry were mis- taken, I have not found it. It is true that M. caroliniensis, as stated on p. 184 of Chevalier’s monograph, had been used by many early authors for the entire complex, ranging from Newfoundland south along the coastal plain into Florida and west along the Gulf Coast into Texas and then north into Arkansas, but that sort of confusion was routinely resolved by Fernald and most other au- thors without abandoning such names. If that were reason enough to routinely drop a name, chaos would reign, as Fernald frequent- ly noted (e.g., 1946, p. 389). Fernald (1935, p. 423) added to the nomenclatural confusion by unequivocally stating without explanation or stated evidence that Myrica cerifera included M. caroliniensis, and this was ac- cepted by Rehder (1949, p. 87b), also without discussion. It should not surprise anyone that, after such a thorough muddling, the binomial M. caroliniensis dropped from botanical usage. In spite of such flagrant abuse, I do not think the binomial irretriev- ably lost. Examination of the protologue of M. caroliniensis, as presented on the first page of this note, in my opinion confirms that those who employed that binomial for the southern bayberry were correct. Myrica caroliniensis (Miller 1768), M. pensylvanica (Mirbel 1804), M. heterophylla (Rafinesque 1838), and M. cur- tissii (Chevalier 1901) are all, in my opinion, synonyms of the eastern bayberry. Those who recognize two species within the eastern bayberries would agree, I believe, that only M. pensyl- vanica ought not be included in that listing. Fernald (1935, p. 423), usually so precise in his bibliographic sleuthing, uncharacteristically misled us in equating Myrica car- oliniensis with M. cerifera and also then followed Chevalier in recognizing the southern bayberry as M. curtissii. Three years ater, Fernald (1938, p. 409-410) took up the earlier M. hetero- phylla for M. curtissii, the bayberry with the more southern range (**?Delaware south into Florida and westward into Arkansas and Texas’’). Rehder (1949, p. 87), in my opinion mistakenly, fol- lowed Fernald (1935) in placing M. caroliniensis unquestioningly in the synonymy of M. cerifera. Thereafter, Miller’s binomial al- most completely disappeared from the botanical literature for the next fifty years, except in synonymy. Fernald (1950, p. 524), in Gray's Manual of Botany, summa- rized his overall unsurpassed knowledge of the flora of north- —_ 38 Rhodora [Vol. 104 eastern North America by recognizing five taxa of Myrica subg. Morella in the Gray’s Manual area: M. cerifera, M. pusilla Rat., M. pensylvanica, and M. heterophylla with its supposed var. cur- tissit (A. Chev.) Fernald. Gleason (1952, 2: 24) recognized only one species of bayberry, which he called Myrica pensylvanica, while placing the earlier M. caroliniensis as employed by Robinson and Fernald (1908), Britton and Brown (1913), and Small (1933) in its synonymy. Myrica heterophylla was appended to the account of M. pensyl- vanica somewhat uncertainly but perhaps as a hybrid. The treat- ment of the northeastern bayberry species was unchanged in Gleason and Cronquist (1963, p. 241) but Cronquist in the second edition (Gleason and Cronquist 1991, pp. 80-81) accepted both M. pensylvanica and M. heterophylla and modified the synonymy of M. pensylvanica by including “‘Cerothamnus caroliniensis of authors, perhaps not of Miller.” It should be noted that Miller’s species was not included in the synonymy of M. heterophylla Where it most certainly belonged. As a synonym of either ™. pensylvanica or M, heterophylla, M. caroliniensis would take pre- cedence due to priority. For simplicity’s sake the case presented here was not further complicated by earlier discussing the species in the genus Morella Lour. to which all waxy-fruited binomials mentioned belong (Baird 1968; Killick et al. 1998; Wilbur 1994). All are agreed that that Myrica sensu lato is divisible into three major taxa: Myrica L. (fruit water-dispersed), Morella (fruit bird-dispersed), and Comptonia L Her. ex Aiton (fruit a nut, possibly small mam- mal-dispersed). That these are meaningful, natural groups seems to be universally accepted even if some still consider them better treated at either sectional or subgeneric ranks. Nearly every in- vestigator in the past eight decades has recognized at least two genera: Myrica and Comptonia, while in recent decades three genera have been increasingly accepted in North America (e.g., Baird 1968; Chevalier 1901; Kartesz and Meacham 1999; Rad- ford et al. 1968; Wilbur 1994). The synonymy of the two species accepted here is restricted to the names applied to the eastern North American representatives (.e., only the eastern United States and Canada). Fortunately the spelling of the binomial “Myrica curtissii’” below is not of press- ing importance since the name is a synonym with little likelihood that it will ever achieve an active role. The specific epithet was 2002] Wilbur—lIdentity of Myrica caroliniensis ee originally published by Chevalier as “‘curtissi,”> who always em- ployed that form in his published work. It often appears as “‘cur- tissu,’ the correction resting no doubt upon the authority of Ar- ticle 60.11. Botanists of the earlier part of the previous century, who knew more Latin than most of us, were far more tolerant of the single 7, actually feeling that in many cases it was superior. In the text I have employed the double i but have used the single i when that was the form there published. Morella caroliniensis (Mull.) Small, Fl. S$. E. U.S. 337 & 1329. 1903. [as Carolinensis | Myrica caroliniensis Mall., Gard. Dict., ed. 8. no. 3. 1768. [LECTOTYPE: Catesby’s Plate 13 in Volume |. 1730. First designated here, as suggested by J. L. Reveal (in litt.). Myrica cerifera B latifolia Aiton, Hortus Kew. 3: 396. 1789. [6B = var.] Myrica cerifera B frutescens Castigl., Viagg. Stati Uniti 2: 302. 1790. [Castighoni cited both Catesby 1: tab. 13 and Myrica caroliniensis Mill. but described in most detail plants from Falmouth in eastern Massachusetts. ] Myrica cerifera B media Michx., Fl. Bor.-Amer. (Michaux) 2: 228. 1803. Myrica pensylvanica Mirb. in Duhamel, Traité Arbr. Arbust. 2: 190. 1804. Myrica heterophylla Raf. in Raf., Alsogr. Amer. 9. 1838. [as hetero- phyla| Myrica sessilifolia Rat., Alsogr. Amer. 10. 1838. Myrica Pee var. latifolia Raf., Alsogr. Amer. 10. 1838. Myrica Curtissitt A. Chev., Mém. Soc. Sci. Nat. & Math. Cherbourg 32: 269. iene. Myric. 185.) 1901. [as Curtissi] Myrica Curtissti var. media (Michx.) A. Chev., Mém Soc. Sci. Nat. & Math. Cherbourg 32: 270. (Monogr. Myric. 186.) 1901. [as Curtis- st iad a ae de ba var. Curtissti (A. Chev.) Fernald, Rhodora 40: 410. . [as Curt a seas (Mill.) Tidestr., Elys. Marian., Ferns 3: 41. Cerothamnus pensylvanicus (Mirb.) Moldenke, Revista Sudamer. Bot. 4: 16 Cerothamnus heterophyllus (Rat.) Moldenke, Phytologia 29: 386. 1975. Morella cerifera (L.) Small, Fl. S. E. U.S. 337 & 1329. 1903. Myrica cerifera L., Sp. Pl. 1024. 1753. Myrica cerifera var. angustifolia Aiton, Hortus Kew. 3: 396, 1789. Myrica cerifera 8 arborescens Castigl., Viagg. Stati Uniti 2: 302. 1790. Myrica cerifera |var.| pumila Michx., Fl. Bor.-Amer. 2: 228. 1803 Myrica pusilla Raf., Alsogr. Amer. 10. 1838. Cerophora lanceolata Rat., Alsog. Amer. || Myrica cerifera B angustifolia C. DC., Prods ae 16(2.1): 149. 1864. 40 Rhodora [Vol. 104 Nom. illeg. (Art. 53.1), non Aiton. Type: Louisiana. prope New Orleans, Drummond s.n. (x, not seen) Myrica pumila (Michx.) Small, Bull. Torrey Bot. Club 23: 126. 1896. Myrica cerifera vat. dubia A. Chev., Mém. Soc. Sci. Nat. & Math. Cherbourg 32: 265. (Monogr. = 181.) 1901. Morella pumila (Michx. ) Small, Fl. S. E. U.S. 337 & 1329. 1903. Cerothamnus arborescens (Castigl.) Se Elys. Marian., Ferns 3: 41. 1910. Cerothamnus ceriferus (L.) Small, Fl. Miami 61 & 200. (26 Apr) 1913. Cerothamnus pumilus (Michx.) Small, Shrubs Florida 8 & 133. (4 Sep) 1913. LITERATURE CITED BAIRD, J. 1968. A taxonomic revision of the plant family Myricaceae of ort a et north of Mexico. Ph.D. dissertation, Univ. N. Carolina, Chapel Hill, NC. BORNSTEIN, A. J. 1997. Myricaceae, pp. 429-434. In: Flora of North America Editorial Committee, eds., Flora of North America North of Mexico, Vol. 3. Oxford Univ. Bees Oxford and New Yor BRITTON, N. L. AND A. Brown. 1913. An Illustrated Figia of the Northern United States, Canada and the British Possessions. Charles Scribner’s Sons, New York. [Myricaceae, |: 584-586] CatessBy, M. 1730-1747. The Natural History of the Carolinas, Florida and the Bahama Islands. 2 vols. Folio. London. [Myricaceae, 1: 13 and 69] CHEVALIER, A. 1901. Monographie des Myricacées, anatomie et histologie, organographie, Classification, et description des especes, distribution geo- graphique. Mém. Soc. Sci. Nat. & Math. Cherbourg 32: 85 341 EWAN, J. 1974. Notes, pp. 89-100. In: The Natural History of Carolina, Florida and the Bahama Islands containing two hundred and twenty fig- ures of birds, beasts, fishes, serpents, insects, and plants by Mi Catesby. Facsimile of the 3rd. ed. (1771). Beehive Press, Savannah, FERNALD, M. L. 1935. Midsummer vascular plants a southeastern Vinginia. Rhodora 37: 378-413, 423-454, plates 384—405. |Myricaceae, pp. 423-— 424] . 1938. Noteworthy plants of southeastern Virginia. Rhodora 40: 364— 424, 434-459, 467-485, plates 509-535. [Myrica, pp. 408-412] . 1946. Types of some American Trees. J. Arnold Arbor. 27: 386-393. [pl Lee | —.. 1950. Gray’s Manual of ane 8th rev. ed. D. Van Nostrand Co., New York. [Myricaceae, pp. 523-525] GLEASON, H. A. 1952. The New : ritton & Brown ee Flora of the Northeastern United States and Adjacent Canada, Vol. e New Yor sien Garden, Bronx, NY. |Myricaceae, 2: 24-25] AND A. CRONQUIST. 1963. Manual of Vascular Plants of Northeastern United sar and Adjacent pe Van Nostrand Reinhold Company, New York. [Myricaceae, pp. 41| AND ———. 199]. Manual - Vascular Plants of Northeastern United 2002] Wilbur—lIdentity of Myrica caroliniensis 4] States and Adjacent Canada, ae ed. The New York Botanical Garden, Bro NY. [Myricaceae, pp. 80-81] bese W. M. 1988. The Bree of section Cerophora of the genus Myrica (Myricaceae) in North America. M.S. thesis, Univ. Georgia, Ath- ens, GA. Howarpb, R. A. AND G. W. STaApLes. 1983. The modern names for Catesby’s plants. J. Arnold Arbor. 64: 511-546. KARTESZ, J. T. AND C. A. MEACHAM. 1999, Synthesis of the North American Flora, CD-ROM version 1.0. North Carolina Botanical Garden, Chapel Hill, NC. Kitiick, D. J. B., R. M. POLHILL, AND B. VERDCOURT. 1998. New combina- tions in African Myricaceae. Kew Bull. 53: 993-995, LINNAEUS, C. 1753. Species Plantarum, Vol. 2 1024] MILLER, a oa Gardener’s Dictionary, 8th ed. Printed for the author, Lon- . Stockholm. [Myricaceae, 2 don, MIRBEL, a “ B. bE. 1804. Jn: H. L. Duhamel du Monceau, Traité des Arbres et Arbustes que lon te en fae Vol. 2. Paris. RADFORD, A. E., H. E. AHLES, AND C. Flora of the es Univ. icaceae, pp. 360-362] RAF Ree Cc . 1968. Manual of the Vascular N. ae Press, Chapel Hill, NC. [Myr . 1838 oa edition 1946]. Alsographia Americana. Ar- old woney Jamaica Plain, MA. [Myricaceae, pp. 8-12] as A. seep auee) of Cult vated Trees and Shrubs. Arnold Ar- boretum, oe Plain, MA. sre ” 87-88 | RENDLE, A. B. 1903. Notes on Myricaceae. J. . 41: 82- ROBINSON, B. L. AND M. L. FERNALD. 1908. ae s New Manual - Botany, 7th ed. American ee Co., New York. [Myricaceae, pp. 329-330] SMALL, J. K. 1933. Manual of the Southeastern Flora. Published = . author. New York. [Myricaceae, pp. 408-410] a R. L. 1990. Identification of the plants illustrated and described in tesby’s Natural History of the Carolinas, Florida, and the Bahamas. Sida 14: 29-48. 1994. The Myricaceae of the United States and Canada: Genera Sabwenerd: and series. Sida 16: 93-107. RHODORA, Vol. 104, No. 917, pp. 42-76, 2002 A FLORISTIC INVENTORY OF MANATEE SPRINGS STATE PARK, LEVY COUNTY, FLORIDA KIMBERELY J. GULLEDGE! AND WALTER S. JUDD Department of aia 220 rome Be a of Florida, ville, all cee he ABSTRACT. — A floristic inventory of the vascular plants of Manatee Springs State Park in Levy County, Florida, was conducted from May 1996 to De- cember 1998. In the 933 ha (2305 acres sh: a total of 360 species was found. The vascular flora sae 8 fe | cycad, 6 conifers, and 345 angiosperm species representing 90 omits ane 241 genera. Twelve natural communities are recognized in the park in addition to ruderal and developed areas: upland mixed forest, xeric hammock, sinkhole, sinkhole lake, swamp lake, basin swamp, bottomland forest, depression marsh, floodplain swamp, floodplain forest, blackwater stream, and spring-run stream. Key Words: Florida, flora, floristic study, vascular plants Manatee Springs State Park is located seven miles west of the city of Chiefland in Levy County, Florida, with the Suwannee River forming its western boundary. The park occupies Sections 13, 23-26, 35, and 36 of Township I] South, Range 13 East. The total area of the park is 933 ha. This includes the Meud-Scot track to the south acquired in 1988 (Department of Natural Re- sources 1989). The park is managed by the Division of Recreation and Parks for public outdoor recreation. The beautiful artesian spring, named for the manatees that take refuge there, is the major recreational feature of the park. Swimmers and divers enjoy a deep blue spring boil surrounded by bald cypress. Camping fa- cilities and nature trails are also provided for exploring the park. The Suwannee River, as well as all other surface waters in the park, 1s designated as Outstanding Florida Waters (Department of Natural Resources 1989), The flora of this area has not received as much attention as that of the panhandle or Southern Florida, and the geographical range of some species is poorly known. Many northern species find their southern limits in the Suwannee River region. This study was conducted in order to provide a detailed checklist of the park’s flora, as well as descriptions of its plant communities, which will be valuable in future management of this state park. 42 2002] Gulledge and Judd—Manatee Springs State Park 43 Climate. Northern Florida typically has a humid, subtropical climate (Winsberg 1990). Positioned in the northwestern portion of the Florida peninsula, Levy County experiences both the warming effect of being south of the jet stream in the winter and the cooling effect of the nearby Gulf of Mexico during the sum- mer. Average annual maximum daily temperatures are 78°—80°F while average minimums are only 55°—56°E Winter is mild with 20-30% of days between December and February receiving temperatures above 75°F Cold fronts from the interior United States regularly affect the temperature, however. Around 40% of days between December and February have min- imum temperatures below 40°F (Winsberg 1990). Most winter rainfall is the result of these fronts, but on average, winter weather brings less rain than summer (Chen and Gerber 1990; Jordan 1985). Spring weather usually arrives in March. The polar jet stream passes farther north, and the days are warm and dry. In May average daily maximum temperatures exceed 88°EK Nighttime temperatures rise, and rainfall increases. Afternoon thunderstorms are common by June. These take place an average of 80 days per year, making summer the wettest season. Annually, this region receives an average of 152 cm of rain (Winsberg 1990). Geology. The park is situated in the Ocala Uplift District as part of the lower basin of the Suwannee River. The Ocala Uplift was formed during post-Oligocene orogeny and has little Mio- cene sediment. However, there are large outcrops of Eocene and Oligocene carbonates present at or near the surface (Brooks 1982; Vernon and Puri 1964). Oligocene deposits, usually Suwannee Limestone, are not present in Levy County (Department of Nat- ural Resources 1989). The oldest tertiary sediments in the area are part of the Paleo- cene Cedar Keys Limestone. This formation is a hard, cream- colored to tan limestone with a thickness of 168 to 183 m and was formed in the open ocean when the coastline was located across what is now Alabama and Georgia (Cooke 1945). Over this layer, several Eocene limestone deposits common to the Ocala Uplift can be found. These limestones can be divided into three age groups: the Wilcox, Claiborne, and Jackson groups from oldest to youngest (Cooke 1945). The Oldsmar Limestone belongs to the Wilcox group and contains gypsum and _ chert $4 Rhodora [Vol. 104 (Cooke 1945). This deposit is 122 to 168 m in thickness. Two deposits of the Claiborne group are present in the area. Lake City limestone 1s the older and consists of both dark-brown and chalky limestones with some gypsum beds. Cooke (1945) reported this layer to be around 152 m thick in Levy County. Above this layer is Avon Park limestone, a cream-colored deposit with some gyp- sum and chert embedded in it. This limestone may be anywhere from 15 to 91 m thick (Cooke 1945). The youngest age-group of Eocene limestones, the Jackson group, 1s represented by the Ocala group, one of three limestone subgroups comprising the layer. These are, from oldest to youn- gest, the Ocala, Williston, and Inglis members. These form a layer that is around 60 m deep. This limestone is exposed around the main spring and is mined for road construction in Levy County (Department of Natural Resources 1989). The entire coastal region of Florida lies in the physiographic region called the coastal lowlands, an area that was covered by the sea during the Pleistocene. The ancient shorelines formed sev- eral terraces, of which the Pamlico is the most extensive (Cooke 1945). This shoreline was located at 8 m above current sea level. Elevations within the park are from 8 m above to I.5 m below e mostly sand, but may also = sea level. The deposits of this age a contain some clay (Cooke 1945). Sinkholes are common in the park. The abundance of limestone underlying the park is primarily responsible for the karst topog- raphy found there. Karst is a landscape formed by the action of dissolving carbonate-rich rocks, which form numerous sinkholes and caves (Myers and Ewel 1990). Because the limestone in the Ocala Uplift District is only thinly covered, solution sinkholes are the most common type formed within the park. Surface water seeps through the rock through cracks and gradually dissolves the surface limestone, forming de- pressions over time. This is in contrast to collapse sinkholes, which form after the underlying bedrock has been dissolved and the roof of the cavern formed collapses under the weight of the overlying soil (Beck and Sinclair 1986). Hydrology. The primary feature of the park is the spring, which empties into the Suwannee River. This area has felt the most impact of human influence. The spring is a popular swim- ming hole, and crowds of visitors are common in the summer 2002] Gulledge and Judd—Manatee Springs State Park 45 months. In addition to swimming, scuba diving is a frequent ac- tivity in the spring boil as well as in Catfish Hotel, a nearby sinkhole, which connects via underground passageways to the main spring (Department of Natural Resources 1989). Manatee is classified as a first magnitude artesian spring. This means that the average discharge must be at least 2.83 m+ s7! (100 ft.2 s-'). Manatee discharges an average of 5.13 m° s"' of hard fresh water into a pool 30 m in diameter and 14 m deep at the center (Rosenau et al. 1977). This water maintains a stable average temperature of 22.0°C year-round (Myers and Ewel 1990). The warm temperature attracts manatees into the spring during the winter months (Department of Natural Resources 1989). The spring water travels 381 m westward and empties into the Suwannee River. Classified as a blackwater stream by the Florida Natural Areas Inventory (1988), the Suwannee forms the western boundary of the park. Blackwater streams are characterized by high levels of tannins, particulates, and organic matter from swamp drainage. The pH is 4.0 to 6.0 unless influenced by groundwater (Florida Natural Areas Inventory 1988). Beck (1965) classified the Suwannee as a calcareous stream, mostly of spring origin, and having a pH level of 7.0 to 8.2. Both classi- fications are probably applicable where the spring run meets the Suwannee. While the river originates from swamp drainage and has a dark tannin color, it also receives heavy influence from Florida springs, such as Manatee, that make it locally more cal- careous and clear. This river discharges into the Gulf of Mexico, located only 24 miles southwest of the park (Department of Nat- ural Resources 1989). The Florida Aquifer is largely uncontained throughout this re- gion, meaning that much of the water is not separated from the atmosphere by impermeable rocks or clay beds (Lane 1986). This contributes to the formation of numerous seeps and flooded sink- holes, which can release water. During floods, however, the aqui- fer may recharge through these openings (Myers and Ewel 1990). There are several permanently flooded sinkholes that provide access to an extensive aquatic cave, which 1s also accessible from the spring. These are Catfish Hotel, Freedman Sink, and Sue Sink. Catfish Hotel is the most commonly used point of entry besides the spring. This sink is 38 m in circumference and 12 m deep. Divers have explored around 3978 m of this system, but further 46 Rhodora [Vol. 104 exploration may be dangerous due to the unstable nature of the caves (Department of Natural Resources 1989). [In addition to flooded sinks, the park also contains several sink- hole ponds and a 3 ha sinkhole lake, Graveyard Pond. Close to Graveyard Pond is a 5 ha swamp lake, Shacklefoot Pond. These are located in the northeastern quarter of the park (Department of Natural Resources 1989), History. “About noon we approached the admirable Manate Spring, three or four miles down the river. This charming nympheum ts the product of primitive nature, not to be im- itated, much less equalled, by the united effort of human power and ingenuity! As we approach it by water, the mind of the inquiring traveller is previously entertained, and grad- ually led on to greater discovery...” (Bartram 1791). The naturalist William Bartram was entranced by the beauty of this spring on the Suwannee River. His admiration was un- doubtedly shared by the many Indians and Europeans who trav- eled by or gathered beside this natural fountain. While there is not much information available on the overall history of this land, evidence suggests that the Manatee Spring region has been in- habited by Indians, visited by early explorers, and settled by Flor- ida pioneers (Gulledge 1999), Manatee Spring was visited by William Bartram in 1774 as he traveled through Florida when it was under British control. He described the flora as being dominated by live oaks, red bay, and magnolias. Manatees, fish, and alligators were abundant in the spring run. Indian activity was noted by the presence of a manatee skeleton on the banks of the spring, indicating that the Seminoles probably valued the area as a source of meat. The flora and fauna does not seem to have changed much in the 200 years since his visit. The flow of water from the spring, however, was quite in- teresting at that time. Bartram’s account is of an intermittent eb- ullition from the spring, which occurred every 30 seconds (Bar- tram 1791). The hydrology has changed such that the water now flows continually. Around the turn of the century, longleaf pine was logged throughout much of the area (Department of Natural Resources 2002] Gulledge and Judd—Manatee Springs State Park 47 1989). The effects of this destruction are still evident in the plant composition of the park. In 1949, the majority of Manatee Springs State Park was ac- quired by the Park Board for use as a state recreational park. Additional land was added up until 1988. While public recreation is the designated use of this park, management has been designed to minimize the impact of humans. The addition of paved walk- ways around the spring, a wooden boardwalk along the spring run, and camping facilities in the park were inevitable, and ac- commodate the many people who enjoy this park (Department of Natural Resources 1989). Over time, however, the area’s status as a protected natural area will help to ensure its lasting natural beauty. PLANT COMMUNITIES Manatee Springs State Park has 13 plant communities, as cir- cumscribed by the Florida Natural Areas Inventory 1988 (Figure 1). Although the overall change in elevation within the park is only a gradual nine meters from the river eastward (Department of Natural Resources 1989), the species composition varies sub- stantially along this gradient. Observations on species dominance within each community were recorded as plant collections were made. Here, each community is described based on personal ob- servations and the ecological literature (Figure 1). The recently acquired Meud-Scot track, a small strip of land bordering the river south of the major portion of the park and encompassing 93 ha, was not included in the management plan’s description of natural areas (Department of Natural Resources 1989). Thus coverage of communities within the rest of the park is given as a percentage of the total land area, excluding the Meud-Scot track. The tract consists predominantly of floodplain swamp with a small strip of floodplain forest and an area of xeric hammock. Upland mixed forest. Around 13% (117 ha) of the park ts upland mixed forest, also known as mesic hammock. Some of the original community has been altered due to the development of camping facilities, but it can also be found scattered in other locations, mostly intergrading with xeric hammock (Department of Natural Resources 1989). This intergradation can be gradual, 48 Rhodora [Vol Suwannee River State Road 320 Ip OTe ie] Xe iock Meud-Scot Track |_| Bottomland Forest [BS| Basin Swamp (SL) Swamp Lake (DM) Depression Marsh Sinkhole Lake sinkhole Spring-run Stream Developed/Ruderal P| Parking Lot SK Sin! Scale in Meters 0 200 400 . 104 Figure I. Plant community map of Manatee Springs State Park (adapted from Department of Natural Resources 1989), 2002] Gulledge and Judd—Manatee Springs State Park 49 and these two plant communities often are arbitrarily delimited (see also Platt and Schwartz 1990). Thus, it is usually impossible to separate this vegetational continuum into distinct, easily de- marcated categories. What is called upland mixed forest in this park is simply one of several transition zones from moist to dry woods. The diversity of tree species is usually high in mesic ham- mocks. Hardwoods such as Magnolia grandiflora, Carya glabra, Liquidambar styraciflua, Ostrya virginiana, Ilex opaca, Quercus virginiana, Q. michauxti, and Persea borbonia dominate. Pinus taeda and P. glabra, however, are also found in this community. Around the camping and picnic areas, Sideroxylon lanuginosum and Tilia americana are common. Toward the northern fence, Diospyros virginiana 1s frequent. Symplocos tinctoria is found more abundantly in the eastern part of the park and near Grave- yard Pond, a sinkhole lake found in the northeast corner of the park. In these patches of upland mixed forest, other commonly encountered trees and shrubs are Celtis laevigata, Juniperus vir- giniana var. silicicola, Osmanthus americana, Prunus carolini- ana, P. serotina, Quercus nigra, Sabal palmetto, Vaccinium ar- boreum, and Callicarpa americana. Various species of Smilax are common vines. Typically, the herb layer is not well developed in upland mixed forest, but many grasses and sedges as well as other herbs such as Galium tinctorium, Amsonia tabernaemontana, and Polygala grandiflora are common. The mesic conditions prevailing in upland mixed forests are normally attributed to the higher clay and organic content in the soil, deeper leaf mulch, and dense canopy that traps humidity. The higher moisture content in these areas makes them less likely to burn than the surrounding pine dominated communities (Flor- ida Natural Areas Inventory 1988). Soils in the park that support upland mixed forest are mostly Otela-Tavares complex, but there is also Jonesville-Otela-Sea- board complex underlying this community southeast of the spring (Natural Resources Conservation Service 1996). Xeric hammock. Xeric hammock replaces upland mixed for- est at higher elevations. Typically, it occupies sandy soils of an- cient dune origin. The canopy can be low or multi-layered, open or closed. The presence or absence of these characteristics can often be attributed to the stage of succession. Hardwoods such as 50 Rhodora [Vol. 104 Quercus geminata, QO. virginiana, and Q. hemisphaerica are the most abundant trees, while the herb layer is sparse (Florida Nat- ural Areas Inventory 1988). Xeric hammock covers 57% of the park, or 533 ha (Department of Natural Resources 1989). Manatee Springs is situated on the northwest corner of what is called the Gulf Hammock area, one of the larger regions in Flor- ida to contain extensive hardwood forests (Myers and Ewel 1990). The management plan for the park defines the upland por- tions of the park as consisting of upland mixed forest, upland pine forest, sandhill, scrubby flatwoods, and xeric hammock (De- partment of Natural Resources 1989). However, with the excep- tion of upland mixed forest, the rest of the upland forests are too uniform and intergrading to be divided into four categories (based upon a subjective assessment by the authors), Probably none of these categories adequately describes the actual pattern of species composition within this community. Xeric hammock is here in- terpreted to cover all of these upland forest categories. There are a large number of pines, including Pinus palustris, P. elliottit, and P. taeda. These could indicate that the area might not always have had dominant hardwoods. Prolonged fire exclu- sion may have caused a succession from a pine dominated com- munity to xeric hammock. The records of extensive longleaf pine logging early in the century would support this idea. In addition, some ecologists theorize that, historically, dominance in these for- ests has shifted several times between pines and hardwoods (My- ers and Ewel 1990). Controlled burns are performed on this land and may serve to eventually change its characteristics (Depart- ment of Natural Resources 1989). Another aspect that varies in xeric hammock is the presence in the park of both closed- and open-canopied forests. Areas with Open canopies may have been interpreted as scrubby flatwoods, but the lack of many characteristic elements, along with the roll- ing topography, does not support this classification. Dominant trees and shrubs are Quercus virginiana, Serenoa repens, Vaccinium arboreum, Quercus geminata, Carya glabra, Quercus incana, Q. falcata, Q. hemisphaerica, Q. myrtifolia, Magnolia grandiflora, Liquidambar styraciflua, Persea borbonia, Lyonia ferruginea, Ilex opaca, Osmanthus americana, [lex vom- itoria, and Gaylussacia dumosa. Solidago odora var. chapmantii and Indigofera caroliniana are characteristic herbs. Otela-Tavares 2002} Gulledge and Judd—Manatee Springs State Park 51 complex soils underlie this community (Natural Resources Con- servation Service 1996). Sinkhole. The karst terrain of the park is marked by numer- ous sinkholes of various sizes with a total area of around 6.5 ha (Department of Natural Resources 1989). The majority of these remain dry for most of the year, draining rapidly after periods of rain. However, if the lower reaches of a sinkhole are located be- low the water level, it remains flooded (Florida Natural Areas Inventory 1988). The majority of dry sinkholes in the park are located southeast of the spring. The surrounding community is primarily upland mixed forest. Vegetation in sinkholes is affected by the steepness of the sides and whether or not sand and soil cover the limestone walls (Flor- ida Natural Areas Inventory 1988). Ferns are common as well as lichens and mosses. Most of the sinkholes in the park have grad- ually sloping sides without much exposed limestone. Sinkhole lake. These are sinkholes that retain water and are therefore constantly flooded (Florida Natural Areas Inventory 1988). There is only one large sinkhole lake in the park, called Graveyard Pond, which occupies about 3 ha in the northeast quadrant of the park. Several small sinkhole ponds can be found as well, which are located near the main spring (Department of Natural Resources 1989). The standing water in these sinkhole lakes and ponds allows for a proliferation of aquatic plants such as Lemna obscura, Landoltia punctata, Wolffia brasiliensis, Wolf- fiella gladiata, Pistia stratiotes, and Salvinia minima. Swamp lake. Shacklefoot Pond, located in the northeast cor- ner of the park, is a 5 ha swamp lake (Department of Natural Resources 1989). Although stumps and trees are found in the lake, it is overall an open, permanent body of water surrounded by a basin swamp. Hydrophilic trees are found both on the fringe of the lake and occasionally emerging in the middle. These include 7Taxodium distichum and Gleditsia aquatica. Throughout the lake itself are many floating and emergent aquatic herbs. The most common of these are Lemna obscura, Wolffia brasiliensis, Spirodela punc- ] 1] tata, Wolffiella gladiata, Limnobium spongia, Ceratophyllum de- N NW Rhodora [Vol. 104 mersum, Utricularia foliosa, Boehmeria cylindrica, and Salvinia minima, Basin swamp. There is a large basin swamp in the northeast corner of the park. This community takes up about 11 ha and surrounds Shacklefoot Pond, a large swamp lake (Department of Natural Resources 1989). This community is often flooded, so species occurring within it must be adapted to a long hydroperiod. The soils found on this site are Placid and Samsula soils, acidic peat over a dark gray sand (Natural Resources Conservation Ser- vice 1996). The dominant tree in this basin swamp community is Taxodium distichum. Other common trees and shrubs are Myrica cerifera, Cyrilla racemiflora, Cephalanthus occidentalis, and Salix caro- liniana. The epiphytes Tillandsia usneoides and T. bartramii are common, as 1s the herb Scutellaria integrifolia. There is a large feral hog population in the park, and hogs are especially active in this area (Department of Natural Resources 1989). The damage these hogs do to the surface of the peat is evident throughout the swamp. As a result, herbaceous plants are less frequent here than might be expected for a typical basin swamp. Bottomland forest. A ribbon of bottomland forest covering about 7 ha surrounds the basin swamp around Shacklefoot Pond (Department of Natural Resources 1989). This is basically a sim- ple transition zone from the constantly inundated pond to the surrounding uplands. Along the slope leading down to the pond, there is a gradual increase in the number of flood-adapted species. The appearance is similar to a floodplain forest, but with a more diverse and abundant herb layer (Florida Natural Areas Inventory 1988). Some of the plants found in this community are Quercus nigra, Sabal palmetto, Magnolia grandiflora, Pinus taeda, Toxicoden- dron radicans, Hypericum galioides, and Scutellaria integrifolia. Depression marsh. One small depression marsh exists with- in the park; it is located in the middle of the park and occupies 0.5 ha. The center of the marsh is flooded and the surrounding soil remains moist year round. During some parts of the year, the pond itself contains aquatic herbs, including Brasenia shreberi. The marsh is dominated entirely by herbaceous species such as 2002] Gulledge and Judd—Manatee Springs State Park =i. Eupatorium compositifolium, Xyris platylepis, Juncus margina- tus, and Cuscuta compacta, except for the common shrub, Ce- phalanthus occidentalis. Floodplain swamp. The entire western edge of the park is a floodplain swamp running along the Suwannee River. The swamp occupies around 158 ha or 17% of the total park area (Department of Natural Resources 1989). Much of the swamp remains inun- dated throughout the year. The soils found here are Chobee-Bra- denton complex, Holopaw-Pineda complex, and Chobee-Gator complex, all frequently flooded soils (Natural Resources Conser- vation Service 1996). By far, the dominant tree in the floodplain swamp is Taxodium distichum, with Nyssa biflora also frequent. Other common plants are Saururus cernuus, Crinum americanum, Cephalanthus occi- dentalis, Samolus valerandi subsp. parviflorus, Proserpinaca pal- ustris, and Senecio glabellus. Floodplain forest. Floodplain forests are transitional from floodplain swamps to upland communities and flood less fre- quently than swamps, typically only during peak water levels. Plants in this community are adapted to only seasonal inundation and cannot survive constant saturation of the soil (Florida Natural Areas Inventory 1988). As would be expected, the floodplain for- est occupies a strip of roughly 92 ha between the swamp and the uplands of the rest of the park (Department of Natural Resources 1989). Much of the soil underlying this strip is either of the Ous- ley-Albany complex, Placid, or Samsula soils (Natural Resources Conservation Service 1996). Typical trees are Quercus laurifolia, Q. lyrata, Fraxinus car- oliniana, Planera aquatica, Acer rubrum, Carpinus caroliniana, Sabal palmetto, and Crataegus spp. Common shrubs include Cor- nus foemina, Serenoa repens, and Sabal minor. Toxicodendron radicans and Ampelopsis arborea are characteristic vines. Herbs include Panicum rigidulum and Amsonia tabernaemontana. In both the floodplain swamp and floodplain forest communi- ties in Manatee Springs, feral hogs are a constant destructive force. Although trapping is an ongoing effort for park rangers, feral hog populations are large, and evidence of foraging is wide- spread. 54 Rhodora [Vol. 104 Blackwater stream. The Suwannee River, which forms the western boundary of the park, is classified as a blackwater stream by the Florida Natural Areas Inventory due to the high tannin levels in the water, which give it a characteristic tea color (Florida Natural Areas Inventory 1988). However, a more popular river classification developed by Beck (1965) ranks the Suwannee as a calcareous stream. Both categories are probably generalizations and do not adequately describe the entire river. Blackwater streams are mostly acidic, originate in swamps, and do not usually have either extensive floodplains or large amounts of submerged aquatics (Florida Natural Areas Inventory 1988). Calcareous streams are fed mostly by springs and are generally alkaline with heavy aquatic plant growth. The Suwannee originates in swamps, but in Levy County, it is fed by several large springs such as Manatee, that influence the river locally with calcareous water (Myers and Ewel 1990). With the exception of the entrance to the spring run, emergent plant growth is sparse along the edge of the river. unas both Senecio glabellus and the exotic pest Alternanthera philoxeroides are common. Spring-run stream. The 381 m stream that carries water from Manatee Spring to the Suwannee River is described as a spring-run stream (Department of Natural Resources 1989). This alkaline stream is an excellent habitat for many aquatic herbs, both emergent and submerged. The water is clear, allowing light to filter to the bottom of the limestone streambed and promoting the growth of Vallisneria americana. Periods of heavy flooding such as during the winter of 1998 can cause tea-colored water from the Suwannee to back up into the spring-run stream (Florida Natural Areas Inventory 1988). The 1998 influx resulted in a partial dieback of submerged aquatics, but a drier than typical spring helped return the stream to its previous state. In the streambed itself, Vallisneria americana is the dominant submerged plant. Along the fringe, however, there are numerous species of both submerged and emergent plants including Cabom- ba caroliniana, Sagittaria kurziana, Nuphar advena, Echinodorus berteroi, and Pontederia cordata. Also present is the exotic pest Hydrilla verticillata. Ruderal and developed areas. Due to the popularity of Manatee Springs as a swimming hole and camping area, devel- 2002] Gulledge and Judd—DManatee Springs State Park 55 opment has occurred. The most frequented area is the spring It- self. A parking lot, bathhouse, and picnic area are present to ac- commodate visitors, as well as a concrete ramp around part of the spring and a small beach area for swimmers to enter the spring with as litthe damage to the remaining edge as possible. A board- walk extends along the spring-run stream to a boat dock on the Suwannee River. There are also two camping areas near the spring and two residences within the park. The development around the spring takes up about 4 ha (Department of Natural Resources 1989). In addition to disturbances such as roads and trails, there is a large borrow pit located southeast of the spring. Disturbed zones are usually dominated by early successional weeds such as Eupatorium compositifolium, Paronychia ameri- cana, Gnaphalium purpureum, G. obtusifolium, and Ambrosia ar- temisiifolia. The roadside is especially diverse in the fall, with numerous composites such as Coreopsis leavenworthii, Liatris elegans, L. graminifolia, Pitvopsis graminifolia, and Solidago odora var. chapmanii. Other notable roadside plants are Dicer- andra densiflora, Trichostema dichotomum, and Arenaria serpyl- lifolia. The borrow pit area contains a large population of the exotic Leonitis nepetefolia, as well as Eupatorium compositifol- ium and Rhynchosia michauxti. MATERIALS AND METHODS Plant collections were made from May 1996 to November 1998. Most of the park could be covered by walking the trail system, with occasional transects into the woods. Exceptions to this were the spring-run stream, riverbank, and floodplain swamp/ floodplain forest boundary. For the spring-run stream and river- bank, a canoe was used to survey the edge, and a mask and snorkel was necessary to find several aquatics growing in the spring. Foot trips were made that generally followed the flood- plain swamp edge and the boundary of the park to assess the species richness in the disturbed vegetation along the fence. Soil maps and previous plant community maps (Department of Nat- ural Resources 1989) were used to identify areas of interest. The plants were initially identified using Clewell (1985), Wun- derlin (1982), and Godfrey and Wooten (1979, 1981). However, after Wunderlin (1998) was published, this guide was used for 56 Rhodora [Vol. 104 most of the remaining identifications. Vouchers were deposited in the University of Florida Herbarium (FLAS). RESULTS The authors found a total of 360 vascular plant species in the park, representing 253 genera and 100 families. The largest fam- ilies were Poaceae (41 spp.), Asteraceae (36 spp.), and Fabaceae (27 spp.). The largest genera were Quercus (13 spp.), Dichan- thelium (8 spp.). Cyperus (6 spp.), Ryachospora (6 spp.), Smilax (5 spp.), /lex (5 spp.), and Vaccinium (5 spp.). The complete annotated list of the vascular plants of the park is found in the appendix. DISCUSSION There were several species of special concern in the park. These were broken down into the following categories: taxonom- ic problems, species at or near their geographical limits, exotic and endemic species, and rare or endangered species. There was an interesting Yucca population in the park that did not seem to fit completely the description for Yucca filamentosa. This entity was a robust plant with stiff leaves up to a meter in length. It bloomed in late July, slightly later than the more com- mon form of this species. This plant may represent a taxonomic entity distinct from the widely distributed form, frequently treated as Y. flaccida. Further study is needed to determine the exact pattern of variation within the Y. filamentosa complex. Several species were at the limits of their geographical ranges. An on-line atlas was used to determine species ranges within Florida (Wunderlin et al. 1997). These plants were divided into several categories for this list (.e., species at their limit and spe- cies near their limit for both northern and southern limits). A species at its southern limit does not occur in any counties south of Levy (and the reverse for a species at its northern limit). A species near its southern limit only occurs one or two counties further south (and the reverse for a species near its northern limit). Ten species were at their southern limit: Pinus glabra, Sium suave, Betula nigra, Quercus lyrata, Dichanthelium oligosanthes, Saccharum alopecuroides, Crataegus aestivalis, C. michauxii, Galium tinctorium, and Planera aquatica. Nineteen species were 2002} Gulledge and Judd—Manatee Springs State Park DT Table |. State-listed endangered (E), threatened (T), and ain exploited (CE) vascular plants occurring in Manatee Springs State Park, fol- lowing Coile (1993). Species Status Asplenium platvneuron T Epidendrum conopseum T Tlex ambigua T Ilex decidua flex opac CE seated cardinalis T telea floridana E pani i alis CE Sabal n T Tille sae ne T Woodwardia areolata T Zamia integrifolia CE near their southern limit: Pinus taeda, Justicia ovata, Sagittaria kurziana, Asimina longifolia, Ostryva virginiana, Triadenum wal- teri, Cuscuta compacta, Cornus asperifolia, Carex dasycarpa, Cyperus plukenetti, Baptisia alba, Desmodium canescens, Les- pedeza stuevei, Quercus michauxti, Carya tomentosa, Fraxinus americana, Halesia carolina, Ulmus alata, and U. crassifolia. Only a few species were at or near their northern limits for Florida. These were Jillandsia recurvata and Senna ligustrina (at northern limit), as well as Zamia integrifolia, Pistia stratiotes, Utricularia foliosa, Cenchrus gracillimus, Phlebodium aureum, and Ulmus crassifolia (near northern limit). Non-native, or exotic, species following Wunderlin (1998) found in the park were Alternanthera philoxeroides, Chenopo- dium ambrosioides, Cyclospermum leptophyllum, Pistia strati- otes, Arenaria serpyllifolia, Cyperus lanceolatus, Crotalaria lan- ceolata, Desmodium canescens, Hydrilla verticillata, Sisyrin- chium rosulatum, Hyptis mutabilis, Leonitis nepetefolia, Brous- sonetia papyrifera, Eremochloa ophiuroides, Lolium perenne, Paspalum notatum, Poa annua, Secale cereale, Sporobolus in- dicus, Richardia brasiliensis, and Xyris jupicai. There were nine Florida endemics or near endemics (see Muller et al. 1989) grow- ing in the park: Aristida patula, Coreopsis leavenworthii, Dicer- andra densiflora, Matelea floridana, Palafoxia integrifolia, Pyc- nanthemum floridanum, Rhynchosia michauxii, Solidago odora var. chapman, and Vicia floridana. 58 Rhodora [Vol. 104 State-listed endangered, threatened, and commercially exploit- ed plants are summarized in Table 1. No federally listed endan- ered species were found in the park. The flora of Manatee Springs State Park is a reasonable rep- resentation of plants that would be expected in natural commu- nities bordering the Suwannee River in Florida. Exotic plants, while common, are still much less prevalent inside the park than in the surrounding areas, proving the benefits of good land man- agement. It is hoped that continued protection will maintain < diverse and historically representative flora of the region. Sf) pad) LITERATURE CITED ADAMS, R. P. 1986. Geographic variation in Juniperus silicicola and Juni- perus virginiana Of the southeastern United States: Multivariate analysis — of morphology and terpenoids. Taxon 35: 61-75. ANGIOSPERM PHYLOGENY GROUP. 1998. An ag c classification for the fam- ilies of Howering plants. Ann. Missouri Bot. rd. 85: 531-55: rns a 1791 [edited by M. van Doren, oo The Travels of William Bartram. Dover Publ., New York. Beck, B. EF AND W. C. SINCLAIR. 1986. Sinkholes in Florida: An introduction. Rep. 85-86-4, Sinkhole pee poe College of Engineering, Univ. Central Florida in coop. with . Geol. Surv., Orlando, FL. Breck, W. M., | 1905. The ae of ee Bull. sete State Mus., ie Sci. 10: 91-126 BENSON, ne The Cacti of the United States and Canada. Stanford Univ. Press, Stato d, Brooks, H., 1982. Guide to the Physiographic Divisions of Florida. Florida oop. - Serv., o Food Agric. Sci., Uniy. Florida, Gainesville, FL. BURKHALTER, R,. E. 1992. The genus Nyssa (Cornaceae) in North America: 2: + 323 A revision. Sida | CAMPBELL, C. S. 1983. Stematie of = Andropogon virginicus complex (Gramineae). J. Arnold Arbor. 64: 171—254. CAULKINS, D. B. AND R. Wyatt. 1990 Variation and taxonomy of Phytolacca americana var. rigida in the soutieasiers United States. Bull. Torrey Bot. Club 117: 357-367. —_ ri AND J. EK GERBER. 1990, Climate, pp. 11-34. /n: Myers, R. L. and ; _ a Ecosystems of Florida. Univ. Central Florida Press, a CLEWELL, A. . ve 985. Guide to the Vascular Plants of the Florida Panhandle. Florida State Univ. Press, Tallahassee, FL. CoiLe, N. C. 1993. Florida’s endangered and threatened plants. Bureau of Entomology, Nematology, and Plant i eerie Section, con- D tribution #26, Florida Dept. Agric. and Consumer Services, Div. Plant Industry, rataene a 2) Gn CooKE, C. W. ee of Florida. Bull. No. 29, Florida Geol. Surv., ea a - 2002] Gulledge and Judd—Manatee Springs State Park 59 DEPARTMENT OF NATURAL RESOURCES. 1989. Manatee Springs State Park Unit Management Plan. Unpubl. ee Florida Dept. Natural Resources, Recreation and Parks, Region 2, Gainesville, FL. FLORA OF NORTH AMERICA EDITORIAL oo E, eds. 1993+. Flora of North America North of Mexico, Vol. ser oar ska and Gymnosperms. Ox- ford Univ. Press, Oxford and Nev v Yor FLORIDA NATURAL AREAS INVENTORY. a Natural Community Classifica- tion ae Element Abstracts. Tallaha L. FURLO . J. 1987. The Carpinus ete He complex in ae fe ee I. A ene analysis of geographical variation. Syst. Bot. 12: 21—40. Goprrey, R. K. AND J. W. WooTEN. 1979. Aquatic and Wetland ere of the Southeastern United States: Monocotyledons. Univ. Georgia Press, Ath- ens, GA. AND ———. 1981. secre and Wetland Plants of the Southeastern United States: Dicotyledons. Univ. Georgia Press, Athens, GA. GULLEDGE, K. J. 1999. A floristic inv ae of Manatee Springs eo Park, a County, a M.S. thesis, Univ. Florida, Gainesville, Hai, D. 1985. Letter to Donald K. Yonker, September 23. cen Dept eee Resources, Div. Recreation os Parks, Region 2, Gainesville, a JORDAN, C. L. 1984. Florida’s weather and climate: Implications for water, 18-35. In: E. A. Fernald and D. J. Patton, eds., Water Resources ia of Florida. Univ. Press Florida, Gainesville, FL, Jupp, W. S., C. S. CAMPBELL, E. A. KELLOGG, AND P. F STEVENS. 1999, Plant Systematics: A Phylogenetic Approach. Sinauer A cace: Inc., Sun- derland, MA. . W. SANDERS, AND M. J. DONOGHUE. 1994. See pau pairs: > Sanne phylogenetic analyses. Harvard Pap. Bot. a): Sls tS KRAL, R. 1997. Annonaceae, pp. 20. In: Flora of North America Editorial Committee, eds., Flora of a pata North of rae Vol. 3. Mag- noliophyta. Oxford Univ. Press, Oxford and New York. LANE, E. 1986. Karst in Florida. nat Geol. Surv. pie Publ. No. 29, Bureau Geol., Florida Dept. Natural Resources, Tallahassee, PL. D. J. CRAWFORD. 1999. Landoltia (Lemnaceae), a new genus 30-533 — L of ee Novon 9: LUTEYN, J. L., W. S. Jupp, S. . VAN DER KLorT, L. J. Dork, G. D. WALLACE, A. Kron, P. E STEVENS, AND S. E. CLEMANTS. 1996. Ericaceae of the mapa so States. Castanea 61: 101-144 MULLER, C. 1970. Quercus, pp. 467-492. In: D. S. Correll and M. C. aman As. Manual of the Vascular Plants of Texas. Univ. Texas at Dallas, Richardson, TX. MuLLer, J. W., E. D. HAarbIN, D. R. JACKSON, S. E. GATEWOOD, AND N. CAIRE. 1989. Summary report on the vascular plants, animals, and gab com- munities endemic to Florida. Nongame Wildlife Progr. Tech. Rep. No. 7, Florida Game and Freshwater Fish Commission, Tallahassee, F Myers, R. L. AND J. J. EWeL, eds. 1990. Ecosystems of Florida. Univ. Central Florida Press, Orlando, FL. NATURAL RESOURCE CONSERVATION SERVICE. 1996. Soil Survey of Levy 60 Rhodora [Vol. 104 County, Florida. U.S.D.A. in coop. with Univ. Florida, Inst. of Food and Agric. Sciences, Agric. Expt. Stations, and Soil and Water Sci. Dept and Florida Dept. Agric. and Consumer Services; U.S.D.A., Washington, Nixon, K. C. AND C. H. Mutter. 1997. Quercus Linnaeus sect. Quercus— White Oaks, pp. 471—S06. /n: Flora of North America Editorial Com- mittee, eds., Flora of North America North of ahi ol. 3. Magno- ee Oxford Univ. Press, Oxford and New PLattr, W. J. AND M. W. ScHWartz. 1990. Temperate Hardwood Forests, pp. ipa 229. . R. L. Myers and J. J. Ewel, eds., Ecosystems of Florida. Univ. Central Florida Press, Orlando, FL. ReEHDER, A. J. 1945. Carya alba proposed as nomen ambiguum. J. Arnold Arbor. 26: ns 483. Rosenau, J. C., G. L. FAULKNER, C. W. HENDRY, JR., AND R. W. Hui. 1977. Springs of lace Bull. No. 31 (revised), Bureau Geol., Florida Dept. Natural Resources, Tallahassee, FL. ScHOLZ, U. 1981. eee ie gattung Oplismenus (Gramineae). Pha- nerogamarum Monographiae Tomus XVIII. J. Cramer, Vaduz, Germany. THORNE, R. FE 1983. Proposed new alignments in the Angiosperms. Nordic J. Bot. 3: 117. VERNON, R. O. AND H. S. Puri. 1964. Geologic Map of Florida. Div. Geol. Map Ser. No. 18, U.S. Geol. Surv. in coop. with Florida Board of Con- aa rea FL. WIERSEMA, J. AND C. B. HeLLQuist. 1994, Nomenclatural notes in Nym- phase or os North American flora. Rhodora 96: 170-178 WINSBERG, M. D. 1990. Florida Weather. Univ. Central Florida Press, Orlan- do, EL. WUNDERLIN, R. P. 1982. Guide to the Vascular Plants of Central Florida. Univ. Press Florida, Gainesville, FL. . 1998. ine to the Vascular Plants of Florida. Univ. Press Florida, ae . B. woe NSEN, AND E. L. BripGes. 1997. Atlas of Florida Vascular Plants. ie Systematic Botany, Univ. South Florida Web Site (http:// www.plantatlas.usf.edu/). APPENDIX ANNOTATED LIST OF VASCULAR PLANTS The species names in this list follow Wunderlin (1998), fern and gymno- sperm family _circumscriptions follow Flora of North ee a North of Mex- ico (Flora of North America Editorial Committee 1993), and pare ies family circumscriptions follow the Angiosperm Phy co Group (1998), cept when stated otherwise. The abbreviations for plant communities are as a UMEF — Upland Mixed Forest; XH — Xeric Hammock; SH — Sinkhole; SKL — Sinkhole Lake; L — Swamp Lake; BS — Basin Swamp; BF — ene Forest; DM — Depression Marsh; FS — Floodplain Swamp; FF — Floodplain Forest; BLS — Blackwater Stream: SR — Spring-run Stream; RU — Ruderal/Developed. Some ~ 2002] Gulledge and Judd—Manatee Springs State Park 61 additional notes may also be given regarding a locations. For abun- ‘ol dance, the lowing abbreviations were used, based on , e collectors’ ob- on of the plant in each community: R — rare (1— peal a I - infrequent (S—9 observations); — occasional es le vations): F — fre- quent (25 or more observations); A — abundant (denotes a plant O; ai dle cei ae Lam. — RU; O; /0, a s Muhl. ex Willd. — RU; F; 4/7, /3/, new. Se album L. — XH; F; 438. E. comp Ae aa Walter — DM ; RU; F; /25, /44. FE. rotundifolium L. — UMF; pe hana Pensstvanicum aa — RU; O; 237. urpu 1L. — RU; O; 239, 306. anaes a area ty (Lam.) Britton & Rusby — RU; I; 58/. Hieracium gronovilt L. — XH & RU; O; 475. Krigia virginica (L.) Willd. — XH & RU; O; 242. Lactuca graminifolia Michx. — RU; O; 88, 287. Liatris elegans (Walter) Michx. — RU; F; /24, new. L. graminifolia (Walter) Willd. — RU; F; /23. L. tenuifolia Nutt. var. tenuifolia — RU; O; 728. Melanthera nivea (L.) Small — FF; R; 488. Mikania scandens (L.) Willd. — FS; I, on river island opposite spring run entrance; //2, //3. Palafoxia integrifolia (Nutt.) Torr. & A. Gray — RU; O; /36. Pityopsis graminifolia (Michx.) Nutt. — RU; A; /37. Pyrrhopappus carolinianus la ad = XH & RU; I; 57, 244, 307. Senecio glabellus FS; Solidago odora in var. anne ce & A. Gray) Cronquist — XH & RU; F; /45, 473. Vernonia angustifolia Michx. — XH; O; 78. BETULACEAE Betula nigra L. — FF; O; 4/0. Carpinus caroliniana Walter subsp. caroliniana — FF & UME; F; 93. (Furlow 1987) Ostrya virginiana (Mill.) K. Koch — UMF; F; /7/. BIGNONIACEAE Bignonia capreolata L. —- UMF & XH; O; 232 Campsis radicans (L.) Seem. ex Bureau — FF & UME; F; 6/2. BRASSICACEAE Lepidium virginicum L. — RU; *Rorippa nasturtium-aquaticum oe ‘ Myc SR; Hall NV. BROMELIACEAE Tillandsia bartramii Elliott — BS & BF; F; /65. T. recurvata (L.) L. —- UME FS, FE BE BS & XH: F; 6/0. 66 Rhodora [Vol. 104 T. usneoides (L.) L. — XH, UME BE FF & BS; A; /S4. BURMANNIACEAE Apteria aphylla (Nutt.) Barnhart ex Small — BS; Hall NV. Burmannia biflora L. — FS; 1; 45, 495, new. CABOMBACEAE (See NYMPHAEACEAE) CACTACEAE Opuntia humifusa (Raf.) Raf. var. humifusa — RU; I; 5. (Benson 1982) CAMPANULACEAE Lobelia cardinalis L. — FS: O, ee se river; /O/7, 102. ee In Triodanis perfoliata (L.) Nieuwl. — CAPRIFOLIACEAE pro parte, (i.e., Viburnum—see ADOXACEAE) CARYOPHYLLACEAE *Arenaria serpyllifolia L. — RU; I; 243a, 546. Drymaria cordata (L.) Willd. ex Schult. — RU; O; 24/7, 568, new. Paronychia americana (Nutt.) Fenzl ex Walp. — XH & RU; O; 7/. P. baldwinti (Torr. & A. Gray) Fenzl ex Walp. — XH & RU; I; 576. Stipulicida setacea Michx. var. setacea — RU; I, 602 CELTIDACEAE Celtis laevigata Willd. — UMF; F; 6/4. CERATOPHYLLACEAE Ceratophyllum demersum L. — SL: 1: 569. ‘HENOPODIACEAE (See AMARANTHACEAE) CHRYSOBALANACEAE Licania michauxti Prance — XH; O; 333. CISTACEAE Helianthemum eae a es Michx. — RU; I; 247. Lechea minor L. — RU; O:; 477, n CLUSIACEAE as um crux-andreae ) Crantz — RU; I, 472, new. H. galioides Lam. — BF; 30, 44, 307, 575. H. hypericoides (L.) Crantz. — UMF & RU: O; 89. 2002] Gulledge and Judd—Manatee Springs State Park 67 H. mutilum L. — BF; O; 27, 346. Triadenum walteri (J. F Gmel.) Gleason — BS & BF; I; 574. COMMELINACEAE Commelina erecta L. — XH: O; S/. CONVOLVULACEAE Cuscuta compacta Juss. — DM; O; 497, new. Dichondra carolinensis Michx. — XH & o O; 250. Stylisma patens (Desr.) Myint — UMF; R; CORNACEAE (incl. NYSSACEAE) Cornus aspertfolia a — FF; O; 74. C. foemina Mill. — FF; O; 253, 452. Nyssa biflora i — FS; F; 459. (Burkhalter 1992) N. sylvatica Marshall var. sylvatica — XH; I, but locally frequent at one site on east side of park; 455, new. CYPERACEAE Bulbostylis eet aces ae XH & RU; I; 379. Carex dasycarpa C. granularis Muhl. ex ae in ae — FF & RU; O, on edge of spring run; 422. C. longiti Mack. — RU; I; 36, 268, ses Cyperus croceus ue — RU; Hall N C. distinctus Steud. F& oy vs on edge of spring run; 423. C. filiculmis Vahl — = & RU; 320 C. flavescens L. — FS, DM & ae “Hall NV. ®C. lanceolatus Poir. in Lam. — BS; I; 353, new. C. plukenetti Fernald — XH; O; 84, new. C. polystachyos Rottb. — FF & DM; Hall N C. retrorsus Chapm. — UME XH & RU; F; on 149, 150. *C. rotundus L. — RU; Hall NV. C. strigosus L. — FF & RU; O, on SS ae os run; 42/, 424. C. tetragonus Elliott — FF & UMP; Eleocharis baldwinii (Torr.) ae = eas oe 3371. E. montevidensis Kunth — Dd, 341. Kyllinga odorata Vahl — BS & DM; Hall NV. Rynchospora colorata (L.) H. Pfeiff. - FF & RU; F; /7. corniculata (Lam.) A. Gray — FS; O; 75, 179. inundata (Oakes) Fernald — FS & RU: I; 466. megalocarpa A. Gray — XH & RU; F; 92, 3/6, 334. microcarpa Baldwin ex A. Gray — DM & BS; O; 335. plumosa Elliott — BS; I; 60. Scleria reticularis Michx. — DM; F; 337. S. triglomerata Michx. — UMF; O; 395, Ais een ae: 68 Rhodora [Vol CYRILLACEAE Cyrilla racemiflora L. — FF; I, 7/4. EBENACEAE Diospyros virginiana L. — UMF; F; 430, 462. ERICACEAE Gaylussacia dumosa (Andrews) Torr. & A. Gray — XH; O; 6/3. Lyonia ferruginea (Walter) Nutt. — XH; F; 382, — Vaccinium arboreum Marshall — UME F & XH; O; 2, 6, 8. V. darrowiti Camp — XH; 1; S88. V. elliottii Chapm. — FF & UMF; F; /, /64, 592. (Luteyn et al. 1996) V. myrsinites Lam. — XH; [; 322, 536 VY. stamineum L. — UMF; F; 9, 446. ERIOCAULACEAE Lachnocaulon anceps (Walter) Morong — DM; A; 328. ESCALLONIACEAE (see ITEACEAE) EUPHORBIACEAE Acalypha gracilens A. Gray — UMF; O; 37 Chamaesyce maculata - .) Small — XH & — , 579. . prostrata (Aiton) nall — RU; . 104 all NV. Cnidoscolus pian eee (ch ) nasi, & A. Gray — XH & RU: F; 236. Croton glandulosus L. — RU; C. michauxti G. L. Webster — ae & O; SO, 373, 435. Phyllanthus ee ae Walter — FS; I; S593. *P. urinaria L. — ull N Stillingia sylvatica Garden ex i — XH; F; &/, 3/7] >, FABACEAE Amorpha fruticosa L. — XH; 1; 434. A. herbacea Walter var. herbacea — XH; O:; 246, new. Baptisia alba (L.) Vent. — UMF; O; 4, /66. B. lecontii Torr. & A. Gray — XH; I; 302. Centrosema virginianum (L.) Benth. — XH & RU; O; 49, 305. Chamaecrista fasciculata (Michx.) Greene — RU: O; /32. Clitoria mariana L. — RU; 1; 304. *Crotalaria lanceolata E. Meyer — RU: O; 477. C. rotundifolia Walter ex J. F Gmel. — RU: O; 50, 237. * Desmodium canescens (L.) DC. — RU: O; 440, new. D. acer (L. z DC. — XH; Hall NV. a tio (L.) DC. — RU; O; /47/. Erythrina ei ea - ; 16. Galactia volubilis (.) Britton — UME a & RU; , 9O, 372, 429. 2002] Gulledge and Judd—Manatee Springs State Park 69 Gleditsia aquatica Marshall — SL; I; 275, 355. Indigofera caroliniana Mill. — XH; . 47. Lespedeza hirta (L. - pine — RU: I; /43. L. stuevei Nutt. — , 409, Medicago ee - - a ; oat Mimosa quadrivalvis L. var. angustata (Torr. & A. Gray) Barneby — XH; O; 62, 314 Rhynchosia difformis (Elhott) DC. — RU; O; 476. R. michauxti Vail — RU; O; 436, 607. Senna ligustrina (L.) H. S. Irwin & Barneby — UMF; I; 4/2. S. marilandica (L.) Link — FF; F; 437. S. obtusifolia (L.) H. S. Irwin & ae — RU; O; 433. Tephrosia chrysophylla Pursh — RU: I; 601. T. florida (EF Dietr.) C. E. Wood — -_ Te 303,.3:76. *Trifolium repens L. — RU: Hall NV. Vicia floridana S. Watson — BS; O; 26, 257. FAGACEAE Quercus austrina Small — UMF; O; 384, new. (Nixon and Muller 1997) . chapmanii Sarg. H: 1; 599. . falcata Michx. — XH: O; 277, 29S, 442, new. . geminata Small — XH; A; 298, 386, 387. . hemisphaerica W. Bartram — UMF & XH; F; /00, 276. (Muller 1970) incana W. Bartram — XH; Hall NV. . laurifolia Michx. — FF; O; ee (Muller 1970) . lyrata Walter — FF; O; 493, 603. margaretta Ashe ex Small — XH; 1; 447. . michauxti Nutt. —- UMF; O; 500. QO. a Willd. — XH; O; 444, 527, 537. QO. nigra L. - UMF & BF; O; 385. Q. belie Walter — XH; O; 5206. Q. virginiana Mill. — XH & UMEF: F; 673. - ee GELSEMIACEAE Gelsemium sempervirens (L.) W. T. Aiton — FF & UMEF; O; 257. GENTIANACEAE Bartonia paniculata (Michx.) Muhl. — BS; R; 496, new. Sabatia calycina (Lam.) A. Heller — BS & FS; O; 28, 66, 344. HALORAGACEAE Proserpinaca palustris L. — FS; O; 63. 70 Rhodora [Vol. 104 HAMAMELIDACEAE (see ALTINGIACEAE) HIPPOCASTANACEAE (see SAPINDACEAEF) HYDROCHARITACEAE drilla verticillata (L. f.) Royle — SR; I; 407. *Hy Limnobium spongia (Bosc) Steud. — SL: 1; 567, new. Vallisneria americana Michx. — SR; A; 479. Hypoxis curtisti Rose — FS; F; 65, /63. IRIDACEAE Sisyrinchium angustifolium oe - — le-235, S. nashii E. P Bicknell — R *§ rosulatum E. P. Bicknell — me - 310. ITEACEAE Ttea virginica L. — FS; Hall NV. JUGLANDACEAE Carya glabra (Mill.) Sweet — UMF & XH; F; 258, 379. C. tomentosa (Poir. in Lam.) Nutt. — UMF & XH; O; 332. (Rehder 1945) JUNCACEAE Juncus dichotomus Elliott — FF & RU; O, on edge of spring run; 420. J. marginatus Rostk. — DM; O; 39, 336, 564. LAMIACEAE Callicarpa americana L. ME; O; S59. Dicerandra densiflora a — RU; O; /26. *Hypris mutabilis (Rich.) Brig. — RU: *PLeonitis nepetefolia (L.) R. Br. in W. T. Aiton — RU; F; 3/2, new. Micromeria brownei (Sw.) ae — RU; O, in lawns around spring; /9, 400. Monarda punctata L. — RU; I; 566. Pycnanthemum floridanum e Grant & Epling — RU; I; 582. Salvia lyrata L. — RU; [; 282. Scutellaria integrifolia L. — BS; F; 25, new. Teucrium canadense L. — BS & BF; O; 327. Trichostema dichotomum L. — RU; O; 1/27. LAURACEAE bes borbonia (L.) Spreng. — pee & XH; F; 4/3, 449. P. palustris (Raf.) Sarg. — BS; 2002] Gulledge and Judd—Manatee Springs State Park 71 LEMNACEAE (see ARACEAE) LENTIBULARIACEAE Utricularia foliosa L. — SL; O; 570. LOGANIACEAE (also see GELSEMIACEAE) Mitreola petiolata (J. F Gmel.) Torr. & A. Gray — FS; F; 64, //8. MAGNOLIACEAE Magnolia grandiflora L. - UME BF & XH; O; 99. MALVACEAE (incl. TILIACEAE) Sida rhombifolia L. - RU & XH; O; S6/. Tilia americana L. var. caroliniana (Mill.) Castigl. — UMF; O; 94. MELASTOMATACEAE Rhexia mariana L. — UMF & XH; O; 46, 325 MORACEAE *Broussonetia papyrifera (L.) Vent. — XH; 1; 598. MY RICACEAE Myrica cerifera L. — BS, FE BF & UMF; F; 96, 294, 297, 299, 573. NYMPHAEACEAE (incl. CABOMBACEAE) Brasenia shreberi J. F Gmel. — DM; F:; 330. Cabomba caroliniana A. Gray — SR: A; //7 Nuphar advena (Aiton) W. 'T. Aiton — BLS; oO 6/5. (Wiersema and Hellquist NYSSACEAE (See CORNACEAE) OLEACEAE Fraxinus americana L. — BS; 447a, new. F. caroliniana Mill. — FS & FF: O; //0, 354. Osmanthus americanus (L.) Benth. & Hook. f. ex A. Gray — UMF & XH; Ope 7: ONAGRACEAE Gaura angustifolia Michx. —- UMF & RU; I Ludwigia repens J. R. Forst. — FS; R: 467. Oenothera laciniata Hill — RU; I; 559. 72 Rhodora [Vol. 104 ORCHIDACEAE Epidendrum conopseum R. Br. — XH; 1; 508. OXALIDACEAE Oxalis corniculata L. — RU: I: 238. PASSIFLORACEAE Passiflora incarnata L. — RU: R: 37/5. P. lutea L. — XH; I; S58. PHYTOLACCACEAE Phytolacca americana L. var. rigida (Small) Caulkins & Wyatt — RU; I; 324. (Caulkins and Wyatt 1990) PLANTAGINACEAE ert ae L. — RU; Hall NV. P. virgi ..— RU; O; 547. POACEAE Andropogon glomeratus (Walter) Britton, Sterns & Poggenb. var. pumilus Vasey — XH; O; 535. SS 1983) A. ternarius Michx. — XH; O; 502 A. virginicus L. var. decipiens C. S. Campb. — XH; O; 7/57, new. (Campbell 1983) A. virginicus L. var. virginicus — XH; F; 503, 505. (Campbell 1983) Aristida patula Chapm. ex Nash — RU; I; 482. Axonopus affinits Chase — BF; O; 55 A. furcatus (Fliieggé) Hitche. — BF: Cenchrus gracillimus Nash — RU; oe on C. tncertus M. A. Curtis » PF; 494. Chasmanthium sessiliflorum (Poir.) Yates —- UMF & RU: F; 38, 56, 82, 339, / 349, 35]. *Cynodon dactylon (L.) Pers. — RU; Hall NV. Dichanthelium aciculare (Desv. ex Poir.) Gould & C. A. Clark — XH: O; 397. D. acuminatum (Sw.) Gould & C. A. Clark var. acuminatum — UME & FF: O; 37, 263 D. commutatum (Schult.) Gould — UMF & FF; F:; 35, 283, 287, 352. D. dichotomum (L.) Gould — FS; I; 464. D. ensifolium (Baldwin ex Elliott) Geula-n es & RU; O; 340. D. oligosanthes (Schult.) Gould — XH: O: D. portoricense (Desv. ex Ham.) B. F a & Wunderlin — XH; O; 285. D. strigosum (Muhl.) Freckmann — BF: O; 269. Digitaria ciliaris (Retz.) Koeler — RU; O; 426. D. serotina (Walter) Michx. — RU: Hall NV. Eleusine indica (L.) Gaertn. — RU: Hall NV. Eragrostis elliottii S. Watson — RU: R; /46. 2002] Gulledge and Judd—Manatee Springs State Park ie FE. virginica (Zucc.) Steud. - DM & RU: Hall NV. *Fremochloa ophiuroides (Munro) Hack. — FF; I; 428. Eustachys petraea ae ek — RU; O; /38, 266. *Lolium perenne L. — , 540. Oplismenus hirtellus : ean subsp. sefarius (Lam.) Mez ex Ekman — UMF&R F; 565. (Scholz 1981) Panicum anceps Michx. — DM, BS & BF; F; 83, /48, 338, 350, 393, 481]. P. rigidulum Bosc ex Nee EF: O; ia 456, 506. *Paspalum notatum Fliepeé —-RU&L , O; 54, 317. P, esos Michx. — ar O: 318. P. repens Bergius — BLS; I; /08. P. oe eum Michx. — RU; ms SS, 425, 439. Piptoc haetium avenaceum ety Parodi — XH: I; 288, 289, 394, new. ww *Poa annua L. — RU; F; Saccharum alopec upoiaes - Nutt. — XH; O; 50/, new. S. baldwinii Spreng. — FF; E on riverbank only; 463. Sacciolepis striata L) re — FS & BS; Hall NV. *Secale cereale L. — ; R; 562. Setaria geniculata aie ) Millsp. & Chase — XH & RU; O; /56, 479. (God- i and Wooten 1979) Sor 2 eee elliottii (C. Mohr) Nash — XH; O; S04. ae nopholis obtusata (Michx.) Scribn. — BS; O; 539. ‘ robolus indicus (L.) R. Br. var. indicus — RU: O; 392. Sanne nan secundatum (Walter) Kuntze — RU; O:; 674. Vulpia elliotea (Raf.) Fernald — RU; 1; 538, new. POLYGALACEAE Polygala grandiflora Walter — UMF & XH: O; 23, 52, 134. POLYGONACEAE Eriogonum tomentosum Michx. — XH: I; 580. Polygonum densiflorum Meisn. — FS, DM & BS; Hall NV. P. punctatum Elliott —- BS & SR: O; 42, 343, 416 Rumex hastatulus Baldwin — RU: I; 2 PONTEDERIACEAE Pontederia cordata L. — SR: O; 104, 105. PRIMULACEAE Samolus valerandi L. subsp. parviflorus (Raf.) Hultén — FS; F; 20, 33. RHAMNACEAE Berchemia scandens (Hill) K. Koch — FS: R: /60. Rhamnus caroliniana Walter — UMF: Hall NV. 74 Rhodora [Vol. 104 ROSACEAE Crataegus aestivalis (Walter) Torr. & A. Gray — FF; O; 4/5, new. C. crus-galli L. — FS; Hall NV. C. marshallii Egg]. — FF; Hall NV. C. michauxti Pers. — XH; 1; 272, S57 Prunus caroliniana (Mill.) Aiton — UMF; O; /2. P. serotina Ehrh. var. serotina — UMF; O; 260, 525. P. umbellata Elliott — XH; I; 380, new. Rubus argutus Link — BS; Hall NV. R. cuneifolius Pursh — XH; O; 329. R. trivialis Michx. — XH; O; /77. RUBIACEAE Cephalanthus Reade L.— BS, FS & DM; O; 326. Diodia teres Walter — QO: /40, 427. D. virginiana L. — FF; ms a. 408, 450. Galium hispidulum sera — XH & RU; O; 249, 474. 7. tinctorium — BS; I; 486 *Hedyotis corymbosa re an — RU; Hall NV. H. procumbens (Walter ex J. KE Gmel.) Fosberg — XH; O; 532. H. uniflora (L.) Lam. — UMF; ey NV. Mitchella repens L. — UME; O; 234 *Richardia brasiliensis Canes — RU: |; 79. RUTACEAE Ptelea trifoliata L. — UMF; Zanthoxylum clava-herculis = cae I. SST. SALICACEAE Salix caroliniana Michx. — FS; O; 259, 4/7. SAPINDACEAE (incl. ACERACEAE and HIPPOCASTANACEAE) Acer rubrum L. — FF & FS; I; 448. Aesculus pavia L. — XH; 1; 432. SAPOTACEAE Sideroxylon lanuginosum Michx. — UMF; O; 497, 530, 587. S. reclinatum Michx. subsp. reclinatum — FF; 1; 490, 492. SAURURACEAE Saururus cernuus L. — FS: A; 18, 106. SCROPHULARIACEAE Bacopa monnieri (L.) Pennell — SR: O:; 453. Gratiola virginiana L. — BS; I, 670, new. 2002] Gulledge and Judd—Manatee Springs State Park Linaria canadensis (L.) Chaz. — RU; O; 543. L. floridana Chapm. — RU; O; 243b. Mecardonia acuminata (Walter) Small — DM; Hall N Micranthemum umbrosum (J. EF Gmel.) S. EF Blake — i > 672, new. Veronica peregrina L. — RU; O; 240, 545. SMILACACEAE Smilax auriculata Walter — RU & XH: 7 se 323, 483, SSS. S. bona-nox L. — FE UME XH & RU: S. glauca Walter — XH; I; 297. S. pumila Walter — UMF & UMF; O; /69. S. smallii Morong — RU & XH; O; 590. STYRACACEAE Halesia carolina L. — XH; I; 3/7, 383, 528. SYMPLOCACEAE Symplocos tinctoria (L.) LHér. —- UMP: A; 280, 387. TETRACHONDRACEAE Polypremum procumbens L. — SL & SKL; Hall NV. TILIACEAE (see MALVACEAE) TURNERACEAE Piriqueta caroliniana (Walter) Urb. - UMF & RU; O; /3, 53, 91. ULMACEAE Planera aquatica Walter ex J. F Gmel. — FS & FR; O; /6/, 604. Ulmus alata Michx. — FS; O; /62 americana L. — FS; O; 254. U. crassifolia Nutt. — FS; O; 454. URTICACEAE Boehmeria cylindrica (L.) Sw. — SL; O; 470, 577. VERBENACEAE *Lantana camara L. — XH & RU; Hall NV. Phyla nodiflora (L.) Greene — RU; F; 402. VIOLACEAE Viola palmata L. — XH; I; 233. V. sororia Willd. — UMF; I; /S68. 75 76 Rhodora [Vol. 104 VISCACEAE Phoradendron leucarpum (Raf.) Reveal & M. C. Johnst. — UMF; O; 274. VITACEAE Ampelopsis arborea (L.) Koehne — FF & UME; O; /07. Parthenocissus quinquefolia ee ) iS aee — FE UME BF & XH; F; 4/7. Vitis aestivalis Michx. — XH; V. rotundifolia Wiehe _ red & on F; 390. XYRIDACEAE *Xyris cee Rich. — FS; O, but locally A on north edge of Meud-Scot » 44]. trac Xx. plaiylepts Chapm. — DM; I; 589. RHODORA, Vol. 104, No. 917, pp. 77-82, 2002 NEW ENGLAND NOTE ANEURA MAXIMA (HEPATICAE: ANEURACEAE) IN NORTON G. MILLER Biological Survey, New York State Museum, Albany, NY 12230-0001 1] sy e-mail: nmiller2 @ mail.nysed.gov Aneura maxima (Schiffn.) Steph. Maine: Kennebec Co., Mud Pond, ca. 5 km SW of Litchfield along Highway 126, 44°12'N, 69°58'W, bottom of an animal run over wet peat, minerotrophic edge of fen mat near pond, 19 Sep 1987, Miller 9497 (Nys). Two species of Aneura are recognized in the North American flora by Schuster (1992), the common and variable A. pinguis (L.) Dumort., and A. maxima, a species only recently discovered to be widespread in eastern North America but previously known in the flora of tropical and temperate Asia. A third species, A. sharpti Inoue & N. G. Mill. (Inoue and Miller 1985) has also been recognized, but in this note I tentatively accept it as a syn- onym of A. maxima, following the circumscriptions and interpre- tations of Schuster (19972). Aneura maxima is based on plants first collected in Java and Sumatra (Schiffner 1898). Its known range was subsequently ex- tended to include other parts of Asia, notably Japan, eastern North America (Schuster 1992), and very recently western and north- western Europe (Finland, Frahm 1997; Belgium, Andriessen et al. 1995; France, Sotiaux and Sotiaux 1996). The pattern of mor- phological variation in North American populations of A. maxima sensu lato is poorly understood, because the species has been collected infrequently so far in our area, and male plants and female ones with mature calyptrae and sporophytes are few or unknown throughout the range of the species. There is only one previous station for Aneura maxima in New England, namely, Rutland County, Vermont, in a fen near the Connecticut River (as A. sharpii; Inoue and Miller 1985). Oth- erwise, the reported North American distribution of A. maxima (incl. A. sharpit) is eastern New York State, central Pennsylvania, West Virginia, Tennessee, Mountain and Piedmont provinces of £4 78 Rhodora (Vol. 104 North Carolina, and Louisiana (Inoue and Miller 1985; Reese and Walters 1987; Schuster 1992). Aneura maxima appears to be un- common in all these regions. Aneura maxima and A, pinguis differ vegetatively in the fol- lowing ways: thallus margins regularly lobate, short, lateral ar- chegonial branches (gynoecia) in most sinuses; unistratose thallus wings 10—20 cells wide, sometimes more; thick, opaque, multis- tratose mid-thallus region narrow, about one-third of the plant width (A. maxima; Figures 1—6), versus thallus margins only sometimes irregularly and unevenly lobate or sinuate, lateral si- nuses bearing archegonial branches scattered; unistratose thallus wings when developed (especially in lax plants from moist or wet habitats) to 10 cells wide but usually fewer; thick mid-thallus region wide, sometimes the entire width of the plant, but usually two-thirds (or more) of the plant width (A. pinguis; Figures 7, 8). Thalli of Aneura maxima are similar to those of Pellia and Moerckia. When present, the short, lateral, ciliate archegonial branches of female plants of A. maxima (visible only from the underside of plants) easily separate species of Aneura from those of the other two genera. In plants of Pellia and Moerckia, sex organs are variously disposed on the upper thallus surface. An- theridial branches of male plants of A. maxima are also short and lateral, but they extend beyond the thallus margins and therefore can be seen from the upper surface of the plant. Too few plants of Aneura maxima with calyptrae and mature sporophytes are known at present in North America to evaluate potential differentiating character states in these life cycle com- ponents. Schiffner (1900), Furuki (1991), and Schuster (1992) stressed that the female inflorescences of A. maxima contain long or very long cilia (paraphyses, sensu Furuki 1991), whereas in A. pinguis they are scalelike (Schuster 1992). However, in young archegonial branches (1.e., those with archegonia cap cells intact) in North American plants I have studied, uniseriate and multis- eriate, scalelike paraphyses are present in both A. maxima and A. pinguis (Figures 9—22). Therefore, at a young stage of develop- ment, it does not seem possible to differentiate between the spe- cies on the basis of paraphysis morphology. However, this may not hold for plants with mature calyptrae. Plants of Aneura pinguis can be highly variable throughout its nearly cosmopolitan range, but one segment of the variation ap- proaches A. maxima in thallus morphology. In a frequently en- 2002] New England Note 79 Figures 1-6. Aneura maxima. \—3, thalli, upper surfaces, note lobate wings and narrow mid-thallus rhizoidal region where the thallus is also thick- est; 4—6, thalli, lower surfaces, note the regular and repetitious occurrence of archegonial branches, which remain small and cushion-like [Maine, Miller 9497 (NYS)]. countered expression of A. pinguis, the plants are an oily or greasy green, compact, brittle, and multistratose to the margins. However, lax plants in wet habitats can have lateral wings similar to those of A. maxima but without the regular lobate configuration of this species. Illustrated in Figures 7 and 8 are the obverse and reverse aspects of the same plant of A. pinguis from a wet, shrub- 80 Rhodora [Vol. 104 Figures 7 & 8. Aneura pinguis, plant from a wet habitat (circumneutral carr). 7, thallus, upper surface, note irregular, mostly nonlobate wings and broad, thick mid-thallus region; 8, same plant, lower surface of thallus, ar- chegonial branches few and irregular in position [Maine, Aroostook Co., Thousand Acre Bog, Crystal, Miller 13262 (Nys)]. by fen margin in north-central Maine. The lateral thallus wings in this plant and others in the collection are up to 10 cells wide and the cells are arranged in fan-shaped tiers, suggesting that they grew out from the edge of the massive tissue in the central part of the thallus. Plants of this morphological type retain the wide, thick central thallus region and irregularly placed archegonial branches typical of compact expressions of A. pinguis in drier Sites: In the northern portion of its known range in eastern North America (New England and New York), Aneura maxima has been found on wet peat in fens beneath a shrub or herb cover and on 2002] New England Note 81 Figures 9-22. Aneura maxima and A. pinguis, filiform and scalelike ar- chegonial paraphyses, all from archegonial branches at the same stage of development. 9-13, A. maxima, showing variation from filiform and simple to multiseriate and branched, archegonia in 9 semidiagrammatic [Maine, Mill- er 9497 (NYS)|; 14-18, - ie from a lax plant in a hygric ee ane: Miller 13262 (Nys)]; 19-22, A. pinguis, from a compact plan a mesic habitat [Michigan, a . shore of Weber Lake, Miller oe (NYS)]. wet organic-rich muck in a Lythrum salicaria L. wetland under a dense, tall, herb overstory. To the south in West Virginia, Ten- nessee, North Carolina, and Louisiana, it grows perhaps exclu- sively on wet rock in streams and over cliff faces, and on stream banks 82 Rhodora [Vol. 104 LITERATURE CITED ANDRIESSEN, L., A. SOTIAUX, C. NAGELS, AND O. SoTIAUX. 1995. Aneura maxima (Schiffn.) Steph. in Belgium, new for the European liverwort flora. J. Bryol. 18: 803-806. FRAHM, J.-P. 1997. A second European record for Aneura maxima (Schiffn.) steph. in Finland. Lindbergia 22: 99 FuruKI, T. 1991. A taxonomical revision of the Aneuraceae (Hepaticae) of japan. 7 a Bot. Lab. 70: 293-397. INOUE, H. AN _G. Mitcer. 1985. A new Aneura (Hepaticae: Aneuraceae) Gor eastern North America. Bull. Natl. Sci. Mus., Tokyo, B. I 1: 95— 101 Reese, W. D. AND D. A. WATERS. 1987. Two bryophytes new to Louisiana. Castanea 52: 147. SCHIFFNER, V. 1898. Expositio plantarum in itinere suo indico annis 1893/94 suscepto collectarum speciminibusque exsiccatis distributarum, adjectis descriptionibus novarum. Series prima. Hepaticarum partem continens. Denkschr. Kaiserl. Akad. Wiss., Wien. Math.-Naturwiss. Kl. 67: 153— . 1900. Die Hepaticae der Flora von Buitenzorg, Vol. 1. Brill, Leiden, Netherlands. SCHUSTER, R. M. 1992. The Hepaticae and Anthocerotae of North America East of the Hundreth Meridian, Vol. 5. Field Museum of Natural History Chicago, IL. SoTiaux, A. O. AND M. Soriaux. 1996. Aneura maxima (Schitfn.) Steph., hépatique nouvelle pour la flore frangaise. Bull. Soc. Bot. Centre-Ouest I. 17: 513-516. RHODORA, Vol. 104, No. 917, pp. 83-85, 2002 NOTE SCIRPUS ANCISTROCHAETUS (CYPERACEAE): FIRST RECORD IN CANADA STUART G. HAY Herbier Marie-Victorin, Université de Montréal, 4101 est, rue Sherbrooke, Montréal (Québec), HI X 2B2, Canada 1 e-mail: stuart.hay @ ca GORDON C. TUCKER Department of Biological Sciences, Stover-Ebinger Herbarium, Eastern Illinois University, Charleston, HL 61920-3099 e-mail: cfgt@eiu.edu A recent revision of material of Scirpus atrovirens sensu lato at the herbarium of the Université de Montréal has turned up an interesting discovery of Scirpus ancistrochaetus Schuyler from the Shawinigan region of Québec (valley of the Riviére Saint- Maurice). This discovery is based on a misidentified collection dating from 1934. It represents the first record of this species in Canada. ECIMEN CITATION: CANADA. Québec: Sainte-Flore (village), comté de Saint-Maurice, lac Mondor, Rive basse, 15 aoat 1934, Gauthier 223] (mr). The approximate coordinates are 46°37'N, 72°44'W. Scirpus ancistrochaetus 1s a relatively unknown bulrush spe- cies that was first described by A. E. Schuyler in 1962 (Schuyler 1962). At the time, Schuyler discovered it in several widely iso- lated localities in the northeastern states of Vermont and Penn- sylvania. Presently, it is known from about 60 localities scattered through the Appalachian region from southwest New Hampshire, adjacent Vermont, and New York to western Virginia. Within this fairly restricted area, it is listed by the United States Fish and Wildlife Service as a Federal Endangered species (USFWS 1991), because it is rare or endangered in all states where it is known to occur (Kartesz and Meacham 1999; Mitchell and Tucker 1997: NatureServe 2000; Royte and Lortie 2000; Strong 1994). This bulrush has sparked considerable interest because of its relatively recent description and its rarity throughout its range. Several studies have been initiated in different states to better 83 84 Rhodora [Vol. 104 evaluate its status and further document sites where it is known to occur (NatureServe 2000). Wherever it occurs, it seems that populations are small and several occurrences are only known historically. The state of Pennsylvania has the highest number of extant populations (Lentz 1998). To explain the isolated occur- rences of this bulrush in the northeastern part of the continent, Schuyler (1962) has suggested that it may be a relict species that is persisting only in pockets of its former range. The Québec specimen was previously identified as Scirpus atrovirens Willd. var. georgianus (R. M. Harper) Fernald. Plants by this name in our area are now referred to as S. hattorianus Makino, as distinct from S. atrovirens sensu stricto (Schuyler 1967). However, a closer examination of the specimen revealed that it was neither S. hattorianus, nor S. atrovirens. In fact, sev- eral well-marked characteristics described by Schuyler (1962, 1967) and Strong (1994) permit us to distinguish this species from the other members of the complex. The inflorescence rays of S. ancistrochaetus tend to droop more at maturity as opposed to the ascending rays of S. atrovirens and S. hattorianus. The rays are also antrorsely scabrous their entire length rather than smooth. The bristles of the achenes are more rigid with sharp- pointed, retrorse teeth that extend nearly to the base, while the bristles of S. atrovirens and S. hattorianus are weaker and have teeth that are finer and concentrated towards the tip of the bristle. To further complicate matters, hybridization has been reported to occur with S. atrovirens and/or S. hattorianus (Schuyler 1962, 1967). In August 2000, a brief attempt was made by Hay to re-locate Scirpus ancistrochaetus at the Lake Mondor locality. The shore- line and adjacent wetland were explored, but although other closely related species such as S. hattorianus (2000-28, 29, MT) and S. microcarpus C. Presl (2000-24, 27, 30, MT) were common, no populations of S. ancistrochaetus were found. The discovery of this species in Québec is a major extension in range from the previous most northern sites known in the Con- necticut River Valley, and thus, continues to raise questions about the status and distribution of this unusual species. As our under- standing of this species improves, further field exploration and a more exhaustive search of other herbarium specimens will likely uncover other new occurrences. Given what we know presently, and particularly because of its rare status throughout its range in 2002 | Note 85 eastern North America, Scirpus ancistrochaetus should be added to the list of rare plants in Québec (Bouchard et al. 1983; Lavoie 1992) and Canada (Argus and Pryer 1990). ACKNOWLEDGMENTS. Mark Strong of the U.S. National Her- barium kindly confirmed the identification of the specimen. Sara Cairns of the State of New Hampshire, Department of Resources and Economic Development made available reports on the status of the species that were done for the New Hampshire Natural Heritage Inventory. LITERATURE CITED ArGaus, G. W. AND K. M. Pryer. 1990. Rare vascular plants in Canada natural heritage. Canadian Museum of Nature; Ottawa, ON, Canada. 19 Our BOUCHARD, A., D. BARABE, M. DUMAIS, AND S. Hay. 3. The rare eer plants of Sa eti ec 48. National Museum of Natural Sciences, Ottawa, - ana KARTESZ, J. T. AND C. x Me ACHAM. 1999, Synthesis of the North American Flora, Cb- FON version 1.0. North Carolina Botanical Garden, Chapel Hill, NC. Lavoir, G. 1992. Plantes vasculaires susceptibles d’étre désignées menacées ou vulnérables au Québec. Direction de la Conservation et du Patrimoine Ecologique, Ministére de |’ Environnement du Québec, Québec, Canada. LENTZ, K. A. 1998. Ecology of endangered northeastern Bulrush, Scirpus rn yen — Ph.D. dissertation, Pennsylvania State Univ., University Park, MITCHELL, R. S. AND . c. TUCKER. oe Revised checklist of New York State plants. Bull. 490, New York State Museum, Albany, NATURESERVE. 2000. An i aie of life [web application], ver- sion 1.1. Association for Biodiversity Information, Arlington, VA. Web Site (http://www.natureserve.org/). Royte, J. L. AND J. PR Lortir. 2000. New records for Scirpus ancistrochaetus 13 in New pare Rhodora 102: 210-2 SCHUYLER, A. E. 1962. A ne ae of Scirpus in the northeastern United States. vee 64: 43-46 1967 yea revision of North American leafy species of Scnpue. ies Acad. Nat. Sci a Areas 119: 295-323. STRONG, M. T. 1994. Taxonomy of irpus, Tric hoon and Schoenoplec- 1s (Cipeuecae) in Virginia. ae 58: 29-68. aeewe (UNITED STATES FISH AND WILDLIFE SERVICE). 1991. Endangered and threatened wildlife and plants; determination of endangered status for Scirpus ancistrochaetus (northeastern bulrush). Fed. Reg. 56: 21091— 21096 RHODORA, Vol. 104, No. 917, pp. 86-91, 2002 NOTE SCHIZAEA PUSILLA IN NORTH CAROLINA RICHARD J. LEBLOND North C arolina Natural Heritage Program, Division of Parks and on PO. =o eee Richlands, NC 285 e-mail: 1 ond@ncf lom.net ALAN S. WEAKLEY NatureServe, 311 Boothe Hill Road, Chapel Hill, NC 27514 e-mail: alan_w eal ley (a PHatubescl Ve.Orge Schizaea pusilla Pursh, the curly-grass fern, has been found growing at a single location in a white cedar forest in Green Swamp in southeastern North Carolina. A survey of nearby suit- able habitat has failed to establish the presence of another pop- ulation, and an analysis of the extant site suggests that the pop- ulation is introduced rather than native, raising problematic con- servation issues. Schizaea pusilla has long been one of the most eagerly sought plants among professional and amateur botanists in North America. Its allure is certainly related to its rarity, its curious disjunct range, and its inconspicuous and unfernlike ap- pearance. The distribution of S. pusilla is centered in the pine- lands of southern New Jersey, but includes nearby Long Island, New York, and Sussex County, Delaware, disjunct populations in Nova Scotia and Newfoundland, and a remarkably disjunct oc- currence in Peru (Montgomery and Fairbrothers 1992; Stolze 1987; Wagner 1993). The North Carolina Natural Heritage Program (Division of Parks and Recreation) has considered Schizaea pusilla among a list of species “‘not currently known to occur in North Carolina, but which are considered to have some possibility of being found in North Carolina, based on their currently known range and hab- itat preferences” (Amoroso 1997). Schizaea pusilla was consid- ered as potentially occurring in the Coastal Plain and fall line sandhills in “‘boggy sphagnous sites associated with white cedar”’ (Amoroso 1997). The suggested possibility of finding S. pusilla 86 2002] Note 87 in North Carolina 1s based on the presence of potentially suitable habitat, and the close biogeographic and floristic relationship be- tween the Coastal Plain Pine Barrens of southern New Jersey and Coastal Plain pinelands of southeastern North Carolina. This re- lationship 1s demonstrated by the many plant species exhibiting a disjunct distribution between the two areas, and often also in- volving other areas, such as the East Gulf Coastal Plain (Florida panhandle, southern Alabama, southwestern Georgia, southern Mississippi, and southeastern Louisiana), New Brunswick, and Nova Scotia. A few examples are Rhynchospora pallida M. A. Curtis, Gentiana autumnalis L., Lophiola aurea Ker Gawl., and Leiophyllum buxifolium (Bergius) Elliott. Current floristic similarities are based on underlying habitat similarities, both areas having strongly acidic sandy soils, abun- dant saturated wetlands, and fire as a frequent and vegetation- shaping natural force. Moreover, these areas have had past con- nections, and during recent glacial periods, plant species now more typical of New Jersey, including Schizaea pusilla, occurred in North Carolina: “At full-glacial time, a continuous coastal plain from Florida to Cape Cod was exposed. At the latitude of the Outer Banks, this plain was an estimated 90 miles in width; its vegetation can be partially reconstructed from palynological studies of the Dismal Swamp, Virginia, and southeastern North Carolina. ... The forests of Virginia were apparently more boreal in aspect—spruce was possibly the dominant tree and fir was probably not uncommon. In southern North Carolina at this time, red or jack pine (perhaps both) were apparently the dominant species; spruce was much less abundant and fir was very uncommon. A number of northern species including Lycopodium lucidulum, L. annotinum, Schizaea pusilla, and Sanguisorba canadensis occurred” (Burk 1968). Such recent vegetational similarities and the prehistoric occur- rence of S. pusilla in North Carolina suggest the plausibility of the presence of relict populations. On June 18, 1997, the first author discovered Schizaea pusilla in a moist, peaty opening in a forest community dominated by Chamaecyparis thyoides (L.) Britton, Sterns & Poggenb. at Green 88 Rhodora [Vol. 104 Swamp in Brunswick County, North Carolina (portion of one individual collected, 18 Jun 1997, LeBlond 4757, NCU). On June 22, 1997, the two authors returned to the site and conducted a careful investigation, and also investigated four other Chamae- cyparis stands in the vicinity. The Chamaecyparis stands them- selves were searched, as well as nearby open habitats, such as bogg areas and moist savanna edges. This strategy was sug- gested by the habitat of the species in New Jersey: “Schizaea occurs in the open bogs, not within dense white cedar forests. Plants are found at bases of young or isolated cedar trees, or stumps or logs, or on edges of peat hummocks including ed of old sand roads” (Montgomery and Fairbrothers 1992). At the discovery site, six individuals of Schizaea pusilla grew on a peat hummock about | m by 0.5 m, and about 3 dm high. The hummock was in one of many small openings in an other- oeS ges yh wise dense Chamaecyparis stand, which is classified as a Cha- maecyparis thyoides/Persea palustris/Lyonia lucida — Hex cori- acea Forest (Weakley et al. 1998) or as Peatland Atlantic White Cedar Forest (Schafale and Weakley 1990). Immediately associ- ated with S. pusilla were Drosera intermedia Hayne, D. rotun- difolia L., D. filiformis Rat., seedlings of C. thyoides, seedlings of L. lucida (Lam.) K. Koch, and Sphagnum spp. More generally associated in the surrounding community were C. thyoides, Cy- rilla racemiflora L., Vaccinium formosum Andr., Gaylussacia frondosa (L.) Torr. & A. Gray sensu stricto, Eubotrys racemosa (L.) Nutt., Persea palustris (Raf.) Sargent, Smilax laurifolia L., Mex myrtifolia Walter, 1. coriacea (Pursh) Chapm., and Myrica heterophylla Raf. Some of these species are frequently associated with S. pusilla in its occurrences in southern New Jersey (D. Snyder, pers. comm., New Jersey Natural Heritage Program), and it is notable that Montgomery and Fairbrothers (1992) state that “the best indicator associates are thread-leaf sundew (Drosera filiformis) and Carolina clubmoss (1; carolinianum).” Four additional white cedar stands and associated open habitats were searched carefully, and although microhabitats similar to those at the first site were seen, no plants of Schizaea pusilla were found. This raised the question of whether the discovered population of S. pusilla is native, or is the result of planting (or the intentional or unintentional scattering of spores by a human). We considered the following lines of evidence: 2002] ie iY Oo = Nn on Note 89 The habitat, location, and associated species are very plau- sible for a native occurrence of Schizaea pusilla in south- eastern North Carolina, showing similarities to its natural habitats in southern New Jersey. The site with Schizaea pusilla is one of the most accessible and well-known white cedar stands in southeastern North Carolina, and has a small trail into it from a nearby road. The additional four stands investigated (and lacking S. pus- illa) have less ready access. The small trail into the Schizaea pusilla site had been flagged relatively recently, and a flagged wooden stake of unknown purpose was in the opening on the hummock sup- porting Schizaea. The second author had searched the site for Schizaea pusilla in late 1980s and did not find any. Of course, S. pusilla is an inconspicuous plant, and the opening which has created apparently excellent conditions for S. pusilla is recent. Growing within a few centimeters of Schizaea pusilla were a few individuals of both Drosera rotundifolia and D. fili- formis. Drosera rotundifolia is not known to occur in south- eastern North Carolina (though it does occur in the moun- tains of North Carolina, with a few disjunct populations in the fall line sandhills). Drosera filiformis (sensu stricto) is a rare plant in southeastern North Carolina, known from eight extant populations. Notably, its habitat in North Car- olina is in open seasonally-flooded depressional wetlands, and it has not been known to occur in, or in proximity to, Chamaecyparis stands in North Carolina. This suggests that both Drosera species were introduced as seeds or small plants along with S. pusilla; it is also possible that D. fili- formis rather than Schizaea was the intentional introduction. An alternative interpretation would be that notably disjunct populations often indicate unusual habitats or relictual con- ditions, and that disjunct populations of other species often co-occur at such sites. The plants of Schizaea pusilla were examined carefully to assess whether they had been transplanted. They appeared to be well established. No apparent discontinuity of soil could be seen; the peaty material at the immediate base of the plants appeared no different than that in the vicinity. If the plants were transplanted, it is likely that they have been 90 Rhodora [Vol. 104 at the site for at least several years, with enough time having passed for the incorporation and intermeshing of soil ma- terial. The authors have seen a privately printed document which reports that four occurrences of Schizaea have been known from Green Swamp since the early 1990s (Murray 1995). The author of this privately printed document is a naturalist familiar with habitats in both the New Jersey Pine Barrens and southeastern North Carolina, but we are not convinced that these reported populations are naturally occurring. In our opinion they are likely based on deliberate introduc- tions. Schizaea pusilla has previously been the subject of a delib- erate introduction to a new state, into an artificial cranberry bog in Massachusetts (B.A. Sorrie, pers. comm., formerly of Massachusetts Natural Heritage Program). = CO The authors conclude that the preponderance of evidence sug- gests that the single site of Schizaea pusilla discovered in North Carolina ts the result of transplantation, but that it is also plausible (though less likely) that this represents a native population. Based on current evidence, S. pusilla is best considered a nonindigenous and marginally naturalized component of the North Carolina flo- ra. Even if this population ts the result of introduction, it remains possible that S. pusilla occurs in North Carolina at undiscovered native populations; botanists should continue to seek S. pusi/la in likely habitats in North Carolina and adjacent eastern South Car- olina and southeastern Virginia. It is unfortunate that the native/introduced status of Schizaea pusilla in North Carolina cannot be determined more definitively. If native, the newly discovered population would warrant consid- erable conservation effort, attention, and resources by conserva- tion organizations and governmental agencies responsible for the conservation of biodiversity in North Carolina; if introduced, it would not. Uncertainty about the native status of populations of plants causes difficulties for scientists, Conservation organiza- tions, and government agencies in determining the native distri- butions of taxa, and the appropriate conservation priorities and actions needed. Plants with high profiles in the amateur botanical world, such as orchids, ferns, and insectivorous species, are par- ticularly likely to be introduced to areas outside their native dis- O02 | Note 9] tributions, and then be found and reported as range extensions. Examples include Dionaea muscipula J. Ellis in Alabama, Flor- ida, western North Carolina, Virginia, and southern New Jersey (all introduced); Drosera filiformis and D. intermedia in West Virginia (considered introduced); Sarracenia leucophylla Raf. in eastern North Carolina (probably introduced); and various Sar- racenia spp. in eastern Virginia and New Jersey (introduced). The authors urge that the introduction of species to natural areas, such as nature preserves or multiple-use public conserva- tion lands, be avoided. It has been abundantly documented that such introduced species can cause unforeseen management prob- lems (though in this case it 1s difficult to imagine curly-grass fern as a pest species outcompeting another species!). Even if the de- liberate introduction does not become a problem, other species are often introduced unintentionally as well, and these may be- come aggressive colonizers. If species are introduced, every effort should be made to document their introduced status in the pub- lished literature to avoid future confusion regarding native distri- butions, and conservation and management priorities. LITERATURE CITED Amoroso, J. L. 1997. Natural Heritage Program List of the Rare Plant Species of North Carolina. N.C. Natural Heritage Program, Div. Parks and Rec- reation, cota 2 NC. Burk, C. J. 1968. A floristic comparison of lower Cape ie petra snl the North orn eas Banks. Rhodora 70: 215 MONTGOMERY, J. AND D. E. FAIRBROTHERS. 1992. New aa Ferns and Fern-allies. ctens ee Press, New Brunswick, NJ. Murray, R. 1995. Tales of the Green Swamp, Brunswick County, North oe Privately printed, copy at N.C. Natural Heritage Program, Ra- leig a M. Pp AND S. WEAKLEY. 1990. Classification of the Rea ese testy of ca Carolina: Third Approximation. N.C. Natural e Program, Ralei Sones . G. 1987. See nasile discovered in Peru. Amer. Fern J. 77: 64-65 WAGNER, W. H., JR. 1993. Schizaeaceae, pp. 112-113. Im: Flora of North nena ae Committee, eds., Flora of North America North of exico, Vol. ace oe and Gymnosperms. Oxford Univ. Press, eae and New WEAKLEY, A. S., K. D. Pow S. LANDAAL, M. PYNE, ET AL. (compilers). 1998. Inter ane Classification of Ecological Communities: Terrestrial Vegetation of the Southeastern United States orking draft of March 1998. The Nature Conservancy, Southeast eee Office, Southern Conservation Sci. Dept., Community Ecol. Group, Chapel Hill, NC. RHODORA, Vol. 104, No. 917, pp. 92-95, 2002 IN MEMORIAM ROLLA MILTON TRYON, JR. 1916-2001 SCIENTIST, TEACHER, AND MENTOR Rolla M. Tryon, Jr. world-renowned pteridologist and long- time member of the New England Botanical Club, died in Tampa, Florida on August 20, 2001 six days short of his 85th birthday. Rolla will be remembered for his scientific contributions to bot- any, for his role as a teacher and a mentor to his students and colleagues, and for his service to many professional organiza- tions, especially to the New England Botanical Club. As a scientist, Rolla authored over 100 articles, papers, and books (Gastony et al. 2001). These ranged from his first, at age 9? 2002] In Memoriam 93 18, on ferns of the genus Osmunda in the Indiana dunes, to tax- onomic revisions of selected fern genera and analyses of fern biogeography, and finally to his comprehensive treatment of the free-sporing vascular plants of the Americas entitled Ferns and Allied Plants, with Special Reference to Tropical America, co- authored with his wife and research partner, Alice E Tryon. Rolla had a deep interest in the geography of organisms. With the advent of chemical and later, molecular methods for probing the secrets of the evolutionary process, some came to think that the only important information was that carried in the molecular warehouse of the cell. Rolla clearly saw the folly of this. He never lost sight of the forest for the trees. He was able to maintain the broad perspective that the process of divergent evolution involves organisms changing through time and space. He was fascinated by the relationship of organisms to space, that is, to their geog- raphy. He understood that organisms may disperse and migrate, but that their geographic range is the direct result of a chain of events leading back in time to the place where divergent evolution occurred. Although that place may never be known, Rolla seemed to have an innate understanding that the evolutionary process ts hugely affected by geography, by that interaction of the geology, climate, and biodiversity of a region. Thus, no evolutionary study was complete without a thorough analysis of what could be learned about an organism’s geography. This was wonderful for his students. We were encouraged to go to the field as much as possible, to see the plants in their natural habitats, and to learn what we could about the natural history of the place where these organisms occurred. Rolla was one of those rare individuals who was awed by the natural beauty that 1s the result of the evolutionary process. For many, it is the human art forms such as sculpture or architecture that bring inspiration. But for Rolla, it was the natural beauty of the free-sporing vascular plants that he found most inspiring. His appreciation of these plants was expressed in many ways. He was a fine artist and won prizes for his watercolor paintings of ferns. He was an avid fern grower and he and Alice always had a back- yard filled with ferns they had transplanted from the countryside. He was especially fond of Botrychium and often, when he re- turned from a visit to the farm in Indiana, brought a potted grape fern for his desk in the office at the Gray Herbarium. But mostly, he expressed his appreciation for these plants by devoting his life 94 Rhodora [Vol. 104 to their study. Rolla truly loved observing, analyzing, and writing about ferns and this was his ultimate form of human expression. For those fortunate enough to have known him well, watching Rolla initiate, fully engage, and focus his attention on a project, and then bring the research to publication was perhaps the ulti- mate academic experience. Rolla had an incredible ability to fo- cus his attention and he had a way of rapidly bringing order to disorder. It seemed that he could see in his mind’s eye, not only the scientific conclusions that could be reached from an analysis of the data, but the entire narrative that would unfold, before he ever began to write. The writing was just the final step of putting what had already been assembled in his mind onto paper. For those of us who struggle with the process of getting our thoughts into coherent form, this was awesome to witness. Students always know when the professor loves his or her sub- ject and those are the courses and experiences that have the great- est impact on students’ intellectual development, even if they do not particularly like the subject that 1s being taught. But when they do like the subject, the professor often becomes an inspira- tion, propelling students on to ideas, to places, and to careers that were never imagined. Rolla was such an inspiration to many stu- dents. This, plus his kindness, his willingness to help even when a student’s ideas were in direct conflict with his own, and his example of a life devoted to an incredibly high standard of work have inspired a generation of pteridologists and countless others who have pursued careers in all walks of life. Rolla received his graduate training at Harvard between 1938 and 1941 where he was mentored by Charles Alfred Weatherby and Merritt Lyndon Fernald, both of whom were influential in Rolla’s early years (Barrington et al. 2001). Following graduate school he held positions at Dartmouth, the University of Wiscon- sin, the University of Minnesota, the Missouri Botanical Garden, and the University of California at Berkeley, finally returning to New England in 1958 where he would spend the rest of his career as Curator and then Professor of Biology at Harvard University until retirement in 1987. While he was at Harvard, Rolla made important contributions to botany in New England. He was a member of the New England Botanical Club for 60 years from 1941 to 2001. During that time he served the Club as Recording Secretary (1964-1968), as an Associate Editor of Rhodora (1961-1977), as Editor-in-Chief of Rhodora (1977-1981), as 2002] In Memoriam 95 Vice President (1984—1986), and as President of the Club (1986— 1988). He also served on the Council for many years. Rolla was also organizer, with Alice Tryon, of the New England Fern Con- ference, an annual regional meeting for pteridologists held at the Harvard Forest in Petersham, Massachusetts for over twenty years (1970-1994), continuing on after his retirement. Rolla had a great aversion to leaving anything unfinished. Fol- lowing his retirement from Harvard University, he and Alice moved to Florida where they became adjunct faculty members at the University of South Florida in Tampa in order to continue their work on ferns. In particular, Rolla wanted to complete the Ferns of Peru, a project begun in the early 1960s. Collaborating with Robert Stolze and others, Rolla saw this huge project to completion between 1989 and 1994, publishing a series of six major papers totaling 842 pages. For many of us, Rolla was and continues to be a role model for balancing careers and lives. We miss his intense academic rigor and his love of picking berries. We miss the attention he gave to training students: the piles of unknown ferns from tropical America we were required to identify, the weekly literature sur- veys and reports we were required to make, and his love of poker. We miss the days when we could wander into Rolla’s office for help with taxonomic or nomenclatorial problems, or to Alice’s office for help with cytology and scanning electron microscopy, and finish off the day at their house for one of Alice’s incredible dinners and a taste of rum with Rolla. Rolla Tryon was deeply loved by his students and he will always be remembered for the incredible difference he made in our lives. LITERATURE CITED BARRINGTON, D. S., D. S. CONANT, AND G. J. GASTONY. 2001. Rolla Milton Tryon, Jr. ees Bull. Brit. espe Soc. 5(6): 350-35 GastTony, G. J., D. S. BARRINGTON, AND . CONANT. 2002. Obituary : Rolla Milton . Jr. (1916-2001). Been 4 92: 1-9, —Davip S. CONANT, Department of Natural Sciences, Lyndon State College, Lyndonville, VT 05851, GERALD J. GASTONY, De- partment of Biology, Indiana University, Bloomington, IN 47405- 3700, and DAvip S. BARRINGTON, Department of Botany, Uni- versity of Vermont, Burlington, VT 05404-0086. RHODORA, Vol. 104, No. 917, pp. 96-99, 2002 BOOK REVIEW Lichens of North America by Irwin M. Brodo, Sylvia Duran Shar- noff, and Stephen Sharnoff. 2001. xxii + 795 pp. illus. color photos, line drawings, and dot distribution maps. ISBN 0- 300-08249-5 $69.95 (hardcover). Yale University Press, New Haven, CT. Lichens, lowly, unlovely lichens are the unlikely topic of the volume at hand. Lichens of North America is a production of pure visual splendor. It brings lichens to life in a manner that will not soon be matched. It is the result of the prodigious effort, sublime artistry, and singular devotion of Stephen Sharnoff and the late Sylvia Duran Sharnoff, whose photographs of lichens are the best I have ever seen. Their co-author, Irwin Brodo, wrote the accom- panying descriptions and commentary for the photographs, nearly every one of which is a small masterpiece. Whether it is a close- up shot or a portrait of lichens in their natural habitat, each image invites the reader to appreciate, to touch, even to smell the li- chens. Having attempted on my own over the past two decades to photograph lichens, I am thrilled with and yes, a little jealous of, the success of the images on these pages. An introduction of over one hundred pages precedes the main (taxonomic) part of the text. Each chapter of the introduction starts with an epigram, sending a potent message that implies the importance of lichens in literature and in natural history. While they are the subject of soliloquies by the likes of Browning and Thoreau, lichens have, in fact, been relatively neglected over the past couple of centuries by the scientific community. The intro- duction attempts to right this wrong with succinct discussions of lichen morphology, chemistry, classification, biogeography, and reproduction. However, the attempt is less than successful, at least from a scientific perspective. The greatest problem is that the issues raised in the introduction are not treated in sufficient depth. The chapters, which are generally well written, correspond to an abbreviated list of references (not cited in the text) that are found at the back of the book. The bibliography is much too short. Little in the way of new literature 1s offered, and many of the sources date from the 1970s and °80s, the height of co-author Brodo’s lichenological career. For example, the chapter on the geography of North American lichens offers less than ten references. The 96 2002] Book Review 97 authors defend this unwarranted brevity with the rationalization that the references are general and that “‘sixty-five other publi- cations were consulted.’ The fact that none of these publications was cited for the benefit of interested readers is a hint that Lichens of North America is less than a serious scientific contribution. One of the high points of the introduction is a chapter offering useful hints for observing and collecting lichens. Another short chapter on human uses of lichens uses winsome photographs to illustrate the text, but much of the chapter, like most of the dia- grams in the book, has been served up in previous works. The too-selective bibliography and the rewarming of a number of old lichen illustrations (Some of which were previously redrawn from even earlier sources) provide further hints that this book fails to attain a certain hoped-for standard of scientific relevance. In all fairness, it should be noted that for the libraries of undergraduate students and for an apparently growing audience of amateur li- chenologists, the book will provide a good reference, or at least a starting point. Why do authors still classify the growth forms of lichens as crustose, foliose, and fruticose? Perhaps it fulfills a human need, articulated by Plato, to construct a world of ideals and essences. Unfortunately, the organismic world defies such a construct, and lichens are more frustrating than most organisms when it comes to pigeonholing their morphological characteristics. Yet, co-au- thor Brodo has attempted just this. In view of his wide experience with lichens, one wishes that he would have offered us a more critical perspective on lichen form, but we are disappointed again by a facile account of the gross anatomy of lichens, and by the authors’ insistence on redrawing figures from earlier texts that are perfectly accessible in their own right. Perhaps the intent was to attract future scholars to the lichen world. Perhaps the authors hoped to engage an audience of amateurs who may lack access to a good scientific library, but with the sumptuous photographs found throughout the text, we fear that Brodo and his co-authors have gilded the lily. It would have been far better to let the orig- inal, beautiful photographs in this text tell their own story of lichen form. Sadly, abbreviation must suffice in this book, which turns out to be more a digest of the North American lichens than a scientific treatise. For example, to find the author of a name of a species, which might give a serious student a handle on taxonomic con- 98 Rhodora [Vol. 104 cepts and history, one has to dig through the index. That problem is surmountable with enough ambition and patience. Other omis- sions are more serious. Abbreviation, especially in taxonomic ac- counts, misleads readers by providing an incomplete picture of the extent and background of the species. The problem is nowhere more palpable than in the taxonomic section of this book, where the authors have simply excluded hundreds of species from the dichotomous keys and their attendant descriptions. [ can only speak to the veracity of treatment given the lichen family Cla- doniaceae, which has been my focus of study for almost twenty years. As elsewhere in the text, the photographs of Cladonia li- chens and their allies range from excellent to breathtaking. How- ever, by excluding over a dozen new species in his account, Bro- do has taken us back taxonomically to 1978, when C. verruculosa was recognized as a distinct entity in the North American lichen flora. The insensitivity of excluding so many species that have been recognized since the late 1970s is inexcusable, notwithstand- ing the fact that I am the author of many of them! Other authors of Cladonia and other genera were also ignored, but Brodo and his collaborators have provided litthe or no hint as to what else is “out there.”” They have apparently been selective about their distribution maps as well. At least in the Cladoniacae, it seems that the authors have chosen to ignore several recent accounts of the biogeography of the group. It will be up to future generations of readers to find the lacunae in this book. They may, however, be distracted by trying to mem- orize the specious, insulting ““common names’’ that have been appended to taxa described within. The authors have done a real disservice to lichenology by Lposiie their cloying appellations on readers. Need I refer to me names “pompon shadow lichen,” “finger-scale foam lichen,” or “changing earthscale’’ to under- stand the crust I see on a rock? Does anyone’s appreciation of nature benefit from the authors’ misleading anthropomorphisms like “rough eyelash lichen” or “‘split-peg soldiers’’? Ultimately, Lichens of North America 1s a scientific disappointment. I hasten to add that not everyone requires scientific accuracy to get a kick out of nature. This handsome volume will be sure to delight the eye of anyone who opens it. It may indeed inspire further nature study, though its large format would tend to take up too much room on a field trip. At just shy of seventy dollars the book is a real bargain, and I suggest you buy it for someone who likes 2002] Book Review 99 more than just pretty pictures of nature. Larger than a stocking- stuffer and eminently more valuable, Lichens of North America will soon take its place as the foremost introductory text to the lichens of this continent. In spite of its shortcomings, it is sure to find its place on the shelves and in the laboratories of lichen- ologists around the world. —SAMUEL HAMMER, College of General Studies, 871 Common- wealth Avenue, Boston University, Boston, MA 02215. RHODORA, Vol. 104, No. 917, pp. 100, 2002 NEW BOOKS Bioconservation and Systematics: Proceedings of the Canadian Botanical Association Conference Symposium in London, Ontar- io, June 2000 by J. B. Phipps and P. M. Catling, eds. 2001. ii + 101 pp. US$17.00 (paperback). Published by the Canadian Bo- tanical Association. [send checks to Dr. Mel Fisher, Box 160, 407 Main St., Aberdeen, Saskatchewan SOK OAO, Canada| Muenscher’s Keys to Woody Plants: An Expanded Guide to Na- tive and Cultivated Species by E. A. Cope. 2001. xi + 337 pp. ISBN 0-8014-8702-1 $50.00 (hardcover), $22.95 (paperback). Cornell University Press, Ithaca, NY. Seventh Catalog of the Vascular Plants of Ohio by T. S. Coop- errider, A. W. Cusick, and J. T. Kartesz, eds. 2001. x + 195 pp. ISBN 0-8142-5061-0 $29.95 (paperback), 0-8 142-0858-4 $65.00 (hardcover). The Ohio State University Press, Columbus, OH. Vascular Plants of Wyoming, Edition 3 by R. D. Dorn. 2001. iv + 412 pp. illus. $20.00 (paperback). Mountain West Publishing, Cheyenne, WY. [distributed by the Rocky Mountain Herbarium, Department of Botany, University of Wyoming, Laramie, WY 82071-3165; please make checks payable to Rocky Mountain Herbarium | 100 RHODORA, Vol. 104, No. 917, pp. 101-105, 2002 NEBC MEETING NEWS October 2001. President Lisa Standley introduced Dr. Kanchi Gandhi, Gray Herbarium Card Index Bibliographer and Database Manager, and Editor of the International Plant Name Index for Harvard University. Gandhi spoke to us on **The Phytogeography of India.” To familiarize the audience with the subcontinent, Gan- dhi presented a series of slides showing the geographical, geo- logical, and political India. British India at one time included Sri Lanka (Ceylon) and Myanmar (Burma), as well as what is now Pakistan, part of Afghanistan, and Bangladesh. In 1907 J. D. Hooker divided what was then India and Malaysia into nine phy- togeographical provinces. Of these, Sri Lanka, Myanmar, and Ma- laysia represent three provinces, whereas the remaining six prov- inces encompass what is now recognized as India: (1) the Eastern Himalayan Province (including Nepal) receives |!Q00—600 in. rain/ year and the vegetation is lush; (2) the Western Himalayan Prov- ince is relatively drier than its eastern counterpart; (3) the Indus Plain is dry, with desert areas and thorny vegetation; (4) the Gan- getic Plain receives moderate rainfall and is characterized by dry deciduous forest; (5) the Malabar Province (Western Ghats) along the southwestern coast receives 75—200 in. rain/year with rainfall declining markedly as one moves east, and it supports a variety of forest types; and (6) the Deccan Province on the eastern side of the Indian peninsula is drier, and is characterized by a dry deciduous forest. Gandhi then showed slides of plants that occur in most parts of India. These included Ficus religiosa, commonly called the Bo-tree because Buddha was sitting under this tree when he re- ceived enlightenment. It is native in the Himalayas but is planted throughout India, especially in temples. Ficus benghalensis, the banyan tree, is a common shade tree; it keeps producing prop roots and can extend over a large area if undisturbed. Others are widely planted because of their economic or medicinal value. Examples included: Azadirachta indica (neem); Mangifera indica (mango); Tamarindus indica (tamarind); Musa (banana); Arto- carpus (Jack fruit); and Moringa oleifera (called the miracle plant because of the high vitamin and mineral content of its leaves and fruits). Gandhi also mentioned several other common plants in- cluding succulent members of the Euphorbiaceae found in the scrub area of the Deccan phytogeographic province and some 101 102 Rhodora [Vol. 104 common aquatics such as Trapa, Nymphaea, Nelumbo, and Ot- telia. Next, Gandhi described some of the regional diversity in India, focusing first on the Eastern Himalayan region and its botanical affinities with China. Some genera the region has in common with eastern Asia are Reevesia, Dillenia, Adina, and Alnus. In contrast, he described a sort of transect of the vegetation in Hassan, which is representative of the diversity in the state of Karnataka on the Arabian Sea. Southwestern Hassan is characterized by moist de- ciduous forest, rainforest, and semi-evergreen forest typical of the Malabar phytogeographic province. This end of the spectrum re- ceives between 10Q—200 in. rain/year and one can find species of Drosera, Garcinia, Costus, Arisaema, and Strobilanthus as well as Piper nigrum and several species of palms. In northeastern Hassan the rainfall is only 15—25 in. per year and the vegetation is similar to that of the Deccan phytogeographic province: scrub and dry deciduous forest. Some notable plants of this area are Gloriosa superba (a lily with tendril-like leaf tips), Dodonaea viscosa (varnish leaf), Prerocarpus marsupium, Tectona grandis (teak), and Santalum album (the fragrant sandlewood tree). Once we had some idea of the diversity of the Indian flora, Gandhi went back to the theme of phytogeography. He stated that the broad divisions of Hooker were modified in 1939 by Chat- terjee and in 1955 by Razi; the latter identified 21 phytogeograph- ic regions within present-day India. Although India is about one third the size of the United States, it has a relatively diverse angiosperm flora of about 17,000 species compared with 25,000 for the U.S. Hooker commented that India was a “‘meeting place” for plants from surrounding regions and suggested that it had no recognizable indigenous species. Subsequent work has shown this to be an overstatement; although India has no endemic families, about 140 genera and 5100 species (ca. 30% of the flora) are endemic. Three areas of endemism are identified, with most of the endemics occurring in the Himalayas (3500 spp.) and the Malabar province (1500 spp.). These two regions of high ende- mism are separated by the largely sedimentary Gangetic Plain, resulting in a second type of unique distribution: disjunct genera. For example, 75 species of /mpatiens are found only in the Mal- abar Province and 100 in the Himalayas, while none occur in the Gangetic Plain. Another disjunct genus is Rhododendron, with One species in the south and over 100 in the Himalayan region. 2002] NEBC Meeting News 103 Gandhi said there were two hypotheses to explain the disjunct distributions: long distance dispersal and Pleistocene glaciation that once covered southern India. Gandhi concluded his presen- tation by showing slides representing families and genera with disjunct or endemic distributions within India. November 2001. The evening’s speaker was Jennifer Forman, a graduate student in the Ph.D. program in the Biology Depart- ment at the University of Massachusetts—Boston and student rep- resentative to the NEBC Council. She presented a talk entitled “Through the Looking Glass: History and Consequences of the Introduction of American Species into Europe.” Jennifer introduced the topic by pointing out that although there was a high level of concern about invasive plants in the United States, many of which were introduced from Europe, few have explored the fate of American introductions into Europe. Jennifer has conducted an extensive literature review and devel- oped a database of 6000 American (North, Central, and South American) plant introductions into Europe to address that issue. Her talk was focused on how the exchange of plant species be- tween Europe and America affected the floras of each region, and on the history and current status of American plants introduced into Europe. In developing her database, Jennifer grouped introduced plants into four categories. In the first category are benign introductions; this group includes plants that cannot grow on their own in the new area. The second group includes casuals and escapes that are occasionally found outside cultivation, but are not able to main- tain their populations. The third group consists of naturalized plants that are able to establish populations and reproduce in the wild. Finally, there are the invasive or weedy species that are established and spreading. Approximately 26% of the flora of North America consists of naturalized plants, with European introductions having a partic- ularly large impact. Most introductions were intentional and fol- lowed colonization, but plants were also introduced accidentally. Currently, about 7% of the North American flora can be consid- ered invasive. Examples of European plants that are now invasive weeds include Lythrum salicaria, Cytisus scoparius, and Vince- toxicum nigrum. As with European introductions to America, most introductions 104 Rhodora [Vol. 104 of American plants into Europe were deliberate. Trees such as Pinus strobus, Picea sitchensis, and Prunus serotina were intro- duced so they could be used in shipbuilding and for fuel. Other plants were sent to physic gardens where they were valued for their medicinal properties (e.g., Sassafras albidum, Podophyllum peltatum) or because of their horticultural interest (e.g., Chryso- lepis chrysophylla, Cypripedium acaule). Many of the prominent names in North American botany, such as Mark Catesby, John Bartram, André Michaux, and John Tradescant, were responsible for introductions through the seed and other plant material they sent back to Europe. For example, Tradescant and his son intro- duced Robinia pseudoacacia, Rhus typhina, and Liriodendron tu- lipifera to England. As in America, other introductions were ac- cidental and arrived in Europe along with textiles, in ship’s bal- last, or with transported animals. Some of the American species introduced into Europe, including the orchid Bletia purpurea and the cactus Echinocereus triglochidiatus remain in cultivation to this day. Others, such as Tradescantia pallida, are occasional es- capes. A few, including Pinus radiata, Lysimachia terrestris, and Mimulus guttatus, have become naturalized. Some of the natu- ralized plants, such as Rhus typhina, Rudbeckia hirta, and Phy- tolacca americana are weedy in the United States. Of the ap- proximately 6000 introductions to Europe from America in her database, about 8% have become either naturalized or weedy in Europe. Jennifer pointed out that there have been a number of expla- nations as to why so many European plant species are invasive in America, but not vice versa. One suggestion is that the Old World species are better weeds in that they grow faster and pro- duce more seeds. A second explanation is related to the fact that immigration rates were much greater from the Old World to the New. It may also be that ecosystem damage due to deforestation and post-colonization grazing facilitated the establishment of in- troduced species. Using contingency tests, Jennifer was able to test several ideas about the species introduced to Europe from America. She was able to show species from some families (e.g., Poaceae and Amaranthaceae) were more likely than those from other families to become weedy. In addition, the latitude of the origin of the species affected the probability that a species would become nat- uralized in Europe. For example, more species from North Amer- 2002 | NEBC Meeting News 105 ica are naturalized in Europe than those introduced from Central or South America. She also showed a very clear positive rela- tionship between the number of methods of introduction and the likelihood that a particular species would become established. Fi- nally, she pointed out that the weediness of a species in America was a good predictor of whether a species would become estab- lished in Europe. She concluded by suggesting that a warning list be made available for the 222 weedy American species intro- duced into Europe that are not yet invasive there. —KAREN SEARCY, Recording Secretary pro tempore. ANNOUNCEMENT INVASIVE PLANT SURVEY OF NEW ENGLAND A CALL FOR VOLUNTEERS The New England Wild Flower Society, Silvio O. Conte Na- tional Fish and Wildlife Refuge, and the University of Connect- icut have recently been awarded a grant from the United States Department of Agriculture to track the distribution and spread of over 100 invasive plant species throughout New England. A corps of volunteers will be trained to identify invasive plants and doc- ument their current range. In 2002 we are seeking to train 25 volunteer participants in each New England state to survey their local area. An additional 50 volunteers in each state will be re- cruited and trained in 2003 and 2004. Trainings will occur in the spring and summer at a series of workshops held in each New England state. Trainings will take two days and include an indoor classroom informational session using slides, herbarium sheets, and other prepared materials, and an outdoor session consisting of field visits to local sites where infestations of invasive species occur. Information collected by volunteers will be entered into the Invasive Plant Atlas of New England (IPANE) at the University of Connecticut. The data in IPANE will be posted on the Internet and used for early detection of problem species, research, and decision making on how to control invasive species to slow their spread and reduce their impact on our native flora. More infor- mation on this project and the survey can be found on the New England Wild Flower Society web site |www.newfs.org] or the web site for the Invasive Plant Atlas of New England lu/invasives/ipane]. People interested in volunteering for the Invasive Plant Survey should contact Bryan Connolly, Invasive Plant Survey Coordi- nator [mailing address: 76 Warrenville Rd., Mansfield Center, Connecticut 06250; phone 860-423-8305 or 508-877-7630 ext. 3209; e-mail bconnolly@newfs.org or connollybryan@hotmail. com]. |www.eeb.uconn 106 CHECKLIST FOR CONTRIBUTORS TO RHODORA Please check items and submit with manuscript. General Instructions Type manuscript on one side only of 8% inch X 11 inch paper. Leave a l-inch margin on all sides. Use a standard 12-pitch font type throughout the manuscript, includ- ing tables and appen Do not justify the right margin. Avoid hyphens or dashes at the right margin. T ices. he manuscript should be fully double-spaced throughout, includ- ing title, authors’ names and ad- dresses, Literature Cited, appendi- ces, tables, and figure Each page of the eres ex- cluding page | uding Lit- erature Cited, ee pene ae and figure legends; should be num- bered in the upper right-hand corner. Correct accents, umlauts, and other diacritical marks should be includ- ed. Where appropriate, multiplica- tion symbol must be used rather than the letter x Only names at the rank of genus and below are italicized or under- lined. If underlining its used, do not underline spaces or paraeee Special typefaces (ita bold) should not be used as where indicated in this checklist Do not italicize common words, abbreviations, G5 et als ee. S10.). Manuscript shoiild be checked for consistency, especially in matters of abbreviation, names of sites or legends. Latin or phrases vegetation. types, spelling = of names, etc. The Chicago Manual of Style, most recent edition, is used as a refer- ence in most matters of style. Refer to recent issues of Rhodora. Assemble the the following order: (1) Introductory material, (2) Text, (3) Acknowl- edgments, (4) Literature Cited, (5) manuscript in 107 ¢ Title should be centered, Appendices, (6) Tables, legends, (8) Figures. sas 7) Figure Introductory Material Running head should be centered, at top of page, in upper and lower case letters. Include author’s sur- name (if two authors use the word “and”; use “‘et al.”” for more that two authors), long dash, and aon title. Total characters, including spaces, must not exceed 4 in uppe and lower case. Only the first word of the title and es nouns should be capitalized. not include au- thors of ee names. Include family name in parentheses unless genus studied is type for the family. * Author(s) name(s) and_ profes- sional address(es) should appear below title, centered, in upper and lower Consolidate lines where possible. Two-letter Joh abbreviations should be used fo states. “Current address:” aon appear on a separate line immedi- ately following address if author has moved, not as a footnote. If more than one author at an addre designate current address of nee who has moved using a ee number. Include e-mail address(es) on a separate line following postal address(es). The first author will be considered the corresponding au- thor unless indicated otherwise by a Rai number. The **Author nce’ statement fol- oe line below the stract must be one Se eee. The ee shoul é ise gs. Do not ae ee or tax- onomic authors, or bbreviations in the abstract. word ‘Abstract’? should be indent- ed, in all capital letters, followed 108 by a period, and should appear on the first line of the abstract Key Words are used in indexing and should be chosen with oe pur- ose in mind. The title should appear at the left followed by a colon. Only Words” margin, proper nouns should be capitalized. Text ¢ The following are examples of ¢ Second-level first-level headings, which should appear centered and in all capital letters: MATERIALS AND METHODS, RESULTS, DISCUS- SION, TAXONOMIC TREAT- MENT. The introduction is not ti- tled in Rhodora. Do not combine results and menage age first consulting with t ~ Do not irate secuion . conclu- summary; these must be incorporated into the discussion, headings should be indented, bold, upper and case, followed by a period, and haute sed on the same line as the subsequent text. The text be written such that addi- levels of headings are not use a sepal sions or lower should tional used. Fach figure and table must be cited in the text in numerical order. The ete Figure’? must be spelled . When citing both together, the ane should be listed first and a semi-colon used to separate the two (e.g., Table 1; Figure 1). Each reference cited in the text must appear in the Literature Cited section and vice versa. Cross-check spelling of author(s) name(s) and dates of publication. Literature is cited in the text as fol- lov on author: Hill (1982) or (Hill authors: Angelo and Bout- 1996) or (Angelo and oe 1996). i) CHECKLIST FOR CONTRIBUTORS 3. More than two authors: Mathie- son et al. _ O) or (Mathieson et al. 2006 4. Note ao ce is no comma separating author and date 5. When more than one paper is cited at a time, they should be listed ape by first au- thor rather than pee aren: C25, ngelo and ford (1996), Hill (1982), i aie et al. (2000)]. 6. Within parentheses, citations should be separated by a semi- .g., Angelo and Bouf- > Hill 1982). 7. Manuscripts accepted but not yet published: Tryon (in press) submited): G. Craw data); G. ow (pers. ¢ x. Crow, pers. comm.): otherwise listed or cited in manuscript or a nati¢ igual know authority, professional affiliation should also be given. > unless the « References to companies manutac- tud turing products used in fats} should not appear in the Literature Cited. Rather, the company name and location should be given in pa- rentheses within the text [e.g., SYSTAT (SPSS, Chicago, Hh- nois)]. Authors of scientific names should be cited for all taxa at the rank of genus and below either at their first usage in the in a table or appendix (e.g., in a flora or table of voucher specimens). It should be indicated which taxonomic ment, revision, text or treat- flora nomencla- hee standard abb author’s names ate in ttp: //www.t rbgke w.org uk/data/ Authors of ie Brummitt and C. ture follow gras a soa or Names by R. K. E. Powell. CHECKLIST FOR CONTRIBUTORS 109 ¢ Names of eer cited in the text should be in itali * Avoid abbreviations in he text un- less indicating measurement, then use a period unless abbreviating a metric term. Other abbreviations should be defined when first used e.g., Scanning Electron Micros- copy (SEM)]. Herbarium acronyms should follow http://www.nybg.ore/ bsei/ith/ or Index Herbariorum, most recent editi ¢ Numbers one ates nine should be written out in the text unless a measurement or part of a taxonom- Ic description. No comma its usec 4-digit numbers. A number should pee precede a decimal oint (e.g., 0.15). . oe parentheses should be avoided by using a semi-colon. Parentheses within parentheses should be avoided by using outer brackets Taxonomic Treatments Use boldface Roman type for new names and new combinations, fol- lowed by “sp. nov.”, “comb. nov. * For nomenclatural history (i.e synonymy and typification) use ne paragraph per basionym [e. Binomial author, literature citation. E: collection aeagnes from least-to-most-specific, collector(s) collection number ‘(Helo ytype: her- um acronym; Isotypes: herbar- ium pene 2 | Exclamation points are used for type specimens examined, and types not seen are indicated as such (e.g., GH!, MO not see Lectotype pee nae are ed together with an indication of where they were designated, what year, and by whom. This reference is listed in the Literature Cited. the author of the paper is making includ- a. eee aerate e designated” 1 ae cited ay as part of no- menclatural history are not includ- ed in the Literature Cited. Books listed here are abbreviated accord- ing to Taxonomic Literature, edi- tion 2, but with initial letters capi- taliz Standard abbreviations for author’s names should be used according to http://www.rbgkew.org.uk/data/au- the phrase ee or Authors of Plant Names by R. K. Brummitt and C. Powell. When dates are given as part of collection information, 3-letter ab- breviations with no period are used for months Use http://www.nybg.org/bsei/th/ or Index Herbariorum, most recent edition, for herbarium acronyms. Designation of a new taxon should include a brief Latin diagnosis, rather than a full Latin description, which sets forth succinctly how the new taxon differs from its conge- ners. A full description, in’ English, should follow. This should be par- allel with other descriptions at the same rank in the paper, and should not repeat information given in any description of the inclusive taxon (i.e., species descriptions should not repeat information characteris- tic of the genus, if also described in the paper). All measurements are metric. Hyphens are used for parenthetical extremes. A multipli- cation symbol is used where appro- priate, rather than the letter Following the description, infor- mation should be given on distri- bution, ecology, uses, and nomen- clature and typification, where ap- propriate. The discussions should be parallel within a given rank. For newly described taxa, this discus- 110 CHECKLIST FOR CONTRIBUTORS sion should explain clearly how the new taxon differs in these charac- teristics from closely related taxa. A high-quality line drawing or ace of the type specimen, illustrating the diagnostic features, should be included for new taxa. Specimen citation should be select- ed critically, especially for com- mon species of broad distribution. A title such as “Specimens exam- ined” or “Representative speci- mens examined” should be indent- ed, in upper and lower case, fol- period. Each country begins a new paragraph. The for- mat of information is as follows COUNTRY. Major political division such as state: smaller political divi- sion such as county, detailed loca- tion, date (e.g., 26 Sep 1950), collec- tor(s) last name(s) collection number or s.m. (herbarium acronym). Keys * Keys are dichotomous and indent- ed. ¢ Leads of each couplet are parallel. Information in the key is consistent with that in descriptions, text, ta- bles, and figures. Data and Voucher Specimens Voucher specimens must be cited in a table or appendix to document sources of morphological or molec- ular data. Format for citation is the same as that for “specimens ex- amined” as part of taxonomic treatments. All sequences used as data must be deposited in one of the internation- al nucleotide sequence databases, and sequence database accession numbers included in the paper (GenBank: gsdb@ gsdb.ncgr.org). All data matrices used in cladistic analyses should be deposited in TreeBASE — (http://www.herbaria. harvard.edu/treebase). Floras * Long lists of taxa are best treated as an Appendix, so that the reada- bility of the text 1s not compro- mised, and so that the list may be used independently by readers. A short introductory paragraph ex- plaining terms or abbreviations used in the list of taxa should follow the Appendix title (see Hickler 1999, Rhodora 101: * Three levels of headings are sible in lists of taxa: centered, all capitals, ANGIOSPERMAE or LIOPSIDA); second-level is cen- tered, all cap ies not bold ee MONOCOTYLEDONEAE or LIL- HDAE); third me is flush a me capitals, bold (e.g., ACORACEAE; this level will be converted to small caps by the printer). * Taxa should be listed alphabetically within each hierarchical category (e.g., species alphabetically within the genus; genera alphabetically within fami Standard abbreviations for authors of binomials should be cording to http://www.rbgkew.org. uk/data/authors.html or Authors of Plant Names by R. K. Brummitt and C. FE. Powell. An indication of ecological prefer- ence, distribution within the area studied, and abundance — be used ac- included, where appro Voucher specimens one be list- ed (collector, collection number, and herbarium acronym; informa- tion common to all or most vouch- ers can be stated in an introductory paragraph). Acknowledgments Acknowledgements should be brief. Information on granting agencies, herbaria from which loans were obtained, artists, and colleagues or advisors who have critically re- CHECKLIST FOR CONTRIBUTORS viewed the manuscript should be included 5 The word ‘Acknowledgments’ should be aoe in all capital letters, followed by a period, and should appear on the first line of the acknowledgments. Literature Cited The Literature Cited contains all references cited in the text and vice versa. The alternative of a general “‘Ref- erences” section requires prior ap- proval by the Editor. All entries should be cross-checked i the text, checking especially or spelling of eee names and — of publica All entries ene be verified ere original sources, checking especially for spelling of authors names and words in languages oth- er than English, exact title, year of publication, and volume and page numbers * Cite references in eee or- = by first author’s last name. En es by a single author should pre- ae multi-authored works with the same first author, regardless of date. List works by the same author chronologically, beginning with earliest date of publication Use long dash when the author(s) is/are the same as in the citation ‘In press” must have been accepted for publication. The name of the journal or book publisher must be inclu Citations of work in progress (1.e., unpublished or not yet accepted for publication) ) should not be listed in the Literature Cited. See format for citation under en riod and a space must be in- serted after each initial of an au- thor’s name. 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Donald Hudson, Jr. Curator of Vascular Plants: Raymond Angelo Assistant Curator of Vascular Plants: Erika Sonder Curator of Nonvascular Plants: Anna M. Reid Librarian: Leshe J. Mehrhoff Councillors: David S. Conant (Past President) David Lovejoy 2002 Arthur V. Gilman 2003 Karen B. Searcy 2004 Jennifer Forman (Graduate Student Member) 2002 Appointed Councillors: David E. Boufford, Associate Curator Janet R. Sullivan, Editor-in-Chief, Rhodora 7 ae Journal of the New England Botanical Club CONTENTS Phylogenetic relationships and ie of Stewartia (Camellioideae, Theaceae) inferred from nuclear ribosomal DNA ITS sequences. Jianhua Li, Peter Del Theaied cee Yang, and Michael J. Donoghue 117 Thomas Walter’s oaks from the coastal region of South Carolina. Robert L. Wilbur 134 Reconstructing the biological invasion of European water-horehound, Lycopus europaeus (Labiatae), along the St. Lawrence River, Québec. 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RHODORA, Vol. 104, No. 918, pp. 117-133, 2002 PHYLOGENETIC RELATIONSHIPS AND BIOGEOGRAPHY OF STEWARTIA (CAMELLIOIDEAE, THEACEAE) INFERRED FROM NUCLEAR RIBOSOMAL DNA ITS SEQUENCES JIANHUA LI' AND PETER DEL TREDICI Arnold Arboretum of Harvard University, 125 Arborway, Jamaica Plain, MA 02130 ‘e-mail: jli@oeb.harvard.edu SHIXIONG YANG Institute of Botany, The Chinese Academy of Sciences, Kunming 650204, Yunnan, People’s Republic of China MICHAEL J. DONOGHUE Department of Ecology and Evolutionary Biology, Yale University, New Haven, CT 06520 ABSTRACT. Sequences of the internal transcribed spacers of nuclear ribo- somal DNA were used to estimate phylogenetic relationships within Stewar- tia. Eighteen samples were included representing two species of Hartia, seven species of Stewartia, and Franklinia alatamaha. Hartia nig and Hf. vil- losa form a clade that is the sister group of Stewartia. Within Stewartia the New World and the Old World species form well-supported clades. The sub- genera and sections of Stewartia proposed by previous authors are not sup- ported by our ITS data. Two clades are recognized within the Old World lineage: S. serrata + S. rostrata and S. pseudocamellia + S. monadelpha + S. sinensis. Southeastern Asia and China may be a recent center of diversi- fication of Stewartia based on the ITS phylogeny and fossil record. Key Words: biogeography. Hartia, nrDNA ITS, phylogeny, Srewartia, Theaceae Stewartia L. comprises 8-21 species (Chang 1998; Li 1996: Spongberg 1974; Yang 1997). Both S. ovata (Cav.) Weatherby and §. malacodendron L. are native to the eastern United States (Figure la; Dove 1981: Kobuski 1951; Wood 1959). Three spe- cies are distributed in southern-central Japan, including S. mon- adelpha Siebold & Zucc., S. serrata Maxim., and S. pseudoca- mellia Maxim., which is also found in eastern Korea (Hara 1958: Lee 1997), while the rest of the species are distributed in central to southeastern China (Figure |b). In China the number of species 117 118 Rhodora [Vol. 104 -95 () 80 ear IO The modern distributi Stewartia and Hartia species a i. Elias 1980: Hara 1958; ee 1993; Lee 1997; Li 1996; Spong- berg 1974). of Stewartia recognized varies from 3 to 16 (Chang 1998; Chang and Ye 1982; Chien and Cheng 1931; Chiu and Zhong 1988; Li 1996; Spongberg 1974; Yan 1981). Stewartia rostrata Spongberg is distributed in Hunan, Jiangxi, and Zhejiang, while S. rubigi- nosa H. T. Chang its endemic to southern Hunan and northern Guangdong. Stewartia sinensis Rehder & E. H. Wilson is wide- spread in central and southern provinces, and its vegetative and floral morphologies are highly variable. Many variants of S. si- nensis have been described either as species or varieties (Chang 1998; Chang and Ye 1982; Chien and Cheng 1931; Chiu and Zhong 1988; Li 1996; Yan 1981). Within Theaceae Srewartia ts generally placed in the taxonom- ically controversial subfamily Camellioideae. Airy-Shaw (1936), based on morphological and anatomical evidence, revised Mel- 2002] Li et al.—Svrewartia Phylogenetics 119 chior’s (1925) classification system of the Camellioideae, recog- nizing two tribes, each with two subtribes. The Gordonieae, to which Stewartia belongs, consists of subtribe Gordoniinae (Gor- donia Ellis, Franklinia W. Bartram ex H. Marshall, and Schima Reinwardt ex Blume) and subtribe Stewartinae (Sfewartia, in- cluding Hartia Dunn). Ye (1990) proposed a 5-tribe system for Camellioideae, but also recognized the tribe Stewartieae, consist- ing of two separate genera, Stewartia and Hartia. While some authors have supported the inclusion of Hartia in Stewartia (Airy-Shaw 1936; Li 1996; Spongberg 1974), others have treated them as separate genera (Chang 1998; Chun 1934; Merrill 1938; Wu 1940; Yan 1981: Ye 1982, 1990). In a recent molecular study of the Camellioideae based on chloroplast DNA sequence data, Hartia was found not to be monophyletic (Prince and Parks 1997). The classification of species within Sfewartia has also been controversial (Table 1). Gray (1849) recognized two subgenera, the first of which, Stuartia (= Stewartia), included two species (S. malacodendron and S. monadelpha). The second subgenus, Malacodendron, consisted of a single species, S. pentagyna L Her. (= S. ovata). Subgenus Stfewartia is characterized by united styles (vs. free styles in subgenus Malacodendron), subglobose capsules (vs. conical capsules), and unwinged seeds (vs. winged seeds). Szyszylowicz (1893) supported Gray’s (1849) groupings but treated them as sections and applied different names (Synstyla instead of Stewartia, Dialystyla instead of Malacodendron). Na- kai (1950) divided Korean and Japanese Stewartia into two sec- tions based on the relative length of sepals and bracts. Section Pseudocamelliae has bracts much shorter than the sepals, whereas section Serratae possesses bracts subequal to, or longer than, the sepals. Spongberg (1974) did not recognize any of these divi- sions. In the most recent treatment of Sfewartia and Hartia, Li (1996) recognized Stewartia s.l., including Hartia and Stewartia, and placed the species of Stewartia s.s. into two subgenera and 4 of the 5 sections of Stewartia s.l. recognized previously. The objectives of this study were |) to estimate interspecific relationships of Stewartia based on DNA sequence data, 2) to test the monophyly of the subgenera and sections that have been pro- posed by previous authors, and 3) to provide possible explana- tions for modern geographic distribution of Stewartia. We chose to use sequence data of the internal transcribed spacers (ITS) of Table |. Previous taxonic treatments of Stewartia species sampled in this study and their groupings in the ITS trees. Szyszylowicz Species Gray (1849) (1893) Nakai (1950) Ye (1982) Li (1996) This Study S. ovata Subg. Malacod-— Sect. Dialystyla N/A Sect. Dialystyla — Subg. Dialystyla New World clade endron S. malacod- dron Subg. Srewartia = Sect. Svastyla N/A Sect. Stewartia = Subg. Stewartia New World clade Sect. Stewartia S. monadel- pha Subg. Stewartia = Sect. Syvnstyla Sect. Serratae Sect. Foliobrac- Subg. Stewartia Sinensis clade tede Sect. Racemosae S. serrata N/A N/A Sect. Serratae Sect. Foliobrac- Subg. Stewartia Serrata clade fede Sect. Serratae S. sinensis N/A N/A N/A Sect. Foliobrac- Subg. Stewartia Sinensis clade teae Sect. Serratae S. rostrata N/A N/A N/A Sect. Foliobrac- Subg. Stewartia Serrata clade teae Sect. Serratae S. pseudoca- mellia N/A N/A Sect. Pseudoca- Sect. Stewartia = Subg. Stewartia Pseudocamellia melliae lia Sect. Pseudocamel- clade ae OCI vAOpoyy OAI PO! 2002] Li et al.— Stewartia Phylogenetics 12] nuclear ribosomal DNA. This is because many studies have shown that sequences of this DNA region are informative in re- solving phylogenetic relationships of plants among genera and species (Baldwin et al. 1995; Li et al. 1999; Li, Boufford, and Donoghue 2001; Li, Davis, Donoghue, Kelley, and Del Tredici Ol). MATERIALS AND METHODS Plant material. Eighteen plants were sampled in this study, representing seven species of Stewartia, two species of Hartia, and the monotypic Franklinia (Table 2). These samples represent all previously recognized subgenera and sections (Gray 1849; Li 1996; Nakai 1950; Szyszylowicz 1893; Ye 1982). Molecular techniques. DNAs were extracted from silica-gel dried leaves using either a standard CTAB DNA extraction meth- od (Doyle and Doyle 1987) or DNeasy Plant Kit (Qiagen Inc., Santa Clarita, CA) following the manufacturer’s protocol with minor modifications. Procedures and protocols for the polymerase chain reactions (PCR), purification of PCR products, and DNA sequencing are described in detail elsewhere (Li and Donoghue 1999). To ex- amine within-individual variation we cloned the ITS regions for Franklinia alatamaha W. Bartram ex H. Marshall, Stewartia ova- ta, S. pseudocamellia, and S. sinensis using standard T-A tail cloning techniques according to manufacturers’ instructions. The pGEM®-T Easy Vector System (cat.# A1360, Promega, Madison, WI) was used to ligate ITS PCR products into pGEM plasmids, which were then transformed into Epicurian Coli® XL1-Blue strain competent cells (cat.4# 200249, Stratagene, La Jolla, CA). Three white colonies for each species were picked and cultured for 17 hours at 37°C, and their plasmids were prepared using a Miniprep Kit (Qiagen, Santa Clarita, CA). A small amount of the prepared plasmid (1 wL) was then digested using GibcoBRL EcoRI restriction enzyme (Life Technologies, Rockville, MD) to check the presence of the ITS inserts. Phylogenetic analysis. Sequences were edited using Se- quencher 3.0 (Gene Codes Corp., Inc., Ann Arbor, MI) to verify Table 2. Species used in this study. Acronyms are as follows: Arnold Arboretum (AA), Jamaica Plain, MA; National Arboretum ), Northhampton, MA; Quarryhill A), Washington DC; Kunming Institute of Botany (KUN), Kunming, China; Smith College (SC Arboretum (QA), CA; University of British Columbia (UBC), Vancouver, Canada. Species Source and Origin GenBank # Srewartia sinensis Rehder & Wilson AA 373-76A: Lushan, Jiangxi, China S. sinensis AA 431-34B; Lushan, Jaingxi, China S. sinensis AA 691-94, Wudangshan, Hubei, China S. pseudocamellia Maxim. QA 89.071; Japan S. pseudocamellia AA 11440A: Korea S. monadelpha Siebold & Zucc. AA 653-74A; Japan S. monadelpha NA 40211; Yakushima, Japan S. rostrata Spongberg AA 769-36A; Lushan, Jiangxi, China S. rostrata Yang 991005; Lushan, Jiangxi, China S. rostrata AA 761-69A: Lushan, Jiangxi, China S. serrata Maxim. UBC Bot. Gard. S. malacodendron L. NA 63252; een i. S. malacodendron SC 07190; Cape Cod, S. ovata (Cav.) Weatherby AA 18847A, eer om S. ovata (Cav.) Weatherby f. grandiflora (Bean) Kobuski AA 18244C, a NC Hartia sinensis Dunn Yang 98913: H. villosa (Merr.) Merr. var. serrata Hu Yang 98924: on Guangxi, China Franklinia alatamaha Bartram ex Marshall AA 2428-2A; Alatamaha, GA AF43 1932 AF431933 AF431936 AF431937 AF339863 AF431934 AF43 1938 AF431935 AF431939 AF43 1941 AF43 1940 AF431943 AF43 1944 APF43 1942 AF339861 AF431946 AF43 1945 AF43 1947 N i eIOpoyYy JOA] bOI 2002] Li et al.—Stewartia Phylogenetics 123 base callings from overlapping sequences and chromatograms generated using different primers. Edited sequences were im- ported into the computer program PAUP* (version 4.063; Swof- ford 2000) and aligned manually. Characters were weighted equally and their states were unordered. Maximum parsimony (MP) analyses were conducted using both gaps scored as missing data and as a fifth character state. Heuristic tree search options included simple sequence addition, TBR branch swapping, Mul- pars on, and steepest decent off. Bootstrap analyses for 300 rep- licates were conducted to evaluate relative support for individual clades (Felsenstein 1985). All of these analyses were conducted using PAUP*. Franklinia alatamaha was included for rooting purposes since several analyses have shown it to be closely re- lated to the clade containing Stewartia and Hartia (Prince and Parks 1997; Tsou 1998; Ye 1990). Maximum likelihood ratio test. To test whether ITS se- quences in Stewartia and Hartia evolved in a clockwise fashion, we conducted maximum likelihood (ML) ratio tests using the HKY85+G model, implemented in PAUP* following Baum et al. (1998). ML analyses included the following options: as-is se- quence addition, TBR (tree-bisection-reconnection) branch-swap- ping, and steepest descent option off. RESULTS Sequence characteristics. Sequences of the entire ITS re- gion of all samples ranged from 646-657 base pairs (bp) in length, excluding Franklinia alatamaha, whose ITS region was 626 bp long. In Sfewartia the lengths of the ITS-1 and ITS-2 were 246-267 bp and 221—229 bp, respectively. In Franklinia the lengths of the ITS-1 and ITS-2 were 242 and 223 bp, respectively. In all samples the sequences of the 5.8S gene were 161 bp in length. The alignment of all sequences produced a data matrix of 678 characters, 65 of which were parsimony informative. Sequence divergence of the ITS-1 ranged from 0—7.3% (mean, or * = 4.4%) among species of Stewartia, from 4.9-8.4% (* = 6.7%) between species of Stewartia and Hartia, from 14.6-21.8% (€ = 16.5%) between species of Sfewartia and Franklinia, and from 17.2— 17.7% (* = 17.5%) between species of Hartia and Franklinia. 124 Rhodora [Vol. 104 Sequences of the ITS-2 diverged from 0.0-8.2% (* = 4.3%) among Sfewartia species, from 2.7—8.2% (* = 5.1%) between species of Stewartia and Hartia, from 16.5-19% (x = 17.3%) between Stewartia species and Franklinia, and from 14.8—15.7% (*¥ = 15.3%) between Hartia species and Franklinia. All sequenc- es have been submitted to GenBank (Table 2), and the data matrix and trees are available from the first author upon request and in TreeBASE (http://www.herbaria.harvard.edu/treebase). Phylogenetic relationships. Parsimony analyses of the ITS data set generated 3 trees of 175 steps when gaps were treated as missing data. The strict consensus (MP-M, maximum parsi- mony-missing) tree is shown in Figure 2 (solid branches, CI = 0.83, RI = 0.83). Species of Hartia form a strongly supported clade (bootstrap, or b = 99%), which is sister to the clade con- taining all species of Stewartia (b = 77%). Within the Stewartia clade, the two North American species, S$. ovata and S. malacod- endron, form a clade (b = 83%), which is sister to the clade containing all of the eastern Asian species (b = 74%). Stewartia serrata and S. rostrata form a well-supported clade (b = 85%), which is sister to the clade containing S. pseudocamellia, S. mon- dadelpha, and S. sinensis (b = %). Accessions of S. pseudo- camellia from Japan and Korea form a clade (b = 100%), which is sister to a clade consisting of S. monadelpha and S. sinensis (b = 96%). When gaps were treated as the fifth character state, the MP analyses produced a single (MP-F, maximum parsimony- fifth) tree of 254 steps (Figure 2, dashed branches; CI = 0.85, RI = 0.82). The MP-F tree is identical to the MP-M tree except that the three accessions of S. sinensis formed a moderately supported clade (b = 70%). The maximum likelihood ratio test indicated that rates of ITS base substitution in the Stewartia and Hartia clade are signifi- cantly heterogeneous (P < 0.05). Thus, we did not attempt to estimate times of divergence for different lineages of Sfewartia. DISCUSSION Monophyly of Stewartia. Hartia was proposed by Dunn (1902) to accommodate a plant collected from Yunnan province (Spongberg 1974). However, it has been debated whether Hartia should simply be included in Stewartia. Some authors have main- 2002] Li et al.—Stewartia Phylogenetics 125 5. Sinensis37376A (C) sinensis43134B (C) . Sinensis691-94 (C) SIN . monadelpha65374A (J) (J) fata) | PSE (J) S. rostrata769-36A (Cc) rostrata991005 (C) SER rtia761-69Ac2 (C) S$. serrataUBC (J) ovata var. (U) . ovatal8847A (U) OVA . malacodendron (U) NA63252 . ma odendron (U) Smi ees sinensis98913 (C) HAR . villosa var. serrata98924 . alatamaha (U) Figure 2. Phylogenetic trees based on maximum ee analyses of sequences of nrDNA ITS: strict consensus of 3 trees of 175 steps treating gaps as missing data (MP-M, not dashed), and the single tree - 254 steps treating gaps as the Sth character state (MP-F, dashed). Numbers above and below the branches indicate ocusete y percentages. Clade denotation: SIN, sinensis; PSE, pseudocamellia; SER, serrata; OVA, ovata; and HAR, hartia. Letters in Saronttieses represent geographic distributions: C for China, J for Japan, K for Korea, and U for the United States. 126 Rhodora [Vol. 104 tained Hartia as a separate genus (Chang 1998; Chun 1934; Mer- rill 1938; Wu 1940; Ye 1982), while others support the inclusion of Hartia in Stewartia (Cheng 1934; Keng 1962; Li 1996; Sealy 1958; Spongberg 1974). In their phylogenetic study of the Theo- ideae based on sequences of the chloroplast gene rbcL, Prince and Parks (1997) concluded that Stewartia might be paraphyletic with Hartia nested within it. However, only three species of Ste- wartia and one species of Hartia were included in that analysis. In our trees (Figure 2), two species of Hartia form a well-sup- ported clade sister to the clade containing species of Stewartia. Hartia and Stewartia have distinct differences in 15 non-molec- ular characters from morphology, palynology, and wood anatomy (Ye 1982). In addition, Hartia and Stewartia also differ in chro- mosome numbers: n = 15 in Stewartia (Santamour 1963) and n = 18 in Hartia (Oginuma et al. 1994). Therefore, our results, together with non-molecular data, suggest that both Hartia and Stewartia are monophyletic genera. Nevertheless, more species of Hartia need to be included in the future to further test this hy- pothesis. Phylogenetic relationships within Stewartia. Although re- lationships within Sfewartia have not been explicitly analyzed prior to this study, previous taxonomic treatments are considered as working hypotheses to be tested. Based on fruit, style, and seed characters, §. ovata has been separated from the rest of the species as either a monotypic subgenus, Malacodendron, or as the section, Dialystyla (Gray 1849; Li 1996; Szyszylowicz 1893). This treatment implies that S. malacodendron, which is the other North American species and has been placed in the Old World group, is more closely related to the Old World species than it 1s to S. ovata. In our ITS trees (Figure 2) S. ovata is linked directly with §. malacodendron. That is, the two North American species form a clade that is the sister group to all of the Old World ~ species. Nakai (1950) recognized two sections mainly based on the rel- ative length of bracts and sepals. Stewartia pseudocamellia differs from all the other Asian taxa in having shorter bracts. On this basis it was treated as a monotypic section Pseudocamelliae, and the rest of the species were assigned to section Serratae. In our ITS trees (Figure 2), S. pseudocamellia is not a sister species to a clade containing the remaining Stewartia species. In contrast, 2002] Li et al.—Stewartia Phylogenetics 127 it forms a strongly supported clade with S. monadelpha and S. sinensis. In his review of Hartia and Stewartia, Ye (1982) recognized three sections within Stewartia. The first section, Stewartia, char- acterized by non-foliaceous bracts and orbicular to obovate se- pals, included S. rubiginosa, S. pseudocamellia, and S. malacod- endron. Although S. rubiginosa was not available for this study, the distant relationship between S$. malacodendron and S. pseu- docamellia (Figure 2) indicates that section Stewartia sensu Ye (1982) is not supported by the ITS sequences. The second section, Foliobracteae, comprising §. monadelpha, S. sinensis, S. rostrata, and S. serrata, was marked by foliaceous bracts and fused styles. In our ITS trees, species of section Foliobracteae form a mono- phyletic group with S. pseudocamellia, which was placed by Ye (1982) in section Stewartia. Thus, ITS sequences indicate that section Foliobractedae sensu Ye (1982) is not monophyletic. Ye’s third section, Dialystyla, was unique in having distinct styles and consisted of three species, S. ovata, S. yunnanensis H. T. Chang, and S. oblongifolia Hu ex S. Z. Yan; the latter two species were transferred by Yang (1997) to the distantly related Pyrenaria Blume. Li (1996) included Hartia within Stewartia and divided Ste- wartia s.1. into two subgenera based on whether the styles are fused (subgenus Sfewartia) or distinct (subgenus Dialystyla). In subgenus Stewartia, he recognized five sections. His first section, Racemosae, consisted of S. monadelpha and six Hartia species. In our phylogenetic trees, however, S. monadelpha is not directly related to Hartia. Li’s second section, Stewartia, included only one species, S. malacodendron. His third section, Serratae, con- tained S. sinensis, S. serrata, and S. rostrata. In our ITS trees (Figure 2), these three species form a clade that also contains S. pseudocamellia of section Pseudocamelliae (see below) and S. monadelpha of section Racemosae. The fourth section, Pseudo- camelliae, consisted of S. pseudocamellia, three Hartia species, S. rubiginosa, and S. damingshanica J. Li & T. Ming. The latter five species were not available for this study, so we are unable to assess the monophyly of this section. Li’s fifth section, Prer- opetiolatae, consisted of four Hartia species. When describing the segregate species, Stewartia rostrata, Spongberg (1974) hypothesized that it was most closely related to S. serrata. Probable synapomorphies of these two lineages in- 128 Rhodora [Vol. 104 clude glabrous ovaries and 2—3 winter bud scales. Recently, Chang (1998) treated S. rostrata as a variety of S. sinensis. In our ITS trees, S. rostrata accessions form a clade with S. serrata with strong support (b = 85%). In contrast, S. sinensis is distantly related to S. rostrata, being most closely related to S. monadel- pha. A close relationship between S. sinensis and S. monadelpha also supports Spongberg (1974), who stated that these two species were so closely related that S. monadelpha could be considered as a subspecies of S. sinensis. In summary, our results indicate that none of the subgenera and sections of Stewartia proposed by previous authors (Gray 1849; Li 1996; Szyszylowicz 1893; Ye 1982) are monophyletic, except possibly for section Pseudocamelliae sensu Ye (1982), whose monophyly we were unable to assess due to insufficient sampling. Evolution of morphological characters. In Sfewartia all species have fused styles except for S. ovata, which has five dis- tinct styles. This condition and a single bract enclosing axillary buds have been used to justify the separation of S. ovata from the rest of the Stewartia species, including the other North Amer- ican species, S. malacodendron (Gray 1849; Li 1996; Szyszylow- icz 1893). In our ITS trees (Figure 2), the two North American species form a well-supported clade, which is sister to the clade containing all of the Old World species of Sfewartia. All species of Hartia have fused styles. Styles are occasionally found to be only half fused in S. sinensis; this condition also appears to be derived within Stewartia. Therefore, having distinct styles may be a derived condition and therefore an autapomorphy of S. ovata. It is interesting to note that the fruits of Srewartia are capsules that split from the top to the bottom loculicidally, releasing seeds. It is possible that the free styles of S. ovata facilitate the release of winged seeds by avoiding the hindrance from the fused styles during the top-to-bottom splitting of the capsules. Field studies could be conducted to compare the seed dispersal efficiency of S. ovata with its sister species S. malacodendron, which has fused styles and unwinged seeds. The bark of Stewartia species is quite variable. Several species develop smooth, mottled bark on the trunks and limbs, resulting in irregularly arranged, buff- or cinnamon-colored patches. These species include S. malacodendron, S. sinensis, S. serrata, S. pseu- 2002] Li et al.—Srewartia Phylogenetics 129 docamellia, and S. monadelpha. Our ITS phylogenies imply that the mottled bark probably evolved several times independently in Stewartia. Stewartia seeds develop a narrow wing around the perimeter in all species except for the North American S. malacodendron and the Japanese/Korean S. pseudocamellia. In addition, species of both Hartia and Franklinia have winged seeds. Therefore, unwinged seeds appear to have been derived twice within Ste- wartia. Biogeographic implications. Species of Stewartia are dis- tributed disjunctly between eastern Asia and the eastern United States (Figure |). This interesting disjunction has long attracted attentions from both systematists and biogeographers (Boufford and Spongberg 1983; Gray 1849; Hong 1993; Li 1952; Li et al. 2000; Tiffney 1985; Wen 1999). Previous hypotheses concerning interspecific relationships of disjuncts have sometimes proven to be erroneous (Gould and Donoghue 2000; Li, Davis, Donoghue, Kelley, and Del Tredici 2001; Wen 1999; Wen et al. 1998: this study). As more phylogenetic studies are conducted some con- gruent patterns are emerging. For example, Xiang, Soltis, and Soltis (1998) have shown that phylogenetic relationships in seven plant taxa, including ferns, conifers, and angiosperms, point to a single phylogenetic split between Old World and New World spe- cies with western North American species being most closely related to eastern North American species. Our results, together with several other recent phylogenetic investigations (Aesculus, Xiang, Crawford, Wolfe, Tang, and DePamphilis 1998; Pachy- sandra, Cuénoud et al. 2000; Torreya, Li, Davis, Donoghue, Kel- ley, and Del Tredici 2001), are consistent with this pattern. To further understand the formation of this disjunction, the fos- sil record of both Hartia and Stewartia should be consulted. In North America, according to Grote and Dilcher (1989), no fossils have been reliably assigned to Stewartia or Hartia. In the Old World, Mai (1975) described fruits and seeds of H. guinquean- gularis Mai from the Upper Miocene of western Europe. Knob- lock and Mai (1986) reported fossil fruits and seeds assigned to the modern Stewartia from the Upper Cretaceous of Europe. Kir- chheimer (1957) and van der Burgh (1978) have found fruits and seeds of S. beckerana (Ludwig) Kirchheimer from central Euro- pean deposits of the Pliocene and Miocene. An amber-embedded 130 Rhodora [Vol. 104 fossil flower was described from the Oligocene deposit in Ger- many as S. kovalewskii by Raiiffle and Helms (1970); however, this assignment is questionable (Mai 1971). Neogene floras of Japan contain two species of Stewartia based on fruit and leaf remains (Tanai and Suzuki 1972). As summarized by Grote and Dilcher (1989), the European late Tertiary sediments include both Hartia and Stewartia, but reliable fossils of Hartia or Stewartia have not been found in either North America or eastern Asia except for Japan. It is possible that species of Hartia and Ste- wartia were absent from China through most of the Tertiary and migrated there relatively recently (Grote and Dilcher 1989). Our analyses suggest that the radiation of Stewartia in China might have taken place rather recently. Neither the Chinese nor Japanese Stewartia species form their own clades in ITS trees, indicating that there has been continuing population exchange between these two areas throughout the Ter- tiary. We did not estimate the time of divergence of Stewartia lineages since the maximum likelihood ratio tests have shown significant rate heterogeneity of the ITS sequences in Sfewartia and Hartia. In these ITS trees (Figure 2), the first branching is between the Old and New World clades, implying that the time of divergence of the New World Stewartia species from the Old World species is earlier than that of the Japanese and Chinese species. 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Thomas Walter was the first post-Linnaean author of a sizable flora in eastern North America. As such, the Flora (€ NG § is an impor- tant hallmark in the botanical history of the United States. S paper is intended to be the first of a series of commentaries on the plants included in Walter’s Flora Caroliniana. The present paper analyzes the thirteen species of oak (Quercus) reported by Walter as occurring in the approximately fifty square miles surrounding his plantation on the south bank of the Santee River some 45 miles northwest of Charleston. Walter’s thirteen oak binomials with their current equivalents are as follows: (1) Q. sempervirens Walter = Q. virginiana Mill.; (2) Q. phellos L. = Q. phellos L.: (3) Q. humilis Walter non Mill. = Q. incana W. Bartram; (4) Q. pumila Walter [There is no type spec- imen and the brief description is in flagrant conflict with the species that has borne the binomial for the past 213 years. A new species (Q. elliottii) is proposed to replace the misapplied name of Walter]; (5) Q. prinus L. [a previously suggested “ambiguous name” soon to pe formally proposed for rejection; Walter’s plant is Q. michauxii Nutt.|: (6) QO. Weta. SENSU Pie non L. = Q. marilandica Miinchh.; (7) Q. ee ‘a Walter = Q. 1 fa re (8) QO. rubra sensu a non L. = Q. falcata sae (9) Q. laevis ate = Q. laevis Heme QO) QO. alba L. = Q. alba L.; (11) O. lyrata Walter = Q. Iyrata Walter; ( on sinuata Walter aan eee (13) QO. villosa Walter = Q. ste ee coe Key Words: Quercus, South Carolina Among the accomplishments of Thomas Walter (c. 1740— 1789), emigrant from England, American patriot, South Carolina planter, merchant, community leader, and landowner (4500 acres), to list merely a sample, was the flora describing in Latin the plants found in the vicinity of his plantation (Rembert 1980). Walter sent his manuscript Flora Caroliniana for publication in England with his friend, the itinerant plantsman John Fraser (1750-1811). Botanists are probably not exhibiting undue paro- chialism in concluding that Walter’s principal claim to fame rests upon his Flora Caroliniana (1788) and that John Fraser’s greatest contribution in all likelihood is in encouraging Walter to bring his floristic investigations to completion as well as providing hun- 134 2002] Wilbur—Thomas Walter’s Oaks ie 3s) dreds of species for inclusion in the Flora that otherwise were unknown to Walter. Walter’s Flora was the first descriptive ac- count of a specific area prepared by a resident of eastern North America appearing after what is accepted as the starting point of botanical binomial nomenclature by Linnaeus in his Species Plan- tarum (1753). Fraser (1789) oversaw the publication of this man- uscript in London and indicated that he had added 420 species, making the total 1060 species treated in Flora Caroliniana. There is no information to my knowledge as to whether (1) Walter and Fraser jointly studied these botanical discoveries from Fraser’s wider exploration and together agreed upon their inclusion, or (2) Walter alone drew up the diagnoses, or (3) the inclusions are the result of only Fraser’s study and incorporation into the manuscript after he had left South Carolina. The third possibility seems the least likely. In any event, all new binomials and genera published in Flora Caroliniana have been attributed only to Thomas Walter. Unfortunately Fraser’s contribution introduced uncertainty as to the area covered by Flora Caroliniana, for Fraser traveled widely in search of horticultural subjects while Walter stated, in the preface of the Flora (Rembert 1980), that all but a few of the plants came from an area no greater than 50 square miles centered on his plantation on the south bank of the Santee River in north- western Berkeley County near the village of St. Stephen’s, about 45 miles north of Charleston. It is impossible to determine from the contents of the Flora if all, or at least most, of the species contributed by Fraser also came from this small area. We certainly know that some species included did not come from the area designated by Walter. Obvious examples would be Magnolia ac- uminata (L.) L. (widespread in eastern North America) and M. fraseri Walter, both included in Walter’s Flora but known only from the mountains of the Carolinas and adjacent montane states. Other examples that must owe their inclusion to Fraser’s travels are Trautvetteria caroliniensis (Walter) Vail (Ranunculaceae) and Frasera caroliniensis Walter (Gentianaceae). Harper (1911) listed twenty-four species included in Walter’s Flora that probably did not “grow within many miles of his home ... and a few that probably have not been seen in South Carolina at all...” More intensive collecting over the past nine decades has very much reduced Harper’s list but there still remain a number of species which are not known from the coastal plain of South Carolina and in all probability never grew there. 136 Rhodora [Vol. 104 Thirteen species of oaks are briefly described in Walter’s Flora Caroliniana, eight of which were first published in that publica- tion in the belief that they were unknown to science, as indicated by their being printed in italics. (Perhaps it should be noted that in practice, Walter’s use of italics was not consistent.) The fate or disposition of all 13 oak binomials included by Walter is dis- cussed in the following paragraphs. Each entry in Walter’s Flora under the generic name consists of three parts: (1) the specific epithet or what Linnaeus referred to as the trivial name, (2) the species number under each genus, and (3) the Latin diagnosis of the species. Ashe (1916), who had much interest in and experi- ence with the southeastern oaks, concluded that considering the brevity of Walter’s descriptions “they are excellent, but each must be considered in connection with the others he describes.”> The ate, astute and careful Howard Rock (1925-1964) noted (1956) that Walter’s descriptive phrases, if rearranged, amounted to a brief key to the species in each genus. — sempervirens 1. foliis lanceolatis perennantibus integerri- mus margine subrevoluto. All commentators noted for the past two centuries are agreed that Quercus sempervirens Walter (1788) is a later synonym of Q. virginiana Mill. (1768). Quercus virginiana Mill., Gard. Dict., ed. 8, Quercus no. 16. 1768. Q. phellos |var.| B L., Sp. Pl. 994. 1753. Q. sempervirens Walter, Fl. Carol. 234. 1788, non Mill, 1768. Q. virens Sol. in Aiton, Hortus Kew. 3: 356. 1789. Q. andromeda Riddell, New Orleans Med. Surg. J. 9: 614. 1853. Q. virginiana var. virescens Sare., Bot. Gaz. 65: 446. 1918. Q. virginiana var. eximea Sare., Bot. Gaz. 65: 447. 1918. Q. virginiana var. macrophylla Sarg., Bot. Gaz. 65: 447. 1918. Q. eximead (Sarg.) Trel., Mem. Natl Acad. Sci. 20: 116. 1924. Phellos 2. foliis deciduis lanceolatis integerrimis seta ter- minatis. The willow oak is abundant in Berkeley County, South Carolina, so we can be confident that it was well known to Walter. However, it seems certain that he may well have compounded with it other similar species that are also frequent in the area, such as Quercus laurifolia Michx. and perhaps Q. hemisphaerica W. Bartram ex Willd. The last two mentioned oaks are apparently frequent in Walter’s area but he obviously did not differentiate them from one another, which is understandable considering that 2002| Wilbur—Thomas Walter’s Oaks ew only in the past half century have botanists made much progress in distinguishing them. Quercus phellos L., Sp. Pl. 994. 1753. humilis 3. foliis lanceolatis integerrimis seta terminatis sub- tus tomentosis. Quercus humilis Walter (1788) is a later hom- onym of Q. humilis Mill. (1768), a European species. Trelease (1924) included Walter’s binomial in the synonymy of the so- called running oak, which has been long referred to as Q. pumila Walter for which the diagnosis of Q. humilis is a better match than that accompanying Q. pumila itself. Walter’s epithet “‘hu- milis” implies that the plant is of humble stature (1.e., a shrub) and its diagnosis stresses the tomentose lower surface of the blade while Walter’s diagnosis of Q. pumila states that the leaves are glabrous and that the lower surface is glaucous. I cannot disprove Trelease’s conviction that Q. humilis Walter is the species in Wal- ter’s Flora Caroliniana that matches the description of the run- ning oak. However, Walter’s protologue of Q. humilis also agrees with the stated characteristics of the species later known by the binomial Q. incana W. Bartram (= Q. cinerea Michx.), the blue- jack oak, except that a tree growing to 10 meters in height, al- though often much smaller, would scarcely be expected to receive the epithet Aumilis. Pursh (1814, p. 625) treated the bluejack oak as Q. phellos B [= var.| humilis citing Catesby’s account (1730, 1: 22. t. 22) and noted that the plant was “‘of low straggling growth.”’ Linnaeus (1753) previously had cited Catesby 1: 22. t. 22 as Q. phellos B. Catesby’s (1730) comments are included in full in the following quotation: Quercus humilior salicis folio breviore The Highland Willow Oak This is usually a small tree, having a dark coloured bark with leaves of a pale green, and shaped like those of a willow. It grows on dry poor land, producing but few acorns, and those small. Most of these oaks are growing at Mr. Fairchild’s. Catesby’s description and plate (1: 22. t. 22) were identified by Ewan (1974) as Quercus laevis while Howard and Staples (1983) and Wilbur (1990) identified it as QO. incana W. Bartram. Quercus incana W. Bartram, Travels Carolina 378. 1791. 138 Rhodora [Vol. 104 Q. phellos 8 ae Lam., Encycl. Méth. Bot. |: 722. 1785. QO. humilis Walter, Fl. Carol. 234. 1788, non Mill. 1768. Q. cinerea Michx.. Hist. Chénes Ameér., Quercus no. 8. pl. 14. 1801. pumila 4. foliis lanceolatis integerrimis glabris subtus glau- cis. As pointed out in the paragraph above, the Latin diagnosis of Quercus humilis matches the species originally proposed by Thomas Walter for what has been called for nearly the past two centuries Q. pumila. The diagnosis provided for Q. pumila by Walter describes a species whose glabrous leaves are glaucous beneath. Walter’s protologue for the running oak, Q. pumila, strik- ingly conflicts with the characteristics of the plant which has borne that binomial for over two centuries. Consequently, a name change is necessary for this very distinctive and familiar dwarf oak that ranges along the coastal plain from southeastern North Carolina southward throughout peninsular Florida and westward into Mississippi. To “retypify’> Q. pumila Walter with a specimen in accord with “current usage” would be in serious conflict with the last three words of the otherwise decidedly uninformative protologue. Quercus elliottii Wilbur, sp. nov., TYPE: UNITED STATES. South Carolina: Hampton Co., pine savanna along NW margin of Pigeys Rd., | mi. W of main office at James W. Web Wildlife Center, 4.2 mi. W of Garnett off secondary highway Rt. 20; 32.6216°N, 81.3213°W, 54 ft. elevation, 13 Oct 2000, Nelson 21668 & Wood (HOLOTYPE: DUKE; ISOTYPES: BKL, BRIT, CU, DLE DUKE, FE FLAS, FSU, FTG, GA, GH, IBE, ILL, LSU, MICH, MISS, MISSA, MO, NCSC, NCU, NLU, NY, TEX, UNA, US, USCH, USK VSC, WIS, WNC). Differt a Quercus incana W. Bartram habitu fruticoso et co- loniali sobolibus, caulibus 1(-2) m altis. Fructus hornotini matu- rescentes, sessiles vel brevipedicelli; cupula 4—5 mm alta, crater- iformis, squamis arcte appressis, cinereis, appresso-pilosis; glans (in cupula) inserta. Shrub, commonly forming extensive clones by subterranean runners; stems O.5—1 (—2) m tall, profusely sprouting from their bases after burning of pinelands, the leaves of sprouts often larger than those of stems unburned for several years. Woody twigs of the season grayish brown, usually much of their pubescence per- sisting through the first year. Winter bud or buds at the tips of twigs ovoid-conic, 3-5 mm long, brown, the scales mostly with 2002] Wilbur—Thomas Walter's Oaks 139 a fringe of minute trichomes around their apical margins. Leaves all deciduous in autumn or a few of them overwintering and falling just before or as new growth commences in spring. Stems of young shoots moderately to densely stellate-pubescent; edges of unfurling leaf blades downwardly curved and recurved cov- ering perhaps as much as half of the lower surfaces, their upper surfaces with sparse, pale, stellate pubescence but eventually gla- brescent; the lower surfaces shortly, densely, and compactly pale- gray, stellate-pubescent. Mature leaves very short-petiolate; pet- ioles stellate-pubescent. Blades mostly 3—10 (—I5) X* 0.7—2 (—5) cm, oblanceolate or spatulate, narrowly elliptic, elliptic-oblong, or lanceolate, usually with a short bristle tip; bases cuneate to narrowly rounded, apices rounded to acute; upper surfaces gla- brous, dark green and lustrous or sublustrous, sometimes dull green, lower surfaces densely and compactly grayish puberulent; flat and with entire margins, sometimes their edges somewhat crisped, only rarely with a few, low, rounded undulations. Fruits maturing in one season, sessile or shortly peduncled, their invo- lucres bowl-like, 4-5 mm deep, embracing about one-third the length of the acorn, scales tightly appressed, grayish brown, broadest basally where many or most of them are humped or bulged, gradually narrowed distally to truncated, flat tips; acorns ovoid, subglobose, or somewhat oblate, 8-12 mm long and broad, basally flat, apically rounded to nearly truncate, outer surfaces light brown, glabrous or faintly and sparsely very short-pubescent near their summits, inner surfaces loosely pale-pubescent near their summits, inner surfaces loosely pale-pubescent, the tri- chomes blond to tawny. ~ It might be argued that all that was needed to rehabilitate no- menclaturally a case like that of Quercus pumila Walter was that a neotype be designated and published, confirming the identity of the plant in the traditional sense and thereby nullifying the questionable phrases in the original diagnosis. However the orig- inal descriptive phrases in Walter’s diagnosis are exceedingly brief. If we were to ignore or delete the questionable last three words from the descriptive diagnosis of the running oak, there would remain very little that was distinctly descriptive, and those three descriptive words exclude the species to which the name has been employed. The preceding entry (i.e., that for Quercus humilis; 3. foliis 140 Rhodora [Vol. 104 lanceolatis integerrimis seta terminatis subtus tomentosis) is clear- ly a much better fit for what has been passing as Q. pumila Walter, than is the descriptive account accompanying Q. pumila itself. That account has been attributed to Q. incana W. Bartram. In any event, Q. humilis Walter (not Q. humilis Mill., 1768) is a later homonym and cannot now be applied to any species named after 768 Prinus 5. foliis ovatis sinuato-serratis, denticulis uniformi- bus. The chestnut oak naturally occurring in the coastal plain of South Carolina is Quercus michauxti Nutt., the swamp chestnut oak. Quercus prinus L. is now most often referred to as Q. mon- tana Willd. Hardin (1979) recommended that the binomial Q. prinus be treated as an ‘“‘ambiguous” name since the lectotypic specimen cannot be conclusively identified because the features displayed are not those that distinguish the two species confound- ed by Linnaeus (1.e., Q. prinus and Q. michauxii) under the bi- nomial Q. prinus. Linnaeus’ binomial has been applied to both species (1.e., to either the chestnut oak or to the swamp chestnut oak, for lengthy periods as shown by Hardin’s table). Fortunately, for the purposes of this paper, the oak in Walter’s area can only be the bottomland swamp chestnut oak, as only that species is known from eastern South Carolina. John Fraser, however, had ample opportunity to observe both species during his extensive travels. It is to be remembered that Sargent (1916) reversed the application of the name Q. prinus from the mountain chestnut oak to the swamp chestnut oak nearly nine decades ago based on his belief that the mountain chestnut oak was not to be found in southeastern Virginia, the presumed “‘type” locality of the Clay- ton specimen described by Gronovius (1739). Sargent’s reversal was generally followed for several decades by American workers and especially by foresters and by E. J. Palmer (1943) whose study convinced him that Sargent was correct in applying Q. pri- nus L. to the swamp chestnut oak. However, additional floristic investigations (e.g., Fernald 1946, p. 391; Harvill et al.1986, pp. 85—86) have demonstrated that both the swamp chestnut oak and the rock chestnut oak are to be found in southeastern Virginia in close proximity to Clayton’s home. In my opinion, the name Q. prinus L. has not yet been formally disposed of and the binomial needs to be either laid to rest by rejection, or epitypified and adopted. A paper proposing the first alternative will soon be sub- 2002] Wilbur—Thomas Walter’s Oaks 14] mitted to Taxon. Quercus michauxii Nutt. is abundant in the bot- tomlands of the Santee River upon whose southern bank Walter’s plantation was located. Quercus michauxii Nutt., Gen. N. Amer. Pl. 2: 215. 1818. Q. prinus L., Sp. Pl. 995. 1753, in part, nom. rej. prop. QO. prinus aes palustris Michx., Hist. Chénes Amér., Quercus no. 5. pl. 6. 1801. [°Q. Prinus taal) Michx.”” QO. prinus a eae Michx., Fl. Bor.-Amer. 2: 196. 1803. Q. prinus var. michauxii (Nutt.) Chapm., Fl. South. U.S. 424. 1860. QO. houstoniana C. H. Mull., Amer. Mid]. Naturalist 2: 743. fig. 1. 1942 nigra 6. foliis obcuneiformibus obsolete trilobis villosis ra- mis inferioribus declinatis, superioribus adscendentibus. The advantage that familiarity with plants in the field provides to the investigator is clearly demonstrated by Walter’s treatment of this species and the next (Walter’s #6 and #7). Walter treated both as species while Linnaeus combined them as varieties of Quercus nigra L. Perhaps it would be more accurate to state that Linnaeus treated as a varietal appendage, the B variety of Q. nigra as the element that became QO. marilandica. Britten (1909) has a detailed explanation of the early travail of the two elements included by Linnaeus within his Q. nigra. Britten there informs us that “*Wal- ter’s herbarium contains a leaf’? of both Q. nigra and Q. mari- landica although neither bears an identification by Walter. Wal- ter’s solution was to remove the Gronovian and Catesbian (1: 20) references as Q. aguatica Walter, leaving the Ray and Catesbian (1: 19) references as QO. nigra L. However, Walter’s solution to Linnaeus’ confusion in placing the water oak and the blackjack oak under the binomial Q. nigra was not the first remedy pro- posed. Miinchhausen (1770, 5: 253) had named the blackjack oak, Q. marilandica, in effect removing the Linnaean £6 variety, leav- ing Q. nigra L. as the binomial for the water oak. The result was to segregate the Gronovian and Catesbian references as Q. nigra and leaving Q. nigra & exemplified by Catesby’s I: 19. t. 19 “Quercus marilandica folio trifida ...” of Ray and Catesby as Q. marilandica Miinchh. The species that Walter retained under the Linnaean binomial, QO. nigra, is now known as Q. marilan- dica. Quercus marilandica Mitinchh., Hausvater 5: 253. 1770. Q. marilandica .. . pete Nat. Hist. Carol. 1: 19. t. 19. 1730. 53 Q. nigra |var.| BL . Pl. 2: 996. 17 142 Rhodora [Vol. 104 YQ. cuneata Wangenh., Beytr. Teut. pocelinge 78. 1787. . ferruginea FE Michx., Hist. Arbr. Forest. 2: 92. pl. 18. 1812. . nigra B quinqueloba Alph. de Candolle, Pate (DC.) 16(2): 64. 1864. nigra y tridentata Alph. de Candolle, Prodr. (DC.) 16(2): 64. 1864. marilandica var. ashei Sudw., Jour. For. (Washington) 20: 167. 1922. . marilandica f. cuneata (Wangenh.) Trel., Mem. Natl. Acad. Sci. 20: 200. 1924 Mower aquatica 7. foliis obcuneiformibus obsolete trilobis submu- cronatis laevibus nitidis, subperennatibus. As explained above, Walter divided Linnaeus’ Quercus nigra into its two component species: Q. nigra was the name retained for the blackjack oak, and the water oak, fittingly enough, was named Q. aguatica Wal- ter. However, Mtinchhausen (1770, 5: 253) had corrected Linnae- us’ confusion earlier by naming the blackjack oak Q. marilandica Miuinchh., which left the binomial QO. nigra L. for the water oak. Quercus nigra L., Sp. Pl. 995. 1753. Q. uliginosa Wangenh., Beytr. Teut. Forstwiss. 80. 1787. QO. nla ‘d ee FI. Carol. 234. Oe QO. nana , Sp. Pl., ed. 4.4(1). 443. 1805. V0. ener Riddell, New Orleans a Surg. J. 9: 614. 1853. Q. aquatica a supttata Alph. de Candolle, Prodr. (DC.) 16(2): 68. 1864. Q. rhombica var. obovatifolia Sarg., Bot. Gaz. 65: 431. 1918. Q. nigra var. tridentifera Sarg., Bar Gaz. BS: 429, 1918. rubra 8. foliis 3 s. 5 lobis obtusis subtus villosis, setaceo- mucronatis glandibus parvis globosis. Totten (Radford et al. 1968) did not map Quercus rubra L. as occurring in the coastal plain of South Carolina although it was well-dispersed throughout the piedmont and mountains of that state. The same source shows it to be widely scattered and apparently rare in the coastal plain of North Carolina. Svenson (1939) and Fernald (1946), among others, have pointed out that many Linnaean species include two or more species, based on the included synonymy according to more recent systematists who have had the advantages of greater familiarity with the plants in the field and/or more extensive col- lections available for comparison. For example, Fernald (1946, p. 391) pointed out that in Species Plantarum (Linnaeus 1753), the name Q. rubra “covered many (if not most) of the eastern species of subgenus Erythrobalanus .’ including the red oak itself. Svenson (1945) concluded “that the Linnaean species from one point of view was the synthesis of all bibliographic citations un- 2002 | Wilbur—Thomas Walter’s Oaks 143 der the species, together with the Linnaean herbarium specimens, whether they were associated with the citations.” Du Roi (1772) was apparently the first to restrict the name Quercus rubra to a single species. That choice determined that the binomial Q. rubra L. thereafter should be reserved for the red oak of northeastern North America as well as covering much of eastern United States and adjacent Canada (see Nixon and Muller 1997, p. 465 for map). However Sargent’s (1915, 1916) own research and sense of propriety convinced him that “the name Quercus rubra belonged to the tree which was later called Q. falcata by Michaux and not to the tree which has always been called red oak in the northern states.”” Sargent admitted that “this change of name is one of the most unfortunate which the study of old specimens of American plants has made necessary .. .”’ Sargent’s prestige was such that many, including most foresters and followers of the American Code of Nomenclature, for the next two decades or so applied the Linnaean binomial Q. rubra to the southern red oak (= Q. falcata Michx.) whose leaves are abaxially densely and perma- nently tomentose beneath. Sargent seemingly attached great im- portance to the first synonym appearing in the Linnaean proto- logue, no doubt influenced by Linnaeus’ own statement that the synonym with the best description should be listed first (see foot- note in Svenson 1939, p. 522). Sargent was also convinced, based on insufficient field experience, that only Q. falcata Michx. of the rubra-complex was to be found in southeastern Virginia, the area in which Clayton and Banister lived and from which they sent collections to European botanists such as Gronovius and Ray. The first synonym listed by Linnaeus, as pointed out by Sargent (1915), is that of Gronovius (1739) based on a collection by John Clayton. Sargent found Clayton’s specimen to be what has been called Q. falcata Michx. and felt that there was no alternative but to apply the name Q. rubra to that element of Linnaeus’ multi- parted concept of Q. rubra. Naturally, applying the binomial Q. rubra L. to two very different species led to confusion, leading Rehder (1938) to propose unsuccessfully that the name be offi- cially declared a nomen ambiguum. Harvill et al. (1986, p. 85— 86) maps show that both the red oak and southern red oak are abundant in southeastern Virginia. Others (e.g., Svenson 1939, 1945; Fernald 1946) took strong exception to Sargent’s retypifi- cation of a species first typified by Du Roi (1772). 144 Rhodora [Vol. 104 Fortunately, for the purposes of this paper there is no problem, as Walter’s descriptive polynomial is explicit for the villosity of the leaf’s undersurface. He clearly was applying the name to the same element that Sargent mistakenly felt obliged to choose (1.e., the element that Michaux called Quercus falcata). The northern red oak has not been found in Walter’s area but the southern red oak is abundant there now, as it surely was in Walter’s time. Quercus falcata Michx., Hist. Chénes Amér., Quercus no. 16. pl. 28. 180] Q. nigra digitata Marshall, Arbust. Amer. 123. 1785. Q. triloba Michx., Hist. Chénes Amér., Quercus no. 14. pl. 26. 1801. QO. oe Willd., hess Naturf. Freunde Berlin Neue Schriften 3: 400. 1801; Sp. Pl. ed. 4.4(1). 444. 1805 Q. falcata B triloba ( Machi, ) Nutt., Gen: N. Amer. Pl. (Nuttall) 2: 214. IS18. Q. carpenterti Riddell, New Orleans Med. .J. 9: 613. 1853. QO. falcata B ludoviciana Alph. de od ‘Prodr. (DC.) 16(2.1): 59. 1864. Q. digitata (Marshall) Sudw., Gard. & Forest 5: 99. 1892. Q. rubra var. triloba (Michx.) Ashe, Proc. Soc. Amer. Foresters. 11: 90. 1916. nom. illegit., Art. 34.1(b). Q. rubra var. leucophylla Ashe, Bull. Charleston Mus. 13: 25. 1917. Q. pagoda var. leucophylla (Ashe) Ashe, J. Elisha Mitchell Sci. Soc. 34: 136. 1918. Q. leucophylla (Ashe) Ashe, Torreya 18: 73. 1918. Q. rubra sensu Sarg., Bot. Gaz. 65: 426. 1918. Q. rubra var. triloba (Michx.) Sarg.. a Gaz. 65: 427. 1918. Q. joori Trel., Mem. Natl. Acad. Sci. 15. 1924. Q. rubra f. gine ak Treh, oa Natl. Acad. Sci. 20: 201. pl. 406, fig Q. rubra ft. “ale ata ae ) Trel.. Mem. Natl. Acad. Sci. 20: 202. pl. 406, fig. 2. 1924. Q. rubra var. triloba (Michx.) Sudw., Check List For. Trees U.S. 89. 1927. Q. rubra var. leucophylla (Ashe) Sudw., Check List For. Trees U.S. 90. Q. rubra var. digitata (Marshall) Cory & Parks, Cat. Fl. Tex. 37. 1937. laevis 9. foliis obtuse sinuatis laevibus setaceo-mucronatis, glandibus magnis depresso globosis calyce subtectis. The syn- onymy of the turkey oak is as follows: Quercus laevis Walter, Fl. Carol. 234. 1788. Q. Catesbaei Michx., Hist. Chénes Amér., Quercus no. 17. pl. 29-30. ISOl. 2002] Wilbur—Thomas Walter’s Oaks 145 Q. flammula W. Bartram, Travels Carolina 228, 344, 359, 403, 470. 1791, nom. nud. alba 10. foliis pinnatifidis laevibus, lobis finus subaequan- tibus, supra saturate viridibus subtus glaucis, glandibus mag- nis ovatis. There seems to be no doubt that Walter’s concept of the white oak, Quercus alba was also that of Linnaeus. This spe- cies is abundant about Walter’s former plantation. Quercus alba: L,.,. sp. PL. 996. 1733. QO. alba frutescens Miinchh.. Hausvater 5: 253. 1770. ?. 5 a pinnatifida Michx., Hist. Chénes Amér., Quercus no. 4. pl. 5, I. 180] Q. Pon 8 repanda Michx., Hist. Chénes Amér., Quercus no. 4. pl. 5, fig. 3. I8OL. Q. alba var. latiloba Sarg., Bot. Gaz. 65: 435. 1918. lyrata 11. foliis lyratis laevibus sinubus obtusissimis lobis remotis inaequalibus, glandibus magnis globosis subtectis. Again, no controversy has yet surrounded the identity of the ov- ercup oak first named and described by Walter. Quercus lyrata Walter, Fl. Carol. 235. 1788. Scolodrys lyrata (Walter) Raf., Alsogr. Amer. 29. 1838. sinuata 12. foliis sinuatis laevibus obtusis supra _ pallidis, subtus subglaucis, glandibus mediocribus globosis calyce sub- plano. Contrary to the lack of debate concerning the identity of such species as Quercus alba and Q. lyrata, there has been much uncertainty about the identity of Q. simuata Walter. This uncer- tainty 1s not lessened by the lack of original material among Wal- ter’s specimens at BM [so reported by Sargent (1918, p. 436) and by Nixon and Muller (1997, p. 497)]. Prior to Camus’ (1939, 2 678), Muller’s (1951), and Dorr and Nixon’s (1985) acceptances of Q. durandii Buckley as a synonym of Q. sinuata Walter, there had been a slowly growing consensus that this was the proper disposition of Buckley’s binomial (see Elias 1971, p. 183). How- ever, I find that the considerable uncertainty as to the identity of Walter’s Q. sinuata prevents me from joining that growing con- sensus. Original specimens representing Thomas Walter’s oak collec- tions are unknown and hence their interpretation must depend upon their original descriptions. Walter’s descriptions, in the judg- 146 Rhodora [Vol. 104 ment of W. W. Ashe (1916) are “excellent” in spite of their brevity, but “each .. . must be considered in connection with the others he describes.”” The description of Quercus sinuata has proven to be most problematic. Both Engelmann (1876, p. 400) and Sargent (1895, p. 144) concluded that Q. sinuata was the hybrid of Q. catesbaei (= Q. laevis) X Q. nigra. Ashe (1916) challenged this interpretation since the hybrid has a deep acorn cup with a rounded base and not the saucer-shaped cup with a nearly flat base described by Walter, and also foliage that “‘is dark green and lucid above and not pale [and] is bright green below and not sub-glaucous.”’ Ashe at first unfortunately confused Q. austrina Small (1903) with Walter’s Q. sinuata, overlooking the fact that Small’s species was described as having both leaf sur- faces bright green and with an acorn cup hemispheric in contrast to the flattened cup described by Walter for Q. sinuata. Ashe (1918, p. 11) unobtrusively admitted his error in placing Q. aus- trina in the synonymy of Q. stnuata and made thereafter no fur- ther pronouncements on the identity of Q. sinuata. Trelease (1924, p. 1O1) however, apparently unaware of Ashe’s retraction, followed Ashe’s earlier opinion in combining Q. sinuata and Q. austrina. Unfortunately Trelease paid little or no attention to Wal- ter’s description as he separated Q. sinuata f. sinuata from f. durandti (Buckley) Trelease by the former’s green lower leaf sur- face in contrast to the pale lower surface of the latter. Palmer (1945) thought Engelmann’s conclusion (1876-1877) that Quercus sinuata Walter was a hybrid between Q. /aevis and Q. nigra was “a more reasonable interpretation” than Ashe’s ear- lier (1916) conclusion that Q. austrina Small was a later synonym of Q. sinuata. Both Trelease (1924) and Muller (1951) accepted the earlier opinion of Ashe (1916) that Q. simuata was an earlier name for Q. durandii. This conclusion was firmly rejected by Ashe (1918). Palmer concluded that Q. simuata had indeed been mistakenly identified as synonymous with Q. austrina Small by Ashe, as Ashe (1918) himself had admitted in an obscure retrac- tion. In his detailed study of the Q. durandii complex, a group restricted in his opinion to the calciphilic soils of the Gulf coastal plain and east Texas, Palmer (1945) maintained that synonymi- zation with Q. sinuata was clearly unwarranted since Q. durandii is not known in Walter’s region, and in no character except pos- sibly in the shape of the leaves could Walter’s description be reconciled with Q. durandii. Palmer felt so strongly about the 2002] Wilbur—Thomas Walter’s Oaks 147 matter that he claimed that “‘until a specimen named by Walter can be seen the name must remain doubtful.’ Palmer treated QO. sinuata as a nomen dubium. Muller (1951), ignoring or at least making no reference to Palmer’s paper, took up Quercus sinuata Walter, including in its synonymy both Q. austrina Small and Q. durandii Buckley, feel- ing that “arguements against identity of this plant with Walter’s name include the inability of contemporary collectors to find the species in Walter’s immediate territory, which is distinctly incon- clusive.”” Walter’s description of Q. sinuata leaves as “‘subtus subglaucis”” and the fruit as ““mediocribus globosis calyce sub- plano” indeed excludes other southeastern oaks and agrees per- fectly with the form with silvery lower leaf surfaces that Buckley named Q. durandii. [One can’t help but point out that Buckley’s only mention of surface features in the original description was ““when mature, smooth on both sides,” which offers little support to Muller’s own description (1951) of the species he called OQ. sinuata (including in synonymy Q. durandti and Q. austrina): “upper surfaces from sparsely minute-stellate becoming glabrate and glossy dark green, lower surfaces persistently pubescent with minute appressed dense stellate hairs strikingly silver or appear- ing green if the pubescence is sparse, occasionally tardily glabrate .’ | Later, Nixon and Muller (1997) recognized Q. austrina as a species separate from Q. sinuata, but only after Dorr and Nixon (1985) accepted Ashe’s (1916) submergence of the two species (.e., QO. austrina within Q. sinuata), making no mention of Ashe’s retraction (1918). Nixon and Muller (1997, p. 498) stated that “the original description of Q. sinuata 1s consistent with the con- cept presented ... by W. W. Ashe (1916) and W. Trelease (1924), and inconsistent with any other oak from the broad area covered by Thomas Walter’s Flora...” This review of the pertinent literature is not one that gives confidence that enough is known about the identity of the types, the morphological limits of the species involved, or their geo- graphic ranges, etc., to be dogmatic as to the application of the binomials of these littke known taxa. The application of the bi- nomial Quercus sinuata Walter is too uncertain, in my opinion, to be adopted at the present time; it very much remains a nomen dubium. More field work and observation are very much needed for many of the southeastern oaks. 148 Rhodora [Vol. 104 villosa 13. foliis obtuse lobatis, supra nitidis subtus villosis glandibus parvis globosis. Although neither Michaux (1803) nor Pursh (1814) placed Walter’s Quercus villosa in synonymy of any species in their early floras of North America, later authors have rather unanimously identified Q. villosa Walter as a synonym of the earlier Q. stellata Wangenh. This post oak 1s common in Walter’s area as well as much of the eastern United States. There is little in Walter’s diagnosis that would have convinced me that the plant described was the post oak, but there is nothing that would cause me to question the identity except that the descriptor “villose” would not have occurred to me as describing the very familar Q. stellata. The pubescence on the stem and leaves of the post oak, in my experience scarcely qualifies as being villose. Quercus stellata Wangenh., Beytr. Teut. Forstwiss. 78. pl. 6, fig. 5. 1787 Q. alba minor Marshall, Arbust. Am. 120. 1785. ullosa Walter, Fl. Carol. 235. : . lobulata Sol. in Smith & Abbot, Insects of S 1: 93. pl. 47. 1797. . obtustloba Michx., Hist. Chénes Amér. pl. 1801. . stellata B floridana Alph. de Candolle, aa (DC.) 16(2): 24. 1864. . minor (Marshall) Sarg., Gard. & Forest 2: 471. 1889. . Stellata var. eg Sarg., Bot. Gaz. 65: 438. 1918. . ashei Ste i Elisha Mitchell Sci. Soc. 37: 178. 1922. . stmilis ae , J. Elisha Mitchell Sci. Soc. 40: 43. 1924. QO. stellata var. i. (Ashe) Sudw., U.S.D.A. Misc. Circ. 92: 107. 1927 oo LITERATURE CITED ASHE, W. W. 1916. Notes on Trees. Bull. Charleston Mus. 14: 9-12. —.. 1918. Notes on Trees. Proc. Soc. Amer. Foresters 11: 88-90. BRITTEN, J. 1909. Pan nigra. J. Bot., British & Foreign 47: 349-350. Camus, A. A. 1934-1954. Les Chénes. Monographie du genre Quercus, Vol. 2. Paul Lechevalier, Paris. [2: 678] CaresBy, M. 1730-1742. The Natural History of the Carolinas, Florida and the oe Islands. 2 vols. Folio. London, U.K. Dorr, L. b K. C. Nixon. 1985. Fypiiston of the Bor nia taxa een by S. B. Buckley (1809-1884). Ta 34: 211-22 Du Ro, J. P. 1772. Harbkesche Wilde Baumzucht Theils cee Sees Ist ee ih senhaus Btichandlung, Braunschweig, Germany. [p. 265| Eun T. S. . The genera of Fagaceae in the ea United States. re pee 52: 152-195. — G. 1876-1877. About the Oaks of the United States. Proc. St. Louis Acad. Sei. 3: 372—400, 539-541. 2002] Wilbur—Thomas Walter’s Oaks 149 Ewan, J. 1974. Notes, pp. 89-100. /n: The Natural History of Carolina, Florida and the Bahama Islands, containing two hundred and twenty figures of birds, beasts, fishes, serpents, insects, and plants, by M. Cates- by. Facsimile of the 3rd. ed. (1771). Beehive Press, Savannah, GA. FERNALD, M. L. 1946. Types of some American trees. J. Arnold Arbor. 27: 386-394. [pl]. 1-3] FRASER, J. 1789. F short history of the Agrostis cornuc sas or the new American grass and also some account of a journey to the Gea Nation, in oe of new plants. 8 page folio. London, U.K. GRONOVIUS, J. F 9 & 1743. Flora Virginica Exhibens Plantae quas V. C, Johannes ie in Virginia observavit atque collegit. 2 parts. Leiden, Netherlands. HARDIN, J. W. 1979. Quercus Prinus L.—nomen ambiguum. Taxon 28: 355— HARPER, R. M. 1911. Early spring aspects of the coastal plain vegetation of South Carolina, Georgia, and northeastern Florida. Bull. Torrey Bot. Club. 39: 223-238. [n.b. footnote on p. 232] Harvi__, A. M., JR., T. R. BRADLEY, C. E. STEVENS, T. EF WieBoLtpt, D. M. E. WARE, AND ‘DW . OGLE. 1986. Atlas of the Virginia Flora, 2nd ed. Virginia Botanical Associates, Farmville, WA. [Quercus, pp. 85-86] Howarb, R. A. AND G. W. STAPLES. 1983. The modern names for Catesby’s plants. J. Arnold Arbor. 26: 482—483 LINNAEUS, C. VON. 1753. Species Plantavita: Vol. 2. Stockholm. [Quercus, 2: 994-997 | MiIcHAux, A. 1803. Flora Boreali-Americana (Michaux), Vol. 2. Paris and eae oo 2: 194-200] MULLER, C. H. 1951. Oaks of Texas. Contr. Texas Res. Foundation, ae TX. a 20-311] MUNCHHAUSEN, O. VON. ae Der Hausvater. 6 vols. Hannover, Ger- any. Nixon, K. C. AND C. H. MULLER. 1997. Quercus L. sect. Quercus, pp. 471- 506. In: Flora of North America Editorial Committee, eds., Flora of North America North of Mexico, Vol. 3. Oxford Univ. Press, Oxford and New York PALMER, E. J. 1943. Quercus Prinus Linnaeus. Amer. Midl. Naturalist 29: 783-784. ——.. 1945. Quercus Durandii and its allies. Amer. Midl. Naturalist 33: —519. PursH, FT. 1814. Flora Americae Septentrionalis; or, a Systematic Arrange- ment and Description of the Plants of North America, Vol. 2. London, K. [Quercus, 2: 625-634] RADFORD, A. E., H. E. AHLES, AND C. R. BELL. 1968. Manual of the Vascular Flora of the Carolinas. Univ. North Carolina Press, Chapel Hill, NC. Quercus, pp. 372-385 REHDER, A., E. J. PALMER, AND L. CRoizat. 1938. Seven binomials proposed as nomina ambigua. J. Arnold Arbor. 19: 282-285. |Quercus rubra L., pp. 283-284] REMBERT, D. H., JR. 1980. Thomas Walter, Carolina botanist. Bull. No. 5, South Carolina Museum Comission, Columbia, SC. Co 150 Rhodora [Vol. 104 Rock, H. EF L. 1956. The eae of Helenium in Walter’s Flora Carolt- niana. ee 58: 311-317. SARGENT, C, Sx. 1895,- The Sava of North America, Vol. 8. Houghton Mifflin and nee ae and New York. . Three of Clayton’s oaks in the British Museum. Rhodora 17: ——. 1916. The name of the red oak. Rhodora 18: 45—46. . 1918. Notes on North American trees. Bot. Gaz. 65: 423-459. SMALL, J. K. 1903. Flora of the amore United States. Published by the author, New York. [Quercus, pp. 348-355] SVENSON, H. K. 1939. Quercus rubra once more. Rhodora 41: . 1945. On the descriptive method of Linnaeus. Rhodora re oe 302, 388. 363- TRELEASE, W. 1924. The American Oaks. Mem. Natl. Acad. Sci. 20: 1-255. WALTER, 7 nee Flora Caroliniana. Privately published by John Fraser, Lon- do K. [Quercus, pp. 234-235] WILBUR, an a 1990. Identification of the plants illustrated and described in Catesby’s Natural History of the Carolinas, Florida and the Bahamas. Sida 14: 29-48. RHODORA, Vol. 104, No. 918, pp. 151-160, 2002 RECONSTRUCTING THE BIOLOGICAL INVASION OF EUROPEAN WATER-HOREHOUND, LYCOPUS EUROPAEUS (LABIATAE), ALONG THE ST. LAWRENCE RIVER, QUEBEC DANIEL LACHANCE! AND CLAUDE LAVOIE Département d’aménagement and Centre de recherche en aménagement en développement, Université Laval, cane roy, i G1K 7P4, Canada I e -mail: d *rad.ulaval.« ABSTRACT. In Québec (Canada), one of the most recently introduced ex- otic wetland plants is European water-horehound (Lycopus europaeus). The first specimens were discovered in 1963 near Montréal. In this study, we used herbarium specimens and conducted field surveys to reconstruct the history * the invasion of European water-horehound in Québec, and to accurately iene its current northeastern distribution. Few European water-hore- hound specimens were collected before 1970. However, between 1970 and 1974, the range of European water-horehound expanded 380 km northeast- ward from Sorel to Trois-Pistoles River. [In , the northeastern distribution limit of European water-horehound was at Bic Pievacil Park, 65 km north- ast of Trois-Pistoles River. Between 1963 and 197 uropean water-hore- eee spread rapidly lene the St. Lawrence River (45 km/yr.), which was probably related to the fact that seeds remain viable after floating. Between 1974 and 1999, it spread more slowly to the northeast of Trois-Pistoles River (3 km/yr.). The limited range a ion of European water-horehound in east- ern Québec between 1974 and 1999 suggests that the salinity of surface wa- ters, and more een . scarcity of coastal or riverine marshes east of Rimouski, prevented populations from establishing in the estuarine part of the St. Lawrence River Key Words: Lycopus europaeus, European water-horehound, Québec, St. Lawrence River, biological invasion, exotic species In North America, invasion by exotic species is considered to be one of the main threats to preserving the integrity of ecosys- tems. In the United States alone, approximately 50,000 nonindig- enous species cause major environmental damage and financial losses totaling US$137 billion per year (Pimentel et al. 2000). More than 5000 introduced plant species are now naturalized in North American ecosystems. Several of these plant species are problematic. For example, control costs and forage losses asso- ciated with purple loosestrife (Lythrum salicaria L.) have been estimated at US$45 million in the United States (Pimentel et al. IS] L352 Rhodora [Vol. 104 2000). It is therefore important to understand the mechanisms underlying successful plant invasions, and to develop models to predict the spread of invaders. Such models are useful for im- proving management plans that have been established to mini- mize the impacts of invasive species on natural ecosystems (Re- jmanek and Richardson 1996). One of the key components in modelling is the rate at which a species spreads, or the distance its range expands each year, indicated by newly established individuals outside the distribution range of a species (Lonsdale 1993). Rates of spread for plants are highly variable, and mainly depend on autecological charac- teristics of species and dispersal vectors. For example, between 1970 and 1984, the sedge Carex praegracilis W. Boott (a native species in North America) migrated eastward from Illinois, In- diana, and Michigan to New England states at a rate of 73 km/ yr. This very rapid expansion rate was probably related to the development of highway networks, since this species is adapted to open and saline habitats commonly found along roads (Rez- nicek and Catling 1987). On the other hand, the rate of spread for the woody weed species Mimosa pigra L. in northern Aus- tralia (an exotic species in that country) was only 0.076 km/yr. between 1979 and 1985. Nevertheless, this rate is considered to be rapid for this pest species in Australia (Lonsdale 1993). Wetland plant species usually spread rapidly because water is an effective dispersal vector (Catling and Porebski 1995; Lons- dale 1993; Pysek and Prach 1995). Wetland exotic plants are also among the most aggressive invaders and have dramatically changed the vegetation of many marshes at temperate latitudes. Several species are known to reduce the biomass of native plants, contribute to filling in small ponds, and form almost monospecific plant communities (Galatowitsch et al. 1999), In Québec (Cana- da), one of the most recently introduced exotic wetland plants is European water-horehound (Lycopus europaeus L., Labiatae). The native distribution range of this species is in Europe and western Asia (Stuckey and Phillips 1970). European water-hore- hound is a medium-sized plant, 0.4—1 m in height, copiously pubescent, with toothed leaves 4—12 cm long and 1.5—5 cm wide (Henderson 1962). This species is very similar to the widespread native American water-horehound (L. americanus Muhl.), but L. europaeus can be easily distinguished by its pubescent leaves (Scoggan 1979). Both species colonize marshes and drainage = 2002] Lachance and Lavoie—Lycopus europaeus 153 ditches, as well as the shores of ponds, lakes, and rivers (Fleurbec 1987; Stuckey and Phillips 1970). The range of American water- horehound extends from British Columbia to Newfoundland, and from James Bay to Texas (Fleurbec 1987). European water-hore- hound was introduced into North America about 1860, probably in Virginia. Populations are now widespread along the Atlantic coast, from North Carolina to Nova Scotia. Numerous popula- tions are also located along the shores of Lake Erie, Lake Ontario, and the St. Lawrence River. The introduction of European water- horehound into the Great Lakes—St. Lawrence River system (about 1903) and along the Atlantic coast of the United States were probably distinct events related to the release of ship’s bal- last (Stuckey and Phillips 1970). The occurrence of European water-horehound in Québec is re- cent. The first specimens were discovered in 1964 near Valley- field (Figure 1; Rousseau 1968). In 1974, Gauthier (1977) estab- lished its northeastern distribution limit at the mouth of Trois- Pistoles River (48°06'N, 69°14'’W). On the shores of the St. Law- rence River, European water-horehound populations have mainly been found in marshes located in the supralittoral zone, and in the upper part of the intertidal zone (Chrétien 1994; Gauthier and Lavoie 1975). Stuckey and Phillips (1970) suggested that this plant was migrating down the St. Lawrence River, but there has been no historical reconstruction of the spread of European water- horehound in Québec that could be used to substantiate this as- sertion. In this study, we used herbarium specimens and con- ducted field surveys to reconstruct the history of the invasion of European water-horehound in Québec, and to accurately deter- mine its northeastern distribution limit. We also calculated the rate at which this species has spread since its introduction into the province. Since water is probably the main dispersal vector for this species (Fleurbec 1987), we hypothesized that European water-horehound has spread along the St. Lawrence River at a rapid and constant rate over the last 35 years. MATERIALS AND METHODS To reconstruct past and recent distribution ranges for European water-horehound in Québec, we gathered information on all her- barium specimens of this species collected in the province. Her- barium specimens were requested from a total of eight herbaria: VK Pp ¥ 46°N BY Day Aaa ' Nags oe Before be! BSE \ 1965 Trois-Pistoles N Saint-Jean-Port-Joli wipe s Before 1975 0 Figure 1. Location of European water-horehound (Lycopus europaeus) herbarium specimens collected in Québec before 1965, 1970. 1975. and 2000. respectively. Black dot: specimen found in herbarium; white dot: specimen collected during this study (August 1999). Subdivisions = municipal regional counties. PSI elopouy POL 19A] 2002] Lachance and Lavoie—Lycopus europaeus i535 CAN, DAO, MT, MTMG, QFA, QSA, QUE, and SFS (Index Herbariorum website: www.nybg.org/bsci/ih/ih.html). Each herbarium speci- men was checked for possible misidentification, and we noted the specimen number, collection location and year, and habitat char- acteristics. Data on specimens were incorporated into a geograph- ical information system to accurately reconstruct the evolution of European water-horehound’s distribution range during the 20th century in Québec. We also examined herbarium specimens of Lycopus americanus trom QFA (which has the biggest collection of L. europaeus and L. americanus from different locations in Québec) to find misidentified European water-horehound speci- mens. To accurately determine the northeastern distribution limit of the species in Québec in August 1999, we surveyed the south shore of the St. Lawrence River between Trois-Pistoles River (the known distribution limit) and Sainte-Anne-des-Monts (49°O8'N, 66°30'W), 300 km to the northeast. This region corresponds to the regional landscape unit of Rimouski (Robitaille and Saucier 1998). The mean annual temperature in the region is 2.5°C, and mean annual precipitation totals 900 mm. The salinity of the St. Lawrence River near Trois-Pistoles River and Sainte-Anne-des- Monts is 24%c and 27%c, respectively. Approximately 67% of the region is comprised of agricultural land (Bourget 1997; Robitaille and Saucier 1998). We did not survey the steep north shore of the St. Lawrence Estuary because there are very few suitable habitats (coastal marshes) for European water-horehound along this shoreline. Furthermore, surface currents, which are likely to disperse plant seeds, flow upstream near the north shore of the St. Lawrence River (Centre Saint-Laurent 1996). Consequently, an expansion of European water-horehound’s distribution range is unlikely to occur along the north shore of the St. Lawrence River Estuary. Between Trois-Pistoles River and Sainte-Anne-des-Monts, sampling points were chosen systematically every 5 km along the shore of the St. Lawrence River. At each sampling point, where access to the shore was possible, we surveyed the shore of the river (the supralittoral zone and the upper part of the intertidal zone) for a one-hour period to detect the presence of European water-horehound populations. Once a population was discovered, the following information was noted: |) the exact location of the population on the shore, 2) the number of individuals in the pop- 156 Rhodora [Vol. 104 ulation, and 3) any associated vascular plant species. Further- more, one or two specimens were collected for further identifi- cation in the laboratory. All collected specimens are stored in the Louis-Marie Herbarium (QFA) at Université Laval. All sampling points located within a 1!00-km distance from the last sampling point with a European water-horehound population were visited. Beyond the 100-km distance, only sampling points with habitats appropriate for the establishment of European water-horehound populations (1.e., marshes located in the supralittoral zone, and/ or in the upper part of the intertidal zone, or the mouth of fresh- water tributaries) were visited. RESULTS One hundred and ninety-nine (199) herbarium specimens (in- cluding those collected in this study) from 101 locations were carefully examined. Data from these specimens were used to ac- curately reconstruct the recent change of European water-hore- hound’s distribution in Québec (Figure |). We discovered that the oldest specimen was not sampled in 1964, but rather in 1963 on the south shore of Ile-des-Sceurs, near Montréal (45°26'N, 73°33'W). This specimen was originally misidentified as Lycopus americanus. Few L. europaeus specimens were collected before 1970; however, between 1970 and 1974, the range of European water-horehound expanded 380 km northeastward from Sorel to Trois-Pistoles River, which represents a very rapid rate of spread (95 km/yr.). Before this study, no specimen of European water-horehound had been collected northeast of Trois-Pistoles River. In August 1999, we discovered three populations beyond Trois-Pistoles Riv- er (Figure 2): 1. Trois-Pistoles Bay (48°07'N, 69°10'W) 2. Anse-des-Riou (48°09'N, 69°O7'W) 3. Anse-a-lOrignal (48°21'N, 68°46'W), in Bic Provincial Park These three populations were located 8, 12, and 65 km north- east of Trois-Pistoles River, respectively. These European water- horehound populations (10-15 individuals) were found in the su- pralittoral zone at the upper edge of Spartina alterniflora Loisel. marshes. They were also located very close to small freshwater n ag 2002] Lachance and Lavoie—Lycopus europaeus ! I 68°W 67°W Ste-Anne- des-Monts Matane Pri Se site with Euro water-horehound | Sampling site witho European water- ae Rimouski ity a ees ° Spartina alterniflora marsh 0 50 aa | Km Trois-Pistoles Figure 2. Sampling sites along the shore of the St. Lawrence River that were visited in August 1999 to determine the northeastern distribution limit of European water-horehound (Lycopus europaeus) in Québec, and location of Spartina alterniflora salt marshes in the area. Subdivisions = municipal regional counties. brooks, and on silt or clay soils. Populations were surrounded by Typha angustifolia L. or Lythrum salicaria communities. Be- tween 1974 and 1999, the rate of expansion of European water- horehound northeast of Trois-Pistoles River was only 3 km/yr. DISCUSSION Between 1963 and 1974, European water-horehound spread along the St. Lawrence River at a rate of 45 km/yr. During the last 25 years, the species spread from Trois-Pistoles River to Bic Provincial Park at a rate of only 3 km/yr. What could explain this difference? First, it is possible that the expansion of European water-horehound’s distribution range northeast of Trois-Pistoles River was limited by the increasing salinity of estuarine waters. For example, upstream from Saint-Jean-Port-Joli (Figure 1; 47°10'N, 70°15'W), where the salinity of the St. Lawrence River surface water is < 1—2%c, European water-horehound populations are located in the supralittoral and intertidal zones (Chrétien 1994; Gauthier 1977; numerous herbarium specimens). Down- 158 Rhodora [Vol. 104 stream from Saint-Jean-Port-Joli, the salinity of surface water in- creases rapidly (from I—2 to 15%e over a 60-km distance), and European water-horehound populations are located only in the supralittoral zone (i.e., outside the zone regularly flooded by brackish water tides; Bourget 1997). Second, herbarium speci- mens and the field survey conducted in 1999 suggest that in the estuarine part of the St. Lawrence River, European water-hore- hound populations are located only in large Spartina alterniflora marshes. These marshes are very small and scarce northeast of Rimouski (Figure 2; Centre Saint-Laurent 1996). Third, the re- construction of the spread of an invading species using herbarium specimens 1s highly dependent on the occurrence of field surveys conducted during different periods, and, in this case, on the ability of botanists to distinguish Lycopus europaeus in the field from the closely related L. americanus. For example, more than 20% of L. europaeus herbarium specimens that were examined in this study were originally misidentified as L. americanus, and were correctly identified only in 1978-1979 (most of them reviewed by J. Cayouette, DAO). However, no L. europaeus specimen was found in the L. americanus collection of QFA (206 specimens). Our reconstruction of the spread of European water-horehound in Québec should nevertheless be considered with some degree of caution. Whatever the exact rate of spread of European water-hore- hound along the St. Lawrence River, our data clearly indicate that the spread of this exotic species was particularly rapid in Québec. For example, the maximum rate of spread of another exotic wet- and plant species, Hydrocharis morsus-ranae L., was only 16 km/yr. between 1939 and 1994 within the Great Lakes—St. Law- rence River system. Seeds of H. morsus-ranae are dispersed by water, birds, and boats (Catling and Porebski 1995). No animal vector 1s known for European water-horehound, but the fact that seeds remain viable after floating for 12 to 15 months (Stuckey 1969) certainly facilitated the rapid spread of this species over long distances in Québec. The limited range expansion of European water-horehound in eastern Québec between 1974 and 1999 suggests that the salinity of surface waters, and more particularly the scarcity of coastal or riverine marshes east of Rimouski prevented populations from establishing in the estuarine part of the St. Lawrence River. How- ever, populations established upstream from Saint-Jean-Port-Joli —_— 2002] Lachance and Lavoie—Lycopus europaeus 159 seem to be expanding, and Lycopus europaeus may eventually replace the closely related native species L. americanus as one of the main species of the supralittoral and intertidal zones (Chré- tien 1994). European water-horehound is still absent from large tributaries of the St. Lawrence River, and its further expansion into these rivers should be attentively surveyed. ACKNOWLEDGMENTS. This research was financially supported (grants to C. Lavoie) by the Natural Sciences and Engineering Research Council of Canada. We thank M. Garneau, R. Gauthier, and C. Roy for field and laboratory assistance, and S. Pellerin and two anonymous reviewers for comments on an earlier draft. LITERATURE CITED BourcetT, E. 1997. Les Animaux Littoraux du Saint-Laurent. Presses de Univ. Laval, Québec, Canada. CaTLING, P. M. AND Z. S. PoREBSKI. 1995. The spread and current distribution of muropean Frogbit, paces morsus-ranae L., 1n North America. Canad. Field-Naturalist 109: CENTRE SAINT-LAURENT. 1996. Rapport re Synthese sur l’Etat du Saint-Laurent, Vol. 1: VP eae du Saint-Laurent. Environnement Canada, Montré- al, and Editions Multimo ndes, Québec, Cana CHRETIEN, L. 1994. Etude taxonomique d’une enceraialic des gréves d’eau douce du Saint-Laurent: Lycopus laurentianus. M.S. thesis, Univ. Laval, Québec, Canada. FLEURBEC. 1987. Plantes Sauvages des Lacs, Rivieres et Tourbieres. Fleurbec, Saint-Augustin, ate da. GALATOWITSCH, S. M., O. ANDERSON, AND P. D. ASCHER. 1999. Invasive- ness in wetland cae in temperate North America. Wetlands 19: 733— 7 GAUTHIER, B. 1977. Recherche des limites biologiques du Saint-Laurent (phy- ogéographie du littoral). Ph.D. dissertation, Univ. Laval, Québec, Can- ada. AND V. LAvolkE. 1975. Limites hydrobiologiques au niveau de I ana de Montmagny, estuaire du Saint-Laurent. Naturaliste Canad. —662. 102 EA N. C. 1962. A see one revision of the genus Lycopus (Labia- tae). Amer. ee Naturalist 68: 95-138. LONSDALE, W. M. 1993. Rates of ae of an invading species: Mimosa pigra in northern Australia. J. Ecol. 81: 513-521. PIMENTEL, D., L. LACH, R. ZUNIGA, AND D. Morrison. 2000. Environmental and omic costs of nonindigenous species in the United States. BioScience 50: 53-65 Pysek, P. AND K. PRACH. 1995. Invasion dynamics of /mpatiens oo A century of spreading reconstructed. Biol. Conservation 74: 41 160 Rhodora [Vol. 104 REJMANEK, M. AND D. M. RICHARDSON. 1996. What attributes make some plant species more invasive? Ecology 77: 1655—1¢ REZNICEK, A. A. AND P. M. CATLING. 1987. Carex praegracilis (Cyperaceae) in eastern North America: A remarkable case of rapid invasion. Rhodora 89: 205-216 ROBITAILLE, A. AND J.-P. SAUCIER. 1998. Paysages Régionaux du Québec Mé- ridional. Les Publications du Québec, Québec, Cand ROUSSEAU, C. 1968. Histoire, habitat et distribution de 120 lanes introduites au Québec. Naturaliste Canad. 95: 49-169. SCOGGAN, H. J. 1979. The Flora of Canada. Part 4. Dicotyledoneae (Loasa- ceae to Compositae). National Museums of Canada, Ottawa, Canada. STUCKEY, R. 1969. The ne aie and spread of Lycopus asper (western water honehonnd) in the western Lake Erie and Lake St. Clair region. Michigan Bot. 8: I11—120. AND W. L. Puitiips. 1970. Distributional history of Lycopus euro- paeus (European water-horehound) in North America. Rhodora 72: 35 1— 369 RHODORA, Vol. 104, No. 918, pp. 161-169, 2002 THE SURVIVAL OF VAUCHERIA (VAUCHERIACEAE) PROPAGULES IN NEW ENGLAND RIPARIAN SEDIMENTS AFTER REPEATED PREEZE/(THAW CYCLES DANIEL C. MCDEVIT AND CRAIG W. SCHNEIDER! Department of Biology, Trinity Colleg 300 Summit Street, Hartford, CT 06106- ar ‘Author to whom reprint request should be addressed nail: cre ug schneider. | @mail.trincoll.edu ACT. Previous studies have demonstrated the ability of the alga Vaucheria to survive prolonged and stressful periods of desiccation and freez- ing. However, even during harsh New England winters, the top few centi- meters of floodplain came or stream bank mud, where Vaucheria is often found, experience repeated thawing and refreezing events uds from two Connecticut riparian sites an to contain propagules from as many as eight species of Vaucheria were collected in spring and summer, then subjected to a varity of freeze/thaw ( ee Aan Six species of Vaucheria—V. aversa, V. frigida, V. geminata, V. , Vv. tavlorii, and V. undulata—have det a atin al eae to amigas F/T cycles of intervals from 1-10 days. Key Words: _ freeze/thaw cycles, propagules, riparian sediments, seed banks, Vaucheria In New England winters, the natural deep-freezing of subsur- face soil for up to four months is common. However, the upper surface of moist stream and riverbank alluvium above the water line rarely remains frozen for extended periods of time. On sunny winter days above O°C, the top few centimeters of mud will thaw, only to refreeze at night. At times, above-freezing winter tem- peratures are maintained long enough to thaw the mud for several days before refreezing. Organisms that live in the top few cen- timeters of mud are subject to these freeze/thaw (F/T) events and therefore must withstand such conditions to survive. The freshwater members of the cosmopolitan yellow-green alga Vaucheria (Vaucheriaceae, Tribophyceae, Chrysophyta) are commonly encountered living in and on riparian muds (Schneider et al. 1999). As the coenocytic siphons of Vaucheria grow, they weave in and out of their mud substrate and often between the siphons of sympatric species, forming what is referred to as a 16] 162 Rhodora [Vol. 104 **felt-like turf.” In order to survive the environmental extremes such as desiccation and freezing that impinge upon it, this alga, like so many others in their specific environments, has developed propagules as resting structures that are deposited in the flood- plain soils or riverbank muds. When environmental conditions are optimal, Vaucheria grows and reproduces forming zygotes (oospores). But when conditions are limited by abiotic compo- nents in the environment, the siphons of Vaucheria can form sporangia or “‘cyst-like” resting fragments (Dunphy et al. 2001). We have recently shown that the propagules of eight species—V. aversa Hassall, V. bursata (O. F Mill.) C. Agardh, V. frigida (Roth) C. Agardh, V. geminata (Vaucher) Alph. de Candolle, V. prona T. A. Chr., V. taylorii Blum, V. uncinata Kiitz., and V. undulata C. C. Jao—were able to survive desiccation in the lab- oratory from 63-383 days (Dunphy et al. 2001). The survival of each of these species was likely due to the deposition of a “seed bank” of propagules left in the mud during or after periods of active growth, and the physiological tolerance of these resting cells to prolonged periods without soil moisture. Since the freeze/ thaw phenomenon is so prevalent in Connecticut riparian muds and such an impoftant physiological stress for Vaucheria, this study examines the survival tolerance of the propagules of these species to repeated freezing and thawing for varying numbers of cycles. STUDY AREAS The two Connecticut collection sites where we have observed *“felt-like” turfs of Vaucheria with high levels of species sym- patry (Dunphy et al. 2001; Schneider et al. 1999) were selected for this study: 1. Nipmuck Trail (NK)—Ashford, Windham County, approx. 3 km from an entrance to the Nipmuck Trail, a portion of the Mohawk ‘Trail system, on Conn. Rt. 74 [41°S51.301'N, 72°12.821'W (Garmin® GPS 12, v. 4.57)]; Scantic River (SCR)—Enfield, Tolland County, floodplain di- rectly beneath the bridge on Conn. Rt. 190 that crosses over the Scantic River near the intersection with Conn. Rt. 19] (41°58.966'N, 72°30.969'W). Eight species of Vaucheria have been found at SCR—V. av- 2002] = McDevit and Schneider—Vaucheria propagules 163 ersa, V. bursata, V. frigida, V. geminata, V. prona, V. taylorii, V. uncinata, and V. undulata—while all of the above except V. taylorii are known from NK (Dunphy et al. 2001). MATERIALS AND METHODS Bulk samples of mud containing propagules from each site were collected on 24.iv.2000, 5.vii.2000, and 23.viii.2000. The procedures for field collection and preparation of a homogenous mud in the lab are described in Dunphy et al. (2001). After being left uncovered for five days, the moist mud slurry was cut into blocks approximately 36 cm’, placed in zippered plastic bags and frozen to O°C. Control mud samples were immediately placed in individual plastic culture dishes (2.3 cm * 8.5 cm) with Bold’s basal medium (Bischoff and Bold 1963), and cultured as in Dun- phy et al. (2001). Bags of mud totaling 33 blocks per collection site and date were removed from the freezer for each F/T experiment. The bags were placed in a growth chamber set at 15°C, and the mud was allowed to thaw for the specified length of time for each exper- iment. Thawing periods of 1—5, 7, and 10 days were used. Three blocks from each collection site were then removed from their bags and placed in individual culture dishes with 4—5 ml of cul- ture medium, labeled, and placed back in the growth chamber. The bags containing the remaining blocks were returned to the freezer for 2 days at which point the cycle was repeated until all of the mud blocks were cultured. Cultures were monitored for signs of Vaucheria siphons using light microscopy. Vouchers of reproductive materials from numer- ous samples were prepared (20% or 40% Karo™ corn syrup, 1% aqueous aniline blue, and | N HCL in a ratio of 20:1:1) and de- posited in Herbarium C. W. Schneider at Trinity College, Hartford. RESULTS Because the three replicate control dishes of mud collected from each site in April, July, and August produced similar spe- cies, survival data for NK and SCR were combined in Table 1. In the control dishes for NK, four species developed: Vaucheria aversa, V. geminata, V. prona, and V. undulata. In the controls for SCR, we discovered V. frigida, V. prona, and V. undulata 164 Rhodora [Vol. 104 Table 1. Freeze/thaw se survival of six Vaucheria species from two Con- necticut riparian sites (NK, SCR), and percent survival in total experimental dishes. Numbers represent . maximum number of F/T cycles a species survived at each site for specified number of thaw days per cycle. Species appearing in control dishes are denoted by asterisks (*). Culture ieee that never produced gametangia, hence remaining unidentified, are noted as V. spp. No. F/T Cycles Survived at Various No. Species Thaw Days/Cycle Occurrence Collection (% of Total Species Site | 2 3 4 5 7 10 Dishes) V. aversa NK* o = 8 4 5 7 4 24.4 SCR l ~ — 4 4 - | 0.05 V. frigida NK = - - - _ _ _ 0.00 SCR - 7 6 - 8 ~ 0.03 V. geminata NK* - - _ 7 2 - _ 0.01 SCR - 7 _ ~ — 3 2 0.02 V. prona NK* - 5 8 - 2 4 0.07 SCR* 9 8 6 4 7 6 5 26.7 V. taylorii NK _ — — — ~ ~ I 0.01 SCR - - _ _ ~ — I 0.01 Vo undulata = NK* — 5 - | ~ 2 0.04 SCR 2 7 6 4 = 5 5 0.07 V. spp. NK 10 9 a 5 7 6 | 12.2 SCR 3 9 8 6 9 ~ 6 15.4 (Table |). Several species appeared in experimental dishes from the two sites despite not appearing in control cultures: in fact, only V. prona and V. undulata were found in the control dishes from both sites. Vaucheria taylorii was found in neither control, yet appeared in one experimental dish from each site (Table 1). This represents the first report of V. tay/orii from NK, thus the same eight species are known from both collection sites. Prob- lems associated with assessing muds containing unknown quan- tities of propagules in the “‘seed banks” are discussed by Dunphy et al. (2001), but the low frequency of appearances of certain species in experimental dishes and the lack of the same species in the controls suggest small quantities of propagules in our col- lected muds. Each of the species found in the control dishes was able to survive multiple cycles of freezing, followed by one or more thaw days in at least some of the experimental dishes (Table 1). In some instances, the survival of a species after experimental treat- ment may in fact result from a single reproductive population in ee 2002] McDevit and Schneider—Vaucheria propagules 165 one of the three replicate dishes for each site. Combined, 45% of the 375 experimental dishes developed Vaucheria—42% of those from NK and 49% from SCR. However, in many of the experi- mental dishes, Vaucheria siphons never became reproductive even after months in culture, thus disallowing species identifica- tions in 12% of NK dishes and 15% of those from SCR. Only two species were found in greater than 1% of all of the experi- mental dishes from a site: V. aversa in 24% from NK, and V. prona in 27% from SCR. Vaucheria aversa appeared more fre- quently in culture dishes in the late winter to early spring re- gardless of collection time or experimental regimen, similar to findings for rehydrated desiccated muds containing V. aversa propagules from a previous study (Dunphy et al. 2001). The other four species were found at a much reduced frequency (Table 1). As noted above, V. taylorii appeared in only two cultures, one from SCR and the other from NK, the latter representing the first report from this site. Three species survived in most of the experimental treatments of 1-10 days thawing after 2 days of refreezing; Vaucheria aversa from NK, and V. prona and V. undulata from SCR (Table 1). Vaucheria prona was found growing in SCR dishes after 9 cycles with | day of thaw. For this species at SCR (the site and species that provided the greatest amount of data), as the number of thaw days increased in the trials, we observed that V. prona survived the greatest number of F/T cycles with the shortest thaw period (1 day; Table 1). The number of F/T cycles that V. aversa and V. undulata survived compared with the number of days thawed shows no obvious trend. The remaining species and sites had less complete survival data, no doubt due to their lesser presence in the ‘seed bank.’ Nevertheless, all six Vaucheria species showed tolerance to F/T stresses more extreme than we suspect they are exposed to at the sites from which they were collected, including the species with a lesser presence. Vaucheria frigida survived 8 cycles with 5 days of thaw (SCR), while V. taylorii (NK, SCR) and V. geminata (SCR) survived | and 2 cyles, respectively, with 10 days of thaw. DISCUSSION Few studies have been made on the effects of freezing and thawing on algae, although several have looked at the survival 166 Rhodora [Vol. 104 of bacteria, often in boreal and arctic soils. Skogland et al. (1988) discovered that a single F/T cycle could kill as many as 50% of a viable soil microbial population, and Schimel and Clein (1996) noted that following these environmental events, the dead cells contributed significant nutrients to the soil for surviving organ- isms. Other studies have focused on morphological or molecular and biochemical responses of cells disrupted by freezing and thawing, from bacteria and fungi to cereal crop protoplasts (Mor- ris et al. 1988; Steponkus et al. 1983). Although many studies have looked at the effects of prolonged freezing and cryopres- ervation in unicellular and filamentous algae (Ginsburger-Vogel et al. 1992; Morris 1978), little is known about their survivability after repeated cycles of freezing and thawing, conditions many stream and riparian algae are exposed to in their native environ- ments. In one study, Hawes (1990) observed that the vegetative cells of a filamentous green alga from Antarctic streams—an un- identified species of Zygnema—could survive the repeated freez- ing and thawing cycles typical of austral summers with little ef- fect, but that prolonged exposure to —20°C winter temperatures caused extensive cell mortality. He concluded that the few winter- surviving cells in filaments became the “‘seed’’ population for summer growth in Zygnema without the involvement of resting spores or other specialized structures typically utilized by fresh- water filamentous green algae to survive stressful environmental events (Coleman 1983). An organism such as Vaucheria, whose propagules can survive over a year in desiccated mud, would be expected to survive other environmental stresses normally encountered in its habitat, such as winter freezing in New England. We have observed that Vaucheria propagules, including all of the species tested herein for survival in repeated F/T cycles, survive in moist, freezing mud for over a year (unpubl. data). In the present study, the propagules of six species of Vaucheria have been shown to sur- vive the stress of multiple F/T cycles (Table 1), conditions these species might normally encounter in a typical Connecticut winter in the upper soil strata of riparian habitats. Vaucheria aversa, V. prona, and V. undulata were the most frequently encountered species in our experimental dishes, showing the greatest survival after repeated F/T cycles. Presumably, these species had left the greatest numbers of propagules in the “seed bank” in our col- lected muds. Despite being only sporadically found in our culture 2002] = McDevit and Schneider—Vaucheria propagules 167 dishes, the remaining three species, V. frigida, V. geminata, and V. taylorii, nevertheless survived experimental treatments in some of the dishes, showing their ability to survive repeated F/T cycles. Although V. bursata and V. uncinata were previously reported from both the NK and SCR sites (Dunphy et al. 2001), neither appeared in any of the control or experimental cultures. It 1s there- fore reasonable to assume that their propagules were not present in the mud collections made for this study. In this and past studies, we have seen Vaucheria siphons ap- pear above the substrate surface in as little as ten days after thaw- ing frozen muds. Propagule germination must therefore occur much earlier, within the first few days post thaw. The refreezing of muds that have been thawed for prolonged periods of time tests not only the ability of propagules to survive refreezing, but the ability of germinated siphons to survive as well. With ten- day thaw intervals, a large percentage of propagules will likely germinate and thus become susceptible to freezing injury with each F/T cycle. Without the cellular partitioning found in_ fila- mentous freshwater algae such as the chlorophyte Zygnema (Hawes 1990), the siphons of Vaucheria would appear to have fewer options for cellular protection and therefore be more sus- ceptible to mortality. If some percentage of the “‘seed bank’”” sur- vives after each F/T cycle, the species with the most numerous propagules should show the greatest success even if its ability to withstand the stress is no greater than any other species. There- fore, it would appear to be important for species to deposit a large number of propagules in the environment to have a greater chance of surviving F/T stress, as it appears that individual mortality must take its toll. Because V. aversa, V. prona, and V. undulata were commonly collected species in a great sampling of riparian Connecticut habitats (Schneider et al. 1999), and therefore could have deposited the most numerous propagules in our NK and SCR samples, it is not surprising that they showed greater success with longer thawing times than the other species (5 cycles with 10-day thaw intervals). Thus, they continue to appear to be eco- logically opportunistic, having already demonstrated survivability after long periods of desiccation—145, 359, and 383 days, re- spectively (Dunphy et al. 2001). Vaucheria prona and V. undulata survived the greatest number of F/T cycles (5) with thaw periods of 10 days. In Connecticut, the surfaces of floodplain alluvium or river banks would rarely, 168 Rhodora [Vol. 104 if ever, thaw once for ten continuous days and then refreeze dur- ing the winter freezing period of December to March. These two species, along with V. aversa and V. geminata, have shown great success in surviving multiple F/T cycles with longer thaw inter- vals. If they can survive such an extreme and repeated stress, unlikely to occur in their natural habitats, it seems probable they can survive any series of F/T cycles that would naturally occur in New England, assuming the propagules have not all germi- nated and died in the young siphonous form. The survival of all six Vaucheria species exposed to the stress of repeated freezing and thawing cycles further demonstrates the ability of this alga to survive severe environmental stress. Survival appears to de- pend not only on the species’ ability to physiologically handle the stress of repeated F/T cycles and the length of thaw intervals, but also upon the abundance of their propagules in a given hab- itat. Long thaw intervals more than likely allow the germination Vaucheria propagules, and it would appear young siphons would be more vulnerable to repetitive refreezing than resting propagules left ungerminated. ACKNOWLEDGEMENTS. We gratefully acknowledge a_ Trinity College summer undergraduate research assistantship to the first author, as well as field and lab assistance from Jonathan Stein. We thank Michael O’ Donnell and Brian Duval for helpful reviews of the manuscript. LITERATURE CITED BiscHorr, H. W. AND H. C. BoLb. 1963. Phycological studies IV. Some soil algae from Enchanted Rock and related algal species. Univ. Tex. Publ., 9318, Austi COLEMAN, A. W. 1983. The roles of resting spores and akinetes in chlorophyte survival, pp. 1-21. fn: G. A. Fryxell, ed., Survival Strategies of the Algae. Cambridge Univ. Press, Cambridge, U.K. Dunpuy, M. E., D.C. McDevir, C. E. LANE, AND C. W. SCHNEIDER. 2001. The survival of Vaucheria (Va age aoa in’ desiccated ew England riparian sediments. Rhodora 103: 416 GINSBURGER-VOGEL, TL, S. ARBAULT, AND R. PEREZ. 1992, Co study of the effect of freezing-thawing on the gametophyte of the brown alga Undaria pinnatifida. Aquaculture 106: 171-181. Hawes, I. 1990. Effects of freezing and ihawane on a species of Zygnema (Chlorophyta) from the Antarctic. Phycologia 29: 326-331 2002]. = MecDevit and Schneider—Vaucheria propagules 169 Morris, G. J. 1978. Cryopreservation of 250 strains of Chlorococcales by the method of 2-step cooling. Brit. Phycol. J. 13: 15-24. : SMITH, AND G. E. COULSON. 1988. A comparative study of the morphology of hyphae during freezing with the a = thawing of 20 species of fungi. J. Gen. Microbiol. 134: 2897— SCHIMEL, J. P. AND J. S. CLEIN. 1996. Microbial response to fates thaw cycles in tundra and taiga soils. Soil Biol. Biochem. 28: 1061—1066 SCHNEIDER, C. W., C. E. LANE, AND A. NoRLAND. 1999. The fr nates spe- cies 6 Vaucheria (Tribophyceae, Chrysophyta) from Connecticut. Rho- dora 101: 234-263. SKOGLAND, T., S. LOMELAND, AND J. GOKSOYR. 1988. Respiratory burst after freezing and thawing of soil: Experiments with soil bacteria. Soil Biol. Biochem. 20: Neteee STEPONKUS, P. L., M. F DowGrErT, AND W. J. GORDON-KAMM. 1983. Desta- bilization of . plasma membrane e isolated protoplasts during a -thaw cycle: The influence of cold acclimation. Cryobiology 20: 448—465. RHODORA, Vol. 104, No. 918, pp. 170-185, 2002 STUDIES IN NEOTROPICAL APOCYNACEAE I: A REVISION OF THE GENUS LAUBERTIA J. FRANCISCO MORALES Instituto Nacional de Biodiversidad (INBio), Santo Domingo de Heredia, Apto 22-3100, Costa Rica e-mail: forse Ou piace ABSTRACT. A synopsis of the three species of the genus Laubertia (Apo- cynaceae, Apocynoideae, Echiteae) is presented here. Keys, descriptions, dis- tributional data, and taxonomic index are provided. Key Words: Gentianales, Apocynaceae, Apocynoideae, Laubertia, Neo- The small genus Laubertia was described by Alph. de Candolle in 1844, with a single species, L. boissieri. It was characterized by a corolla tube having a conspicuous annular corona, but with- out free corona lobes within, and by eglandular sepals without basal colleters. Following the classification of Endress and Bruyns (2000), within the tribe Echiteae, Laubertia is closely related to Hylaea J. EF Morales and Prestonia R. Br. The three genera are characterized as having in common features such as corolla tubes with conspicuous annular coronas or with five free corona lobes within. However, Laubertia is easily distiguished by its eglandular sepals, without basal colleters within, and corolla tubes without free corona lobes within. Because of the confusion concerning generic limits in the family, the species of this genus were described or placed in other genera (e.g., Echites, Haemad- ictyon). Other workers, such as Miers (1878) or Hemsley (1881), reduced the genus to the synonymy of Exothostemon G. Don and Prestonia, respectively. In 1897, unaware of the main features of Laubertia, Greenman described the monotypic genus Strepto- trachelus. The next and most recent treatment of the genus was that of Woodson (1936). He considered Laubertia to comprise four species, reduced Streptotrachelus to synonymy, made two new combinations, and described one new species. Since then, there has been no comprehensive treatment of the genus. The description of several taxa in the last 60 years, new synonymy, and the necessity of taxonomic changes after the examination of types in European herbaria convinced me to update the genus. 170 2002] Morales—Revision of Laubertia 17] Specimens of Laubertia are usually rare in herbaria: of the three species here recognized, only two are known from more than one collection, while the third one is known from just the type collection. For this review, about 136 collections from 26 herbaria were examined. As a reference for infrageneric classifi- cation within the Apocynaceae, I used the work of Endress and Bruyns (2000). NOTEWORTHY MORPHOLOGICAL FEATURES The main morphological characters are described in the taxo- nomic treatment. However, several features that deserve more de- tailed commentary are described below. Leaves. The leaves of Laubertia are distinctive because of the presence of very diminutive cavities at the junction between the midvein and the secondary veins (Figure |). These structures are very similar to domatia, but are formed by the disconnection of the side vein from the lamiar tissue, thus they do not represent that feature exactly. These inconspicuous structures are not pre- sent in every vein axil and may be lacking in some leaves, but are totally absent in the related genera Hylaea and Prestonia. Domatia are foliar structures not very common in the family. In the neotropics they are present in several species of the genera Forsteronia G. May, Malouetia Alph. de Candolle, and Tintin- nabularia Woodson. Sepals. The sepals in Laubertia are eglandular (Figure 2), that is without basal colleters, while Hy/aea and Prestonia always have calycine colleters that are entire to variously lacerate. De- spite the fact that there are genera in other tribes of different subfamilies (e.g., Rauvolfioideae, Plumeriae) in which the sepals are variously glandular or eglandular (e.g., Al/amanda), in the subfamily Apocynoideae the presence or absence of calycine col- leters in the sepals is a very helpful feature for generic delimi- tation. Corolla and corona. The corolla tube in Laubertia lacks free corona lobes within (Figure 3c). In Prestonia, these lobes [called epistaminal appendages by Woodson (1936)] are present and ob- vious in most of the species, reduced to callus ridges in some Ly2 Rhodora [Vol. 104 =m Figure 1. Laubertia boissieri (Neill 10087, inp). Midvein, showing cav- ities in their junction with secondary veins. others, or are totally absent in just a few species. In Hylaea, the corona lobes are always present and totally exserted, but that genus lacks an annular corona, a character always present in Pres- tonia and Laubertia. TAXONOMIC TREATMENT Laubertia Alph. de Candolle, Prodr. 8: 486. 1844. Type: L. bois- siert Alph. de Candolle. Morales—Revision of Laubertia 2002] ® 3 on e 3° se". ee “SURE ate bate y 1) . . . : el 2.5 mm Figure 2. Laubertia boissieri (Neill 10087, inp). Calyx with eglandular sepals. 174 Rhodora [Vol. 104 6cm | mm Figure 3. Laubertia boissieri (Neill 1008/7, NB). A. Habit; B. Sepal; C. Corolla; D. Fruits; E. Seed. 2002] Morales—Revision of Laubertia [75 Echites P. Browne, Civ. Nat. Hist. Jamaica. 182. 1756, in part. Prestonia R. Br.. Mem. Wern. Soc. 69. 1809, in part. Haemadictyon Lindl., Trans. Hort. Soc. London 6: 70. 1825 (1826), in part. Exothostemon G. Don, Gen. Hist. 4: 70, 82. 1837, in part. Streptotrachelus Greenm., Proc. Amer. Acad. Arts 32: DO8. 1897. TYPE: S. pringlei Greenm. |= Laubertia contorta (M. Martens & Galeotti) Woodson]. Fruticose or suffruticose lianas. Stems terete to subterete, var- iously puberulent when young, usually glabrous to glabrate at maturity. Leaves opposite (very rarely ternate), petiolate, petioles slightly fused at the base, mostly glandular in the axils, with several inconspicuous and diminutive fusiform or conic colleters; blade glabrous, glabrate to very minutely puberulent, eglandular, without basal colleters adaxially. Inflorescence a scorpioid cyme, sometimes reduced and appearing simple or umbelliform, axil- lary, few- to many-flowered, glabrous or glabrate to minutely puberulent, pedunculate, bracts scarious, inconspicuous. Sepals 5, essentially equal, barely imbricate basally, without basal adaxial colleters within; corolla salverform, very minutely puberulent abaxially; tube straight to conspicuously twisted around the sta- mens, with an annular corona, without free corona lobes within, the limb 5-parted, actinomorphic, dextrorsely convolute; stamens 5, usually somewhat exserted, inserted in the upper part of the corolla tube; anthers connivent and adnate to the pistil head, con- sisting of 2 parallel, uniformly fertile thecae borne adaxially near the apex of an enlarged, peltate connective; auricles short, acute; carpels 2, united at the apex; pistil head fusiform or subcapitate; ovules numerous, multi-seriate, borne on an axile, biseriate pla- centa; disk glands 5, separate to very inconspicuously concrescent at the base, entire, distinct. Follicles 2, apocarpous, moniliform to more rarely continuous, glabrous, glabrate to very minutely puberulent, dehiscing along the ventral suture; seeds numerous, dry, truncate, comose apically, usually minutely rugose. The genus comprises three species: one found in México; the second in Guatemala and Belize; and the third in South America in Colombia, Ecuador, Peru, and Bolivia. KEY TO THE SPECIES OF LAUBERTIA 1. Corolla tube straight, not twisted around the stamens; plants 176 Rhodora [Vol. 104 from Colombia, Ecuador, Peru, and Bolivia ........ i Ga ae eg ae ee De ER 1. L. botssieri 1. Corolla tube twisted around the stamens; plants from México, CMinieimale ano BeNIZe o.cet co ee be eewwemes sence . Corolla purple to lilac, the tube 18-23 mm; anthers 6. a eo OO IOS IMIGKICO oon cena tateeoes 2. L. contorta Corolla white, the tube 10-14 mm long; anthers 5 mm long; Guatemala and Belize .......3. L. peninsularis 1. Laubertia boissieri Alph. de Candolle, Prodr. (DC.) 8: 487. 1844. TYPE: ECUADOR. Locality lacking, 1778-1788 (fl), Pa- von §.n. (LECTOTYPE selected here: G-BOIS!; ISOLECTOTYPES: F!, G-BOIS!, G-DC!, photograph Field negative 34137 at F!, INB!, MO!, NY ex G-DC!). Figure 3. Echites dichotoma Kunth in Humboldt et al., svi. nov., Nov. Gen. Sp. 217 (ed. qui), 9 Jul 1819 [1818], non Thunberg, 21 Apr 1819. Mesechites gees apa alg — cyn. S. Am. 233. 1878. TYPE: COLOMBIA. Vaupés nd Amazon River, Aug Cee asertes ds are 2 oe 3627 (HOLOTYPE: P-HB!, hotograph at INB! Echites sanctae-martae Racy: Descr. S. Amer. PI. 85. 1920. Laubertia sanctae-martae (Rusby) Woodson, sym. nov., Ann. Missouri 8: 555 i ot. Gard. 18: 555. l . TYPE: COLOMBIA. Maedalena: Above Jir- acasaca, 3000 ft., 25 Aus 1898-18 a r), Smith 2525 (HOLO- YPE: NY!; ISOTYPES: BR!. CM!, F!. G! i sheets], GH!. K! [2 sheets], MICHI. Mo!, P! [2 sheets], photocopy at INB ex BR!: photograph Field negative 56466 at INB ex F!). Echites eggersii Marker., Notizbl. Bot. Gart. Berlin-Dahlem 9: 78. 1924. PE: ECUADOR. Manabf: near El Recreo, 30 Apr 1897 (fl), Eggers 15684 =e OTYPE: B destroyed: LECTOTYPE selected here: 0!: ISOLEC- TOTYPES: C!. F!, K! [2 sheets], M! MO! NY!. OP! S!. photocopy at INB ex O!, photograph Field negative 56465 at INB ex F!). Liana; branchlets terete to subterete, very minutely and incon- spicuously brownish puberulent to ferrugineous-puberulent, gla- brous to glabrate at maturity; nodal colleters inconspicuous, ca. 1 mm long. Leaves usually opposite, very rarely ternate; petioles 5—17 mm long; blade 5.2—14 (16) X (1.5) 2—5.6 cm, membra- naceous to firmly membranaceous, elliptic to ovate-elliptic, very sparsely or minutely and inconspicuously puberulent when young, usually glabrous to glabrate on both surfaces at maturity, acuminate or short-acuminate to narrowly acute apically, obscure- ly cordate to more or less obtuse basally. Inflorescence conspic- uously longer than the subtending leaves, axillary, very minutely 2002] Morales—Revision of Laubertia Lay and inconspicuously ferrugineous puberulent to glabrate, few- to many-flowered; peduncle 22—80 mm long; pedicels 8-18 mm long; bracts 1-3 X 0.5—l mm, scarious. Sepals 2-6 X 1—1.5 mm, narrowly ovate to narrowly linear-ovate, long-acuminate, very minutely and sparsely ferrugineous puberulent to glabrate; corolla reddish-pink, or reddish-purple to purplish, very minutely brown- ish puberulent without, tube 12-27 x 3-5 mm, conspicuously inflated basally, straight, not twisted; lobes 10-18 xX 6—11 mm, narrowly obovate to narrowly elliptic, spreading; anthers 5-6 mm long, glabrous to very minutely puberulent dorsally; ovary |.5— 2 mm long, glabrous to glabrate; style head 2—2.5 mm long; disk glands about as long as the ovary. Follicles 25-75 xX 2—4 mm, glabrous or glabrate to very inconspicuously, minutely, and sparsely puberulent, obscurely moniliform; seeds 15—19 mm long, glabrous, glabrate, to minutely papillate, coma 2.5—4.8 cm long, creamish to tannish. DISTRIBUTION AND PHENOLOGY. The species is found in north- ern Colombia, southern Ecuador, Peru, and Bolivia at 200—1600 m elevation. It flowers and fruits all year, but mostly July to February. Laubertia boissieri is easily recognized by its straight corolla tube and distribution disjunct from the other two species of the genus. Echites dichotoma was included in the synonymy of Mesechi- tes trifida (Jacq.) Mill. Arg. by Woodson (1936). However, it is obvious that he never saw the type, because it is obviously con- specific with the type of Laubertia boissieri. Laubertia sanctae-martae is here relegated to the synonymy of L. boissieri. Woodson (1936) separated these taxa based on the inflorescence structure and sepal shape. At the time of his revi- sion, only five collections were available from these two species. Since then, many further collections reveal that sepal shape can vary from ovate and acute apically to very narrowly ovate and long acuminate. Corolla length is also very variable in the spec- imens examined and does not warrant the distinction of these taxa. Regarding inflorescence structure, Woodson cited “‘Inflores- cence rather obscurely compound to essentially simple” for ZL. sanctae-martae, however, the Brussels (BR) isotype, which was not examined by Woodson, shows a conspicuously compound inflorescence. Therefore, all supposed differential characters are 178 Rhodora [Vol. 104 ineffective, so L. sanctae-martae is relegated to the synonymy of L. boissiert. SPECIMENS EXAMINED: — BOLIVIA. La Paz: Inquisivi, Lakachaka, mouth of the Rio Aguilani, 21 Sep 1991 (fl), Lewis 40477 (LpB. Mo); NorYungas, Rio Un- duavi valley, 6 Sep 1987 (fl, fr), Serdel & Vargas 1/03 (LeB, Mo); Sud Yungas, E of Puente Villa, road to Chulumani, 28 Sep 1985 (fl, fr), Solomon & Nee 14272 (1NB, MO COLOMBIA. Magd 1948 (fl), Romero-Castanieda 762 (COL. 1898-1899 (fl), Smith 1643 (Gc). ECUADOR. Esmeraldas: de Bilsa, E of San José de Bilsa, 20 Jan 199] (f1), Gentry et al. 72942 (mo). Los Rios: Rio Palenque Science Center, be- tween Santo Domine and Quevedo, 16 Jul 1986 (fl, fr), Gentry & Dodson S4859 (mo [2 sheets], WAG). Napo: Jatun Sacha Biological Reserve, near Puerto Misahualli, 8 Nov 1987 (fl, fr), Cerén 2628 (mo, usr); Jatun Sacha Biological Station, Rio Napo, E of ae 17 Feb 1988 (f1), Cerén 3687 (mo, USF); Orellana, Pompeya, 5 fr), Neill [OOST (NB. MO, QCNE). Orellana: Yasunt National Park, 3 ae ca 8 - fr), Burnham 1794 (NB. MICH, 10). Province unknown: San José, Chimborazo, Jul 1876 (fl), André 4057 (« oy alena: flanco N de la Sierra Nevada de Santa Marta, 3 Mar Mo); Sierra Nevada de Santa Marta, — [2 sheets]). PERU. Cajamarca: San Ignacio, Chirinos, Mandinga, 5 Feb 1996 (fl, fr), Cam- pos & Diaz 24/2 (NB, Mo); San Ignacio, Huarango — San Martin, 15 May 1996 (fr), Vasquez & Vdsquez 20860 (NB, Mo); Pucara, 14 Apr 1960 (fl), Woytkowski S680 (G. Mo). Junin: Chanchamayo, La Merced — Villa Rica Road, between Puente Paucartambo and Rio Colorado, 6 Jan 1984 (fl, fr), Smith et al. 5625 (mo, usr); Yaupi, 23 Jun 1961 (fr), Woytkowski 6326a (mo), 30 Jun 1961 Cf, fr), Woytkhowski 6353 (mo [2 sheets}). San Martin: Muna, 23 May— 4 Jun 1923 (fl), Machride 3902 (hus); La Merced, Aug 1923 (fl, ft), Machride 5473 (EK, US). 2. Laubertia contorta (M. Martens & Galeotti) Woodson in Brit- ton, N. Amer. Fl. 29: 187. 1938. Figure 4. Haemadictyon contortum M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 11: 360. 1844. Exothostemon contortum (M. Martens & Galeotti) Miers, Apocyn. S. Am. 241. 187 a eel Sy Martens & Galeotti) Hemsl., Biol: Cent.-Amer., Bot. 2: 311. 18 TYPE: MEXICO. Oaxaca: Zacatepec, date ae (1), Galeotti 1588 (HOLOTYPE: BR!). Streptotrac ae pringlei Greenm., Proc. Amer. Acad. Arts 32: 298. 1897. Laubertia pringlei (Greenm.) Woodson, Ann. Missouri Bot. Gard. 18: 555. 1931. Type: Mexico. Morelos: lava beds near Cuer- navaca, 23 Sep 1896 (fl), Pringle 6554 (HOLOTYPE: GH! ISOTYPES: F!,G!. GH!. Kk! [2 sheets]. Mo! [2 sheets], Ny!. P!. S!). Prestonia fe oan Standl., Contr. U.S. Natl Herb. 23: 1159. 1924. YPE: MEXICO. Michoacan: La Correa, 50 m, 8 Oct 1898 (fl). Lang- lassé 435 je us!; ISOTYPES: G! [2 sheets], Git. K!. Pt). 2002] Morales—Revision of Laubertia 179 3cm 15cm Figure 4. Laubertia contorta (Martinez & Stevens 23849, inp). A. Habit; B. Corolla; C. Fruits; D. Seed 180 Rhodora [Vol. 104 Liana; branchlets terete to subterete, very inconspicuously to sparsely and minutely puberulent when young, glabrate at ma- turity; nodal colleters 1-2 mm long. Leaves opposite; petioles 10-32 mm long; blade 4-9 X* 2.5—5.7 cm, membranaceous, el- liptic or ovate-elliptic to narrowly ovate, very sparsely and in- conspicuously puberulent to more commonly glabrous or glabrate on both surfaces, acute to shortly and abruptly cuspidate to acu- minate apically, obtuse to inconspicuously or conspicuously cor- date basally. Inflorescence variously shorter or longer than the subtending leaves, axillary, densely and minutely puberulent, many-flowered; peduncle 25—65 mm long; pedicels 7-27 mm long; bracts |-1.5 * 0.5 mm, scarious. Sepals 3-5 * I-1.5 mm, narrowly ovate, acuminate to long-acuminate, densely and mi- nutely puberulent, corolla purple to lilac, moderately to sparsely puberulent without, tube 18-23 x 3—5 mm, conspicuously in- flated basally, twisted around the stamens; lobes 7-10 * 3.5—5 mm, narrowly obovate, spreading; anthers 6.5—7.5 mm long, mi- nutely puberulent dorsally, rarely glabrate, the tips exserted; ova- ry ca. 1.5 mm long, densely hirtellous; style head 2.5—3 mm long disk glands about as long as the ovary. Follicles 27-31 * 0.2— 0.4 cm long, very minutely and densely puberulent, moniliform; seeds 14-16 mm long, very minutely papillate puberulent, coma 2.7—3.5 cm long, tannish. DISTRIBUTION AND PHENOLOGY. Laubertia contorta is endemic to central and southern Mexico, at 50-1550 m elevation. It flowers from June to October. Fruiting collections are from September to December. This species is somewhat related to the South American Laub- ertia boissieri, from which it can be distinguished chiefly by the twisted corolla tube and its disjunct geographical distribution. SPECIMENS EXAMINED: MEXICO. Chiapas: along road from Tuxtla Gutiérrez to the Chicoasen, San Fernando, 9 Sep 1976 (fl), Breedlove 39960 (mo); E of Motozintla, road to Frontera Comalapa, Amatenango, 18 Sep 1988 (fl, fr), Martinez & Stevens 23849 (INB, MEXU); Tuxtla Gutiérrez, 5 Jul 1990 (f1), Reves et al. 1755 (BM, INB, MEXU). Guerrero: Temascaltepec, Ixtapan, 23 Jul 1932 (fl), Hinton 1156 (G,K,Mo); Chorrera, Temascaltepec, 24 Jun 1933 (fl), Hinton 458] (k); Naranjo, Temascaltepec, 17 Oct 1933 (fl, se Hinton 5009 2 sheets], Mo); Ixtapan, 24 Jun 1935 (fl), Hinton 7919 o); Placeres, Mina, 31 Jul 1936 (fl), Hinton 9/83 (K, ee Montes de Oca, | a 1937 (fl), Hinton 10544 («, Mo [2 sheets], TEX); Atoyac, Galeana, 12 Aug 1937 (fl), Hinton 11005 Gey: Petatlan, Acapulco — Zihuatanejo Road, 22 Oct 1983 (fl), Martinez 7 = Ba 2002] Morales—Revision of Laubertia 181 & Silva 5898 (INB, MEXU). Jalisco: Estacion Chamela, Arroyo Colorado, Aug 1985 (fl), Avala 1/5 (mMexu, uO: Estaci6n Biol6gica Chamela, 13 ae 1983 (fl), Lott & Herndndez 1484 (mexu. MO). México: near acre i NW of Iguala, 6 Jul 1982 (fl), Soto & rane 3969 (MEXU, MO). Nayarit: SW of Jests Maria, road to La Mesa del Nayar, 28 Jul 1990 (fl), Flores et al, 2/27 (MEXU, MO). Oaxaca: Chinantla, 1840 (fl), Galeotti 1596 (G), Galeotti 1600 (G, P); La Gritona, SW of Putla to Pinotepa Nacional, 5 Apr 1982 (fl), Torres & Tenorio 230 (MEXU, MO). San Luis Potosi: San Luis Potosi, Huasteca Potosina, date lacking (fr), Villa s.n. (clipIR, INB). Sinaloa: Concordia, Maza- tlan — Durango Road, 6 Dec 1982 (fr), Aguilar et al. 102 (INB, MEXU); Rosario, NE of Chilillos, 26 Jul 1983 (fl), Martinez et al. 4067 (mMEXxU, MO); Sierra Madre, near Colomas, Jul 1897 (fl), Rose 17/6 (mo, us). Data lacking: Sessé y Lacasta & Mogifio 5175 (ma, photograph Field negative 41244 at Np). 3. Laubertia peninsularis Woodson, Ann. Missouri Bot. Gard. 23: 374. 1936. TYPE: BELIZE. Undesignated locality near Belize— Guatemala boundary, date lacking (fl), Schipp s.n. (HOLO- TYPE: MO!). Figure 5. Laubertia gentlei Lundell, Wrightia 5: 256. 1976. TYPE: BELIZE. Toledo: Edwards Road beyond Columbia, 12 Apr 1948 (fl), Gentle 6505 (HOLOTYPE: TEX!; ISOTYPES: F!, MO!, S!, photograph Field negative 61421 at INB ex F'). Liana; branchlets terete to subterete, densely ferrugineous- tomentulose, sparsely puberulent at maturity; nodal colleters less than 1 mm long, inconspicuous. Leaves opposite, petioles 9-36 mm long; blade 4.2—11.5 (-13.2) X 1.5—4.7 (—6.8) cm, membra- naceous, elliptic or narrowly elliptic to narrowly ovate-elliptic, very sparsely puberulent above, densely ferrugineous-puberulent beneath, turning glabrate at maturity, acuminate to caudate-acu- minate apically, obtuse or rounded to very obscurely cordate ba- sally. Inflorescence variously shorter or longer than the subtend- ing leaves, axillary, densely and minutely ferrugineous-puberu- lent, many-flowered, the flowers agglomerate at ends of the branches; peduncle 41—60 (—155) mm long; pedicels 7—11 mm long; bracts 2-4 mm X 0.5—1 mm, scarious. Sepals 5—10 1.5— 2 mm, narrowly elliptic to linear, acuminate, ferrugineous-puber- ulent within and without. Corolla white, tube 10-14 <* 2 mm, conspicuously inflated basally, twisted around the stamens; lobes mm, narrowly obovate, spreading; anthers ca. 5 mm long, the tips exserted; ovary |.5—2 mm long, glabrous; style head ca. 2.5 mm long; disk glands somewhat shorter than the ovary. Follicles unknown. DISTRIBUTION AND PHENOLOGY. Known only from type collection, E 182 Rhodora [Vol. I¢ lcm lcm Figure 5. Laubertia peninsularis oo 6505, Mo). A. Habit; B. Corolla and calyx; C. Leaf pubescence (abaxia 2002] Morales—Revision of Laubertia 183 this species is restricted to eastern Guatemala and Western Belize, below 200 m. It flowers in April. This very distinctive species is poorly known and it has not been collected since the type collection. The main distinguishing character is the small corolla tube, which is twisted around the stamen attachment. The characters used to distinguish Laubertia gentlei Lundell from L. peninsularis are spurious, as was shown by Morales (1999). EXCLUDED SPECIES Laubertia laxiflora Rusby, Bull. New York Bot. Gard. 4: 408. 7. TYPE: BOLIVIA. Data lacking, Bang 2056 (HOLOTYPE: NY; ISOTYPES, NY, US, photocopy at INB ex NY) = Odontadenia lax- iflora (Rusby) Woodson. ACKNOWLEDGMENTS. I thank the curators and directors of BM, BR, C, CIIDIR, CM, COL, E G, G-BOIS, G-DC, GH, K, LPB, MEXU, MICH, MO, NY, O, P, P-HB, QCNE, S, US, and USF for providing specimens on loan. LITERATURE CITED ENpRESS, M. E. AND P. BRuyns. 2000. A revised classification of the Apo- cynaceae s./. 7 Bot. Rev. 66: 1-56. GREENMANN, J. M. 7. Descriptions of new and litthe known plants from exico, Proc. nee Acad. Arts 32: mee HEMSLEy, W. B. 1881. Apocynaceae, Vol. 2. /a: EF D. Godman and O. Slavin, eds., Biologia Centrali-americani, us London, U. Miers, J. 1878. On the Apocynaceae of South America. Williams and Nor- gate, pees K. Moraes, J. E 1999. Miscellaneous notes on Temnadenia and Laubertia ees Novon 9: 240. Woopson, R. E. 1936. Studies in the Apocynaceae. IV. The American genera of Echitoideae XX VI. Ann. Missouri Bot. Gard. 23: 169-438. APPENDIX | INDEX TO NAMES IN SYSTEMATIC TREATMENT Accepted names in italics. Echites Jacquin E. dichotoma Kunth (= L. boissieri) E. eggersii Marker. (= L. boissieri) 184 Rhodora [Vol. 104 E. sanctae-martae Rusby (= L. boissieri) Exothostemon G, Do E. contortum (M. arene & Galeotti) Miers (= L. contorta) Haemadictyon Alph. de Candolle H. contortum M. Martens & Galeotti (= L. contorta) Laubertia Alph. de Candolle L. boissieri Alph. de Candolle L. contorta (M. Martens & Galeotti) Woodson gentle: Lundell ae L. acne bs; Seba Wood: el (Greenm.) oe adson (=. age Sane ae: martae (Rusby) Woodson (= L. boissieri) Prestonia R. Br. P. contorta (M. Martens & Galeotti) Hemsl. (= L. contorta) P. langlassei Standl. (= L. contorta) enor Greenm. glei Greenm. (= L. contorta) — APPENDIX 2 INDEX TO EXSICCATAE Aguilar, R. et al. 102 (2) André, E. 4051 (1) Ayala, M. et al. 115 (2) Breedlove, D. 39960 (2) Burham, C. 1794 (1) eee . & O. Diaz 2412 (1) JC. os (1): 3687 (1) Se . 15684 (1) Flores, G. et al. 2127 ( Galeotti, H. G. 1588 (2): 1596 (2); 1600 (2) Gentle, . 6505 (3) Gentry, A. & C. Dodson as (1) Gentry, N et al. 72942 Hinton, G. 1156 (2); per ); 5009 (2); 7919 (2); 9183 (2); 10544 (2); 11005 (2) Humboldt, F W. H. A. & A. J. A. Bonpland, 3627 (1) Langlassé, E. 435 (2) Lewis, M. 40417 (1) ott, E. & R. Hernandez 1484 (2) Macbride, J. 3902 (1); 5473 (1) Martinez, E. & N. Silva 5898 (2) Martinez, E. & W. Stevens 23849 (2) Martinez, E. et al. 4067 (2) Neill, D. 1OO81 (1) Pavon, J. A. s.m. (1) — 2002] Morales—Revision of Laubertia Sits 6554 (2) Rey . et al. 1755 (2) ae oe R. 762 (1) Rose, J. | : Schipp, W. s.7 Seidel, R. & E. vargas 1103 (1) Sessé y Lacasta, M. & J. Mocino 5175 (2) Smith, D. N. et al. 5625 (1) Smith, H. 1643 (1); 2525 (1) Solomon, J. & M. Nee 14272 (1) Soto, J. & E. Martinez 3969 (2) Torres, E. & P. Tenorio 230 (2) Vasquez, R. & R. Vasquez 20860 (1) Villa, J. s.n. (2) Woytkowski, F 5680 (1); 6326a (1); 6353 (1) 185 RHODORA, Vol. 104, No. 918, pp. 186-200, 2002 STUDIES IN NEOTROPICAL APOCYNACEAE II: A REVIEW OF THE GENUS FERNALDIA J. FRANCISCO MORALES Instituto Nacional de Biodiversidad (INBio), Santo Domingo de Heredia, Apto 22-3100, Costa Rica e-mail: fmorales @inbio.ac.cr ABSTRACT. A synopsis of the three species of Fernaldia, a genus of Neo- tropical Apocynaceae restricted to Central America, is presented here. A sum- mary of descriptive morphology, specific relationships, and synonymy ts pro- vided. Key Words: Gentianales, Apocynaceae, Apocynoideae, Fernaldia, Neo- tropics Fernaldia Woodson is a genus of vines characterized by eglan- dular leaves that lack colleters along the midrib adaxially, sepals with a single colleter within, racemose inflorescence, corolla tube without annular corona or free corona lobes within, and corolla lobes usually villose adaxially. The genus was first proposed by Woodson in 1932 in honor of Merrit Lyndon Fernald (1873-— 1950), of the Gray Herbarium (GH), Harvard University. It occurs from Mexico to northern Panama. Despite their ornamental and edible features, these plants are rarely collected and very few herbarium specimens exist. Fernaldia was last treated by Wood- son (1936), when two species were known. Several new names for species or varieties have been published since then, but no recent revision exists. Therefore, a revision of the genus is pre- sented here. A key to the species, along with descriptions, illus- trations, and citations of selected specimens are given below. Specimens from St. Petersburg (formerly Leningrad), Russia (LE), cited here were examined at the Missouri Botanical Garden (Mo), where some material is on loan. NOTEWORTHY MORPHOLOGICAL FEATURES Sepals. In Fernaldia, the sepals are further solitary, truncate, and sometimes very deeply lacerate apically (Figure |). The se- pals are characterized by having a single colleter within adaxially. Within the subfamily Apocynoideae, the sepal colleters can be a very helpful character to distinguish genera. 186 2002 | Morales—Review of Fernaldia 187 2mm Figure 1. Fernaldia sepals. A. F. speciosissima (Morales 7131, inp); B. F. pandurata (Morales 3074, Np), C. FF. asperoglottis (Mexia 8751, 1x). Corolla pubescence. When Woodson described Fernaldia (1932), one of the features mentioned by him to distinguish the genus was the conspicuously arachnoid-villous corolla lobes. Af- ter careful field study of the three species of the genus, it is suitable to note that the pubescence is restricted mostly to the corolla throat and just at the base of the lobes. The hairs are long (2—3 mm), always conspicuous in fresh material, and invariably white to greenish-white. In related genera (e.g., Echites, Temna- denia), the upper part of the corolla tube and the mouth are gla- brous to glabrate, and never with long hairs. Following Morales (1999), in the key and species descriptions in this paper, the lower part of the corolla tube is measured from the base of the corolla up to the position where it is expanded abruptly (stamens attachment). The length of the upper part is measured from this point to the base of the lobes. Anthers. The anther shape in Fernaldia is a very important feature to recognize this genus from other closely related genera such as Echites and Temnadenia. In Fernaldia, the anther auricles are usually obtuse to almost rounded basally, while in the other genera the auricles are conspicuously acute to acuminate; only F. pandurata (Alph. de Candolle) Woodson has auricles very shortly and broadly acute. However, the most striking difference is the 188 Rhodora [Vol. 104 6 mm Figure 2. Fernaldia anthers. A. F. asperoglottis (Mexia S751, INB); B. F. pandurata (Morales 3074, INB); C. fF. speciosissima (Morales 7131, NB). presence of a hyaline border in the anthers of Fernaldia (Figure 2), a character never present in the three other related genera. TAXONOMIC TREATMENT Fernaldia Woodson, Ann. Missouri Bot. Gard. 19: 48. 1932.TyYPE: F. pandurata (Alph. de Candolle) Woodson. Echites P. Browne, Civ. Nat. Hist. Jamaica. 182. 1756, in part. Mandevilla Lindl., Edward's Bot. Reg. 26: t. 7. ahaa in mene nom. CONS. lea alee Mull. Arg., Fl. Bras. (Martius) 6(1): 1860, in part. Urechites Mill. Arg., Bot. Zeitung (Berlin) 18: 22. a in part. nena Miers, ee S. Am. 173. 1878, in part. Suffruticose lianas. Stems terete to subterete, glabrous or gla- brate to very minutely and variously puberulent; nodes with few intrapetiolar conical to conical-fusiform inconspicuous colleters. 2002] Morales—Review of Fernaldia 189 Leaves opposite, petiolate, petiole slightly fused at the base; blade glabrous or glabrate to variously puberulent beneath, eglandular, without basal colleters adaxially. Inflorescence racemose, axillary, usually many-flowered, very minutely puberulent to glabrous or glabrate, pedunculate, bracts scarious, inconspicuous. Sepals 5, essentially equal, very slightly imbricate basally, with a single colleter within, truncate, entire to variously and very minutely erose or fimbriate; corolla infundibuliform, glabrous or glabrate to very minutely and variously puberulent without, the lobes var- iously pubescent adaxially (very rarely almost glabrous in Fer- naldia pandurata) with long white hairs, sometimes these re- stricted mostly to base of the lobe or around the corolla mouth; tube straight, without annular corona or free corona lobes within, the limb 5-parted, actinomorphic, dextrorsely convolute; stamens 5, included, inserted in the upper part of the corolla tube; anthers connivent and adnate to the pistil head, conformed by 2 parallel, uniformly fertile thecae borne adaxially near the apex of an en- larged, peltate connective; auricles almost inconspicuous, broadly rounded basally; carpels 2, united at the apex; pistil head fusiform or subcapitate; ovules numerous, several-seriate, borne on an ax- ile, biseriate placenta; disk annular, usually 5-lobed. Follicles 2, apocarpous, continuous, glabrous to glabrate, dehiscing along the ventral suture; seeds numerous, dry, truncate, comose apically, usually minutely rugose. The genus comprises three species, ranging from Mexico to northern Panama. KEY TO THE SPECIES OF FERNALDIA 1. Lower part of the corolla tube 2—5 mm long; corolla lobes 8— [29 5 6 oases ee eee Rs 1. F. asperoglottis |. Lower part of the corolla tube 18—30 mm long; corolla lobes D4 A SIDI: 66st bdo ee end eae ewes Bae eX 2. Peduncles 2—4.5 cm; upper part of the corolla tube 10—15 mm long; anthers 5-6 mm long; dry forest ..... Tee eee eee See eee eee eens eee ee 2. F. pandurata 2. Peduncles 15—23 cm; upper part of the corolla tube 19—24 mm long; anthers | 1.5—12.5 mm long; wet forest .... Da F. SPECEOS ESS UTECE 1. Fernaldia asperoglottis Woodson, Ann. Missouri Bot. Gard. 26: 96. 1939. TYPE: MEXICO. Guerrero: Sierra Madre del Sur, N 190 Rhodora [Vol. 104 of Rio Balsas, 5 Nov 1937 (fl), Mexia 8757 (HOLOTYPE: MO!: ISOTYPES: ARIZ not seen, CAS!, F!, G!, GH!, NY!, photograph Field negative 56468 at INB ex F!). Figure 3. Liana; branchlets sparsely puberulent to glabrate; nodal colle- ters inconspicuous or absent, only intrapetiolar colleters present. Leaves: petiole (0.3—) 0.6—2.1 cm; blade |.8—-8 (—I1) X 0.8-6.8 cm, membranaceous, ovate, narrowly ovate to elliptic, caudate- acuminate at the apex, obtuse, attenuate to obscurely cordate ba- sally, glabrate above; densely puberulent to glabrate beneath, more rarely glabrous. Inflorescence lax, longer than the subtend- ing leaves, many-flowered, puberulent; peduncle 2—16 (—20) cm; pedicels 4—9 mm; bracts 1.5—2 * | mm, scarious; sepals I—1.5 x I—1.5 mm, ovate to narrowly ovate, acute, sparsely puberulent, colleters ca. 0.8 mm long, variously erose apically to conspicu- ously lacerate; corolla white to white-yellow, sparsely puberulent to glabrate without; lower part 2—5 * 2—3 mm; upper part broadly conical to conical-campanulate, 9-19 * 7-12 mm in diameter at the orifice; lobes 8-12 * 7—9 mm, obovate to narrowly-obovate, spreading; anthers 4.5-6 mm, glabrous; ovary |.5—2 mm, gla- brous; style head ca. 1.5 mmy; disk ca. 1.5 mm long, annular, irregularly lobed. Follicles 20-23 * 0.5—0.6 cm, smooth and gla- brate; seeds I—-1.2 * 0.2 cm, rugose, very minutely and incon- spicuously puberulent, coma 3—3.5 cm, creamish. DISTRIBUTION AND PHENOLOGY. This species ts restricted to Mexi- co, in Guerrero, Michoacan, and Mexico states, at 200—1350 m. Fernaldia asperoglottis flowers September to March. Fruits are borne from October to April. Fernaldia asperoglottis is distinguished from other species in the genus by having a conspicuously short corolla tube. Further- more, the corolla lobes are shorter than in any of the other spe- cies, only 8-12 mm long. This species may also resemble some species of the genus Mandevilla, more specifically the complex around M. convolvulacea (Alph. de Candolle) Hemsl. and M. an- drieuxtt (Mill. Arg.) Hemsl., with which it shares a similar leaf shape and inflorescence structure. However, F. asperoglottis 1s easily recognized by its eglandular leaves, without colleters along the midrib adaxially. SPECIMENS EXAMINED: MEXICO. Guerrero: Temascaltepec, Guayabal, 11 Feb 2002] Morales—Review of Fernaldia 19] 1.5 cm 4cm Figure 3. Fernaldia asperoglottis (Mexia 8751, Mo). A. Habit. B. Calyx and corolla. C. Fruits. D. Seed. 192 Rhodora [Vol. 104 1933 (fl, fr), Hinton 3372 (« [2 sheets]); Pungarabato, Coyuca, 23 Feb 1934 (fr), Hinton 5683 (« [2 sheets]); Temascaltepec, Guayabal, 16 Jan 1935 (fl, fr), Hinton 7239 (k [2 sheets]); Coyuca, Quebradas, 22 Jan 1935 (fr), Hinton 7261 (kK [2 sheets]); Placeres, Cigarillo, 11 Mar 1936 (fl. ey Hinton 9786 (kK); N of La Uni6n, road to Coahuayatla, 24 Oct 1983 (fl), Soto & Nun 6067 (MEXU, MO). Mexico: San Antonio Tlatlaya, 25 Jan (f1), ee et al. 28003 (MEXU, MO); between Sultepec and Amatepec, 31 Dec 1953 (fl), Matuda 30097 (MEXU. MO); Los Bejucos, Teyupilco, 27 Aug 1954 (fl), Matuda al. 31389 (MEXU. MO); Pete eo del Bravo, 5 Sep 1954 (f1), Matuda et al. 31416 (Mexu, MO); La Junta, Valle del Bravo, 11 Sep 1954 (fl), Matuda et al. 31649 (MEXU, oi: ene are Paso Tierra Caliente, 10 Mar 1938 (fl, fr), Hinton 13308 (K.Mo, Ny); Ajuage, Apatzingan, 13 Oct 1939 a Hinton 15329 (G. K. NY. P): Puente Las Pilas, road Zitacuaro-Huetamo, 3 Dec 983 (fl), Lott 2757 UNB. MEXU. MO): SW of La Huacana, 31 Dec 1977 (fr), Soto 592 (MEXU, MO); NW of La Eréndina, road to Casacuaro, 6 Sep 1981 (f1), Soto 3076 (MEXU, MO); San Jeronimo, road Huetamo San Jeronimo, 8 Oct 1981 (fl), Soto 3227 (MEXU. MO); Tumbiscatio, road Nueva Italia Playa Azul, 28 Oct 1981 (fl, fr), Soto 3586 (MEXU,. MO) 2. Fernaldia pandurata (Alph. de Candolle) Woodson, Ann. Mis- sourt Bot. Gard. 19: 48. 1932. Figure 4. Echites pandurata Alph. de Candolle, Prodr. (DC.) 8: 458. 1844. Am- ot aan pandurata (Alph. de Candolle) Mill. Arg., Linnaea 30: 448. 1860. Angadenia pandurata (Alph. de Candolle) Miers, Apo- cyn. ‘ Am. 182. 1878. TYPE: MEXICO. Oaxaca: San Dionicio, Aug 1832 (fl), Andrieux 245 (LECTOTYPE selected here: G-bc!; ISOLEC- TOTYPES: K!, photograph Field negative 7559 at F!. INB!, MO! NY! US ex G-DC!). Urechites karwinskii Miill. Arg., Linnaea 30: 440. 1860. TYPE: MEXICO. Tamaulipas?: ““Huefulta,” 1841-1842 (fl), Karnwinsky 474 (LECTO- TYPE selected here: Le!). Echites barbata Sessé & Moc., Naturaleza (Mexico City), Ser 2, 2 pp.): 45. 1893, non Desvaux ex Hamitten 1825, nec. D. Dietrich 1839. Type: Mexico: Data lacking (fl in August), Sessé y Lacasta & Mogino 5671 (HOLOTYPE: MA not seen; ISOTYPE: F!). a iMa velutina K. Schum. m Engl. & Prantl, Nat. Pflanzenfam. 2): 171. 1895. Type: costa rica. Data lacking (fl), Hoffmann 710 eee B-destroyed, photograph Field negative 4533 at F!, INB! MO!, US!); COSTA RICA. Guanacaste: Nandayure, Pacifico Norte, Be- juco, Cerro La ine, 24 Aug 1994 (fl), Estrada & Rodriguez 193 NEOTYPE selected here: INB!; ISONEOTYPES: CR!, M Mandevilla potosina Brandegee, Univ. Calif. Publ. ee 4: 276. 1912. Type: Mexico. San Luis Potosi: Rascon, Aug 1911 (fl, fr), Purpus 5408 (HOLOTYPE: UC!; ISOTYPES: F!, BM!, MO! Ny!, photograph Field negative 5 eX F!). Echites pinguifolia Standl. Publ. Field Columbian Mus., Bot. Ser. 8(1): 35. 1930. TYPE: Mexico. Yucatan: Izamal, 1895 (fl, fr), Gawmer 815 (HOLOTYPE: F!; ISOTYPE: MO!; photograph Field negative 56462 at INB ex BF!) 2002 | Morales—Review of Fernaldia 193 4cm Figure 4. Fernaldia pandurata (Morales 3074, inp). A. Habit; B. Fruits: C. Seed. 194 Rhodora [Vol. 104 Fernaldia brachypharyax Woodson, syn. noyv., Ann. Missouri Bot. Gard. 19: 380. 1932. Type: GUATEMALA. Along the road from Escuintla to the port of San José de Guatemala, 23 Aug 1860 (fl), Hayes s.n. (HOLOTYPE: GH!). Fernaldia pandurata var. glabra Ant. Molina, Ceiba 3: 95. 1952. Fer- naldia glabra (Ant. Molina) Lundell, Wrightia 5: 256. 1976. Type: HONDURAS. Cortés: faldas de la Montana Santa Ana, Rio Santa Ana, 6 Dec 1950 (1), Molina 3640 (HOLOTYPE: EAP!; ISOTYPES: F!, pho- tograph Field negative 56469 at INB ex F!). Liana; branchlets sparsely puberulent to glabrate; nodal colle- ters inconspicuous or absent, only intrapetiolar colleters present. Leaves: petiole 0.9—4 cm; blade 5—14 (17) * 4—11 cm, membra- naceous, ovate, ovate-elliptic, narrowly elliptic to variously pan- durate, acuminate to caudate-acuminate at the apex, rounded, ob- tuse to obscurely cordate basally, glabrous to glabrate above, densely puberulent to glabrate or more rarely glabrous beneath. Inflorescence agglomerate, usually shorter than the subtending leaves, rarely longer, few- to many-flowered, glabrate to very minutely and densely puberulent; peduncle 2—4.5 cm; pedicels 4— 12 mm; bracts |1.5—3 & I-—1.5 mm, scarious; sepals 1.5-3 X 1.5 mm, ovate to narrowly ovate, acute, very minutely and sparsely puberulent, colleters ca. 0.5 mm long, apex variously lacerate; corolla white to greenish-white or creamish, glabrous to glabrate or very sparsely and minutely puberulent to glabrate without: lower part 18-22 * 2-3 mm: upper part conical, more rarely broadly conical, 10-15 xX 8-11 mm in diameter at the orifice; lobes 9-14 * 9-11 mm, obovate to narrowly-obovate, spreading and distally reflexed: anthers 5-6 mm, glabrous to glabrate, rarely minutely puberulent; ovary 1.5—2 mm, glabrous; style head ca. 2.5 mm; disk I—1I.5 mm long, 5-lobed to variously lobed. Folli- cles 21-36 X 0.5—0.6 cm, rugose, glabrous to glabrate; seeds 1.4— 1.6 X 0.2 cm, minutely rugose, coma 4—4.5 cm, creamish. DISTRIBUTION, PHENOLOGY, AND LOCAL NAMES. México to north- western Costa Rica, mostly in dry forest or in open and second- growth forest, O-1200 m. Flowering June to December. Fruiting March to January. Known locally as Loroco (Jutiapa, Guatemala; Cortez, Honduras; Ahuachapan, El Salvador). The available specimens of Fernaldia pandurata, the most common and widespread species, show that leaf shape and pu- bescence are morphological features that are highly variable with- in this taxon. Although there are slight differences scattered 2002] Morales—Review of Fernaldia 195 throughout the plant’s geographic range, they are mostly sporadic in ocurrence and do not merit taxonomic recognition. Within the Apocynaceae subfamily Apocynoideae, the acceptance of species defined only by pubescence is unwarranted (Morales 1997, 1999), Therefore, /. pandurata var. glabra and the subsequent combi- nation based on this name are not recognized here Fernaldia brachypharynx is here considered a synonym of F. pandurata, showing only slight differences in corolla length. Ad- ditional specimens examined since Woodson’s monograph reveal that this feature is variable within the geographical range. In northern Mesoamerica, the flowers of this species have been used as a flavoring for rice (Woodson 1936). For further refer- ences, see the work of Morton et al. (1990). Mandevilla velutina K. Schum. is neotyfied here. No other du- plicates were found in BM, Cc, or K, where Hoffmann specimens are located. SPECIMENS EXAMINED: COSTA RICA. Guanacaste: Canas, La Pacifica, 10 Nov 1969 (fl, fr), Daubenmire 256 (& usi), 2 Mar 1970 (fl, fr), Daubenmire 592 USD; pines La Pacifica, 3 Sep 1972 (fl), Heithaus 350 (mo); Parque pier Palo Verde, 30 July 1994 (fl, fr), Morales 3074 (NB); La Pacifica, NW Canas, 20 Nov 1972 (fl), Opler 1568 (cr. & Mo [2 sheets]); Palo Verde nae Park, 11 Dec 1996 (fl, fr), Redrrguez et al. 1529 (NB, MO); Palo Verde, 7 Sep 1973 (fl, fr), Solomon 62] (CR, E USI). / SALVADOR. Ahuachapan: San Benito, E of San Alfonso, El Imposible, | Jun 1993 (fl), Sandoval & Sandoval 1309 (B, LAGU, MO); EI se posible National Park, . Alfonso, 10 Jul 1990 bee Sermeno 232 (B. LAGU, GUATEMALA. Chiquimula: between Ramirez and Cumbre de ‘errata. 15 Oct 1940 (fl), Standley 74496 MO). Jutiapa: vicinity of Jutiapa, 1940 (f1), Standley 75297 (& Mo). Petén: exact locality lacking, 12 Aug 1967 (fl), Con- treras 7008 (k, LL). Santa Rosa: vicinity of Oa. 1940 (fr), Standley 79683 (EMO). ONDURAS. Cortés: Santa Ana mountain, Rio Santa Ana, 6 Dec 1950 (fl ate 3640 (BM. EAP. MO). Morazan: Villa San Roque, Sep 1948 (fl), Lee Se 26257 (BM, F). MEXICO. Chiapas: near Chiapilla, 14 Nov 1980 (f1), iat 47493 (MO); along the road from Acala to Venustiano Carranza, 25 Oct 1966 (fl), Laughlin 2669 (Mo); Ocozocoautla, 19 Sep 1988 a Reyes & Urquijo 1015 (BM,MEXU Guerrero: Sierra Madre. 8 Nov 1898 (fl. fr), Langlassé 597 (G [3 sheets]. p); Acapulco, Oct 1894 — Mar 1895 (fl), ie 259 («). Jalisco: Tonala, ia Cruz, Barranca de la Cruz, 9 Jan 1975 (fr), Diaz 5484 (eNcB, INB). Nayarit: SE of Ahuacatlan, 2 Jan 1986 (fr), Téllez 9371 (NB. MEXU). Oaxaca: exact locality lacking, 1834 (fl), Andrieux 246 (G-bc. kK); W of Tuxtepec, along road to Ixcatlan, 7 Aug 1971 (fl), Stevens 7397 (mo). San Luis Potosf: Tamazun- chale, 7 Oct 1937 (fl), Taylor 479 (Mo, TEX) Aeracialions: Sierra de Tamauli- _~ — 196 Rhodora [Vol. 104 pas, region of Rancho Las Yucas, NNW of Aldama, 27 Jul 1957 (f), eet 2037 (Mo). Veracruz: Laguna Encantada, NE of San Andrés Tuxtla, 2 Nov 1971 (fr), Beaman 5225 (MeExu, MO); El Salto de Eyiplanta, near Sih, 9 Oct 1974 (fl), Calzada 1567 (mo): Bafios del Carrizal, Aug 1912 (fl), Purpus 6020 (BM. MO); Banos del Carrizal, Aug 1912 (fl), Purpus 6232 (Mo). State unknown: Boca del Monte, date lacking (fl), Andrieux s.n (G); 1833 (fl), Andrieux 399 (G. kK). Data lacking: (f1), Coulter 958 (« [2 sheets]); Sessé y Lacasta & Mogino SO80, (MA: photograph Field negative 41240 at INB). NICARAGUA. Boaco: San José de los Remates, N of Teustepe, Cerro Alegre, 10 Oct 1982 (fl), Sandino 3680 (mo). Chinandega: along road Somotillo — Cinco Pinos, 11 Oct 1993 (fr) “Riteda & Dolmus 1170 (mo); Volcan San Cristobal, N of Chinandega, 23 Aug 1984 (fl), Soza & Grija ba 166 (MO). Esteli: San Juan de Limay, Valle La Cascada, 1 Sep 1980 (fl), Moreno 1893 (Mo); Paso Leén a Estelf, 23 Oct 1983 (fl), Moreno 22327 (mo). Leén: La Paz Centro, road to Momotombo, 13 Jul 1981 (fl), Moreno 9834 (mo): along Rio Sinecapa, 15 Sep 1977 (fl), Stevens 3565 (mo), S of Estelr, road to Es- tanzuela, 11 Aug 1978 (fl), Stevens 9960 (mo). Matagalpa: Rancheria, NE of Muy Muy, 20 Aug 1984 (fl), Moreno 24434 (mo). Nueva Segovia: N of edge of Ocotal, Quebrada El Nancital, 7 Aug 1977 (fl), Stevens 3057 (BM. MO). Rivas: Isla Ometepe, Volcan Concepcion, San José del Sur, 12 Dec 1984 (ff), has /566 (mo), SE of San Juan del Sur, NW of Rio La Flor, Playa El “oco, Il Sep 1977 (fl), Stevens 3865 (BM, MO). jae 3. Fernaldia speciosissima Woodson, Ann. Missouri Bot. Gard. 26: 300. 1939. TyPE: PANAMA. Chiriqut: Rfo Chiriquf to Re- medios, 11 Jul 1938 (fl), Woodson et al. 1179 (HOLOTYPE: MO!, photograph at INB!). Figure 5. Liana; branchlets glabrous; nodal colleters inconspicuous, ca. 1 mm long. Leaves: petiole 2.5—6 cm; blade 9.5-14 * 4-10 cm, membranaceous, elliptic to broadly elliptic, glabrous, shortly acu- minate to caudate-acuminate at the apex, obtuse to rounded ba- sally. Inflorescence lax, longer than the subtending leaves, many- flowered, glabrous; peduncle 15-23 cm; pedicels 10-18 mm; bracts 1.5—2.5 * | mm, scarious; sepals 3-5 * 1.5—2 mm, ovate, acute to obtuse, glabrous; colleters ca. | mm long, apex scarcely and very minutely fimbriate; corolla creamish to creamish-white, glabrous; lower part 20—30 * 1.5—2 mm; upper part 20-30 X 6-— 8 mm in diameter at the orifice; lobes 19-24 * 1O—15 mm, ob- ovate to narrowly-obovate, spreading and somewhat reflexed; an- thers 11.5-12.5 mm, glabrous; ovary 3 mm long, glabrous; style head ca. 3 mm; disk ca. | mm long, inconpicuously 5-lobed. Follicles 33-34 * 0.6—0.7 cm, smooth, glabrous to glabrate; seeds unknown. DISTRIBUTION, HABITAT, AND PHENOLOGY. This species 1s restricted 2002] Morales—Review of Fernaldia 197 3cm 4 cm 3 mm re 5. Fernaldia er Ss oe 71317, INB). A. Habit; B. Co- ee ie C. Ovary and disk . Fruit 198 Rhodora [Vol. 104 to wet forest or seasonal wet forest in southwestern Costa Rica and northwestern Panama, at 90-600 m. Until recent flowering collections from Costa Rica were made, the species was known only from the type collection. Fernaldia speciosissima flowers in July and fruits from December to January. Fernaldia speciosissima was described by Woodson based on just three fallen corollas found in Chiriqui, Panama in 1938. Al- though he was exasperated to base a new species on such limited material, Woodson (1939) said that “‘The anthers, stigma and arachnoid internal villosity of the corolla are all unmistakable characters,” which was nicely confirmed with the Costa Rican flowering material. The flowers are very fragant, with a smell similar to crushed fig leaves (Ficus carica L., Moraceae), and upon tasting they produce a very sweet flavor. However, none of the local people from the collection locality know of any use for the plant. SPECIMENS EXAMINED: COSTA RICA. San José: Acosta, Fila Aguabuena, Rio Tiquires, on road to Zoncuano, 11 Jul 1999 (fl), Morales 7131 (CR. INB. Mo. K). PANAMA. Panama: SE side of Madden Lake, near Puente Natural, | Jan 1975 (fr), Nee & Hansen 14056 (mo ACKNOWLEDGMENTS. — I thank the curators and directors of ARIZ, BM, BR, C, CHDIR, CM, CR, ENCB, FE G, G-BOIS, G-DC, GH, K, MEXU, MICH, MO, NY, O, P. P-HB, S, US, USE USJ, and UVAL for providing specimens on loan. LITERATURE CITED MorAtes, J. F 1997, = synopsis of the genus Macropharynx (Apocynaceae). Rhodora 7 252-262. . 1999. A synopsis of the genus Odontadenia (Apocynaceae). Jn: A. J. M. Leeuwenberg, ed., Series of Revisions of Apocynaceae XLV. Bull. 3 Morton, J. ev AL. 1990. Loroco, Fernaldia pandurata (A. DC.) pica ar A popular edible flower of Central America. Econ. Bo 44: 310 re - E. 1932 . New or otherwise Seis Apocynaceae of Trop- ical America II. A in. Missouri Bot. Gard. I¢ —76. 936. Sai in the Apocynaceae. [IV The es genera of Echi- ies XXV . Missouri Bot. Gard. 169-438. 1939. aoe or iene noteworthy oe naceae of Tropical Amer- ica VIL Ann. Missouri Bot. Gard. 26: 95-98. 2002] Morales—Review of Fernaldia 199 APPENDIX | INDEX TO NAMES IN SYSTEMATIC TREATMENT Accepted names in italics. Amblyanthera A. pandurata (Alph. de Candolle) Mill. Arg. (= F. pandurata) Angadenia A. pandurata (Alph. de Candolle) Miers (= F. pandurata) Echites E. barbata Sessé & Moc. (= F. pandurata) E. pandurata Alph. de Candolle (= fF. pandurata) E. pinguifolia Kunth Standl. (= F. pandurata) — Fernaldia e. asperoglottis Woodson F. brachypharynx Woodson (= F. pandurata) F. pandurata (Alph. de Candolle) Woodson F. speciosissima Woodson Mandevilla +8 M. potosina Brandegee (= F. pandurata) M. velutina K. Schum. (= F. pandurata) Urechites U. karwinskii Mill. Arg. (= F. pandurata) APPENDIX 2 INDEX TO EXSICCATAE Andrieux, G. s.n. (2); 245 (2): 246 (2); 399 (2) Beaman, J. 5225 (2) Breedlove, D. 47493 (2) Calzada, J. 1567 (2) Contreras, E. 7008 (2) Coulter, E. 958 (2) Daubenmire, FE 256 (2); 592 (2) Diaz, C. 5484 (2) Dressler, R. 2037 (2) Estrada, A. & A. Rodriguez 193 (2) Gaumer, G. 815 (2) Hayes, S. s.n. (2) Heithaus, E. 350 (2) Hinton, G. 3372 (1); 5683 (1): 7239 (1): 7261 (1); 9786 (1); 13308 (1); 15329 (1) Hoffman, C. 710 (2) Karwinsky, W. F 474 (2) 200 Rhodora [Vol. Langlassé, E. 597 (2) ae N R. 2669 (2) , E. 2157 (1) ae. E. 30097 (1) Matuda, E. et al. 28003 (1); 31389 (1); 31416 (1); 31649 (1) Mexia, Y. 8751 (1) Molina, A. 3640 (2) Morales, J. EF 3074 (2): 7131 (3) Moreno, P. 1893 (2); 9834 (2); 22321 (2): 24434 (2) .M. & B. Hansen 14056 (3) Opler, P. 1568 (2) Palmer, E. 259 (2) Purpus, C. 5408 (2); 6020 (2); 6232 (2) Reyes, A. & G. Urquijo 1015 (2) Robleto, W. 1566 (2) Rodriguez, A. et al. 1829 Rueda, R. & R. Dolmus 1170 (2) Sandino, J. 3680 (2) Sandoval, E. & M. Sandoval 1309 (2) Sermeno, A. (2) Sessé y Lacasta, M. & J. Mocino 5080 (2), 5671 (2) sie be J. 611 (2) Soto, J. 592 (1): 3016 (1): a. (1); 3586 (1) Soto, , & - Nunez 6067 ( Soz A. Grijalva 166 i: ee P. 26257 (2): 74496 a 75297 (2); tas (2 Stevens, D. 1391 (2); 3057 (2): 3865 (2): 9960 ( Taylor, M. 749 (2) Téllez, O. 9371 (2) Woodson, R. et al. 1179 (3) 104 RHODORA, Vol. 104, No. 918, pp. 201-204, 2002 NEW ENGLAND NOTE A NEW NATIVE PLANT FOR MASSACHUSETTS, CAREX BACKII (CYPERACEAE) ROBERT I. BERTIN Biology Department, College of the Holy Cross, Worcester, MA 01610 -matl: rbertin@holycross.edu KAREN B. SEARCY Biology Department, University of Massachusetts, Amherst, MA 01003 PAUL SOMERS Natural Heritage and Endangered Species Program, Division of Fisheries and Wildlife, Westborough, MA 01581 Carex backii Boott is one of two members of the section Phyl- lostachyae of the genus Carex found in New England. It is dis- tinguished from the other species, C. willdenowti Schkuhr ex Willd., by having a lower pistillate scale that is wider than the perigynia and concealing them (Catling et al. 1993). It occurs from the Gaspé Penninsula, Québec south through New England, and west to British Columbia, Wyoming, and Colorado (Saarela and Ford 2001). It formerly occurred but has not been found recently in New Jersey and Pennsylvania (Kartesz and Meacham 1999), and appears to have a patchy distribution in its current range. It is uncommon in New England, previously having been reported from Maine, New Hampshire, Vermont, and Connecti- cut. It is most common in Vermont, with a ranking of S3, cor- responding to 21—100 occurrences. Its rank is undetermined in New Hampshire (where it 1s being reviewed for state listing), and S| and Endangered in both Connecticut and Maine, with one and several occurrences, respectively (Connecticut Department of En- vironmental Protection 1998; Maine Department of Conservation 1999; A. Haines, New England Wildflower Society, pers. comm.). The species 1s a new addition to the native flora of Massachusetts, Where it is listed as Endangered. This note reports on the two known Massachusetts occurrences. One population was discovered in 1997 in open woods on an east-facing slope of Wachusett Mountain in Worcester County 201 DOF Rhodora [Vol. 104 (Bertin 1587, 24 Jun 1997, MAss). The population consisted of about 42 clumps, each containing multiple shoots of this “tufted” (Gleason and Cronquist 1991) species. The shallow rocky soil overlies bedrock mapped as biotite granodiorite to tonalite gneiss (Zen 1983). The tree canopy consisted of Fraxinus americana L. and Quercus rubra L., with a few Carya ovata (Mill.) K. Koch. The shrub layer included Acer pensylvanicum L., Crataegus sp., Q. rubra, Prunus serotina Ehrh., P. virginiana L., and Betula lenta L. The herb layer was relatively dense, and was dominated by Polygonum cilinode Michx. Other herbaceous species included Carex communis Bailey, C. pensylvanica Lam., Deschampsia flexuosa (L.) Trin., Parthenocissus quinquefolia (L.) Planch., Fes- tuca Subverticillata (Pers.) E. B. Alexeev, Elymus hystrix L., Cir- caea lutetiana L., Maianthemum racemosum (L.) Link, Poa sp., and Viola sp. The second population was discovered during 2001 in the Hol- yoke Range in Hampshire County (Searcy 403, 19 Jun 2001, MASS). Approximately 18 widely separated clumps, each sup- porting 4—38 culms, were found near the summit of Long Moun- tain in shallow soil on a steep north-northeast-facing slope of the basalt ridge that makes up the crest of the range. Based on tests conducted by the Soil Testing Lab at the University of Massa- chusetts, the pH of the A horizon in these soils was 4.7—5.0. Calcium concentrations were high (ca. 2200 ppm), at least an order of magnitude higher than in soils overlying nearby sedi- mentary rock. As with the first population, the forest canopy was relatively open. Woody species included Betula lenta, B. papyr- ifera Marshall, Acer rubrum L., Ostrya virginiana (Muill.) K. Koch, Carya glabra (Mill.) Sweet, Tsuga canadensis (L.) Carri- ere, Hamamelis virginiana L., and Viburnum acerifolium L. Con- spicuous herbs included Dryopteris marginalis (L.) A. Gray, Par- thenocissus quinquefolia, Carex pensylvanica, and one or more Carex in the section Laxiflorae. Carex backii is sometimes considered a calciphile (Scoggan 1950; M. Lapin, consulting ecologist, pers. comm.). The one Con- necticut site 1s a marble ridge in Canaan, Litchfield County (Mehrhoff 1995; T. Rawinski, Massachusetts Audubon Society, pers. comm.). Many of the Vermont records are from soils derived from limestone, dolomite, or other calcareous rocks (M. Lapin, pers. comm.; T. Rawinski, pers. comm.). Maine occurrences seem to span a wider range of soil types. Dibble (1993) reported the 2002] New England Note 203 species from a rocky bluff with oak-hornbeam forest along with Hepatica nobilis Mill. A second Maine location also supports associates that suggest non-acid conditions [e.g., Carex platy- phylla Carey, Woodsia obtusa (Spreng.) Torr., Aquilegia cana- densis L., Ranunculus fascicularis Muhl. ex Bigelow, Arabis mis- souriensis Greene; Rawinski, pers. comm.]. However, other Maine sites are in red oak-northern hardwoods forests on appar- ently acid soils (Haines, pers. comm.). Neither Massachusetts site is basic, though they may be less acid than most Massachusetts soils. We have no information on the Princeton soil type, though areas within several hundred meters downslope support Adiantum pedatum L., Sanguinaria canadensis L., Actaea rubra (Aiton) Willd., Geranium robertianum L., and Caulophyllum thalictro- ides (L.) Michx. The New Hampshire site supports a soil with pH of 6-7 on calcite-rich diorite/granodiorite. Associated species include Cypripedium calceolus L., Carex platyphylla, Cynoglos- sum virginianum L., and Dryopteris goldiana (Hook. ex Goldie) A. Gray (E. B. Engstrom, consulting ecologist, pers. comm.). Carex backti is a relatively inconspicuous plant and rarely seems to occur in extensive populations. This 1s reflected in the fact that although both Massachusetts and Connecticut are well botanized, the first records from these states are from the last 15 years. Several of the Maine and Vermont records were also added during this period. It seems likely that additional populations of the species occur in New England, and further botanizing on neu- tral and alkaline soils during the late June to early July fruiting period will reveal some of these. ACKNOWLEDGMENTS. We thank Lois Somers and Ann E. Sil- veri for help with field observations and the following individuals for sharing information on occurrences of Carex backii in other states: E. Brett Engstrom, Mare Lapin, Les Mehrhoff, Bill Nich- ols, Tom Rawinski, and Dan Sperduto. Two anonymous reviewers made helpful suggestions on the manuscript. LITERATURE CITED CATLING, P. M., A. A. REZNICEK, AND W. J. CRINS. 1993. Carex juniperorum (Cyperaceae), a new species from northeastern Dae a with a key to Carex sect. Phyllostachys. Syst. Bot. 18. 496 CONNECTICUT DEPARTMENT OF ENVIRONMENTAL PROTECTION. (oe Connecti- 204 Rhodora [Vol. 104 cul’s endangered, threatened and special concern a State of Con- necticut, Dept. Environmental Protection, Hartford, Dipsie, A. C. 1993. Back’s sedge, Carex backii od. Maine Natu- ralist 1: eee GLEASON, H. A. b A. Crongouist. 1991. Manual of Vascular Plants of Northeastern cae States and eQiecent Canada. 2nd ed. The New York Botanical Garden, Bronx, New Karrtesz, J.T. AND C. A. MEACHAM. 1999, The ecg of the North Amer- ican Flora. meg Carolina Bot. Gard., Chapel Hill, NC. MAINE DEPARTMENT OF CONSERVATION. 1999, Maine’s rare, threatened and endangered rae Natural Resources Information and — Center, Bureau of Geology ol ve ee: AYE zusta, ME. Web Site (http:// ww.state.me | index.htm) MEHRHOFF, L. J. 1995, Additions ie the preliminary checklist of vascular flora of Connecticut. Rhodora ie SAARELA, J. M. AND B. A. Fo ok ‘aeeenoa of the Carex backit com- plex (section Phyllostac ie Cyperaceae). Syst. Bot. 26: 704-721. SCOGGAN, H. J. 1950. The flora of Bic and the a Peninsula, Québec. Bull. 115, National Museum of Canada, Ottawa, ON. ZEN, E.-AN. 1983. Bedrock Geological Map of is U.S. Geolog- ical Survey, Washington, DC RHODORA, Vol. 104, No. 918, pp. 205-207, 2002 NEW ENGLAND NOTE OCCURRENCE OF SC/RPUS GEORGIANUS (CYPERACEAE) IN MAINE ARTHUR HAINES 1568 US Route |, Freeport, ME 04032 e-mail: arthur.haines @ worldnet.att.net Scirpus georgianus R. M. Harper is a perennial, grass-like herb of wetland communities. It is closely related to S. atrovirens Willd., and the two are part of a group of five morphologically similar species in North America [e.g., S. atrovirens, S. flaccidi- folius (Fernald) Schuyler, S. georgianus, S. hattorianus Makino, and §. pallidus (Britton) Fernald]. The S. atrovirens complex 1s recognized by: trifid styles; mucronate scale apices: relatively straight perianth bristles with thin-walled, round-tipped retrorse barbules confined to the distal % of the bristle; and a haploid chromosome number of 7 = 25—28 (Schuyler 1967; Schuyler and Whittemore, in press). Identification of Scirpus georgianus relies primarily on peri- anth bristle morphology. This species frequently lacks bristles altogether. When bristles are present, they number 1—3 per flower, are typically very short (rarely up to 0.75 times the length of the achene), and are smooth or have a few retrorsely oriented bar- bules near the very tip of the bristle. All other species of the S. atrovirens complex have 5 or 6 perianth bristles that are usually more than 0.75 times as long as the achene and are retrorsely barbellate in the distal % or more of the bristle. Further, the hap- loid chromosome numbers of S. georgianus (n = 25, 26, and 27) are relatively unique in this complex (Schuyler and Whittemore, in press). Only the n = 27 cytotype is shared with another spe- cies—S. flaccidifolius of the mid-Atlantic and southeastern United States. The taxonomic boundaries of Scirpus georgianus have been interpreted differently by different authors over the years. Fernald (1921) reduced this species to a variety as S. atrovirens var. geor- gianus (R. M. Harper) Fernald. He recognized this taxon by its shorter perianth bristles and lower leaves with fewer cross-septae. The illustration that accompanies the description in Gray's Man- 205 206 Rhodora [Vol. 104 ual of Botany, (page 274; Fernald 1950) clearly shows four peri- anth bristles (only one side of the fruit is visible) that are nearly as long as the achene. This indicates that Fernald included within S. atrovirens var. georgianus a plant considered to be a different species by Schuyler (1967)—S. hattorianus. Cronquist (in Glea- son and Cronquist 1991) went a step further and included all the species in the complex into a large, variable S. atrovirens. While reviewing specimens of Scirpus georgianus at the Har- vard University Herbaria, I discovered a specimen annotated by A. E. Schuyler as S. georgianus from Maine. This was the first voucher known to me of this species from Maine. Unfortunately, the label did not contain detailed locality information. SPECIMEN CITATION: UNITED STATES. Maine: York Co., North Berwick. springy, grassy bank, local, 22 Jul 1899, Parlin 1194 (NEBC). On 5 August 2001, Lisa Kuronya and I performed a vehicle survey of rural roads in North Berwick for Scirpus georgianus. Species of this complex routinely occur in human-disturbed hab- itats such as low areas in fields, ditches, and on farm pond shores (Schuyler and Whittemore, in press). A small colony of S. geor- gianus was discovered in a wet ditch on a narrow, gravel road in the eastern half of the township. Seventeen stems were counted ina3 X | m area. Associated species included Viburnum den- tatum L. var. lucidum Aiton, Glyceria striata (Lam.) A. S. Hitche., Carex projecta Mack., C. scoparia Schkuhr ex Willd., Juncus effusus L., Euthamia graminifolia (L.) Nutt., and Sym- phyotrichum lanceolatum (Willd.) G. L. Nesom. The site occurred at 62 m above mean sea level. A few stems had been cut or knocked over by mowing for road maintenance. SPECIMEN CITATION: UNITED STATES. Maine: York Co., North Berwick, road- side ditch, E side of Billy Lane, at 62 m elevation, with Viburnum dentatum ar. lucidum, Glyceria striata, Carex scoparia, Juncus effusus, and Euthamia eraminifolia 5 Aug 2001, Haines & Kuronya s.n. (MAINE) The occurrence of Scirpus georgianus in Maine is not surpris- ing given that it occurs on Prince Edward Island and in Strafford County, New Hampshire (Schuyler and Whittemore, in press). Though this species 1s widely distributed over much of the eastern half of the United States, it is rare and disjunct in the northern part of its range, including New England (Schuyler 1967). Scirpus 2002] New England Note 207 georgianus is a target species of the Herbarium Recovery Project. This two-year project, directed by the New England Wild Flower Society, is collecting information on some of New England’s rar- est and/or poorly known native taxa through herbarium survey. Information gathered from this research will be used to direct conservation efforts in New England. ACKNOWLEDGMENTS. A portion of the information presented herein was collected during review of specimens for the Herbar- ium Recovery Project. The New England Wild Flower Society is thanked for permission to use this information. Alfred Schuyler is also thanked for permission to use the draft Scirpus contribu- tion for the Flora of North America North of Mexico. LITERATURE CITED FERNALD, M. L. 1921. The Gray Herbarium expedition to Nova Scotia. Rho- dora 23: 134. 1950. Gray’s Manual of Botany, 8th ed. Von Nostrand Reinhold Co., New York. GLEASON, H. A. AND A. C. Cronguist. 1991. Manual of Vascular Plants of Northeastern Unie States and Adjacent Canada, 2nd ed. The New York Botanical Garden, Bronx SCHUYLER, A. E. 1967. F taxonomic revision of North American leaf species of Scirpus. Proc. Acad. Nat. Sci. ee 119: 295-323. WHITTEMORE. In press. Scirpus. In: Flora of North America Editorial Committee, eds., Flora of North America North of Mexico, Vol. 23. Oxford Univ. Press, Oxford and New Yor RHODORA, Vol. 104, No. 918, pp. 208-211, 2002 BOOK REVIEW Seventh Catalog of the Vascular Plants of Ohio by Tom S. Coop- errider, Allison W. Cusick, and John T. Kartesz, eds. 2001. x + 195 pp. illus. map. ISBN 0-8142-5061-0 $29.95 (soft- cover); ISBN 0-8142-0858-4 $65.00 (hardcover). Ohio State University Press, Columbus, OH. The Seventh Catalog of the Vascular Plants of Ohio augments major works published since 1961 that focus on Ohio vascular plants (Andreas 1989; Braun 1961, 1967; Cooperrider 1995; Cus- ick and Silberhorn 1977; Fisher 1988; Weishaupt 1971). Tom Cooperrider first conceived of the new catalog in 1960; in the 1970s and 1980s he drafted a preliminary checklist and began planning the Seventh Catalog. John Kartesz independently pre- pared a preliminary checklist for Ohio, and in 1994 suggested that he and Cooperrider combine their efforts. Five additional contributors were enlisted to help compile the Seventh Catalog. Included among the seven authors are some of Ohio’s foremost floristic botanists of today. There has been profound need for the Seventh Catalog. A pe- riod of seventy years has elapsed since publication of Schaffner’s (1932) catalog of Ohio vascular plants. Since then, many taxa have been discovered to occur in Ohio, and marked changes have been made in taxonomy and nomenclature. The Seventh Catalog reflects these developments. According to the Statistical Summary of the Seventh Cata- log, Ohio has 2716 species of vascular plants and 139 inter- specific hybrids: 108 pteridophytes, 17 gymnosperms, 1994 di- cotyledons, and 736 monocotyledons. An additional 143 infra- specific taxa are also listed. Approximately 34% of species, 17% of interspecific hybrids, and 17% of infraspecific taxa are alien to Ohio. The Seventh Catalog has nine main parts, numbered here for convenience: (1) Introduction, (2) Natural History of the Ohio Flora, (3) Catalog of Vascular Plants, (4) Appendix 1: Statis- tical Summary, (5) Appendix 2: Deletions, (6) Literature Cited, (7) Index to Scientific Names, (8) Index to Common Names, and (9) Contributors. Authors include Tom Cooperrider (Part 1) and Guy L. Denny and Cooperrider (Part 2). Part 3 has four main sections: Pteridophytes (Allison Cusick), Gymnosperms 208 2002] Book Review 209 (Cusick), Dicotyledons (Cooperrider, John J. Furlow, and Cus- ick), and Monocotyledons (Barbara K. Andreas, Cooperrider, Cusick [Cyperaceae], and John V. Freudenstein [Orchidaceae ]). Authors are unspecified for Parts 4—9, which represent joint contributions. The actual Catalog of Vascular Plants (Part 3; 79 pages) is the major part of the book. The nomenclature, circumscription, and sequence of suprageneric taxa are based on Cronquist (1981), Gleason and Cronquist (1991), and/or Flora of North America Editorial Committee (1993, 1997). Nomenclature and circum- scription of taxa below the rank of family follow unspecified sources. Genera, species, and interspecific hybrids are listed al- phabetically within families. The following information is given for each species or hybrid: Latin name and author(s); status as native, naturalized, or adventive (or otherwise not established in the flora); and common name. Provided for selected taxa are syn- onym(s) and sometimes additional information crucial for under- standing circumscription. Varieties are given for some species. Interspecific hybrids are listed by the hybrid name, if available [e.g., Asplentum Xinexpectatum (E. L. Braun ex Friesner) C. V. Morton], followed by parentage (e.g., Asplenium rhizophyllum xX A. ruta-muraria). No illustrations or keys are provided, although they are nonessential for this work. Clearly, great effort was required to compile this Seventh Cat- alog. In addition to the numerous taxonomic and nomenclatural decisions involved, I know personally that Tom Cooperrider was determined that listed taxa be correctly identified. Comparison of Cusick’s treatment of Ohio pteridophytes (Part 3 of the Seventh Catalog) with that of Weishaupt’s (1971) Vascular Plants of Ohio, Third Edition exemplifies the extent of contribution of the Seventh Catalog in updating available resources. First, Cusick’s study resulted in the listing of 87 species and 21 hybrids, com- pared with Weishaupt’s 70 species and 2 hybrids; this includes the deletion of four species and one hybrid. Second, Cusick’s list reflects the considerable changes in pteridophyte nomenclature since 1971 (following Kartesz 1994). Among the many examples that could be given here: he listed eight families in place of the more broadly circumscribed Polypodiaceae; the four currently recognized genera (Diphasiastrum, Huperzia, Lycopodiella, and Lycopodium sensu stricto) in place of Lycopodium, and the three currently recognized genera (Athyrium sensu stricto, Diplazium, 210 Rhodora [Vol. 104 and Deparia) instead of the more broadly circumscribed At/yr- ium. Assembly of the Seventh Catalog must have demanded ex- tensive research, meticulous organization, and, ultimately, inten- sive and prolonged proofreading. The Seventh Catalog, however, has two unfortunate deficien- cies. There are no indications of which taxa and how many taxa are newly added to the known flora of Ohio. Thus, considerable burden is placed upon users to extrapolate this information from additional sources. In contrast, Cusick and Silberhorn (1977) pro- vided a list of such taxa. Neither voucher specimens nor herbaria are cited for any of the listed taxa; this would have been espe- cially desirable for new Ohio records. Thus, any misidentifica- tions, however improbable their existence, are likely to go un- corrected for years to come. In contrast, for rare Ohio species Andreas (1989) and Cusick and Silberhorn (1977) cited voucher specimens and herbaria. It is also surprising that some taxa listed for Ohio by Kartesz and Meacham (1999) are neither included in, nor listed as dele- tions from, the Seventh Catalog. Examples include Alopecurus geniculatus, Cardamine Xmaxima, Phellodendron amurense, and Tagetes patula. One wonders if authors of these two works, re- spectively, employed different standards of proof for the occur- rence of taxa within Ohio. A minor complaint relates to the nonalphabetical organization within the Seventh Catalog of taxa of higher rank, particularly of orders and families. Readers unfamiliar with Cronquist’s (1981) system may have difficulty locating taxa. More efficient would have been a strictly alphabetical listing of families, as was pro- vided by Andreas (1989). One recalls favorably the strictly al- phabetical arrangement of taxa within Swink and Wilhelm’s (1994) Plants of the Chicago Region, a feature contributing sub- stantially to that volume’s ease of use. Overall, however, the Seventh Catalog of the Vascular Plants of Ohio represents a most welcome and necessary contribution for persons seriously interested in the Ohio flora. As indicated earlier, reference to the Seventh Catalog reveals that the most recent manual of Ohio vascular plants (Weishaupt 1971) is very out-of-date. One hopes that among Ohio’s floristic botanists there are or will be one or more individuals who will properly revise Weishaupt’s standard work. 2002] Book Review 21) LITERATURE CITED ANDREAS, B. K. 1989. The vascular flora of the glaciated oo Plateau region of Ohio. Ohio Biol. Surv. Bull. New Series 8: I-I¢ Braun, E. L. 1961. The Woody Plants of Ohio: Trees, Sh tie = Woody Climbers Native, Naturalized, and Escaped. Facsimile edition. Hafner Press, New York. . 1967. The Mion oc oly iedonede: Cat-tails to Orchids. Ohio State Univ. Press, Columbus, OH. COOPERRIDER, T. S. 199 5. The oo of Ohio. Part 2: Linaceae through Campanulaceae. Ohio State Univ. § ie abe: OH CRONQUIST, A. 1981. An integrated oe - Ree of Aceeane aa Columbia nae Press, New CusIc . W. AND G. M. SILBERHORN. 1977. The vascular plants of ungla- pe Ohio. ran ae Surv. Bull. New Series 5: 1-157 FISHER, T. R. 1988. The Dicotyledoneae of Ohio. Part 3: Asteraceae. Ohio State Univ. ae Columbus, OH FLORA OF NORTH AMERICA EDITORIAL COMMITTEE, EDS. 1993. Flora of North America North of Mexico, Volume 2. Pteridophytes and Gymnosperms. Oxford Univ. Press, Oxford and New York. 1997. Flora of North America North of Mexico, Volume 3. Mag- noliophyta: Magnoliidae and Hamamelidae. Oxford Univ. Press, Oxford and New York . GLeason, H. A. AND A. Cronguist. 1991. Manual of Vascular Plants of Northeastern ae States and Adjacent Canada, 2nd ed. The New York ee aera Bronx, NY. KARTESZ, J. 1994. A Synonymized Checklist of a oe Flora of the United a. net and Greenland, Vols. & 2, 2nd ed. Timber Press, Portland, OR. AND C. A. MEACHAM. 1999. Synthesis of the North American Flora, version 1.0. North Carolina Bot. Garden, Chapel Hill, N SCHAFEFNER, J. H. 1932. Revised Catalog of Ohio Vascular Plants. Ohio Biol. Surv. Bull. 25: 87-— 215. SWINK, E AND G. WILHELM. 1994. Plants of the Chicago Region, 4th ed. Indiana Academy of Science, Indianapolis, IN. WEISHAUPT, C. G. 1971. Vascular Plants of Ohio, 3rd ed. Kendall/Hunt Pub- lishing Co., Dubuque, IA. —GEORGE J. WILDER, Department of Biological, Geological, and Environmental Sciences, Cleveland State University, 1983 Ea. 24th St., Cleveland, OH 44115-2403. RHODORA, Vol. 104, No. 918, pp. 212-214, 2002 BOOK REVIEW Bioconservation and Systematics: Proceedings of the Canadian Botanical Association Conference Symposium in London, Ontario, June 2000 by James B. Phipps and Paul M. Catling, eds. 2001. 101 pp. ISBN 0-9689565-0-5 US$17.00, CAN$23.00 (softcover). Canadian Botanical Association. [for copies contact Paul M. Catling, catlingp@em.agr.ca] This report comprises seven papers that provide a useful pic- ture of how current trends in systematics and taxonomy affect plant conservation in the Canadian setting. Since Canada, despite its geographical size, has a relatively small flora and a relatively high number of taxonomists and herbaria (Parnell 1993), one might assume that sufficient systematic resources exist to support plant diversity studies and conservation biology. This volume provides interesting examples of systematic research for conser- vation, but also suggests that here, as elsewhere, there are not enough skilled taxonomists, and there is an inadequate infrastruc- ture for collection, management, and use of systematic data. In the first paper, ““A never-ending role for biosystematics in the protection of vascular plant diversity in Canada,”’ Catling dis- cusses the several contributions that taxonomy and systematics make to conservation biology, with plentiful Canadian examples. In addition to re-analyses of nomenclature and the study of spe- cific taxa of known concern, new taxa are being added to the flora, such as Platanthera praeclara, an orchid newly described in 1986. Canada’s 147 endemics remain understudied as well, as do the successive waves of invasives. These and other tasks re- quire more sophisticated information tools, including the devel- opment of national and local taxonomic databases, and their in- terfacing with global and regional data systems like the Integrated Taxonomic Information System for North America. At the other end of the volume, Anton Reznicek (*‘Can sys- tematists help conserve rare plants in the twenty-first century?” ) argues that there 1s a growing dearth of systematists acquainted with plants in their ecological settings, and that this reflects both the increased emphasis on molecular techniques and on land- scape-level ecological study. This means a lack of information needed for good conservation decisions, and it 1s related to the at 2002] Book Review 240 impoverishment of systematic collections, which has been noted for other groups as well (e.g., Winker 1996). Oldham and Sorrill (*“The role of conservation data centres in the conservation of Canada’s flora’’) describe the relatively recent establishment of Conservation Data Centres, or Natural Heritage Programs—the first being set up in Québec in 1988. The authors point out that the Centres’ work is hampered and perhaps threat- ened by the paucity of biologists trained in field identification and in the use of (and contribution to) reference collections. Another aspect of the biodiversity infrastructure in Canada, the Committee on the Status of Endangered Wildlife in Canada (CO- SEWIC) is described by Catling in “Protecting vascular plant biodiversity in Canada: Progress and problems with the taxon approach.” Once again, the basic challenge of up-to-date, com- prehensive, and reliable catalogs of taxa and their status is urgent, at a time when government policy on biodiversity is evolving. Roberts (“Planning with plants in Hlinois’’) contributes a de- scription of the interconnected efforts that have contributed to the evaluation and protection of centers of biodiversity in that inten- sively settled and studied state. Brouillet (“‘Floristics and conser- vation: An example from Newfoundland”) points out that while a basic inventory of the boreal flora may be nearly achieved, much remains to be known about the distribution of the taxa across the huge expanse of the biome. He describes three com- plementary surveys undertaken in Newfoundland, at three differ- ent scales, and demonstrates that such studies provide much new information about species distribution and abundance. He also shows that the electronic cataloguing and management of floristic data are essential elements of basic floristic studies of this sort, as well as being essential to management policy. Finally, Husband and Burgess discuss “‘Evaluating hybridiza- tion as a cause of species endangerment: A role for systematics in plant conservation.” Specifically, they discuss studies that evaluate the impact of hybridization of the rare red mulberry (Morus rubra) with the introduced white mulberry (M. alba). Here is an interesting case in which some hybrids are morpho- logically identifiable, but molecular studies reveal much more hy- bridization than hitherto suspected. There are no breakthrough papers in this collection, but botan- ical libraries should have it on hand. This will be a useful volume for those with an interest in the current state of plant conservation 214 Rhodora [Vol. 104 in Canada. For those with a more general interest in the role of systematics in the protection of biodiversity, the articles provide an interesting patchwork of issues and examples very practically grounded in the current science and policy climate of Canada. Almost every page reemphasizes the urgent need for taxonomists skilled in the field and the herbarium, and the papers provide concrete examples of how this need affects the progress of plant conservation. LITERATURE CITED PARNELL, J. 1993. Plant taxonomic research, with special reference to the tropics: Problems and potential solutions. Conservation Biol. 7: 809— 814. WINKER, K. 1996. The crumbling infrastructure of biodiversity: The avian example. Conservation Biol. 10: 703-707. —BRIAN DRAYTON, TERC, 2067 Massachusetts Ave., Cambridge, MA 02140 RHODORA, Vol. 104, No. 918, pp. 215-217, 2002 NEBC MEETING NEWS February 2002. Vice President Paul Somers introduced the evening’s speaker, Dr. Scott Bailey, U.S.D.A. Forest Service. Bai- ley began his talk, ““Case studies in Geobotany: Refining our understanding of the influence of substrate on plants,” by men- tioning that though his degree was in geology, he has always had a strong interest in botany. After winning over the crowd with this confession, he launched into a discussion of water and nu- trient movement in forests. Watershed studies in the eastern U.S. examined how nutrients accumulate and predicted future changes in storage. While nitrogen had a net accumulation, mostly due to acid deposition, phosphorus and potassium storage have changed very little. Calcium (Ca) and magnesium (Mg) supplies have de- creased substantially, a cause for great concern. Soil exchange sites store nutrients as cations, but there is a question as to wheth- er mineral weathering can keep up with nutrient losses. Weath- ering occurs at widely varying rates (e.g., a small amount of calcite can have a much larger impact than the very common plagioclase feldspar, because calcite weathers 100,000 times fast- er). The potential for air pollution and land management to change the balance between mineral weathering and cation stor- age has renewed interest in the roles of Ca and Mg in plant distribution and health. The first of three case studies presented was conducted on the Allegheny Plateau (NY, PA), a region that has experienced ex- tensive mortality of sugar maple (Acer saccharum) since 1980. Maple death was attributed to Multiple Stress Syndrome (MSS). As its name indicates, MSS can have many causes, and in this case was due to low soil Mg levels (below 0.03 cmol+/kg) and multiple insect defoliation events during the 1970s. In the absence of defoliation, stands tolerated lower Mg levels, and with high soil Mg, stands could withstand several defoliation events. Ex- perimental liming application in 1985 produced a positive re- sponse in sugar maple, though other species such as beech (Fagus grandifolia) and black cherry (Prunus serotina) showed no re- sponse. The second study was an investigation of landscape patterns found in nutrient availability. Two adjacent unglaciated stands, one on a summit with low pH and Ca, the other on a mid-back- slope with 100 times more Ca, illustrated the effect of physical 215 216 Rhodora [Vol. 104 geography on herb diversity and MSS. Bailey and his colleagues discovered that the soil at the mid-backslope site was influenced by groundwater seepage from the underlying bedrock. Although dominated by quartz, the sandstone bedrock contained 10% cal- cite. Bailey suggested that acid rain played a role in MSS by increasing the portion of the landscape with nutrient levels under the threshold necessary to support healthy maple. An expansion of the study suggested that poor base cation supply is just as common in New England. After noting the wide difference in plant diversity, Bailey and colleagues surveyed the flora with the idea of creating an indi- cator system for site nutrient status. Canonical correlation anal- ysis, used to evaluate relationships between floral composition and environmental conditions, identified four species groups: _~ ) Strong Indicators—confined to sites with the highest pH, Ca, and Mg; (2) Medium Indicators—prefer higher pH and base cations but also influenced by organic matter and moisture: (3) Weak Indicators—prefer better sites but occasionally found at nutrient-poor sites; (4) Cosmopolitan Species—no site preference. No species re- liably indicated acidic or nutrient-poor conditions. Current efforts to explain spatial patterns in site quality involve analyzing and predicting bedrock and soil composition. While attempting to deal with these issues in northern hardwood eco- systems, Bailey simplified things by studying species that grow directly on rocks. Epipetric (rock-loving) ferns turned out to be the perfect candidates for the third case study, based on several cliffs in New Hampshire. His study showed that fern species cat- egorized as “‘calcicoles”’ are often found on rock types considered to be Ca-poor. Three hypotheses could explain this: (1) Plants may be rooted in Ca-rich organic matter that accu- mulates on rocks; (2) The rocks have atypical mineral content, such as sandstone containing small amounts of calcite; (3) Lengthy hydrologic flowpaths carry Ca-rich water to the ferns. Bailey concluded his talk with suggestions for the better un- derstanding of the influence of substrate on plants. Researchers 2002] NEBC Meeting News 217 should focus on mineral content rather than the general lithology and should look at horizontal movement of water, rather than focusing on vertical movement. Also, GIS data should be used with discretion, because they are generally compiled on a large scale. As his research has shown, many site-specific “quirks”’ in soil development and hydrologic flowpaths may turn up only in a close examination. —JENNIFER FORMAN, Recording Secretary pro tempore. ANNOUNCEMENT MERRITT LYNDON FERNALD AWARD Merritt Lyndon Fernald was born in 1873 in Orono, Maine. In 1891 he enrolled in Harvard University and started working at the Gray Herbarium, both of which he remained associated with until his death in 1950. During those 60 years he intensively studied the flora of eastern North America, made numerous field expeditions throughout the northeastern United States and south- eastern Canada, and authored over 800 papers on floristically re- lated subjects. Two of his most important contributions were: Per- sistence of Plants in Unglaciated Areas of Boreal North America (1925) and Gray’s Manual of Botany, 8" Edition (1950). Fernald served as an Associate Editor of Rhodora, Journal of The New England Botanical Club from its inception in 1899 to 1928, and as Editor-in-Chief from 1928 until his death in 1950. He was an active member and promoter of the Club. The Council of the New England Botanical Club has decided to honor Fernald’s exemplary contributions to the botany of northeastern North America through a new award, the Merritt Lyndon Fernald Award. The award will be given annually, if deemed appropriate, to the author(s) of the best paper published in each volume of Rhodora that has made use of herbarium spec- imens and/or involved fieldwork. Topics to be considered include, but are not limited to, biogeography, floristics, life-history stud- ies, Monographs, and revisions. Papers on vascular or nonvascular plants, lichens, fungi, and algae will be considered. The compe- tition is not limited to a particular geographic area, but is open to studies in any part of the world. Recipients of the Fernald Award will receive $1000.00 and a certificate acknowledging their achievement. The award will be presented when the New England Botanical Club hosts its annual Distinguished Speaker. ee BS THE NEW ENGLAND BOTANICAL CLUB Elected Officers and Council Members for 2002—2003: President: Paul Somers, Massachusetts Natural Heritage and Endangered Species Program, | Rabbit Hill Rd., Rt. 135, Westborough, MA 01581 Vice-President (and Program Chair): Arthur V. Gilman, Wm. D. Countryman Envi tal Assessment and Planning, 868 Winch Hill Rd., Northfield, VT 05663 Corresponding Secretary: Nancy M. Eyster-Smith, Department of Natural Sciences, Bentley College, Waltham, MA 02154- 4705 Treasurer: Harold G. Brotzman, Box 9092, Department of Biology, Massachusetts College of Liberal Arts, North Adams, MA 02147-4100 Recording Secretary: Neal W. Anderson Curator of Vascular Plants: Raymond Angelo Assistant Curator of Vascular Plants: Erika Sonder Curator of Nonvascular Plants: Anna M. Reid Librarian: Leslie J. Mehrhoff Councillors: Lisa A. Standley (Past President) Judy Warnement 2003 Kanchi Gandhi 2005 Jennifer Forman (Graduate Student Member) 2004 Appointed Councillors: David E. Boufford, Associate Curator Janet R. Sullivan, Editor-in-Chief, Rhodora RHODORA Journal of the New England Botanical Club CONTENTS The flora of Penikese Island, Massachusetts: The fifth survey (1998-1999), with emphasis on the woody vegetation. Richard H. Backus, Pamela Polloni, Brian L. Reid, Paul Somers, and Theodore O. Hendrickson 219 Allozy id for the hybrid origin of Desmodium humifisum (Fabaceae). Jay A. Raveill 253 Varieties of Astragalus pulsiferae (Leguminosae). Stanley L. Welsh, Robin Ondricek, and Glenn Clifton 271 The pare OF see nate on seedling recruitment in sea lavender (/ lumbaginaceae). Jennifer L. Baltzer, Heather L. Hewlin, Edward G. eves Philip D. Taylor, and J. Sherman Boates 280 NEW ENGLAND NOTES A new combination in Lycopodiella (Lycopodiaceae). Arthur Haines... 296 Recent plant collections—Massachusetts. Tad M. Zebryk .............. 299 Cotoneaster divaricatus (Rosaceae) naturalized in Massachusetts. Pefer / 302 NOTES New Scla of Poaceae in the rocky substratum of Municipality of Perote, Veracruz, Mexico. Teresa Mejia-Saulés, Gonzalo Castillo- Campos, ae pa ergio Avendano Reves 304 Turion production by Ruppia maritima in Chesapeake Bay. Michael S. Rosenzweig and Bruce C. Parker 309 BOOK REVIEW Flora of New Brunswick, Second Edition: A Manual for Identification of the Vascular Plants of New Brunswick 12 NEW BOOKS 316 NEBC MEETING NEWS Sie, ANNOUNCEMENT NEBC Graduate Student Research Award 323 Order form for Index to Volumes 76—100 324 NEBC Officers and Council Members inside back cover Vol. 104 Summer, 2002 No. Issued: October 24, 2002 919 The New England Botanical Club, Inc. 22 Divinity Avenue, Cambridge, Massachusetts 02138 RHODORA JANET R. SULLIVAN, Editor-in-Chief Department of Plant Biology, University of New Hampshire, D ‘ 24 e-mail: janets@cisunix.unh.edu ANTOINETTE P. HARTGERINK, Managing Editor Department of Plant Biology, University of New Hampshire, ham, NH 03824 e-mail: Ahartgrink@aol.com Associate Editors ROBERT I. BERTIN STEVEN R. HILL DAVID S. CONANT THOMAS D, LEE GARRETT E. CROW THOMAS MIONE KANCHI GANDHI—Latin diagnoses and nomenclature RHODORA (ISSN 0035-4902). Published four times a year (January, April, July, and October) by The New England Botanical Club, 810 East 10th St., Lawrence, KS 66044 and printed by Allen Press, Inc., 810 East 10th St., Lawrence, KS 66044-0368. Periodicals postage paid at Lawrence, KS. POSTMASTER: Send address changes to RHODORA, P.O. Box 1897, Lawrence, KS 66044-8897. RHODORA is a journal of botany devoted primarily to the flora of North America. Monographs or scientific papers concerned with systemat- ics, floristics, ecology, paleobotany, or conservation biology of the flora of North America or floristically related areas will be considered. SUBSCRIPTIONS: $80 per calendar year, net, postpaid, in funds paya- ble at par in United States currency. Remittances payable to RHO- DORA. Send to RHODORA, P.O. Box 1897, Lawrence, KS 66044- 8897. MEMBERSHIPS: Regular $40; Family $50; Student $25. Application form available at http://www.huh.harvard.edu/nebc/ NEBC WEB SITE: Information about The New England Botanical Club, its history, officers and councillors, herbarium, monthly meetings and special events, annual graduate student award, and the journal RHO- DORA is available at http://www.huh.harvard.edu/nebc/ BACK ISSUES: Questions on availability of back issues should be ad- dressed to Dr. Cathy A. Paris, Department of ies any, University of Vermont, Burlington, VT 05405- 0086. E-mail: )z00.uvm.edu ADDRESS CHANGES: In order to receive the next ainiber of RHO- DORA, changes must be received by the business office prior to the first day of January, April, July, or October. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). RHODORA, Vol. 104, No. 919, pp. 219-252, 2002 THE: PLORA ‘OP PENIRESE ISLAND, MASSACHUSETTS: THE FIFTH SURVEY(1998—1999), WITH EMPHASIS ON THE WOODY VEGETATION RICHARD H. BACKUS Woods Hole Oceanographic Institution, Woods Hole, MA 02543 Current Address: 244 ee Hole Rd., Falmouth, MA 02540 il: d2backus @aol.com PAMELA T. POLLONI Acting Curator, Marine Biological Laboratory Herbarium Woods Hole, MA BRIAN L. REID Division of Biological Sciences, University of Montana 32 Campus Drive #4824, Missoula, MT 59812-4824 PAUL SOMERS Massachusetts Natural Heritage and Endanget red Species Program, Westborough, MA O15S8 THEODORE O. HENDRICKSON PO. Box 460732, Ft. Lauderdale, FL. 33346 STRACT. Five vascular plant surveys have been made between 1873 and 1999 on Penikese, one of the Elizabeth Islands (Massachusetts). The five surveys have noted a total of 326 species, the most recent survey, 218 species. hace half of the species noted are alien on all five survey lists. Four rare (state-listed) native species were found in 1998-1999, The most ney change in the island’s vegetation over 125 years is the great increase in woody vines and shrubs following cessation of the farming that stripped the island of its presettlement forest. Fifteen woody species, some of them recent intro- ductions, are thought to be increasing. Two of the island’s ponds—Tubs and South—are brackish, supratidal pools without vascular North, Leper, Tern, Typha—are fresh, shallow, and usually dry up an- nually, at which time their bottoms aie a dense, diverse flora. species and numbers are fewer than forme plants. Four ponds— Salt marsh '. There is evidence that the island as plant habitat is drier than in the past, Bae as a result of the increase in woody vegetation. For instance, ferns, once Common on Penikese , are now almost wholly absent. Certain species common on nearby islands are missing: for instance, no blueberries or other ericads are found on Penikese. In the absence of further disturbance, it is possible that Penikese will again become forested with red cedar (Juniperus virginiana) as a presettlement account of 219 220 Rhodora [Vol. 104 1602 describes it, but island-wide burns are suggested for destroying invasive woody plants and encouraging native grasses. Such burns might also restore former tern-nesting sites to usefulness. Key Words: alien species, Juniperus virginiana, Penikese Island, prescribed burns, rare species, red cedar, woody vegetation The first botanical survey of Penikese was made in 1873 by David Starr Jordan, who spent the summer on the island as a student of Louis Agassiz at the latter’s Anderson School of Nat- ural History (Jordan 1874). Jordan preserved no specimens of vascular plants, but listed 114 species using the fifth (1867) edi- tion of Gray’s Manual (Fogg 1930). Six species were restricted to the little satellite, Gull Island, which now, at high tide, shows only as a heap of rocks. Bartholomew Gosnold had visited Pen- ikese in 1602 when it was “full of cedars”’ (Archer 1625 as quot- ed in Quinn and Quinn 1983), but after many decades of culti- vation and grazing, it was, in 1873, “absolutely treeless and near- ly shrubless ... about as barren looking a pile of rock and stone as one could well imagine” (Jordan 1874). Settlers had cut trees and grazed sheep there as early as 1675 (Buckley 1997). Early history of the island is also given by Howland (1964). The second botanical survey was made from Woods Hole in 1923 by the Marine Biological Laboratory (MBL) and the Fish- eries Biological Station of the U.S. Bureau of Fisheries on the 50th anniversary of the founding of the Anderson School (Lewis 1924). The island had served the Commonwealth of Massachu- setts as a leper colony from 1905 to 1921, during which period gardening was encouraged (Buckley 1997). The leper colony kept sheep for some of its years, but grazing was likely reduced or intermittent from about 1865 or 1870 to about 1915, when it ceased altogether. Lewis (1924) said of the second survey that “one day was devoted to collection, July 24, and casual visits in August added a few observations.”” Four people observed the vegetation, eight others collected vascular plants. The final list of the latter was provided by John M. Fogg, Jr., then at work on his Ph.D. dissertation on the flora of the Elizabeth Islands under M. L. Fernald. One hundred fifty-nine species of vascular plants were listed. Specimens were deposited in the herbarium at the MBL (SPWH). When Fogg published his thesis (Fogg 1930), 19 addi- tional species were noted for Penikese. Altogether, 90 species not 2002] Backus et al.—Flora of Penikese Island 221 seen in 1873 were recorded. while 40 species seen in that year went unreported. e third survey was conducted from the MBL in 1947. The list of vascular plants was prepared by Edwin T. Moul (1948). He and five colleagues collected on July 6, July 31, and August 3. Specimens were deposited in sPpwH. Moul noted that the asters recorded in earlier surveys were missing or “‘were overlooked because of their late summer flowering.’’ Moul listed 156 plant species, 24 of which had not been reported earlier, while about 90 seen previously were not found. Moul (1961) records a return visit to the island. During much of the interval between the second and third sur- veys (1923-1947), the Commonwealth had used the island as a game farm and wildlife refuge. Annual reports (Massachusetts Division of Fisheries and Game 1925-1939) mention much that is relevant to the natural history of the island. The following were noted (by common names, as given here) as having been planted for wildlife food or cover: arbor vitae, bayberry, beach plum, blueberry, buckthorn, Carolina poplar, inkberry, Japanese barber- ry, laurel, mulberry, Norway spruce, privet, rose (native), sago pondweed, Scotch pine, sumac, viburnum, and widgeon grass. The fourth survey was made in 1973 by botanists from Smith College as a part of the M.A. thesis research of Scott D. Lauer- mann under C. J. Burk (Lauermann 1974; Lauermann and Burk 1976). Some or all of five people collected on June 12, July 14 and I[5, August 8, 9 and 13, and September 20. By 1973, the island had been uninhabited for about 40 years and ungrazed for at least 50. Twenty-nine species not reported earlier were noted, while 109 species listed earlier were not found. Specimens were placed in the Smith College Herbarium (SCHN). Also in 1973, the Penikese Island School was established on the island, bringing new gardeners with new plants. Altogether, it is clear from Pen- ikese’s history that there have been waves of plant introduction and extirpation as land use has changed. SITE DESCRIPTION Penikese Island (41°27'N, 70°55'W) lies 19 km from Woods Hole, Massachusetts at the southern extremity of Buzzards Bay in the Town of Gosnold, Dukes County. The island consists of a fragment of the now partly submerged Buzzards Bay Moraine of ee) Rhodora [Vol. 104 Wisconsinan glaciation (Zinn and Kahn 1972). Save for Penikese, the Elizabeth Islands lie in a straight northeast-southwest string from Woods Hole, with Cuttyhunk at the southwest end. Penikese is out of line with this string, being one mile north of Cuttyhunk, the land nearest to it. Penikese, totalling about 185 hectares (75 acres), consists of two hilly parts connected by a narrow, flat strip of land called “the Isthmus” or, in the past, the ““Neck”’ or “Causeway” (Figure 1). The maximum elevation, 25 m, 1s found on the greater part; the smaller portion, known as Tubs Point, is a few meters lower. The Isthmus is formed from the coalescing upper parts of two back-to-back beaches that head embayments indenting the eastern part of the island one from the south. The beach on the south side of the Isthmus is wide and sandy, and there are sandy stretches of shore south along the east side of the island almost to its southern extremity, South Point; otherwise the perimeter of the island is a jumble of cobbles and boulders. The New England hurricane of 1938 is estimated to have reduced the island by about 25 hectares (10 acres; Massachusetts Division of Fisheries and Game, Annual Re- port for 1938). The “Soil Survey of Dukes County, Massachusetts”” (Fletcher and Roffinoli 1986) describes the Elizabeth Islands as having “very deep ... well drained, sandy and loamy soils formed in reworked glacial outwash or in glacial tll.’ Most Penikese soil is of the Eastchop-Montauk complex (EnC) or the Plymouth- Montauk complex (PtC and PtD). These soil-map units are de- scribed as rolling or hilly, very or extremely bouldery, and consist of loamy sands or sandy loams. Soil permeability is mostly mod- erate to rapid, and available water capacity is moderate to very a shallow indentation from the north, a deep low. Nothing appears to have been published regarding the 1s- land’s soil chemistry. Edgartown, Martha’s Vineyard, about 32 km to the east-south- east of Penikese, is thought to have a climate similar to the lat- ter’s. Climatic averages for Edgartown for the period 1961—1990 are as follows: annual rainfall, 45.25 inches: wettest month, No- vember, 4.45 inches; driest month, July, 2.92 inches; annual tem- perature, 49.7°F; coldest month, January, 29.2°F; warmest month, July, 69.8°F (Northeast Regional Climate Center, 1123 Bradfield Hall, Cornell University, Ithaca, NY). The island is the nesting site for gulls and terns. In 1999, there 2002] Backus et al.—Flora of Penikese Island 228 aw Leper cemetery Rankin Pond Tern Pond Penikese Island School | | Pier North | | South Pond | | 0 0.125 0.25 mile Figure |. Penikese Island; insert showing its location off the Massachu- setts coast; from Buckley (1997). 224 Rhodora [Vol. 104 were about 1000 gull nests on the main part of the island (about 87% herring gull, Larus argentatus, and 13% great black-backed gull, L. marinus) and a tenth that number of tern nests (almost wholly common tern, Sterna hirundo, with a few arctic terns, S. paradisaea) divided among three spots—on the Isthmus, on the south shore of Tubs Point, and at South Point (Blodget 1999). Most of the food of these birds comes from the surrounding sea or from places remote from Penikese, with much excretion and egestion occurring on the island. Thus, since there is little export of organic material, Penikese would seem to be accruing an ever- larger supply of plant nutrients. Penikese is streamless, but has several shallow ponds. Except for a few planted trees and gardens near existing buildings and the vegetation at the shore and around the ponds, the rolling is- land is best thought of as long-abandoned pasture and cropland covered with grasses or grasses mixed with low shrubs or vines. Here and there are individual tall shrubs or patches of the same, some of the patches being of many square meters (Figure 2). MATERIALS AND METHODS Plants were collected on Penikese in 1999 by R. H. Backus, T. O. Hendrickson, P. T. Polloni, B. L. Reid, and Jessica Schultz. One to three of this group worked on April 28, May 14-17, June 11— 13, July 9-12, August 20—23, September |8—21, and October 15— 17. The whole island was walked over repeatedly. Estimates of plant cover are visual ones based on these explorations, and state- ments of abundance are subjective. About 430 specimens were pressed, then studied in the herbarium at the MBL (sPpwH), where R.H.B. and PT.P. prepared and deposited about 230 sheets. Much of the 1999 material was identified by the last two, although B.L.R. identified most of the grasses, graminoids, and goldenrods. Paul Somers made some identifications and verified others. Paul and Lois Somers and Jeanne Livingston collected on the northern two- thirds of the island on June 24—26, 1998. Among the 95 specimens collected by the Somers party were three species not found in 1999—Amaranthus blitoides, Scleranthus annuus, and Agrostis hyematlis; these are included on the list for 1999. A few observa- tions were made by R.H.B. on July 8-11, 2000 and PT-P. made a few on February 7, 2001. Plant names have been brought into conformance with Sorrie and Somers (1999). 2002] Backus et al.—Flora of Penikese Island 225 Figure 2. Looking east-southeast to Tubs Point from the northeast slope of the main part of Penikese Island. RESULTS The vascular plant species found at Penikese in 1998 and 1999 are listed in the Appendix together with the results of the earlier surveys. Certain groups of garden plants observed in 1998—1999 are not listed. One group, just north of the Schoolhouse, contained three apple trees of cultivated varieties, three trees of a Prunus sp., probably a plum, and one white spruce, Picea glauca. These plants were overgrown to varying degrees with Asian bittersweet, sumac, and Japanese honeysuckle. A catalpa stood nearby at the southeast corner of the Schoolhouse. Another group, called the Lower Garden, was about 300 yards north of the House (the residence and principal building of the Penikese Island School). It was planted with nursery stock and contained the following in July 2000: two apples, one pear, four blueberries, four grapevines, one nectarine, one peach or nectarine, one cherry, one plum or cherry, one Rose-of-Sharon, and eight of a horticultural variety of Juniperus virginiana. No annual flowers or vegetables in gar- dens near the House have been listed, although the weeds of these gardens have been included. 226 Rhodora [Vol. 104 DISCUSSION Alien species. The five surveys together report 326 species, one of which is represented by two varieties, for a total of 327 taxa. Of the species on the composite list for the five surveys, 48% are alien (155), the same ratio as in the most recent survey (105 of 218 species). The percentage of alien species for each of the earlier four surveys between 1873 and 1973 are 44, 48, 43, and 48, respectively. Although the percentage of aliens on Peni- kese is close to that reported by Sorrie and Somers (1999) for the entire Commonwealth of Massachusetts (45% of 2814 spe- cies), Massachusetts itself is high in aliens in comparison with New England as a whole and with the other New England states for which numbers are available. A recent summary (Mehrhoff 2000) shows that 31% of New England’s 2882 species are alien, for Connecticut 35% of 2625 species, for Maine 30% of 2103 species, and for Rhode Island 24% of 1618 species. The follow- ing smaller New England areas for which floral lists have recently been prepared can be compared with Penikese with respect to percentage of alien species (arranged in order of diminishing size of flora): Berkshire: Caunty, Mass: whose plants were well on their way to making seed. The smallest species were Eleocharis parvula and Limosella subulata. Described as “‘a giant” among them was 2002] Backus et al.—Flora of Penikese Island 255 Chenopodium glaucum, only 8-10 cm high. Another diminutive species was Rumex maritimus var. fueginus. The parts of pond bottom that had been dry longer held taller species dominated by Pluchea odorata. By our visit on August 20, much of the pond was pink-purple with the last, and it and other tall species had succeeded the tiny flora noted in July. By our September 18 visit, the pond again held a few centimeters of water in its center. Conspicuous among the pond-bottom flora then were Cyperus diandrus, Cc. erythrorhizos, and Cc. filicinus. Other conspicuous species in or at the edges of North Pond were Angelica lucida, Aster novi-belgii, A. subulatus, Bidens con- nata, Carex lurida and other Carex spp., Cuscuta polygonorum, Gnaphalium uliginosum, Hibiscus moscheutos, Hypericum mutil- um, Impatiens capensis, Iris versicolor, Juncus effusus var. pylaei, Lycopus americanus, L. uniflorus, Mentha arvensis, Polygonum pensylvanicum, P. lapathifolium, P. punctatum, Scutellaria gal- ericulata, Scirpus pungens, S. tabernaemontani, Sparganium eu- rycarpum, Spartina patens, and Xanthium strumarium. Beach plants. In 1999, common species of the rocky shore were Achillea millefolium, Anagallis arvensis, Bromus tectorum, Lathyrus japonicus, Leucanthemum vulgare, Oenothera biennis, Raphanus raphanistrum, Rumex crispus, Solanum dulcamara, Solidago sempervirens, and Verbascum thapsus. Common species of the sandy shore, where they grow particularly strongly in piles of decaying Zostera, were Ambrosia artemisiifolia, Atriplex spp.. Chenopodium macrocalycium, Datura stramonium, and Erechti- tes hieractifolia. Glaucium flavum was very conspicuous in the wrack on the Isthmus and at South Point when it was in flower in June. The loss of ferns. Ferns, once common on Penikese, were very rare in 1999. Dennstaedtia punctilobula, the only fern re- ported by Jordan (1874), was included by Lewis (1924) in a list of “the more common plants of the grassland area.”” Lewis also listed Athyrium filix-femina, the only Penikese survey to do so, (“South end of island”) and Thelypteris palustris (‘Low wet places, Typha and Tub Ponds’’). Moul (1948) said for Dennstaed- tia, “Grassy hillside, n.w. of the reservoir,” then said that in 1960 both this species and 7. palustris “are no longer growing at their former sites” (Moul 1961). With the 1973 survey by Lauermann 234 Rhodora [Vol. 104 and Burk (1976) 7. palustris dropped from the list of Penikese plants, and they noted but ‘‘a single specimen” of hay-scented fern. In 1999, we noted only a single poorly growing plant of the latter species, curiously, at the mouth of a petrel burrow in the rock retaining wall near the House. The only other fern reported from Penikese is Onoclea sensibilis, found in 1999 in a tangle of other plants at the north end of North Pond. It may have been a recent arrival on Penikese or, judging by the difficulty we had in re-locating the few fronds we had found earlier, simply over- looked by the other surveys. Why have ferns been lost to Penikese? Is it simple competition with species that are spreading such as the woody species noted earlier, or is the island drying a little superficially, perhaps also attributable to the spread of (deeper-rooted) woody species and a consequent increase in evapotranspiration? The label of a sheet of Thelypteris palustris collected in 1923 (Fogg 460, SPWH) says, “Low wet places. All parts of is.” With the exception of the ponds, there were, in 1999, no places on the island that could be called low and wet. The total disappearance of Dry Pond and the dryness of Rankin Pond lend additional support to the notion that Penikese as plant habitat was somewhat drier in 1999 than for- merly. Plants not found. Conspicuous among woody shrubs and vines common on the Elizabeth Islands as a whole (Cherau 1998; Fogg 1930), but missing on Penikese in all surveys, were mem- bers of the family Ericaceae. Aside from the planted blueberries in the Lower Garden (growing very poorly), we know of but a an old and over- single ericaceous plant on the island in 1999 grown specimen of Vaccinium corymbosum on the west bank of Typha Pond, perhaps a survivor of blueberries planted by the Commonwealth about 1930. Moul (1948) listed V. fuscatum say- ing, “Rare” and Kalmia angustifolia, ““One colony in grassland.” These, too, were probably survivors of plantings made around 1930. These are the only records of ericads for Penikese. We suppose that edaphic factors explain the lack of these plants. Penikese and Cuttyhunk compared. O'Neill (1981) de- scribed the vascular flora of Cuttyhunk for 1974 and compared it to the one described by Lauermann and Burk (1976) for nearby Penikese for 1973. O'Neill calculated the Simpson Index of Re- 2002] Backus et al.—Flora of Penikese Island 2309 semblance, 100c/n, (where c is the number of species common to the two floras and n, is the number of species in the smaller flora), to be 67.6, Penikese (163 species) and Cuttyhunk (264 species) having 110 species in common. The more diverse Cut- tyhunk flora was attributed to the island’s greater size and con- sequent greater diversity of its plant communities. O'Neill re- ported an increase in the number of species of shrubs from the eight given for Cuttyhunk by Fogg (1930) to 40 for 1974 and also found a recent general increase in the island’s shrubby veg- etation. The floral future on Penikese. Earlier writers on the Peni- kese flora have usually speculated as to whether the island will regain the forest that once covered it, but often with ill-founded assumptions about what that presettlement forest was like. Jordan (1874), without citing any authority, said, ““When Penikese was first known it was covered with a growth of trees said to be similar to those now found on Martha’s Vineyard and Naushon. Among these may be mentioned the red cedar, pitch pine, red maple, shag bark etc.” Lewis (1924) said, “The original vege- tation, like that of neighboring islands, is said by Jordan to have been of a forest type, with pitch pine, red cedar, red maple, shag- bark etc.”’ and “‘As the early records of the island mention trees belonging to forests of an advanced type, it is possible that such a forest may again develop.”” Moul (1948) said, “the original climax of forest mentioned by Dr. Jordan may return,” but the same author (1961) said, “In 1948, I expressed the belief that the original tree cover, mentioned by Gosnold’s naturalists in 1602, might return, but today the evidence indicates that a grass ‘sub- climax” may persist into the future.” Fogg’s (1930) consideration of the question is more thoughtful. He argued that the post-glacial forest of the Elizabeth Islands (including Penikese) developed when sea level was much lower than at present so that the shoreline was then many miles south of what are now the islands. Thus, the current regrowth of the islands’ forests must occur under a much harsher set of conditions than those prevailing during their original growth and “it would seem futile to hope that the devastated areas can ever regain their former wooded luxuriance” (Fogg 1930) Gabriel Archer’s and John Brereton’s (1625 and 1602, respec- tively, as cited in Quinn and Quinn 1983) descriptions of a di- 236 Rhodora [Vol. 104 verse oak-hickory forest (including cedars) for Cuttyhunk, where the Gosnold party camped, must ultimately be the source of Jor- dan’s remarks about nearby Penikese’s presettlement forest. But the forest on Penikese was different from the one on Cuttyhunk as indicated by the facts that the Gosnold party, which visited Penikese several times, described the latter as “‘full of Cedars” and came there especially to cut a cargo of that tree for taking back to England (“Captain Gosnoll fell downe with the ship to the little Het of Cedars etc.”; Archer 1625 as quoted in Quinn and Quinn 1983). This seems to indicate that the pre-settlement forest on Penikese was dominated by red cedar. (We take ‘‘cedar”’ to be Juniperus virginiana. Both ‘‘cedar’? and “‘cypresse’” are mentioned in accounts of the voyage, although only ‘“‘cedar’’ is attributed to Penikese. We take “‘cypresse”’ to be Chamaecyparis thyoides. ) By 1930 Juniperus virginiana was rare in the Elizabeth Islands. Fogg (1930) recorded it only for Naushon, where it was ‘‘Plen- tiful in the woods near the East Gutter.’’ But seven decades later, Cherau (1998) found many “in all parts of Naushon.”’ Probably the species is generally increasing in the Elizabeth Islands. — Thus, while two arborescent species, Prunus serotina, black cherry, and Juniperus virginiana, red cedar, seemed to be slowly increasing on Penikese in 1999 as noted above, we subscribe to Fogg’s (1930) argument as to the difficulty of reforestation of the island and believe that a hardwood forest such as has been de- scribed for presetthement Cuttyhunk and assumed for presettle- ment Penikese will not come about. A regrowth of the red cedar that the Gosnold party found in 1602 is quite possible, as this species’ increase on Penikese suggests. Why red cedar, an early successional species, might have dominated on Penikese when Gosnold visited remains an interesting question. We can only sug- gest that this dominance may have resulted from deliberate or accidental burns by the aboriginal Pokanokets who, according to the Gosnold reporters, were seen on Penikese, but did not have a settlement there. Management. G. GRAY. 1967. Hybridization between legumes: Des- eee intortum, “ uncinamim, and D. sandwicense. J. Austral. Inst. . Sci. 33: 122- a: B. Cc AND D. BLOGG. 1983. Variability in isozyme gene frequency in the tropical pasture legume, “Greenleaf? Desmodium. Trop. Agric. (Trin- idad) 60: 193-196 = 2002]. Raveill—Hybrid Origin of Desmodium humifusum 269 IsELY, D. 1953. een panic ue (L.) DC. and D. viridiflorum (L.) DC. Amer. Mid]. Naturalist 49: —93 83. T | The Se ie panic ate e ) DC. (Fabaceae) complex revisited. Sida 10: 142-158 | . Vascular Flora of the Southeastern United States, Vol. 3, Part 2 pene (Fabaceae). Univ. North Carolina Press, Chapel aa NC. 98. Native and Naturalized Leguminosae (Fabaceae) of the United States. Brigham Young Univ. Press, IT. 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Measures of genetic similarity and genetic distance from a small ee of individuals. Studies in Genetics, Univ. Texas Publ. 7213: 145-153. Rorar, P. PR AND K. H. Cuow.1971. Morphological variation and interspecific hybridization among Desmodium intortum, D. sandwicense, and ae uncin- atum. Hawai Agric. Exp. Sta. Univ. Hawai Techn. Bull. . 82, 1-28. S.J. Park, A. BROMbEP, AND U. URATA. 1967. ae aa flow- ering behavior in Spanish Clover, Desmodium sandwicense, and other 270 Rhodora [Vol. 104 Desmodium species. Hawaii Agric. Exp. Sta. Univ. Hawaii Techn. Pro- gress Rep. 164: I-13 an Urata. 196 57. — studies in the genus Desmodiui: Some dine one counts. . Bot. 54: 1-¢ SCHUBERT, B. G. 1950a. bone pp. 91S 923, In: M. L. Fernald, Gray’s ee of Botany, 8th ed. American Book Co., New York. Ob. ee Preliminary studies IIl. Rhodora 52: 135-155. SMALL, : ‘i 1933. Manual of the Southeastern Flora. Published by the author, York. SOKAL, R. R. AND E J. ROHLP. 1981. Biometry. W. H. Freeman & Co., New York Soutis, D. E., C. H. HAUFLER, D.C. DARROW, AND G. J. GASTONY. 1983. Starch gel electrophoresis of ferns: A een of grinding pee gel and electrode ee ers, and staining schedules. Amer. Fern 9-27 STACE, ae A. 1987. Hybridization and the plant species, pp. a In: K. ian ed., Differentiation Patterns in Higher ae Academic ane New STEYERMARK, J. A. 1963. Flora of Missourt. ee, State Univ. Press, Ames, IA, SWOFFORD, D. L. AND R. B. SELANDER.1981. BIOS YS-1I: A FORTRAN pro- gram for the aoe analysis of sector data in popula- tion genetics and systematics. J. Heredity 72: 2 TURNER, B. L. AND O. S. FEARING. 1959. C aie Auber in the Leg- uminosae. II]. Amer. J. Bot. —57. VaIL, A. M. 1892. A preliminary list of the species of the genus Meibomia Heist., occurring in the United States and British America. Bull. Torrey Bot. Club 19: 107-118. Voss, E. G. es Michigan Flora, Part II. mee (Saururaceae—Cornaceace ). aa ee Institute Science, ers eae WENDEL, J. 1 WEEDE . Vi es and interpretation of plant 1 see op. 5-45. In: = Mice - . Soltis, eds., Isozymes in Plat lology. ae aan Portland, WERTH, C. - 1985. a an isozyme ern at a field station. Virginia J. Sci. 36: 53-76 1989, The use of isozyme data for inferring ancestry of polyploid pteridophytes Biochem. Syst. & Ecol. 17: 117-130. ——. 19 ymecicles: Pre-prepared frozen isozyme assays. Isozyme Bull. se 109. WuHITHAM, T. G., P. A. Morrow, AND B. M. Ports. 1991. Conservation of ae a Science 254: 779-780. WIEGAN 1935. A taxonomist’s experience with hybrids in the wild. ati Sl: 16 166. Youna, J. O. 1940. Cytological investigations in Desmodium and Lespedeza. 30t. Gaz. (Crawfordsville) 101: 839-850. RHODORA, Vol. 104, No. 919, pp. 271-279, 2002 VARIETIES OF ASTRAGALUS PULSIFERAE (LEGUMINOSAE) STANLEY L. WELSH S. L. Welsh Herbarium, Life Science Museum, Brigham Young University, Provo, UT 84602 e-mail: SLSLWELSH @aol.com ROBIN ONDRICEK California Department of Fish and Game, Central Valley Bay-Delta Branch, 4001 North Wilson Way, Stockton, CA 95205-2486 GLENN CLIFTON 910 Sanitarium Rd., Deer Park, CA 94576 ABSTRACT. Described as new is Astragalus pulsiferae var. coronensis. This new variety is distinguished by its superficial root crown and longer pod trichomes, as well as by more subtle differences in the type of internode pubescence. In addition, the stipules in the new variety are all distinct, cor- relating with the above-ground stem. Key Words: Astragalus, new variety, taxonomy, California Gray (1874) named Astragalus pulsiferae, ascribing it to “Pha- ca, Inflati.”. Subsequently a second plant, A. suksdorfii, was de- scribed by Howell (1893). Barneby (1958) treated the later-named plant at varietal rank within A. pulsiferae. The species as broadly interpreted by him (Barneby 1964: 965) is an **... enigmatically variable species with decidedly bladdery pods more often than not unilocular but sometimes provided with a rudimentary septum; its lower stipules vary from free to connate and its root-crown from superficial to buried.”” Barneby (1964: 969) also noted: “Occasional populations found in the same area [as var. pulsiferae|, at least sometimes in stiffer soils, combine the characteristic vesture and calyx with a superficial root- crown and stipules all free to the base; samples from these, judged in isolation from the whole species, cannot be ex- cluded on technical grounds from sect. /nflati.”’ From the 1964 tentative placement with sect. Monoenses, the species was realigned, in its entirety, to sect. Humistrati subsect. 21] 272 2 Rhodora [Vol. 104 Micromerii (Barneby 1984: 171), where it was compared to A. tigchmu Barneby. That placement was followed subsequently in the treatment of the Fabales in the /ntermountain Flora (Barneby 1989), and tentatively by Welsh (unpubl. manuscript). Plants with a superficial root crown but with spreading stem pubescence of var. pu/siferae were regarded, at least tentatively, by Barneby (1964: 972), who had seen the plants in the field and of which he had made a collection, as being products of habitat differences; “*... but plants from stiffer soils, which are com- posed of sand compacted with basalt pebbles, have a superficial root-crown and stipules not or at least less strongly connate.” These latter plants, which Barneby included within var. pu/sifer- ae, differ in other respects, also. The plants tend to be erect and tufted, not prostrate-reclining from where they protrude from the soil, and while they have the spreading or spreading-ascending pubescence on stems and petioles, the pod trichomes are decid- edly longer than in those plants with a subterranean caudex. Ad- ditionally, the stipules in those plants with the superficial root crown are all distinct. Despite the differences, a part of which might still be related to different microhabitats, the relationship of the tufted versus prostrate plants is clearly evident. Co-authors Ondricek and Clifton have studied the plants in the field for several seasons; Welsh made observations on some pop- ulations during June of 2001. These observations, added to those made by Barneby and to information derived from the exami- nation of a rather more complete set of specimens at BRY, CAS, ORE, RSA, UC, and WTuU from throughout the range of the species, give indication that there are at least three taxa in the pulsiferae complex, as discussed below. In addition to the representative specimens listed below, there are specimens of all three taxa at CHSC. Astragalus pulsiferae A. Gray, Proc. Amer. Acad. 10: 69. 1874. TYPE: U.S.A. California: Sierra and Plumas Cos., Pulsifer- Ames & Lemmon s.n. [LECTOTYPE designated by Barneby (1964: 972): California: Plumas Co., Aug 1874, Pulsifer- Ames 33 GH!]. Perennial, caulescent, (4) 10-25 (30) cm long, from a branching subterranean (or nearly or quite superficial) caudex, the branches slender. Pubescence basifixed, strigose-strigulose, vil- lous-hirsute, or villosulous. Stems slender, prostrate to decumbent 2002] Welsh et al.—Astragalus pulsiferae 245 or erect or erect-ascending, buried for a space of (0) 2—9 cm. Leaves 1—4.5 (5.5) cm long; leaflets (3) 7-13, 2-12 mm long, 1- 4 (4.8) mm wide, oblanceolate- or obovate-cuneate, retuse or truncate and more or less apiculate, almost flat to loosely folded, rather thinly villous below, less so above; stipules 1—4.5 (5) mm long, either all distinct or some of the buried ones connate. Pe- duncles 0.4—2.5 cm long, very slender, shorter than the leaf; ra- cemes (2) 3—13 flowered, the flowers spreading at anthesis, the axis (2) 4-12 mm long in fruit; bracts 0.8—2.4 mm long; pedicels 0.7-1.8 mm long, disjointing in age; bracteoles O; calyx 3.2—-5.8 (6.2) mm long, the tube |.3—2.6 mm long, shallowly campanulate, villous or villosulous, the teeth |.4—3.6 mm long; flowers (5.2) 6—8.5 mm long, whitish, the banner lilac-veined and keel tipped with lilac, the banner abruptly recurved through 90—100°; ovules (3) 5-9. Pod spreading or declined (often humistrate), sessile, 8— 20 mm long, (5) 6-11 (13) mm thick, bladdery-inflated, some- what dorsiventrally compressed, half-ovoid or ovoid-ellipsoid, unilocular or subunilocular, strigulose to thinly villous, villosu- lous, or pilosulous. The species is confined, except for an outlier in Klickitat County, Washington, to the adjacent Shasta, Lassen, Modoc, Plumas, and Sierra Counties, California, and Washoe County, Nevada. KEY TO VARIETIES OF ASTRAGALUS PULSIFERAE 1. Caudex subterranean or less commonly superficial, the stems foliose only to ground level, the subterranean caudex branches lacking leaves; stems strigulose or villous to vil- lous-hirsute; stipules all distinct or the lowermost connate around the stem; pods strigulose or hirtellous, the hairs OA es IOUS 2 eens eee a eee ee ERE PR REE (2) 2. Stem (at least distally), leaf-rachis, and peduncle villous or villous-hirsute with widely spreading or spreading-as- cending hairs; stems subterranean for 6-10 cm ...... GM e eee oe ene eee ee var. pulsiferae 2. Stem, leaf rachis, and peduncle strigose to loosely strigu- lose with ascending and subappressed, sinuous hairs: stems subterranean for (0) 1—-2.5 (4) cm ........ var. suksdorfit 274 Rhodora IVol. 104 1. Caudex superficial, the stems foliose to the base; stems vil- losulous; stipules all distinct; pods villosulous, the hairs eel ae 0080 a 2. 0 ee ee var. COronensis Astragalus pulsiferae var. pulsiferae Tragacantha pulsiferae (A. Gray) Kuntze, Rev. Gen. Pl. 947. ie Phaca pulsiferae (A. Gray) Rydberg, N. Amer. Fl. 24: 357. 19 Plants with root crown commonly subterranean. Stems mostly buried for a space of (0) 2—9 cm, commonly branched at emer- gence from soil, the foliose internodes villous-hirsute. Calyx teeth (1) 1.4—3.6 mm long. Pod pubescence 0.6—-0.9 mm long. Flowering May to August. Loose sandy sites and interdune valleys, often with sagebrush, on the east side of the northern Sierra Nevada. Mostly on sand derived from weathered granitic rocks at 1310-1798 m in Lassen, Plumas, and Sierra Counties, California, and Washoe County, Nevada. This variety has a rather narrow geographic distribution, from Sierra Valley (Plumas County) and Long Valley (Lassen and Si- erra Counties), California, and generally due east about 16 km in Washoe County, Nevada (Antelope and Red Rock Valleys). The individual plants appear as tufts arranged in a circle around central area filled level with sand. This circular pattern is not evident at sites where the plants are in competition with Bromus tectorum L. The tufts arise from the ends of prostrate, subterra- nean, naked caudex branches, which arise from a central, deeply set taproot. The longest hairs (these spreading or spreading-as- cending) of stems and foliage are more than 0.7 mm long, and with pod hairs less than | mm long. REPRESENTATIVE SPECIMENS: California: Lassen Co., ca. 34 mi. SW of Hallelujah jaan 2 Jul 1999, Ondricek-Fallscheer 195 ake Beckwourth Pass (E side), 19 Jul 1955, oa 50,818 (ORE); Plumas Co. .S mi. ESE of Frenchman Lake and 3 mi. due NE of Beckwourth, 8 Jul (999, pace icek- Fallscheer 197 (sry); Beckwourth Pass, W side, 13 Jun 2001, Welsh & At- wood 28,/20 (Bry); Beckwourth Pass, 19 Jul 1955, Rose 55,152 (BRY): Sierra “o., Long Valley, 1874, Lemmon 515 (photo at seps). Nevada: Washoe Co., Red Rock Valley, 21 mi. N of Red Rock exit from Reno, | Jun 1982, Lavin, Williams & Barneby 4125 (Bry). Astragalus pulsiferae var. suksdorfit (Howell) Barneby, Aliso 4: (31; 1O56. — ee Howell, Erythea 1: 111. 1893. Type: Washington: Falcon Valley, 3 Jun, 21 Jul 1883, Suksdorf [Lecrorype designated WN 2002] Welsh et al.—Astragalus pulsiferae an by Barneby (1964: 971): 3 Jun 1883, Sudsdorf 4S8/ ORE!; ISOTYPES: GH!, NY!, US, WS]. Phaca suksdorfii (Howell) Piper, Contr. U.S. Natl. Herb. 11: 369. 1906. Plants with caudex commonly subterranean for (0.5) 1.5—2.5 cm, or the caudex rarely exactly oe cance Stems mostly simple, sometimes branched or spurred at | or 2 nodes preceding the first peduncle, the foliose internodes strigose-strigulose. Calyx teeth 1.4—2.5 mm long, subequal to the tube. Pod pubescence 0.4—0.7 mm long. Flowering May to July. Open pine forest in loose volcanic substrates at 1380-2005 m, in northwest Plumas and adjacent Lassen and Shasta Counties, California, and also in Falcon Valley, Klickitat County, Washington, at approximately 605 m. Materials from the main body of the variety in northeastern California tend to average smaller, especially in overall stature (7-10 vs. 20—33 cm tall) and leaf (1.3-—2 vs. 3.5—4.7 cm) and leaflet size (1.5—5 vs. 4—12 mm; and the leaflets are more defi- nitely conduplicate) than those in the disjunct type locality Klickitat County, Washington. Additionally, the California rep- resentatives appear to have a more definitely subterranean caudex than those from Washington. The size of the vegetative parts ap- pears to be definitive. However, the floral measurements appear to be identical, and the pod size seems to form a continuum. The difference in size between the disjunct plants in Washington ver- sus those in California within var. suksdorfii is matched by similar size range within individuals of var. pulsiferae, a main difference being the geographic disjunction of specimens from the type locality of var. suksdorfii in Washington and the body of he variety in northern California. The species is evidently miss- ing in Oregon. Despite qualitative differences, it seems best at the present to maintain both of the morphological variants from Washington and California within the concept of var. suksdorfii. _—s REPRESENTATIVE SPECIMENS: — California: Lassen Co., ca. 6.5 mi. W of Cra- ter Mt. and 7.5 mi. due ENE of Lassen Volcanic National Park, 14 Jul 1999, Ondricek-Fallscheer 205 (Bry); Plumas Co., gravelly plain about the airfield west of Chester, 22 Jun 1938, Heller s.n. (RSA); Shasta Co., Bunchgrass Val- ley, i Aug 1911, Eggleston 7537 (Ny); Bunchgrass Valley, 6 mi. due N of jet. Hwy. 44 and 89 (ct. 1s near 7 corner of Lassen Volcanic Se ae 14 Jul 1999, ease Fallscheer 202 (Bry). gay Kl Co., W of Coney e National Wildlife Refuge, 7 Jul 2006 ee Fallscheer 208 (Bry); vn Valley, 16 Jul 1908, Suksdorf ae (ORE). — 276 Rhodora [Vol. 104 Astragalus pulsiferae var. coronensis Welsh, Ondricek & Clif- ton, var. nov. Type: California. Lassen Co., E of Hwy. 395, rd. to Rams Horn Spring Campground, 40°41.500'N, 120°16.931'W, silty sand, in juniper, sagebrush, and Purshia community, at 1540 m (5050 ft.), 14 Jun 2001, Welsh & Atwood 28,158 (HOLOTYPE: BRY; ISOTYPES: CAS, ISC, NY, POM, UC, and others to be distributed). Figure |. Similis var. pudsifera et var. suksdorfo in habitu generali sed in caudicibus superficialibus internodiis villosulosis et pilorum leg- uminibus longioribus differt. Plants with root crown superficial. Stems branching at soil lev- el, foliose to the base, the internodes villosulous. Calyx teeth |.5— 2.5 mm long. Pod pubescence I—!.7 mm long. The new variety is named for the type locality near the Rams Horn Spring campground, the Latin corona being one possible translation of “horn.” Plants appear as low tufts, with no hint of a subterranean caudex. The branches arise from the root crown where it emerges from the soil. The presence of the superficial caudex, a subtle difference in internode pubescence, and definite- ly longer pod hairs are evidently diagnostic. Perhaps of less im- portance are the free stipules in this variety. Union of lowermost stipules in plants with a subterranean caudex is a common con- dition. HABITAT, DISTRIBUTION, AND PHENOLOGY. Astragalus pulsiferae var. coronensts flowers May through July, and is found growing in sandy silt, friable at the surface, hard-packed beneath, among basalt cobble and gravel with juniper, sagebrush, bitterbrush, and Jeffrey pine at 1345-1890 m. Plants of var. coronensis grow on the Modoc Plateau in Modoc and Lassen Counties and on vol- canic inclusions in the Sierra Nevada Range in Plumas County, California. It is evidently rare in Washoe County, Nevada, ap- proximately 30 mi. (ca 42 km) east of the California border. Discussion by Barneby (1964: 969) of the racial subunits of this complex species aggregation is pertinent. After delimiting the typical phase of the species, he points to “Occasional populations found in the same area, at least sometimes in stiffer soils, combine the characteristic vesture and calyx with a superficial root-crown 2002] Welsh et al.—Astragalus pulsiferae eee BS {7 = (Se ee “9a ee) nN Nine y, aN Ss Vere “\ Lg a a py ioe GM Gates Mills; HH, Highland Heights; HV, Hunting Valley: I, ea L, Lake- wood: Li, Lindale: M, Mayfield; MaH, Mayfield Heights; MH, Maple Heights; MiH, Middleburg Heights; MoH, Moreland Hills; MV, Mayfield Village: NO, North Olmsted; NR, North Royalton; OF Olmsted Falls: OT, Olmsted Township; P, Parma; PH, Parma Heights; PP, Pepper Pike; Rocky River; S, Solon; SE, South Euclid; SeH, Seven Hills; SH, Shaker 360 Rhodora [Vol. 104 Heights; St, Strongsville; UH, University Heights; VV, Valley View: W, West- lake; WaH, Warrensville Heights; WH, Walton Hills. Dash(es) between ab- biacone signifies(y) collection(s) made by the boundary or boundaries be- tween municipalities. PTERIDOPHYTES DRYOPTERIDACEAE ystopteris tenuis (Michx.) Desv. — Vertical rock outcrop; //679, Be. Dryoptet ‘is Xboottii (Tuck.) Underw. — Swamp within gorge; 5455, B cae clintoniana (D.C. Bato Dowell — Endangered; vertical rock outcrop; side of creek; //680, ein tne drvopteris (L.) a man — Threatened; vertical rock outcrop; EQUISETACEAE Equisetum Xferrissti Clute — Wetland and slope (both habitats by RR tracks); 4508, C LYCOPODIACEAE Lycopodium hickeyt W. H. Wagner, Beitel & Moran — Woods: fields; /0349, HH (B, BV-W, GH-VV MONOCOTYLEDONS ALISMATACEAE Alisma triviale Pursh — Near creek; water along railroad tracks; in ditch; 6066, se ¢ ree be AMARYLLIDACEAE *Galanthus elwesti Hook. f. — SR. Along trail on forested ridge; /2723, C. * Narcissus Sincomparab P. Mill. — SR. Along railroad tracks; woodlands; 13158, C (CIH. MoH). *Narcissus Xmedioluteus P. Mall. — [Narcissus ll W. Curtis] SR. B railroad tracks; field; woodland; /3309, P (Bk-C, CIH-EC). *Narcissus poeticus L. — Along RR tracks; clump in woodland; /065/, BV- W (GM). COMMELINACEAE Tradescantia ohiensis Raf. * Tradescantia virginiana L. — SR. Three formas are represented, as follows a. Forma with blue nacre comparable in size to flowers of typical 7. ohiensis and T. virginiana, by RR tracks; along alley by RR tracks: 7090, C b. Forma with blue flowers much larger than those of typical 7. ohiensis 2002] Wilder and McCombs—New Records for Ohio 361 and 7. A apparently a garden escape; along railroad tracks; 11959, Ro Forma ok whine petals, green sepals, and blue stamen hairs, the flower size comparable to that in typical 7. ohiensis and T. virginiana; behind urban cemetery and by RR tracks: //995, C O CYPERACEAE Bulbostylis capillaris (L.) Kunth ex C. B. Clarke — In highly insolated cinder along tracks and where RR tracks were removed; //523, C Carex albicans Willd. ex Spreng. var. albicans * Carex umbellata Schkuhr ex Willd. — . albicans var. albicans = Carex artitecta Mack.| SR. Upland bordering Rocky River; /0740, NO. Carex oe ens Schwein. — Sone land-locked region between RR tracks; meadow; swamp; 5163, E(B; Carex atherodes Spreng. — Potentially ene insolated swamp; /4857/, MiH. Carex careyana Torr. ex Dewey — Woods; 10733, B. alone road: 11947, WH Ge. CH) Carex hitchcockiana Dewey — Near path in woodland; /4658, BP-NO-OT. Carex molesta Mack. ex Bright — Wetland along RR tracks; abundant in field: 4056, C (I, MH). Carex pellita Muhl. ex Willd. — [C. lanuginosa Michx.] Wet ditch along road; 4274 Carex texensis (Torr.) Bailey — Base of shaded hill; /4752, B. Carex viridula Michx. — Potentially threatened; grassy area by railroad tracks; 13814, C Cyperus houghtonti Torr. — SR. Presumed a a in highly insolated dry substrate along RR tracks; //530, C-E Cyperus —— L. — [C. aristatus Rottb. and C. inflexus Muhl.] Cracks in pavement of parking lot: /4/46, C. Eleocharis ae (Torr. ) ae — Field; /4096, Be. Rhynchospora capitellata (Michx.) Vahl — Wet meadow; 5642, HH. Scirpus acutus Muhl. ex Bigelow — In ditch: 38/3, B DIOSCOREACEAE *Dioscorea oppositifolia L. — [D. batatas Decne.] Ravine; disturbed urban land; 7564, CIH-EC LEMNACEAE Wolffia brasiliensis Wedd. — |W. papulifera C. H. Thomps.] Beaver pond: lagoon; ///68, GM-M ae NO-OT; I). Wolffia columbiana H. Karst. — Pond; of lake; //287, NR (SH). Wolffia punctata Griseb. — er 140023, 362 Rhodora [Vol. 104 LILIACEAE *Allium cepa L. — Embankment along RR tracks; 4244, C (Wal). — schoenoprasum L. — In garden debris dumped within woodland; /3547, we uscari armeniacum Leichtlin ex ioe — SR. In piles of dirt and debris in vacant lot and along RR tracks; 7935, C (MH). *Scilla siberica Haw. ex Andr. eee edge of gorge: 7963, vicinity GM (CIH-SH NAJADACEAE Najas guadalupensis (Spreng.) Magnus — Pond; /4892, | ORCHIDACEAE Spiranthes ochroleuca (Rydb.) Rydb. — Shaded area along power lines; por- tion of old field bordering forest; //776, I (S) sila ovalis Lindl. — Potentially threatened; > 11554, S (B). jon disturbed land: field by RR ur rac POACEAE “Aegilops cylindrica Host — Ballast between RR tracks: along road: eroded slope beneath terminus of RR bridge; 4027, C (BkH). *A eropyron desertorum (Fisch, ex Link) J. A. Schult. — Clearing by terminus of railroad bridge; /3978, H *A erostis stolonifera L. — In park; 2933, CIH-EC. *Alopecurus geniculatus L. — Dense keane in oe grassy area along parkway; insolated land by power lines; /072/, St (Bk-C, P). *Alopecurus pratensis L. — Fields, swamp, ek along RR tracks; 12153, W (B, BH, CH, S, SH). *A pera ath te (L.) P. Beauv. — Insolated urban field; dry, highly insolated subst along RR tracks; //050, C (B, BkKH-C). onus a us Vahl — SR. Eroded slope at terminus of railroad bridge: 3639, BkH *Bromus racemosus L. — Disturbed, insolated urban land by Cuyahoga River; entrance ramp to I-90; along RR Baan oo bridge; on jetty extending nO Lake Erie; along trail; 3972, C ( ¢G) ris verticillata Nutt. — Along ie eee and tree oo 4831, k — eas compressa Austin ex Pec Field, woods; /43/7/7, P (CIH-EC; BC, BA): Deschampsia flexuosa (L.) Trin. — Potentially threatened; promontory in woods: 4420, CIH-EC. Eragrostis capillaris (L.) Nees — Along RR tracks; along alley; vacant dis- turbed urban land; by wall in disturbed area; 2537, C (BeH: EC: MH). *Eragrostis curvula (Schrad.) Nees — Urban field bordering RR tracks: //583, C-EC. *Fragrostis pilosa (L.) P. Beauv. — Vacant, highly pearee is land; tree lawn; overgrown garden; ballast between RR tracks; 4838, C. 2002] Wilder and McCombs—New Records for Ohio 363 onacuin brachyantherum Nevski — SR. Meadow along RR tracks; /4778, Bk- Rerdann pusillum Nutt. — Solitary plant between RR tracks; 3930, C. *Hordeum vulgare L. — Terminus of railroad bridge; /3SO9, BkH. *Microstegium vimineum (TVrin.) A. Camus var. tmberbe (Nees) Honda — Shaded roadside; /4028, P. *Miscanthus sinensis Andersson — Second-growth woodland; 8982, BH. Muhlenbergia tenuiflora Se Britton, Sterns & Poggenb. — Eroded slopes in woods; //376, B-BH ( Panicum tmplicatum ok: _ Dichanihelicn acuminatum (Sw.) Gould & ar. fasciculatum (Torr.) Freckmann]| Field, clearing beneath powerlines, insolated slump, woodland, along railroad tracks, trailside, swamp; /3729, P (B, i. Bk, C, CIH-EC, E, FP-RoR-W, MH, OF). Panicum lindheitmeri Nash — |Dichanthelium acuminatum (Sw.) Gould & C. A. Clat . lindheimert (Nash) Gould & C. A. Clark] Endangered; dis- turbed aa 13730, P. *Panicum miliaceum LL. creek bed; along RR tracks: 79/4, C (Br, CH Paspalum laeve Michx. — In insolated lawn; //5/9 *Poa nemoralis L. — Two formas are represented, che es forma in which green and a forma with blue-green living shoots. Woods, EC Cin. Cay — Along roads; under oo dried portion of living shoots are ¢g by Rocky River; 3/56 (C, CIH, C * Saccharum ravennae (L.) L. = |krianthus ravennae (L.) P Beauv.| SR. long railroad tracks; /4062, C * Secale cereale L. — Eroded slope at terminus of railroad bridge; /3654, BkH. *Sorghum bicolor (L.) Moench — Along roads; field; lawn; urban waste areas: among boulders; dried-up Rene of creek bed; along RR tracks; 4903, C (Be, BV-W, CIH-EC, GH, *Vulpia ssa (L.) K. C. a — Along RR tracks; disturbed, insolated, urban land; ///74, L (Bk-C, BkKH-C; C, CIH-EC). Vulpia oc sae (Walter) Rydb. — Between RR tracks; 3937; C. *Zea mays ase of embankment along RR ae disturbed land by bank of ea ahee: River; //779, G-S (BkH). TYPHACEAE Typha X glauca Godr. — Urban wetland along RR tracks; /0258, C. DICOTYLEDONS ACERACEAE “Acer campestre L. — Along railroad tracks, in open sunlight; woodlands; 1). 10652, RoR (Bt, ClH ACTINIDIACEAE SR. Abundant climber *Actinidia arguta (Sicbold & Zucc.) Planch. ex Miq. — SSSO, on trees at forest edges (along road and by swamp in woodland): H (CH) 364 Rhodora [Vol. 104 AMARANTHACEAE *Amaranthus blitum L. — Waste area; overgrown urban bce bare dirt; ed see ons of beds of Cuyahoga River and of creek; by creek and river: 9 railroad tracks: S238, Bt (C, CH, CIH . I, PR. RoR). nee powellti S. Watson — Waste area beneath railroad bridge; /40/2, BkH. Amaranthus tuberculatus (Moq.) Sauer — Urban flower bed: large flower pot retained outdoors; 9040, C (CIH APIACEAE *Anethum graveolens L. — Edge of parking lot at urban farmers’ market; 11425, EC *Anthriscus sylvestris (L.) Hoffm. — Along Big Creek and Cuyahoga River; 13383, C Chaerophyllum tainturiert’ Hook. var. tainturiert — SR. Dry, highly insolated substrate along railroad tracks; //047, BV AQUIFOLIACEAE *Hex crenata Thunb. — Woodlands (including woodland land-locked between R tracks); 6548, E Ilex opaca Aiton — Wosdiands: field: 2086, NO-W (HH, P). ARALIACEAE *Hedera helix L. — In woods; /0368, BV (Bc, Bt, OF CIH). ASCLEPIADACEAE Cynanchum laeve (Michx.) Pers. — [Ampelamus albidus (Nutt.) Britton] On hedge along sidewalk; /3003, C ASTERACEAE Aster subulatus Michx. — Along roads ES Omenmes ina ea under bridge; disturbed urban land near RR tre S856, C (B, B Bidens aristosa (Michx.) Britton — Fields (along RR tracks ad not so); 975, (MH). Brachyactis ciliata (Ledeb.) Ledeb. — [Aster brachyactis Blake] Beneath Hee nae roads; along parking lot; disturbed land beneath power lines; 1/1648, Be (B, Bc, BH, C, FP. I, NR, P). o officinalis L. — Among boulder by Lake Erie; 8/8, C. *Centaurea debeauxil Gren. & — [C. pratensis Thuill.] Along railroad tracks: insolated waste area; aon a bordering disturbed, insolated land: 14431, C (BP. ae P). Crepis setosa Haller f. — SR. Large iy aaa in overgrown lawn; 14081, UH. *Dyssodia papposa ( ae A. S. Hitche. — Eastern edge of I-71; 9297, St. Helianthus hirsutus Rat. — Slump me ae land along RR trac a 11497, B(C Helianthus mollis Lam. — Threatened; disturbed area of railroad land; /3948, 2002] Wilder and McCombs—New Records for Ohio 365 *Hieracium piloselloides Vill. — Lawn; along and between RR tracks; inso- lated disturbed areas s Gneluding slope); in shade at farmers’ market; /0924, O = 5). Iva xanthifolia Nutt. — Ballast along RR tracks; highly insolated dirt along urban road; 4980, C. *Leontodon taraxacoides (Vill.) Mérat — Two formas are represented, as fol- low a. Typical forma with yellow ligulate corollas; lawns, cemetery; 4445, B (C, CIH, GM-M, MV, SE). b. Forma with cream-colored ligulate corollas: lawn bordering road: 4447, B. Liatris pycnostachya Michx. * Liatris spicata (L.) Willd. — SR. Field; /4/01/, noes kia fulgida Aiton — Fields; wetland; /908, MH (NR-P, SH). betes patula L. — mee are along railroad tracks; exposed portion creek bed; //623, BV-W (P). BERBERIDACEAE *Berberis vulgaris L. — In woods; 7/73, GM-M. * Mahonia lee (Pursh) Nutt. — [Berberis aquifolium Pursh] Along rail- road tracks; /06 Bk-C (Bc BETULACEAE *Betula pendula Roth — Meadow; wasteland bordering railroad tracks; /3/55, C (Bc) BIGNONIACEAE *Catalpa bignonioides Walter — Along alley; along RR tracks; edge of field; 6954, C (P). Catalpa speciosa (Warder) Warder ex Engelm. — Along RR tracks; 6958, CIH-EC (MH) BORAGINACEAE *“Asperugo procumbens L. — Beneath high bridge; 7024, C-FP. *Heliotropium europaeum L. — Highly insolated urban waste land near Cuy- ahoga River; 4427, C. *Myosous arvensis (L. ) Hill — Flood ae or erga River; 7/77, *Myosotis stricta Link ex Roem. & J. — Urban and rural wns 6794, C (B). BRASSICACEAE *Brassica juncea (L.) Czern. — Lawn an eroded slope beneath RR bridge over Cuyahoga River: 11816. CH (BkH *Cardamine bulbifera (L.) Crantz — SR. oa and disturbed area; /09/5, Bt. *Cardamine flexuosa With. — Weed in flower bed: by picnic area; /3/25, C 366 Rhodora [Vol. 104 Cardanine XMaXUNG te ) Wood — [Dentaria maxima Nutt.| Woodland by Chagrin River; *Cardamine pratensis L. var. pratensis — Lawns; land by shore of Cuyahoga River; 5/04, EC (C . SH). *Chorispora tenella (Pall.) Alph. de Candolle — Along and between RR tracks; 6507, C Descurainia pinnata (Walter) Britton — Threatened; dry, fine ballast along RR tracks; /0666, BV-W. *Descurainia sophia (L.) Webb ex Prantl — Between and near RR tracks: land beneath terminus of RR ees over Cuyahoga River; 4967, C *Erucastrum gallicum (Willd.) ¢ . Schulz — Alo — tracks (sometimes in railroad ballast); rocky il ea MH (C, G *Erysimum cheiranthoides L. uck of exposed ore of bottom - be es ahoga River; along and et RR tracks; weed in flower bed; //252, “H (BV-W, C, E). *Erysimum repandum L. — Along and between RR tracks; along roads; cleared land beneath power lines; dump; 5240, C (B, Bk-C, Br, G, vicinity VV). Be eerad ruderale LL. — Insolated, recently planted lawn; insolated barren and; emersed portion of stream bed; 5409, C (Br, P). cn maritima (L.) Desv. — Exposed portion of creek bed: dumped debris along RR tracks; along curb in hess area; Ae BV-W (C *Lunaria annua L. — Escape in lawn along 1-90; 6832 Rorippa sessiliflora (Nutt.) A. S. Hitche. — ae . Cc ae River, /4898, *Sinapis alba L. — [Brassica hirta Moench] Disturbed land by terminus : railroad bridge; /3626, BKH *Sisvaibrium loeselit L. — Edge of apparently vacant building within urban area; 8217, C. *Thlaspi alliaceum L. — Edge of entrance ramp onto I-90; 53/7, C. BUDDLEJACKAE *Buddleja davidii Franch. — Ballast along railroad tracks; /4/24, MH. CACTACEAE Opuntia humifusa (Rat.) Raf. — Potentially threatened: in ditch near railroad tracks: 1/4/64 CALLITRICHACEAE C es he terrestris Rat. — More-or-less bare soil of parks and picnic areas: , EC @, SI CAPRIFOLIACEAE *Lonicera Xminutiflora Zabel — SR. Along RR tracks; 9627, L *FLonicera ruprechtiana Regel — [Lonicera 4.0 m were collected using 2002] Scribailo and Alix—Rare Aquatic Plants 377 SCUBA. By employing multiple methods, a larger proportion of the lake could be comprehensively inventoried and the likelihood of sampling rare species was greatly increased. Voucher speci- mens have been deposited in either the Aquatic Plant Herbarium of Purdue University North Central (indicated here as PUNC) or the Friesner Herbarium at Butler University (BUT). Percent seed set was determined by visual inspection of in- fructescences and estimations of the number of mature seeds per spike versus the total number of flowers. This nondestructive method was used because of concerns over species rarity. RESULTS AND DISCUSSION Bidens beckiti Torr. Bidens beckii, often cited as Megalo- donta beckii (Torr.) Greene, is commonly referred to as the water marigold. It is the only submersed aquatic member of the genus Bidens L., which comprises some 200 species (Gleason and Cron- quist 1991), many of which are emergent wetland species. The water marigold exhibits a heteroblastic sequence of development, culminating in the production of emergent leaves associated with flowering. The emergent leaves are simple, opposite, and sessile, having either toothed or serrated margins. In deeper water, plants often lack the bract-like emergent foliage and fail to flower (pers. obs.). The submersed leaves of B. beckii are characterized by having finely dissected leaves crowded at nodes with varying degrees of similarity to the leaves of several other aquatic species in the genera Cabomba Aubl., Myriophyllum L., Ranunculus L., and Utricularia L., with which it 1s often confused (Peattie 1930; Voss 1996). Voss (1996) described the leaves as being opposite, but branching, giving the foliage a whorled appearance. Flowers are borne in emergent heads, having the typical yellow ray florets that are characteristic of the genus. Some authors believe that differences in the morphology of the florets, awn lengths of the achenes, and chromosome number warrant the segregation of the water marigold from Bidens into the genus Megalodonta Greene (see Roberts 1985). Although Bidens beckii is found throughout the Midwest, it is state-listed in over 50% of the Great Lakes states (Table |). His- torical records are somewhat vague as to the distribution and abundance of this species in Indiana. Pepoon (1927) and Peattie (1930) suggested that this species was quite rare in northwestern 378 Rhodora [Vol. 104 Indiana during the early 1900s, having been collected or observed from only a few small intradunal ponds in Lake County. Deam (1940) reported this species from seven counties in northern In- diana and suggested that it was once probably found throughout most of northern Indiana, but had likely been destroyed by lake- front setthkement. Swink and Wilhelm (1994) reported B. beckii from LaPorte County, based on a single collection from Stone Lake in August of 1983 (Rowlatt 1297, MOR). This is the last known collection for the state prior to the current study. Indiana EORs simply refer to the original site records from Deam (1940). During the fall of 1997, we found Bidens beckii along the northwestern littoral zone of Stone Lake, LaPorte County, in wa- ter 1.0 m deep with the entire population consisting of less than 60 plants. Plants did not have the emergent foliage associated with flowering. Plants growing in association with B. beckit were Myriophyllum sibiricum Kom., Potamogeton robbinsii Oakes, P. zosterformis Fernald, Ranunculus aquatilis L. var. diffusus With., and Vallisneria americana Michx. This site was revisited during the summer of 1998. The number of individual plants had de- clined to twelve. Signs of habitat degradation were apparent, re- sulting from landowner development of the shoreline. Myrio- phyllum sibiricum, P. robbinsti, and R. aquatilis var. diffusus also exhibited a substantial decrease in their population sizes. A search of the sheltered, undisturbed backwaters of Stone Lake revealed no additional plants of B. becki. Despite the unsuccessful attempt to locate additional beds of Bidens beckii in Stone Lake, an investigation during the summer of 2000 of the aquatic plant communities in Pine Lake, which ts located directly north of and connected to Stone Lake, resulted in the discovery of another population of the water marigold. This population was larger than the one previously reported for Stone Lake, yet it too was threatened by shoreline development along the northwest shore of the highly populated island. Other species growing in association with B. beckii included Ceratophyllum demersum L., Myriophyllum spicatum L., Najas flexilis (Willd.) Rostk. & W. L. E. Schmidt, Stuckenia pectinata (L.) Borner, and Zosterella dubia (Jacq.) Small. In June of 1998, a population of Bidens beckit (> 500 plants) was found at a second site in LaPorte County in a bay along the southwest shore of Hudson Lake. Plants were found growing in water up to 1.5 m in depth. This population was extensive, form- 2002] Scribailo and Alix—Rare Aquatic Plants oye, ing large dense stands encompassing most of the area within the bay. Like the Stone Lake plants, these plants had neither produced emergent foliage nor flowered. Hudson Lake is a marl lake that is largely dominated by the growth of Chara globularis Thuill., which at the time formed large beds that covered much of the littoral zone. Other common submersed species found with B. beckti included Elodea nuttallii (Planch.) H. St. John, Potamo- geton crispus L., P. gramineus L., Utricularia purpurea Walter, and Zosterella dubia. In late August of 1998, the first flowering population of Bidens beckii was found in 0.5 m of water off the western shore of Wauhob Lake, Porter County. Yellow emergent flowers could be seen among the floating leaves of Brasenia schreberi J. F Gmel. and Nymphaea odorata Aiton subsp. tuberosa Wiersema & Hellq. At the time, only seven plants had flowered, but others had floral buds. Although the number of plants present was quite low (< 30), the plants appeared to be secure and quite vigorous in this habitat. Discussions with the lakefront property owner on the sta- tus of the species resulted in an agreement by the owner not to rake ““weeds”’ from the littoral zone in this area. VOUCHER SPECIMENS: Indiana: LaPorte Co., LaPorte. Stone Lake. 41°36'43"N, 86°45'03"W, SW of oe channel, 10 Jun 1998, Altx s.n. (BUT); Lake ce Hudson Lake, 41°42'43"N, 86°33’00"W, in water 1.5 m deep off SW shore, 22 Jun 1998, as & Alix 138 (puNC); LaPorte, Pine Lake, 41°37'40"N, 86°45'07"W, S shore of upper Pine Lake towards N shore of island and E of intersection of Holton Rd. and Island Dr., 7 Sep 2000, Alix s.n. (BUT); Porter Co., Valparaiso, Wauhob Lake, 41°32'01”"N, 87°02'40"W, in water 0.75 m deep near boat rental dock, 21 Aug 1998, Scri- bailo 155, 156 (PUNC) Lemna valdiviana Phil. Lemna valdiviana (pale duckweed) is one of eight Lemna species known to occur in Indiana. Its common name is derived from the pale green appearance of its fronds. The intensity of green color is dependent upon growth conditions, the thickness of the frond, and the content of chlo- rophyll in the different cell layers (Landolt 1986). The fronds of L. valdiviana are often asymmetrical, giving them a distinctive falcate shape (Mohlenbrock 1970), resembling the sole of a shoe. Each frond has a single vein; a character shared with only one other North American Lemna species, L. minuta Kunth, which also occurs in Indiana. Lemna valdiviana can be distinguished 380 Rhodora [Vol. 104 from L. minuta by the greater length of the vein, which in the former species distally exceeds the extension of air space tissue (Landolt 1986, 2000). Distinguishing between these two species can be very difficult and often requires the clearing and subse- quent examination of fronds using light microscopy (Landolt 1986). Lemna valdiviana was first recorded for Indiana as L. cyclos- tasa (Elliot) Chev. by Deam (1932), who collected this species from Noble County. Hicks (1937) summarized Deam’s locality data for this species in connection with the upcoming flora. In his Flora of Indiana, Deam (1940) reported L. valdiviana as be- ing “local in the lake area.”” Deam referred to map 582 in the text of his flora regarding the distribution of L. valdiviana, how- ever, the correct map citation is actually 578. Without the erratum pamphlet that was subsequently provided for the flora, or the correct distribution map in Hicks (1937) for comparison, it is not possible to determine whether the number cited in the text, the maps themselves, or the species designations are in error. Map 578 indicates L. valdiviana from Lagrange, Noble, and Wells Counties in the northeastern part of the state. An examination of herbarium specimens from Indiana University (IND) indicated that this species had been collected from Noble County in 1931 as discussed above (Deam 50405), and Lagrange County in 1933 (Deam 54088). Swink and Wilhelm (1994) did not report this species for the northwestern counties of Indiana. There is only one EOR for this species, which reports its occurrence from Bea- ver Dam Lake fen in Steuben County in 1974. This site was visited in 2000, but we were unsuccessful in locating a population of this species. Lemna valdiviana is listed as extirpated in 50% of the Great Lakes states (Table On September 22, 1997, the pale duckweed was found in the eastern backwaters of Long Lake, Porter County, Indiana. Plants of Lemna valdiviana were typically found in small tangled masses just below the water’s surface. This habit is one quite often ob- served in the related species L. trisulca L. Colonies were rare to occasional and were restricted to the northeastern shoreline and a small backwater pond off of the lake, which largely consisted of patchy stands of Cephalanthus occidentalis L. Other associated plant species included Ceratophyllum demersum, L. minor L., L. trisulca, Proserpinaca palustris L., and Utricularia vulgaris L. On October 4, 2001, an additional population of Lemna val- 2002 | Scribailo and Alix—Rare Aquatic Plants 381 diviana was discovered at the north end of Kaiser Lake beside the public boat launch in Kosciusko County in northeastern In- diana. Plants were abundant, either floating on or submersed just below the surface, and intermixed with L. minor, L. trisulca, Spi- rodela polyrrhiza (L.) Schleid., and Wolffia columbiana H., Karst. Clumps of this species were also found submersed and entangled with the leaves of Ceratophyllum demersum and Myriophyllum spicatum. Lemnid species covered over 50% of the lake’s surface area and harbored an extensive population of L. valdiviana, com- prising an estimated 30% of the mat. VOUCHER SPECIMENS: Indiana: Porter Co., Valparaiso, Long Lake, 41°31’ SUN, 87°02'49"W, in eastern backwaters of the lake, 22 Sep 1997, pees & Alix 13/7 (PUNC); Kosciusko Co., Yellowbanks, Kaiser Lake, 1°18'59"N, 85°39'42”"W, large mat E of public access site, 4 Oct 2001, Scri- a & ree 506 (PUNC). Myriophyllum pinnatum (Walter) Britton, Sterns & Pog- genb. Myriophyllum pinnatum is one of five perennial water- milfoil species occurring in Indiana. It is common throughout the Midwest, though endangered in Indiana (Table |). This species is quite variable in its morphology and habitat, producing whorled pinnatifid leaves on elongated stems when submersed, and branching more freely with scattered leaves when it is anchored along the shoreline (Correll and Correll 1975). Myriophyllum pin- natum is more easily recognized as a terrestrial species (Aiken 1981) and can be confused with M. heterophyllum Michx., an- other native species in Indiana that will often produce a short terrestrial form on exposed mudflats as summer lake levels de- cline. When available, mature fruits provide the best characters for distinguishing between the two species. Although the fruits of both species are 4-angled and have dorsal ridges, these ridges are smooth in M. heterophyllum, but are tuberculate in M. pin- natum. Historical records indicate Myriophyllum pinnatum was. col- lected by Deam (1940; as M. scabratum Michx.) from Jasper County, Indiana, approximately 0.8 km west of the Teft Bridge in the Kankakee River. Swink and Wilhelm (1994) reported the Deam specimen in Plants of the Chicago Region, but did not report any new locations for this species in the northwestern counties of Indiana. An EOR reveals that M. pinnatum was col- lected from Fishtrap Lake, LaPorte County, in July of 1985 by 382 Rhodora [Vol. 104 J. Aldrich and J. Wilhelm (the latter possibly Gerould Wilhelm), who stated that this species was abundant at the time of collection and was found intermixed with plants of M. heterophyllum. A comprehensive inventory of Fishtrap Lake in 1998 by the current authors did not result in the discovery of this species, but only verified the presence of M. heterophyllum. Because M. pinnatum and terrestrial variants of M. heterophyllum can be morphologi- cally similar with pinnate leaves, it is our opinion that the report from Aldrich and Wilhelm may be erroneous. No herbarium spec- imens could be located to evaluate the record from Aldrich and Wilhelm. In 1997, a small population of Myriophyllum pinnatum (20 plants) was found along the northern shore of a small cove of Loomis Lake, Porter County, Indiana, which is the first county directly north from where Deam (1940) collected his specimen. Plants of M. pinnatum were found on the muddy shoreline in shallow water (0.20 m). Associated species included Ceratophyl- lum demersum, Elodea nuttallii, Lemna minuta, L. trisulca, Po- lygonum amphibium L., Spirodela polyrrhiza, Wolffia borealis (Engelm.) Landolt, W. brasiliensis Wedd., and W. columbiana. Although most plants found at this site were terrestrial, some were submersed with emergent flowering and fruiting spikes. The fruits were diagnostic in identification and had very distinct tu- berculate dorsal ridges, a purple tinge, and were subtended by coarsely toothed bracts (Aiken 1981). Two additional populations of Myriophyllum pinnatum were subsequently discovered in adjoining counties. The first popula- tion was found in 1998 at Chamberlain Lake, St. Joseph County, where both flowering and fruiting plants were present. Like the aforementioned population, plants were found either growing as terrestrial variants exposed on mud flats or in their submerged form in shallow water along the sheltered northwest shore. The most common species associates were Ceratophyllum demersum and Nuphar advena (Aiton) W. T. Aiton, the latter extending over half the distance across the lake. The second population was dis- covered in the fall of 2000 on the southern shoreline of Stone Lake, LaPorte County. The population at this site was primarily terrestrial and strictly vegetative, sprawling across the muddy shoreline for nearly 3 m. Species associated with M. pinnatum at this site included Pontederia cordata L. and Sagittaria latifolia Willd. 2002] Scribailo and Alix—Rare Aquatic Plants 383 VOUCHER SPECIMENS: Indiana: Porter Co., Valparaiso, Loomis Lake, 41°31'04"N, 87°03'28"W, northern shore of small cove. 8 Aug 1997, Alix 67 (PUNC); St. Joseph Co., West Field, Chamberlain Lake, 41°39'22’N, 86°22'00"W, in shallow water along north shore, 7 Sep 1998, Scribailo 184 (pUNC); LaPorte Co., LaPorte, Stone Lake, 41°36'34"N, 86°44'39"W, on shore across from intersection of Lakeshore Dr. and Craven Dr., 15 Sep 2000, Alix 341 (PU Najas gracillima (A. Braun ex Engelm.) Magnus. Najas gracillima (thread-like naiad) is morphologically similar to WN. flexilis and N. minor All., which also occur in Indiana. These species are easily confused because of the presence of extensive plasticity in vegetative characters. The most definitive characters for positive identification of naiad species are seed shape and seed coat reticulation patterns, which can only be determined micro- scopically. Fruits of N. graci/lima have an off-center style at their apex, and the seed coats have areoles that are much longer than broad (Haynes 1979, 2000). Najas gracillima is a relict coastal plain aquatic plant species in the Great Lakes region (Peattie 1922; Reznicek 1994) and 1s particularly common in the New England states (Haynes 2000; Stuckey 1983). Stuckey (1983) has commented on the rarity of this species in Ohio, Hlinois, and [Indiana and has cited macro- fossil records (Watts 1970; Wright and Watts 1969) as indicating that the species was once far more common in this region. Stuck- ey postulated that the xerothermic period (beginning 8000 Y BP) that advanced a ‘‘Prairie Peninsula’’ (Gleason 1923; Transeau 1935) eastward for a period of some 3000 years would have re- sulted in the loss of extensive wetland and aquatic habitat that could have contributed to the loss of populations of this species. Recent evidence however, questions the true extent of the “*Prairie Peninsula” xerothermic in the Midwest (Baker et al. 1996). More paleobotanical studies utilizing aquatic plant macrofossils are needed to further determine the impact of the postulated xero- thermic period on aquatic plant distributions, particularly in the eastern states of the Midwest. It is our contention that the apparent scarcity of Najas gracil- lima in the Great Lakes region, and particularly in Indiana where it is endangered (Table 1), may actually be an artifact of under- collecting. Najas flexilis is very common in Indiana and ts often found growing in abundance with occasional plants of N. gracil- lima, so that the latter may easily be overlooked. 384 Rhodora [Vol. 104 We are aware of a single EOR from Pulaski County in north- west Indiana collected by Kriebel in 1938, although it has been collected from two southern counties within the state. Deam (1940) reported the 1935 specimen collected in Lawrence County (Kriebel 3477, IND). Wentz and Stuckey (1971) reported the spe- cies from Knob Lake, Jackson County (7 Sep 1958, Starcs 2/23 BUT; 19 Jul 1970, Starcs 3/00, BUT). We first identified this slender and delicate species from the shallow waters of the eutrophic Mink Lake in Porter County. Eighty percent of the water’s surface area was choked by Nuphar dadvena and the mean water depth within the littoral zone was less than 1.0 m. Plants of Najas gracillima were common, grow- ing along the northeastern and northwestern shorelines at a depth of 0.5 m. Plants had a definite red tinge to their leaves and stems. We have observed distinctive red and green color morph variation in this species at many localities, but do not know the cause of this variation. This red coloration was also observed in plants of Elodea nuttallii. Other associated species included Ceratophyllum demersum, C. echinatum A. Gray, Lemna trisulca, Najas flexilis, Nuphar advena, Nymphaea odorata subsp. tuberosa, Potamoge- ton amplifolius Tuck., P. crispus, P. pusillus L. subsp. tenuissimus (Mert. & W. D. J. Koch) R. R. Haynes & Hellq., Utricularia gibba L., and U. vulgaris. Although it has been suggested that Najas gracillima may not have the ability to withstand eutrophic waters (Haynes 1979; Wentz and Stuckey 1971) this does not appear to be the case at the Mink Lake site. The population at Mink Lake could be near its tolerance limits to eutrophication and may decline or disappear with time. Fertilizer runoff from an adjacent golf course is the primary source of nutrients contributing to the eutrophication. In the latter part of August of 1998, Najas gracillima was found in Silver Lake, LaPorte County. This population was quite extensive, forming dense patches from the eastern to the southern shoreline. The plants had a lime-green coloration and many had set fruit. Plants growing in association with N. gracillima were N. flexilis, Nuphar advena, Nymphaea odorata subsp. tuberosa, Potamogeton amplifolius, P. diversifolius Rat., P. epthyvdrus Rat., P. pusillus subsp. ftenuissimus, and Zosterella dubia. VOUCHER SPECIMENS: Indiana: Porter Co., Valparaiso, Mink Lake, 41°31'49"N, 87°02'14"W, in water 0.5 m deep off the northeastern shoreline, 2002 | Scribailo and Alix—Rare Aquatic Plants 385 10 Jun 1997, Scribailo & Alix 60 (PUNC): LaPorte Co., Rolling Prairie, Silver Lake, 41°41'25”"N, 86°35'45"W, in shallow water along the southwestern shoreline, 29 Aug 1998, Scribailo & Alix 179 (PUNC); 7 Aug 2000, Alix s.n. (BUT). Potamogeton bicupulatus Fernald. Potamogeton bicupula- tus (snail-seed pondweed) is one of the most diminutive and del- icate pondweeds of North America. Like Najas gracillima, this species 1s another of several coastal plain submersed aquatic plant species represented in the northern Great Lakes region (Peattie 1922; Reznicek 1994). It is one of only three linear-leaved pond- weed species in North America having dimorphic inflorescences and embryos with more than one complete spiral; it 1s restricted to acidic waters (Haynes and Hellquist 2000). Potamogeton bi- cupulatus 1s morphologically similar to P. diversifolius, which occurs in many of the southern and central counties of Indiana, as well as a few localities in northwestern Indiana. Peattie (1922, 1930) actually noted P. hybridus (referenced in brackets in his works as P. diversifolius) as being found in the intradunal pond region of Indiana. These specimens, which were collected from Dune Park in Porter County (4 Jul 1906, Hill 732, F; 18 Sep 1903, Hill 156, F), were subsequently annotated as P. bicupulatus by both Barre Hellquist and Robert Haynes and are the only known historical records for the species in the state. We are un- aware of any EORs for this species. In midsummer of 1998, four plants of Potamogeton bicupu- latus were located in a shallow drainage ditch, which flowed into Chamberlain Lake, St. Joseph County. The culvert appeared to have been recently installed and sand within the ditch was part of a pile on top of the pipe. It 1s quite possible that the sand may have been brought in from another area that contained a small seed bank of this pondweed. Only one of four plants had set seed and seed set for that individual was estimated at 50%. A survey of Chamberlain Lake yielded no other plants of P. bicupulatus. Associated species included Ceratophyllum demersum, Nuphar dadvena, and Sagittaria latifolia. VOUCHER SPECIMEN: Indiana: St. Joseph Co., West Field, Chamberlain Lake, 41°39'21"N, 86°21'57"W, near drainage ditch, 7 Jul 1998, Alix /43 (PUNC). Potamogeton epithydrus Raf. Potamogeton epihydrus (rib- bon-leaved pondweed) is one of the most distinctive and easily 386 Rhodora (Vol. 104 recognized North American species of pondweed. Its thin, trans- lucent, strap-like submersed leaves and elliptic floating leaves readily identify this species. Unlike many pondweeds, there are no intergradations between leaf types present. The only North American species morphologically similar to P. epihydrus is P. tennesseensis Fernald, which is distinguished by having long-ta- pering apices in the submersed leaves (Haynes and Hellquist 2000), however the latter species is not found in Indiana. Potamogeton epifydrus is rare in the lower midwestern states and common in Michigan, Wisconsin, and the eastern states (Stuckey 1983). It is very rare in Indiana where it is listed as endangered (Table |). This species has previously been collected from only one location (Deam 1940; Tryon 1937; Swink and Wilhelm 1994), at State Line Creek in LaPorte County (8 Aug 1936, Tryon 30593, BUT; 0742/2, ND). Although a review of EORs indicates that this species has been recorded from Ridinger Lake in Kosciusko County and Loon Lake in Steuben County, no herbarium specimens are available to corroborate these re- ports. In addition, repeated visits by the authors to both of the aforementioned lakes yielded no populations of P. epihydrus. Ex- tensive populations of P. natans L. were present at both lakes. Observed plants had both smaller floating leaves and longer stip- ules than are typically representative of this species in Indiana. The combination of these two characters at the two locations might have resulted in the erroneous reporting of the latter species as P. epihydrus. It is noteworthy that these lakes are marl lakes, typically of higher pH and alkalinity, and would not likely sup- port the growth of a softer water species like P. epihydrus. In 1997, Potamogeton epihydrus was discovered at Silver Lake in LaPorte County where it grew along the shallow sandy banks of the eastern and southeastern shores. In Silver Lake, the ribbon- leaved pondweed formed small patchy beds in water from 15 to 50 cm deep. Seed set was close to 100% (n = 40) on plants from this population. Associated species included Najas gracillima and P. diversifolius. In the summer of 1999, Potamogeton epihydrus was found at two additional locations in LaPorte County approximately 1.5 km west of Silver Lake in two small ponds behind the Rolling Prairie Elementary School. The species was also discovered at Clear Lake in Porter County. Plants at Clear Lake were found at greater water depths (1.25 m) than at the LaPorte locations. Seed set of 2002] Scribailo and Alix—Rare Aquatic Plants 387 plants at this location was approximately 50% (n = 20). Waters of the LaPorte County lakes were more turbid than those of Clear Lake, which may have inhibited the establishment of the ribbon- leaved pondweed at greater depths. According to Hellquist (1980), Nichols (1999), and Stuckey (1983) Potamogeton epihydrus is a species of circumneutral to soft water lakes and ponds. Habitats of this type are limited in Indiana, where calcareous groundwaters and marl deposits have produced a majority of lakes of alkaline pH. Stuckey (1983) rec- ognized P. epihydrus as an additional example of an aquatic plant that may have once been more extensive in range prior to the postulated xerothermic period. VOUCHER SPECIMENS: Indiana: LaPorte Co., Rolling Prairie, Silver Lake, 41°41'27"N, 86°35'33"W, in shallow water of southeastern littoral zone, 23 Sep 1997, Scribailo & Alix 132, 133 (puNC); 7 Aug 2000, Alix s.n. (BUT); Prairie Pond, 41°40'57"N, 86°36'34"W, in shallow water off S shore directly in front of observation deck, 7 Aug 1999, Alix 255 (puUNC):; Porter Co., Jack- son Township, Clear Lake, 41°33'11”N, 86°55'55”W, in shallow water of small bay near W side of lake, 21 Jun 1999, Scribailo & Alix 203 (PUNC). Potamogeton pulcher Tuck. Potamogeton pulcher (spotted pondweed) is a coastal plain species with a predominantly south- eastern range (Haynes and Hellquist 2000), and is not only en- dangered in Indiana, but is also one of the rarest pondweeds found in the Great Lakes region (Table 1). The localities of this species in Indiana are few and include: Jasper County (Welch 1931): Sullivan County (Deam 1940); Pine Station in Lake Coun- ty (Hill 1885); Miller Woods Pond in Lake County in 1982 (Si- monin 26, MOR); intradunal swales west of Miller in Lake County in 1985 (Wilhelm 12955, MOR) and 1991 (Wilhelm & Wetstein 19722, MOR); and Little Lake in Porter County in 1991 (Plampin & Newgent 1-/99/7, Mor). In July of 1997, we revisited the Little Lake site in the Indiana Dunes National Lakeshore and found only a single plant of P. pulcher growing within a monospecific stand of cattails. In September of 1997, a small population (< 10 plants) of Potamogeton pulcher was found off the eastern shore of Fishtrap Lake in LaPorte County. The black-spotted stems and _ petioles along with the cordate-based floating leaves were quite prominent in these plants and are distinctive features of this pondweed spe- cies. The submersed leaves were lanceolate, up to 9.0 cm in 388 Rhodora [Vol. 104 length and averaging 1.2 cm wide, having wavy margins and tapering at the base to petioles 1.0 cm long (e.g., Beal 1977). No flowering or fruiting was observed in these plants. The population of spotted pondweed was 3 m from the eastern shoreline, growing in water 0.75 m in depth. Associated species included Nuphar advena, Potamogeton robbinsii, and Utricularia vulgaris. This site was revisited in July of 1998, but no plants of P. pulcher were tound. In September of 1999, an extensive population of Potamogeton pulcher was found in sloughs along the north and south sides of County Road 700 North within the Jasper-Pulaski Fish and Wild- life Area in Jasper County. Although the population consisted of hundreds of plants, none had flowered. Associated species in- cluded Elodea nuttallii, Myriophyllum heterophyllum, and Nym- phaea odorata subsp. tuberosa. In the summer of 2000, two additional populations of this spe- cies were found. The first population was located in shallow for- mer sand-mining ponds located in the dune and swale topography of Lake County in an area known as the Bonji Tract. This pop- ulation consisted of approximately 30 plants growing in associ- ation with Proserpinaca palustris in water only 0.25 m deep at the base of a small stand of Scirpus pungens Vahl. The second population was found within a remnant area of the Great Marsh in the Nature Preserve portion of the Indiana Dunes State Park in Porter County. This population consisted of less than 10 plants growing with Ceratophyllum demersum, Nuphar advena, Peltan- dra virginica (L.) Schott, and Sparganium americanum Nutt. The multiple leaf types observed in Potamogeton pulcher largely agree with the descriptions by Robbins (1867), who iden- tified the presence of three kinds of leaves in this species: floating leaves roundish-ovate, cordate, or ovate-oblong, all alternate; up- per submersed leaves usually lanceolate, acute at base and with very long acuminate tips, very thin, undulate, short-petioled; low- est submersed leaves thicker, flat, oval or oblong with a rounded base, or spatulate-oblong, on longer petioles. Although we com- pletely agree with these descriptions for floating and upper sub- mersed leaves, we found the lowermost submersed leaves to be either lanceolate or spatulate, but not ovate. In addition, the low- ermost submersed leaves of plants at the Jasper-Pulaski Fish and Wildlife Area had distinctive dentate margins with less than ten 2002] Scribailo and Alix—Rare Aquatic Plants 389 teeth per side. This observation has not been previously reported for this species. VOUCHER SPECIMENS: Indiana: Jasper Co., Jasper-Pulaski Fish and Wildlife Area, 41°09'30"N, 86°56'24"W, in road ditch next to gravel pit fishing area, 7 Sep 1999, Scribailo & Alix 301, 302, 303 (puNC); 21 Jul 2000, Alix sn. (BUT); Lake Co., Gary, Bonji Tract, 41°37'12”N, 87°23'38"W, in shallow water directly east of Clark and Pine Nature Preserve ca. 100 m after crossing Clark St., 5 Aug 2000, Scribailo & Alix 315 (PUNC); LaPorte Co., LaPorte, Fishtrap Lake, 41° a 03"N, 86°43'44”"W, near E shore, 29 Sep 1997, Alix 134 (PUNC): ii Co., Indiana Dunes National Lakeshore, Great Marsh in Dunes State ark, 41°39'29"N, 87°02'49"W, in shallow water next to bridge/observation ana on trail #2, 5 Aug 2000, Scribailo & Alix 328 (PUNC) Potamogeton vaseyi J. W. Robbins. Potamogeton vaseyi ap- pears to be a species of softer waters (Hellquist 1980; Hellquist and Crow 1980; Hopkins 1919; Nichols 1999). The scarcity of softer water lakes in Indiana is likely a factor contributing to the rarity of this species in the state. This species is state-listed in six of the eight states in the Great Lakes region (Table 1). Small populations of Potamogeton vaseyi (< 50 plants) were discovered at Round and Wauhob Lakes, Porter County, in the middle of June 1997. Potamogeton vaseyi was common off the northwest shore of Wauhob Lake and rare in Round Lake. As- sociated species included Ceratophyllum demersum, Myriophyl- lum heterophyllum, M. verticillatum L., Potamogeton amplifolius, P. richardsonti (A. Benn.) Rydb., Utricularia gibba, U. purpurea, and U. vulgaris. During the summer of 1997, two additional populations of Po- tamogeton vaseyit were discovered in LaPorte County. The first population was found along the eastern shoreline of Fishtrap Lake, LaPorte County, and was comprised of four beds, having a total coverage area of over 120 m?*. Unlike the plants of Round and Wauhob Lakes, these plants were robust and many were ei- ther in flower or had fruited profusely. Seed had set on emergent spikes of individuals closer to shore before the inflorescences on deeper plants had emerged from the water. Individual plants from Fishtrap Lake produced multiple infructescences, whereas indi- viduals from Round and Wauhob Lakes each produced only a single infructescence. Associated species included P. pulcher, P. pusillus subsp. tenuissimus, and P. robbinsii. The second population from LaPorte County was discovered 390 Rhodora [Vol. 104 in Saugany Lake (Alix and Scribailo 1998), approximately 16 km northeast of the Fishtrap population. Plants were few in number (< 25) and were found in a small bay of the lake near the west beach. A majority of these plants were growing in relatively shal- low water no deeper than 1.5 m. Although plants at this site were in flower and had produced turions, no seeds were observed. As- sociated species included Myriophyllum spicatum, Potamogeton amplifolius, P. foliosus Rat. subsp. foliosus, P. praelongus Wul- fen, Ranunculus aquatilis var. diffusus, Utricularia vulgaris, and Zosterella dubia. VOUCHER SPECIMENS: Indiana: LaPorte Co., LaPorte, Fishtrap Lake, 41°38'04"N, 86°43'48"W, eastern shoreline directly NE of VFW dock, 22 Jun 1997, Alix s.n. (BUT); Birchim, Saugany Lake, ina ee 86°35'13"W, in naan water next to W beach area, 13 A 997, Alix s.n. (BUT); Porter Valparaiso, Wauhob Lake, 41 °39'00'N, ae 40"W, off W shore near posi rental dock in water 0.75 m deep, 15 Jun 1997, Alix s.n. (BUT); Round Lake, 41°31'57”"N, 87°02'27"W, collected from mouth of channel, 15 Jun 1997, Scribailo 66 (PUNC). CONSERVATION ISSUES There are many concerns regarding the conservation of rare aquatic plant species in the state of Indiana. As previously noted, lakes of the state are typically small and shallow, making them more susceptible to anthropogenic impacts. Although efforts have been made to improve land use by adopting better management practices such as no-till farming, many lakes continue to suffer from an accelerated rate of eutrophication due to nutrient loading from agricultural runoff. Many of Indiana’s lakes have extensively developed shorelines consisting of cottages with concrete retaining walls and mowed lawns to the water’s edge. Buffer strips, which would protect shorelines, provide habitat, and intercept land runoff, are under- utilized in the state. Because of the importance of farming in Indiana, many farmers view ditches and creeks primarily as drain- age and irrigation sources without concern for the possible impact of these practices on the waterways involved. A major problem in assessing the status of rare aquatic plant species in Indiana is the almost complete lack of baseline histor- ical surveys on which to base evaluations of rarity. As a result, it is difficult to ascertain whether species are rare because of 2002] Scribailo and Alix—Rare Aquatic Plants 391 habitat destruction, because lake quality has been degraded, or because they have always been rare. Issues of rarity are of par- ticular concern with the coastal plain species discussed here. Pop- ulations of these species in the Great Lakes region are widely disjointed from their predominantly eastern ranges and may rep- resent unique genotypes of importance to conservation (Reznicek 1994). Most assessments of aquatic plant communities in Indiana are carried out through the Lake and River Enhancement (LARE) Program of the Indiana Department of Natural Resources (IDNR), which awards lake diagnostic grants to private consulting firms. Aquatic plant inventories are required as part of these surveys. The consulting firms involved often do not have the personnel or time to comprehensively survey and identify each species. As a result, these surveys are of limited value because they are often based on one day of collecting, during which only the most com- mon species are identified. In addition, voucher specimens are rarely prepared or deposited for proper documentation. Condon- ing the preparation of voucher specimens of course assumes that the collector would assess the abundance of rare species prior to collection and possibly provide GPS coordinates as an alternative for species that are exceptionally rare. Personnel from the IDNR are often asked to make decisions on lake projects involving such diverse issues as shoreline de- velopment, herbicide application, and dredging that may be based on the aforementioned data. An additional problem is that IDNR biologists do not typically have the necessary training or time to identify populations of rare species that might be impacted. Where herbicide application permits are required and evaluated by IDNR for public lakes, it is often the herbicide applicator that has conducted the aquatic plant inventory, raising the question of vested interest. Herbicide treatment of aquatic plants in Indiana is a multimil- lion-dollar industry and is by far the most prevalent form of aquatic plant control utilized in Indiana. Unfortunately, there are no state permit requirements for application of herbicides in pri- vate lakes. Approximately 50% of Indiana Lakes are considered public because there is public access of some kind. It is important to note though, that since the larger lakes tend to be those that have public access, approximately 80% of the total acreage of Indiana lakes could probably be deemed as public (Carol New- 392 Rhodora [Vol. 104 house, Indiana Department of Environmental Management, pers. comm.). As of July 1, 2002, changes to Indiana state law will reduce the extent to which a riparian resident can treat aquatic vegetation in a public lake without a permit. Formerly, it was not to exceed 50% of the aquatic vegetation area or one-half acre, whichever was less. The new ruling specifies not to exceed 25 ft. along the legally established, average, or normal shoreline and out to a water depth of 6 ft., which now limits the area of treatment to 625 ft’. It is important to note that the old ruling only applied to chemical treatment, whereas the new ruling governs any type of control method. Although this change represents a positive step favoring conservation of aquatic plants, the regulation still allows multiple residents along a shoreline to eradicate major portions of the aquatic vegetation without the need of a permit. Even if permits are not required for small-scale herbicide treatment in lakes, prior notification of local DNR biologists should be re- quired of all proposed applications, so they can assess if there might be negative impacts on rare aquatic plant species. A major issue concerning the success of aquatic plant conser- vation in Indiana is the apparent lack of understanding by the public of the role of aquatic plants in lake ecosystem dynamics. People in Indiana, as in other states, typically want beaches that look like swimming pools. Few lake residents seem to make the connection between aquatic plants and the health of fish popu- lations. This lack of understanding leads to the misconception that the aquatic plant beds can be eradicated without any effect on desirable game fish or other aspects of lake health. Lakeside res- idents are also seemingly unaware of the importance of aquatic plant beds in reducing shoreline erosion and trapping sediments and nutrients. Education programs are needed to enhance public awareness of the importance of aquatic plants in the maintenance of lake quality. Fortunately, Indiana recognizes many of these issues as problems, and has a variety of education and volunteer programs designed to enhance the understanding of lake resi- dents. The state is also currently drafting more rigorous require- ments for the sampling of aquatic plants during lake quality sur- veys. ACKNOWLEDGEMENTS. The authors thank two anonymous re- viewers for comments on the manuscript. The authors wish to 2002] Scribailo and Alix—Rare Aquatic Plants Bie. thank the curators and staff members at BUT, F, IND, MOR, and ND for their assistance in transcribing herbarium specimen label data. In addition we are grateful for access privileges to Silver Lake granted to us by Amy Huber of the Silver Lake Conservation Committee. We greatly appreciate the comments and suggestions made by Dr. C. Barre Hellquist on an earlier version of this paper. 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Report on the Historical Remains, Scenic Places and Natural Monuments in Kyoto Prefecture 18: 1-127. [In Japanese] MUENSCHER, W. C. 1944. Aquatic Plants of the United States. Comstock Publ. *o., New York. Ouwl, J. 1953. Flora of Japan. Shibundo Co., Ltd., Publishers, Tokyo. [p. $26] 2002] Yu et al.—Revision of Myriophyllum 42] ——.. 1975. Flora of Japan, new ed. Shibundo Co., Ltd., Publishers, Tokyo. Po > M. KiraGawa. 1992. New Flora of Japan. Shibundo Co., Ltd., econ Tokyo. [p. 1098] ORCHARD, A. E. 1979. Myriophyllum (Haloragaceae) in Australasia. I. New —— A revision of the genus and a synopsis of the family. Brunonia 247-287. . 1990, Pee ce ic, pp. 5-85. In: A. S. George, ed., Flora of Aus- tralia, Vol. ce oe Publ. ets Canberra PATTEN, B. C., is ah The status of some American species of Myrio- phyllum as revealed by the discovery of intergrade material between M. exalbescens Fern. and M. spicatum ie in New Jersey. Rhodora 56: “O13 225: PRAGLOWSKI, J. 1970. The pollen morphology of the Haloragaceae with ref- erence to pean Grana 10: 159-239 PRESTON, C. D. A .M. ae T. 1997. Some Plants in Britain and Ireland. Harley oe B. H. & A. Harley Ltd., Colchester, U.K. PuGsLey, H. W. 1938. A new variety = Myriophyllum alterniflorum DC. J. Bot. 76: 51-53. ReGeL, E. 1861. Tentamen Florae Ussuriensis, oder Versuch einer Flora des Ussuri-Gebietes. Nach den von Herrn R. Maack gesammelten Pflanzen bearbeitet. Mém. Acad. Imp. Sci. Saint-Pétersburg VII. Ser. 4(4): 60 TAKHTAJAN, A. L. 1978. Floristicheskye oblasty Zemli (Floristic Regions of the World). Nauka, Leningrad. [In Russian] 2000. Haloragaceae, pp. 135-140. In: C. J. Chen, ed., Flora _Reipubticae Popularis Sinicae, Vol. 53. Science Press, Beijing. WANG, N. Z., FS. ZHANG, H. R. HUANG, and E L. Ma. 1983. The Illustrated Flora a Aquatic Wicealae Plants of China. Hubei People’s Press, Wuhan, ae WEBER, J. A. AND L. D. Noopen. 1974. Turion formation and germination in ire verticillatum: Phenology and its interpretation. Michigan Bot. 13: IS1I-158. YAN, S. Z. 1983. The illustrated flora of higher aquatic plants of China. Chinese Academic Press, Beijing Yu, D. 1992. Aquatic plants of Northeast China. Unpubl. Ph.D. dissertation, Nomen Forestry Univ., Harbin, Chin RHODORA, Vol. 104, No. 920, pp. 422-428. 2002 NEW ENGLAND NOTE NEW RECORDS FOR CHENOPODIUM FOGGII IN NEW ENGLAND ARTHUR HAINES 44] Dead River Road, Bowdoin, ME 04297 e-mail: arthur.haines @ worldnet.att.net Betsy W. NEWCOMER 1044 Middle Road, Parsonsfield, ME 04047 e-mail: newcomer @psouth.net Chenopodium foggii Wahl is a rare annual herb native to east- ern North America. It currently possesses a global rank of G3Q (fewer than 100 world occurrences, questionable taxonomy: Pennsylvania Natural Diversity Inventory 2001). Unlike many fa- miliar species of goosefoots, such as C. album L., it occurs in non-anthropogenic habitats. Chenopodium foggti is frequently lo- cated on rock outcrops, at cliff bases, and along sparsely wooded slopes (Wahl 1954). It is closely related to, and sometimes in- cluded in, western C. pratericola Rydb. (Clemants 1992; Gleason and Cronquist 1991). Chenopodium pratericola, however, is ad- ventive in the east where it is found in open, disturbed, often saline soil (e.g., coastal beaches, salted roadsides; Seymour 1982). Although Basset and Crompton (1982) recognized C. fog- gil in their review of the genus in Canada, they apparently con- fused important morphological characters, as suggested also by Clemants (1992). Chenopodium foggti will be recognized as a distinct species in the upcoming Flora of North America contri- bution (Clemants and Mosyakin, in prep.). This paper presents results of recent field and herbarium surveys for C. foggii in New England. Chenopodium foggii is a relatively recent addition to the flora of North America. It was described by Wahl (1954) during a North American revision of the genus. Chenopodium foggii 1s typically a short, sparingly branched plant with moderately fari- nose surfaces, keeled sepals, horizontally oriented fruits, and a loose or irregularly rupturing, minutely echinate pericarp that de- taches from the body of the lustrous black seed. It shares these 422 2002] New England Note 423 character states with C. pratericola. Chenopodium pratericola, however, has thicker, less often toothed, and narrower leaves than C. foggil. Historic New England occurrences. Along with his descrip- tion, Wahl] (1954) documented nine occurrences of Chenopodium foggii in New England (summarized in Table | and discussed below). Sites were identified by herbarium specimen review and not by field survey. Since its description, C. foggii has not been reported from New England. Both of the specimens Wahl cited from Maine are old records (i.e., prior to 1900) and were found in areas of the state that have been heavily developed since. Furthermore, the herbarium label data are vague and no detailed location information was recorded. Historic New Hampshire stations of Chenopodium foggti were mainly in open, rocky woods and cliff bases. Three of the sites reported by Wahl are in the northern half of the state, and one collection (Walpole, Chesire County) 1s from extreme southwest- ern New Hampshire. The most recent collection reported by Wahl was from 1920. Examination of specimens at NHA by the first author yielded three additional collections of C. foggii from Mount Stanton, in Bartlett (6 Jul 1965, Hodgdon et al. 14504; 2 Aug 1960, Steele s.n.; 26 Aug 1954, Steele /53/). All three col- lections had been misidentified as C. boscianum Mogq., a fre- quently used and inappropriate name for the eastern C. standley- anum Aellen. No extant sites of C. foggii are known from the state, and recent surveys of the Harts Ledge have not re-located the species (Bill Nichols and Dan Sperduto, New Hampshire Nat- ural Heritage Inventory, pers. comm.). Wahl cited a single record from Vermont, and no other occur- rences are known (Bob Popp, Vermont Nongame and Natural Heritage Program, pers. comm.). Vermont is currently the only state in New England to list this plant as a species of conservation concern (Vermont Nongame and Natural Heritage Program 2000), though it is listed incorrectly under the name of Chenopodium desiccatum A. Nelson. Wahl listed only two sites for Massachusetts, though a third site is known from Hampden County by a 1916 collection an- notated by Wahl in 1963 (Karen Searcy, University of Massa- chusetts Herbarium, pers. comm.). Weatherbee (1996) considered Chenopodium foggti to be uncommon in Berkshire County. Bruce Table 1. Collections of Chenopodium foggti attributed to New England by Wahl (1954). Collection numbers are not provided in the table as they were not cited by Wahl and Harvard University Herbaria collections were not available during research for this manuscript. County Town Location Date Collector and Herbarium Massachusetts Berkshire New Marlboro 28 Aug 1920 Hoffman (NEBC) Berkshire Mount Washington Bash Bish Falls 9 Sep 1919 Hoffman (NEBC) Maine Androscoggin Auburn 21 Jun 1896 Merrill (NEBC) 3 Sep 1898 Parlin (GH, NEBC) South Berwick Pease (NEBC) York New Hampshire Carroll Bartlett Whites Ledge & Sep 1915 Cheshire Walpole Fall Mountain 31 Jul 1900 Fernald (GH) Coos Hadleys Purchase Harts Ledge 9 Sep 1915 Pease (NEBC) Grafton Haverhill 18 Aug 1917 Fernald (NEBC) Vermont range Fairlee 4 Aug 1928 Pease (NEBC) LIOPOYY TOA] vO! 2002] New England Note 425 Sorrie (formerly of the Massachusetts Natural Heritage and En- dangered Species Program, pers. comm.) considered this species to be very rare in Massachusetts and did not encounter it during floristic work in the state. It is surprising to note that given the lack of current records for this species in New England, Chenopodium foggii has re- ceived very little conservation focus and has been formally listed by only one of the six states. Confusion with the adventive C. pratericola has likely contributed to its being overlooked in the northeast. Field observations. On 3 October 1999, the primary author visited Bartholomew’s Cobble in Sheffield, Berkshire County, Massachusetts. This well-known feature adjacent to the Housa- tonic River comprises low outcrops of dolomitic marble (De- Lorme 1998). Both mesic and xeric substrates occur, supporting a large number of calciphilic plants. A relatively small Cheno- podium was observed on a dry, open terrace with southwest as- pect. Morphology, in particular keeled, moderately farinose se- pals, small leaves (less than 4 cm long) with few or no teeth, and horizontally oriented fruits in the calyx, suggested the population could be C. foggii. Examination of the fruits at 20 confirmed this, and the identification was verified by Steven Clemants (Brooklyn Botanic Garden). The site was characterized by ex- posed bedrock and sparse, stunted Juniperus virginiana L. As- sociated species included Aqguilegia canadensis L., Rubus occi- dentalis L., Schizachyrium scoparium (Michx.) Nash, Carex ce- phaloidea (Dewey) Dewey, Woodsia obtusa (Spreng.) Torr., Hy- pericum perforatum L., and Achillea millefolium L. The location was approximately 195 m above mean sea level. This is the only known extant site in Massachusetts. Bartholomew’s Cobble is owned by the Commonwealth of Massachusetts and managed by the Trustees of Reservations. State employees have been made aware of the occurrence of Che- nopodium foggii and its rarity in New England. The specimen, which includes a color image of the plants im situ, has been de- posited at the New England Botanical Club Herbarium. VOUCHER SPECIMEN: Massachusetts: Berkshire Co., Sheffield, Bartholo- mew’s Cobble, 3 Oct 1999, Haines s.n. (NEBC). On 21 July 2000, we visited a Polygonum douglasti Greene 426 Rhodora [Vol. 104 station on Cedar Mountain in Parsonfield, York County, Maine. The site occurs on a small, open bald of Devonian-Silurian lime- stone (Osberg et al. 1985). We observed a relatively small Che- nopodium in flower that did not appear to match any species known to be extant in the state. The plants were generally shorter than 30 cm with moderately farinose surfaces. The leaf blades did not exceed 4 cm in length and were essentially entire. Though characteristics of the sepals could be observed, such as a well- formed keel, fruit size and details of the pericarp were not as- sessable. The flowering morphology and associated natural com- munity suggested this plant could be C. foggii. As this species had not been seen in Maine for 102 years, a return trip was made by the second author to collect a fruiting stem and confirm the identification. The specimen collected from Cedar Mountain demonstrated the pericarp morphology for Chenopodium foggli (e.g., minutely ech- inate texture, non-adherant). Chenopodium foggti is similar, 1 _ 71 regard to the freely separable pericarp, to another uncommon eastern forest species that is historically known to occur in Maine, C. standlevanum. The keeled sepals and farinose habit, however, distinguished the Cedar Mountain plants from C. standleyvanum, which has unkeeled sepals and nearly glabrous herbage. The iden- tification was confirmed by Steven Clemants. The Cedar Mountain site is the only known extant station of Chenopodium foggti in Maine. Associated species include Polyg- onum douglasti, Carex backit Boott, Poa compressa L., Rumex acetosella L., Aquilegia canadensis, Corydalis sempervirens (L.) Pers., Saxifraga virginiensis Michx., Stellaria graminea L., and Dryopteris marginalis (L.) A. Gray. The station occurs at ca. 260 m elevation and has southern aspect. The property owners are aware of the plant and plan to conserve the area. The specimen has been deposited in the University of Maine Herbarium. VOUCHER SPECIMEN: Maine: York Co., Parsonfield, Cedar Mtn., 18 Sep 2000, Newcomer s.n. (MAINE). Herbarium survey. An herbarium survey was initiated by the New England Wild Flower Society to collect information on rare and poorly known native species. The goal of this research, called the Herbarium Recovery Project, is to verify the accuracy of collections in regional museums and gather label information 2002] New England Note 427 for 532 species in New England. Chenopodium foggil is a target species of this project. While examining material at the Harvard University Herbaria, the primary author annotated three speci- mens as C. foggtt. The New Hampshire specimen was collected while in flower, and although the morphology and habitat matches that of C. foggii, it cannot be identified with certainty. However, the specimen is an apparent duplicate of one cited by Wahl (1954). Steven Clemants has also reviewed these sheets and con- curred with the determinations. This represents the first report of C. foggti from Connecticut. SPECIMENS EXAMINED: oe ee New Haven Co., aaa Haven, East Rock, dry rocky wooded waste, 14 Sep 1932, Eames 1/488 (Gu). New Hamp- shire: Cheshire Co., eae Fall Manon rocky woods, 31 7 1900, Fer- 425 (GH nont: aie Co., West Rutland, Twin Mountains, 15 pee 1900, ee 2077 ( lot Chenopodium foggii is a poorly known and overlooked species in New England. The premature inclusion of this species in the synonomy of C. pratericola has likely reduced the intensity of field efforts that may have resulted in its earlier rediscovery. Che- nopodium foggii fits criteria for a Division | species in New Eng- land (globally rare with fewer than [OO world occurrences; Brum- back and Mehrhoff et al. 1996). Field surveys should be directed toward locating new and historic populations, particularly in high pH bedrock regions. ACKNOWLEDGMENTS. The following people contributed to this note and are thanked: Steven Clemants: Bill Nichols, Dan Sper- duto, Bob Popp, Karen Searcy, Bruce Sorrie, Thomas Vining, and Pamela Weatherbee. The New England Wild Flower Society is also thanked for permission to use data collected during the Her- barium Recovery Project. LITERATURE CITED Bassett, I. J. AND C. W. Crompton. 1982. The genus Chenopodium in Can- ada. Canad. J. Bot. 60: 586-610 BRUMBACK, W. E ofa Dre es HRHOFF ET AL. 1996, Flora Conservanda: New England. Rhodora 98: 233-361. CLEMANTS, S. E. 1992. Crenopadiaea eo Amaranthaceae of New York St New York State Museum Bull. 485, Alb State. DELORME. 1998. Massachusetts ce and ae oe a ME. 428 Rhodora [Vol. 104 GLEASON, H. A. AND A. CRONQUIST. 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd ed. The New York Botanical Garden, Bronx, NY. OSBERG, P., A. Hussty, AND G. Boone. 1985. Bedrock Geologic Map of Maine. Maine Geological Survey, Augusta, ME. PENNSYLVANIA NATURAL DIVERSITY INVENTORY. 2001. Biota of Concern in aceasta Plants. Pennsylvania Dept. Conservation and Natural Re- sources, Harrisburg, PA. SEYMOUR, ¥ Cc. 1982. The Flora of New England. Phytologia Mem VERMONT NONGAME AND NATURAL HERITAGE PROGRAM, 2000. Vermont s Un- common and Rare Plant Species. Vermont Dept. Fish and Wildlife, Wa- ter T, WAHL, HL. - 1954. A preliminary study of the genus Chenopodium in North America. Bartonia 27: 1—46. WEATHERBEE, P. B. 1996. Flora ue ‘coum County, Massachusetts. The Berkshire Museum, Pittsfield, RHODORA, Vol. 104, No. 920, pp. 429-432, 2002 NOTE THE DELETION OF CYPERUS HERMAPHRODITUS (CYPERACEAE: TETRAGONI) FROM THE LOUISIANA FLORA DAVID J. ROSEN U.S. Fish and Wildlife Service, 17629 EI] Camino Real, Suite 211, Houston, TX 77058-3051 STANLEY D. JONES Herbarium, Botanical Research Center, PO. Box 6717, Bryan, TX 77805-6717 Fieldwork in Louisiana produced collections of Cyperus thyr- siflorus Jungh. (Rosen 789, 1084, NO). We reviewed Thomas and Allen (1993) for information on the distribution of C. thyrsiflorus in Louisiana and found this species synonymized under C. her- maphroditus (Jacq.) Standl., a putative, yet largely allopatric Neo- tropical ally of the poorly circumscribed Section Tetragoni (Cart- er and Jones 1997). This classification problem also occurs in the Manual of the Vascular Plants of Texas (Correll and Johnston 1970), erroneously extending the range of C. hermaphroditus into the southeastern United States. In his revision of the Mexican species of Cyperus, Tucker (1994) also included Texas in the distribution of C. hermaphroditus. Preparation of the Vascular Plants of Texas by On et al, (1997) brought to light that the name C. hermaphroditus had been misapplied to specumens On, thyrsiflorus, thus eas the deletion of C. hermaphroditus from the Texas flora. In Horvat’s (1941) revision of the subgenus Mariscus, she reported a collection of C. hermaphroditus trom Arizona, and an evident “fugitive” from Alabama, apparently the only collections for North America. The remaining collections are from Central America, South America, and Mexico. We have examined specimens identified as C. hermaphroditus from Lou- isiana (NLU, NO) and found them to be C. thyrsiflorus and C. pilosus Vahl. We, herewith, wish to continue the clarification of the taxonomic confusion associated with these two species in the southeastern United States by proposing the deletion of C. her- maphroditus from the Louisiana flora. 429 430 Rhodora [Vol. 104 The following key separates Cyperus ee from C. her- maphroditus and C. pilosus. Though C. hermaphroditus does not occur in Louisiana, it is included for comparison. Cyperus pilosus apparently has often been mistaken for C. thyrsiflorus in Lout- siana, and therefore is included in the key. Pertinent synonymy following Jones et al. (1997), a brief description modified from Tucker (1994) and Carter and Jones (1997), and an illustration of C. thyrsiflorus are also provided. — [Rees BIADOUS: 45-44 eeeone ekese wee hus ea eee eee ees (2) 2. Spikelets remote, 7—9 per 5 mm rachis span in proximal half of rachis; achenes narrowly elliptic to oblong, 0.5— 00 5 MAN Wide so bd. ba at eee eke C. thyrsiflorus 2. Spikelets more congested, 18-26 per 5 mm rachis span in distal half of rachis; achenes elliptic to oblong to nar- rowly obovate, 0.6—0.8 mm wide C L J | peer a ve FECEPIUCEPATED UCECL MY li; RAGHIS BUITONSG]Y SINCOSS: occ: 55.5664 aes C. pilosus Cyperus thyrsiflorus Jungh. Linnaea 6: 24. 1831. Figure I. = C. anceps Liebm., C. SS C. Nees Torr, C. pallens (Lieb) Standl. & Steye . regiomontanus vat. es (Liebm.) Ktik., C. tribrachiatus (L. rie ) Ktik., Mariscus dissitiflorus (C. Nees ex Torr.) C. B. Clarke, M. pallens Liebm., M. tribrachiatus Liebm. | Rhizomatous perennial, 20-40 cm tall. Mid-culm diameter 0.5—1.5 mm, trigonous, smooth. Leaves 0.8—2.8 (—3.0) mm wide. Inflorescence rays (2—) 3—6; peduncles conspicuous; inflorescence bracts 5—7. Spikes oblong to subglobose; spikelet length 3.4—7.4 (-17.0) mm, mostly divaricate; scale length 2.0-3.0 mm long, veins and margin whitish. Stigmas 3, stamens 3, mature achenes 1.8-2.1 mm long, 0.4—0.45 mm wide, trigonous, brown. Infre- quent in dry-mesic woods, more common in waste places and disturbed areas from Florida west to Texas, the Caribbean, Mex- ico, and South America. SPECIMENS EXAMINED: Loutstana: Ascension Parish, 18 May 1999, Rosen 789 (NO); Avoyelles, 11 Oct 1985, Thomas et al. 9408S (NLU): — aie 6 Sep 1984, Thomas et al. QOSOS (NLU); East pee Rouge Parish, Jul 1934, Chilton & Trotter 104 (NL er Parish, 22 Jun 1983 “Tham & “a oy 54393 (NLU); ie Gon ek Il Sep 1980, Dane 1922 (NLU) 2002] Note 431 Figure 1. Cyperus thyrsiflorus. A. spikelet showing overlapping fertile scales (bar = | mm); B. habit (bar = | cm); C. spike showing oblong shape and remote, mostly divaricate spikelets (bar = | mm). Drawn from Rosen 789. 432 Rhodora [Vol. 104 Orleans Parish, 28 Jul 1974, Thomas et al. 40637 (NLU); Plaquemines Parish, 4 Sep 1978, Fleming 395 (NO); St. Bernard Parish, 17 Jun 1960, Lemaire 628 NO); St. pe Parish, 28 Apr 2000, Rosen /O84 (NO); Terrebonne ce 16 Jun 1991, mas et al. 123938 (NLU); Vermilion Parish, 11 Jul 1989, ee 997 (NLU); West Feliciana Parish, 14 Aug 1972, Curry ef 7 469 NLU). aS ACKNOWLEDGEMENTS. Our appreciation 1s extended to the her- barium staff at Tulane University (NO) and Northeast Louisiana University (NLU) for their prompt response to our specimen loan requests. Two reviewers provided useful comments on an earlier version of this manuscript. Mr. Eddy Dawson prepared the illus- tration. LITERATURE CITED CARTER, R. AND S. D. JONES. 1997. Notes on the Cyperus retroflexus rai Cyperaceae) with ies nomenclatural et ae Rhodora 99: 319 CorRELL, D. S. AND M. C. JOHNSTON. 197( anual of the Vascular Plants of Texas. Texas Raclack Foundation, ee TX. Horvat, M. L. 1941. A revision of the subgenus Mariscus found in the United States. Catholic Univ. Amer., Biol. Ser. 33: 1-147. Jones, S. D., J. K. Wiprr, AND P. M. MONTGOMERY. 1997. Vascular Plants ¢ DA Ca Checklist including Synonymy, iblioumohy: pa aaa Univ. Texas Press, Austin, TX. THOMAS, R. D. AND C. M. Att 1993. Atlas of the Vascular Flora of Lou- isiana, Vol. 1: Fern Allies, Conifers, and Monocotyledons. The Natural Penns Program, Wildlife Div., Louisiana Dept. Wildlife and Fisheries, Baton Rouge, LA TucKER, G. C. 1994. Revision of the Mexican species of Cyperus (Cypera- ceae). Syst. Bot. Monogr. 43: 1-213 RHODORA, Vol. 104, No. 920, pp. 433, 2002 NEW BOOKS Annotated Checklist of the Vascular Plants of the Washington— Baltimore Area: Part Il, Monocotyledons by Stanwyn G. Shetler and Sylvia Stone Orli. 2002. 95 pp. (softcover). Published by Botany Section, Department of Systematic Biology, National Mu- seum of Natural History, Smithsonian Institution, Washington, DC. [complete checklist is available in pdf format at www.nmnh.si.edu/botany/projects/dcflora | Field Guide to Liverwort Genera of Pacific North America by W. B. Schofield. 2002. viii + 228 pp. illus. line drawings. ISBN 0- 295-98 194-6 $25.00 (softcover). Published by the Global Forest Society in association with the University of Washington Press, Seattle and London. Plant Systematics: A Phylogenetic Approach, Second Edition by Walter S. Judd, Christopher S. Campbell, Elizabeth A. Kellogg, Peter F Stevens, and Michael J. Donoghue. 2002. xvi + 576 pp. illus. line drawings; black & white photos. ISBN 0-87893-403-0 $86.95 (hardcover). Sinauer Associates, Inc., Sunderland, MA. [companion CD-ROM with over 2200 color photographs includ- ed] The Wild Orchids of Arizona and New Mexico by Ronald A. Coleman. 2002. xiii + 248 pp. illus. 32 plates of color photos; dot distribution maps. ISBN 0-8014-3950-7 $39.95 (hardcover). Cornell University Press, Ithaca, NY. RHODORA, Vol. 104, No. 920, pp. 434-438, 2002 NEBC MEETING NEWS September 13 Field Trip. Fourteen NEBC members assembled in a light, misty rain at Kettle Pond in Groton State Forest, Ver- mont, for a leisurely field trip. Art Gilman introduced the area and pointed out the salient landscape features. Groton State Forest is the largest state-owned parcel in Vermont with nearly 26,000 acres of managed forest lands. The area is underlain by the gra- nitic Knox Mountain pluton, which outcrops in the numerous hills, and the soils are acidic and relatively nutrient-poor, being derived from glacial till of mostly local origin. Leaving the parking lot, the first item of interest was a severe gall problem noted on the leaflets of Rhus typhina; these large (marble-sized) hollow galls were filled with insects that Don Mill- er tentatively identified as Homoptera (Ap/is). Further along the trail, the ericaceous shrub community dominant along the shore- line of Kettle Pond included Kal/lmia angustifolia, Vaccinium myr- tilloides, Chamaedaphne calyculata, Rhododendron groentandi- cum, and R. maximum. Also present were typical associates such as [lex verticillata and Nemopanthus mucronatus, the latter in particularly handsome fruiting condition. The numerous shrubs of R. maximum were observed in healthy condition despite their lo- cation near the eastern wind-exposed shore of the pond. They bore numerous capsules and had obviously flowered abundantly this year. Here and at other stations in Groton State Forest the species 1s disjunct from its main range by approximately 100 miles. The group next crossed Route 232 to the old railroad bed, now a popular hiking trail. Underneath a large granite boulder along the side of the trail was a small stand of the uncommon luminous moss, Schistostega pennata. Due to drought conditions, the typ- ically reflective protonemal mat could not be observed, but the tiny feather-like fronds were readily observed with a hand lens. A short hop by car brought the group to Owl’s Head, by which time the rain had stopped and the clouds lifted to provide excel- lent views of Kettle Pond and the southern portions of the Forest. The bald granite knob, although highly trampled by hikers and sightseers, nevertheless provided numerous items of botanical in- terest. Potentilla tridentata and Solidago simplex subsp. randti var. randii were evident, and various shadbushes (Amelanchier spp.) were discussed without reaching consensus. A highlight for 434 2002] NEBC Meeting News 435 many was a small tree of the high-elevation Sorbus decora (here at 1900 ft.) with large orange fruit and short blunt leaflets. This was easily compared to an adjacent specimen of S. americana with smaller, slightly redder fruit. A brief search for Rhododen- dron canadense, although known from Owl’s Head, failed to re- veal this emblem of the Club’s official publication. On the short hike down to the parking lot, Melanie Schori pointed out script lichen (Graphis scripta) on the bark of several trees, and Don Lubin was able to find a small stand of Diphas- tastrum habereri. At the end of the trip, the skies promptly cleared to bright sunshine as members returned to their cars for the trip to St. Johnsbury for the evening meeting. September 2002. The evening meeting was held at the Fair- banks Museum and Planetarium in St. Johnsbury, Vermont. Vice President Arthur Gilman introduced Marcia Spencer-Famous, who spoke to the Club on *‘The Feasibility of Peatland Resto- ration.”’ Marcia and her husband, Norm Famous, have teamed to study the possibility of restoring raised peatlands following ex- traction, or mining, of the peat. This issue has become of special interest because extraction of horticultural and fuel peat using processes that drain and remove peat over large areas started in the twentieth century. In North America, most of such activity is in Canada, with only limited extraction in the United States. Marcia began by reviewing the formation of raised peatlands (raised bogs), stressing that the hydrologic regime of these sys- tems results from a peat accumulation process, which takes thou- sands of years and is an integral part of the resulting ecosystem. Because horticultural peat, being largely the partially decomposed remains of Sphagnum, retains water in large amounts, such sys- tems are similar to saturated sponges with the upper layers above the regional groundwater level. They can range from relatively simple systems to large complexes that are a mosaic of multiple domes, secondary ponds, and a variety of other wetland types, as Marcia amply illustrated with aerial photographs. In addition to Sphagnum, raised bogs host a suite of plants adapted to acidic conditions, low nutrient availability, and saturated organic soils. In addition to woody ericads, some plants commonly found in raised peatlands include Rubus chamaemorus, Geocaulon livi- dum, Calopogon tuberosus, and Eriophorum vaginatum var, spis- sum. 436 Rhodora [Vol. 104 Production of horticultural peat involves developing a bog by excavating perimeter (primary) ditches, installing cross-drains called field (secondary) ditches, removing vegetation over large areas, and crowning the areas between the field ditches to form mining fields. During the summer each year, the surface is sear- ified to promote airdrying and the top 4 to % inch of peat is mechanically vacuumed or removed using a milling process. Typ- ically, up to 4 inches of peat is removed per year. Until the last two decades, in-kind restoration of peatlands abandoned after mining was not a priority, but today’s environmental laws and ethics are forcing a new look at the situation. When merely abandoned, mined bogs present a variety of en- vironmental problems that make reestablishment of any wetland vegetative cover, not to say restoration to original community, extremely difficult. High soil acidity, low and/or changed nutrient levels, changes to the soil structure and the hydrologic regime, drought-like surface conditions (caused by drainage and crowning of the fields) alternating with seasonally flooded conditions, wind erosion, Water erosion during storm events, hydrophobic surface crusting, and frost heaving all are difficult to overcome. Investigations into natural recolonization patterns have found that plant succession does not follow the pattern of the original bog development. Typical pioneers are cotton grasses (Eriopho- rum spp.) and birches (Betula spp.). Cotton grasses typically die after 10-15 years, but their tussocks form moist microniches, sometimes aiding the slow return of Sphagnum. However, Sphag- num, SO critical to the community, is not typically a pioneer genus and may not colonize for several decades. Under good conditions, bog species such as crowberry (Empetrum nigrum), leatherleat (Chamaedaphne calyculata), and other ericads, or larch (Larix laricina) colonize eventually, but total cover may not happen for an extended period. For example, after 20 years poorer sites may have only 5—10% cover, while sites with better growing condi- tions may achieve 50-75% cover. The best sites may achieve 100% vegetative cover, but even when this occurs the results are usually not equivalent to the original peatland community. For example, one abandoned extraction area developed a complete cover of leatherleaf (C. calyculata), but still had no Sphagnum established within it. To obtain a self-sustaining wetland plant community, a number of conditions, especially soil saturation, are required. Rewetting = + 2002] NEBC Meeting News 437 is sometimes achieved by blocking drainage ditches, leveling crowned fields, creating berms to retain precipitation on site, and flooding, where possible. Even with such manipulation, sites are often not wet enough to support establishment of Sphagnum. Studies of a particular system in England found that the com- munity established today, 500 years after the initial extraction of peat, was still dissimilar to the original community. Thus, while plant communities can eventually become established on peat ex- traction sites, restoration to a state equivalent to the original peat- land is not likely to be achieved in the short term, especially if plans for development are not made with restoration principles in mind. Much research has been conducted over the past two decades in Europe, Canada, and, to a lesser extent, the United States. Investigations into the recolonization of Sphagnum, rewetting techniques, and edaphic changes from drainage, among other top- ics, have led to a better understanding of mined peatland man- agement. This knowledge has facilitated the development of man- agement recommendations for restoring mined peatlands to func- tional ecosystems, 1f not to their original condition. —ARTHUR V. GILMAN, Recording Secretary pro tempore. October 2002. President Paul Somers introduced Past-President C. Barre Hellquist who spoke to the Club on **Dodging crocodiles in tropical Australia for aquatic plants.” Barre spoke at length of his tenth trip to Australia since the 1981 Botanical Congress (Syd- ney), which served as a follow-up trip to his 1997 sabbatical research. Like that past sabbatical endeavor, this two-month ex- pedition included teaming up with Surrey Jacobs of the Royal Botanic Garden — Sydney. This fieldwork focused primarily on the aquatic genus Nymphaea, the water-lilies. Barre’s quests for aquatic plants took him from the northern- most point on the mainland, the tip of the Cape York Peninsula in Queensland, through the rugged Kimberley at the northern end of Western Australia. In all, this venture carried him by train, plane, automobile, and helicopter to some of the most remote places for fieldwork. Australia is home to numerous plants adapted to its permanent ~~ or temporary freshwater bodies, and serves as the center of di- versity for several groups. The essentially cosmopolitan Menyan- 438 Rhodora IVol. 104 thaceae is centered there with three of its five aquatic genera, including Nymphoides (water snowflakes). This genus is most diverse in Australia with 20+ species, including N. exigua, N. cristata, and N. indica. The often-aquatic Haloragaceae 1s also most diverse in Australia with about 20 species of Myriophyllum (water mil-foils) alone, including M. latifolium and M. verrucos- um. Australia also boasts the world’s largest water-lilies and argu- ably some of the most beautiful tropical water-lilies. The genus Nymphaea (Nymphaeaceae) is well represented (subgenera Anec- phya, Brachyceras, and Lotos) in the country and often presents itself as a taxonomic challenge. Unusual flower colors and mor- phological variations are plentiful, and upon further study may be the basis for the naming of new species or hybrids. For ex- ample, the marked floral variation found in N. violacea calls into question its current taxonomic limits. Typically this species has fragrant blue flowers with short stipules. However, atypical white flowered, long-stipuled populations have been found lacking fra- grance. Other populations have exhibited unusual purple-striped sepals and peduncles, yet with otherwise typical flowers. Low Lake in Queensland, which serves as a dumping ground for troublesome crocodiles, hosts a remarkable population of Nymphaea atrans. This is typically a “changeable” species, in which the flower color gradually changes over the course of blooming from bluish-white to pink to dark red. At this locale, however, the flower color remains constant during the days of anthesis. Other unusual variations include an odorless night- blooming N. pubescens; a diminutive, faint-smelling, day-bloom- ing N. noucheli; and a white-flowered N. tmmutabilis. There was a special variant discovered in Queensland with less bronze-col- ored foliage and purple flowers that may be described as a new species. In the Kimberley region, one of Australia’s last frontiers, other notable Nymphaeaceae were observed. An unusual population of “Nymphaea immutabilis” was found as well as typical Ondinea purpurea. Ondinea is the only monotypic genus in the water-lily family and is endemic to Australia. Attempts to cultivate this genus, as well as other Australian water-lilies, have been largely unsuccessful. —DONALD J. PADGETT, Recording Secretary pro tempore. REVIEWERS OF MANUSCRIPTS 2001-2002 The Editor-in-Chief of Rhodora is grateful to the members of the editorial staff and to each of the following specialists for their participation in the review process. Their conscientious and thor- ough evaluation of manuscripts helps to maintain the quality of this journal. Susan Aiken Kelly Allred Ihsan Al-Shehbaz Connie Asmussen Theodore Barkley David S. Barrington David E. Boufford Steven Broich C. John Burk Richard Carter Steven Clemants Brian Drayton Debra A. Dunlop Brian Duval Mercedes S. Foster Susan M. Galatowitsch Lara Gengarelly Gerald Guala Steven N. Handel Charles R. Hart Robert R. Haynes C. Barre Hellquist R. James Hickey Matthew Hickler Charles N. Horn John Kartesz Helen Kennedy Thomas G. Lammers Donald H. Les Leonard Lord Paul J. M. Maas James Macklin Arthur C. Mathieson Leslie J. Mehrhotf J. K. Morton Glenn Motzkin Bengt Oxelman Judith Pederson John Pruski A. A. Reznicek Nur P. Ritter Craig W. Schneider Paul Somers Bruce A. Sorrie Lisa A. Standley Edmund W. Stiles Anne Stork John L. Strother John W. Thieret Gordon C. Tucker Michael A. Vincent Edward G. Voss Robert T. Wilce Michael J. Wynne 439 INDEX TO VOLUME 104 New scientific names are in bold face. Alien species 219-252, 350-372 Alix, Mitchell S. 373-395 Allozyme evidence for the hybrid or- igin of Desmodium humifusum (Fabaceae), 253-270 Allozymes 253-270 Aneura maxima (Hepaticae: Aneura- ceae) in Maine, U.S.A. 77-82 New England Note) Aneura maxima, A. pinguis 77-82: morphology 78-81 Aneuraceae 77-82 ANNOUNCEME rs: Invasive Plant Survey of New England: A call for volunteers 106 Merritt Lyndon Fernald Award 218 NEBC Graduate Student Research Award 323 Apocynaceae 170-185, Apocynoideae 170-185, Aquatic plants 373-395 Aristida purpurea vat. 304—308 Asteraceae |—I- Astragalus pulsiferae 271-279; var. pulsiferae 274, var. suksdorfti 274-275; var. nov. 276-278 273-274 186-200 186-200 curvifolia coronensis, var. Backus, Richard H., Pamela T. Pol- . Brian L. Reid, Paul Somers, and Theodore O. Irickson. The dee of Penikese ae Mas- fifth urvey emphasis on sachuse The (199 oy. with the woody vegetation. 219—2 Baltzer, Jennifer L., Heather L. Hew- lin, Edward G. Reekie, Philip D. Taylor, and J. Sherman Boates. The impact of flower on seedling recruitment in sea lav- : key to varieties of harvesting (Limonium Plumbaginaceae). 280—29° Barrington, David S. 92-95 Bertin, Robert I. Losses of plant) species from Massachusetts. 325—: Bertin, Robert Karen B. Searcy, and Paul Somers. A new native plant for Massachusetts, Carex 201-204 ender carolinianum, 5 native Worcester, 9 backti (Cyperaceae). (New England Note) Bidens beckii 373-395 Biodiversity 325—349 Biogeography 117-133 Biogeography of Sfewartia (Camel- lioideae, The ribbsomal DNA Phylogenetic ships and, 117-133 oe invasion 151—160 3oates, J. Sherman 280-295 Boltonia decurrens \—\3 BOOK REVIEWS: Bioconservation and Systematics: Proceedings e Canadian Botanical Association ence Symposium in London, Ontario, June 2000. 212-214 Flora of New Brunswick, Second Edition: A Manual for Identifi- cation - the Vascular Plants of >w Brunswick. 312-31: Lichens of North America. 96—99 Seventh Catalog of the Vascular Plants of Ohio. 208—21 1 Marlin L., Karel A. Lawrence W. Zettler, and Tonya Wilson Delaney. Crossing effects and experimental iceae) inferred from nuclear ITS se- quences. relation- Confer- Bowles, Jacobs, ed viabilit germination of the Federal Threat- ened Platanthera ee (Or- on se chidaceae). 14—3¢( Breeding system ee 30 Bromus eae 304-308 440 2002 | peat pringlet 304—308 Californ —279 Canada 33 —§5, 151-160 Carex backit 201-204 Castillo-Campos, Gonzalo 304—308 Cawly, John |-1- Checklist for contributors to Rhodo- 07-113 pe al as pl foggit 422-428 aera Bay 309-311 ees 396-42 | Clifton, Glenn 271-279 Conant, David S., Gerald J. Gastony, id S. Barrington. Rolla Milton Tryon, Jr. 1916-2001: Sci- entist, Teacher, and Mentor. 92—95 (In Memoriam) Connecticut 161—169 onservation 14—30, 373 Cotoneaster divaricatus naturalized in Massachusetts. 303 (New England Note) Cotoneaster divaricatus 302—303 Crossing effects on seed viability and ee germination of Federal Threatened Platanthera leucophaea a 14—30 Cuyahoga County, Ohio 350—372 —395 (Rosaceae) 302— =n a se aes 83-85, 201-204, 205- , 429-432 oo hermaphroditus, deletion from the Louisiana flora 429-432 key to separation of C. heme: phroditus, C. pilosus, and C. thyr- siflorus 430 Dan, Yu, Wang Dong, Li Zhen-yu, and A. M. Funston. Taxonomic re- vision of the genus Myriophyllum (Haloragaceae) in China. 396—42 Del Tredici, Peter 117—133 Delaney, Tonya Wilson 14—30 Deletion of Cyperus hermaphroditus Cyperaceae: Tetragoni) from the The, 429-432 Louisiana flora. (Note) Desmodium ere D. panicu- al tundifolium 253-270 Dimorphic aa 1-13 ~ ~ ~ Index to Volume 104 Dong, Wang 396-421 Donoghue, Michael J. 117-133 Duckweeds 373-395 Effect of achene morphology and mass on germination and seedling erowth of Boltonia decurrens (As- teraceae), a threatened floodplain species. 1-13 Padansered mneraphytes 373-395 Endangered S: oo 201-204; Indiana 373-3 Erioneuron avenaceum var. avena- ceum 304—308 Erysimnum hieractifolium, new record 299-301 European water-horehound 151—160 Exotic species 151-160 Fabaceae 253-270 Fernaldia 186— 200; F. marae 96; 189-192; F. pandurata 192— F. ee ey 196-198; i 189 Floodplain 1-13 Flora of Penikese Island, Massachu- setts: The fifth survey (1998— 1999), with emphasis on_ the woody vegetation. The, 219-252 Flora: Florida 42—76; Indiana 373-— 395; Ohio 350-372; Penikese Is- land 219—252; Worcester 325—349 Florida 42—76 Floristic inventory of Springs er Park, Levy Florida. 6 Floristic me oe Flower harvesting 280—295 Freeze/thaw cycles 161—169 Funston, A. M. 396-421 Manatee County, Gastony, Gerald J. 92—95 Gentianales 186— a Germination 1-13, 14-30 Gulledge, ee J. and Walter S. Judd. A floristic inventory of Manatee Springs State Park, Levy County, Florida. 42—76 442 Habitat destruction 325—349 Haines, Arthur. Occurrence of Scir- pus georgianus (Cyperaceae) in Maine 207 (New England Note) Haines, Arthur. A new combination in Lycopodiella (Lycopodiaceae). 296-298 (New England Note) Haines, Arthur and Betsy W comer. New records for Chenopo- dium foggeit in New England, 422— 428 (New England Note) Sane 396-42 Hartia 117-133; distribution 118: phylogeny 124-128 Hay, Stuart G. and Gordon C. Tuck- er. Sci (pus ANCISIFOC haetus (C ye record in Canada. map in) ee 83-85 Helic Ses pubescens, new re- cord 299-301 Hendrickson, Theodore O. 219-252 oo - wlin, Heather L. 280-295 Hybriivation, Desmodium humifus- wn 253-270 Hybrids 350-372 Identity and history of Myrica caro- liniensts (Myricaceae). The, 31-41 Impact of flower harvesting on seed- ling recruitment in sea lavender (Limonium slag ort Plum- baginac 8 IN MEMORIAM: Rolla Milton Tryon, Jr 1916— Scientist, Teacher, and Mentor. 92— Indiana 373-395 Invasive species 151—160 ITS (Internal transcribed spacers) —133 Jacobs, Karel, A. 14-30 Judd, Walter, S. 42—-7¢ Juniperus virginiana 219-252 Lachance, Daniel and Claude Lavoie. Reconstructing the biological in- Rhodora [Vol. 104 vasion of European water-hore- hound, Lycopus europaeus (Labia- ae), along the St. Lawrence River, Québec. 151-160 Laubertia 170-185; L. boissieri 76-178; L. contorta 178-181; Lh peninsularis 181-183; key 175— - 176 Lavoie, Claude 151—160 Leguminosae 271—279 Lemna valdiviana 373-395 -vy County, Florida 42—76 Li, ae Peter Del Tredici, iong Yang, and Michael J. ghue. Phylogenetic relationships and biogeography of Stewartia (Camellioideae, Theaceae) in- ferred from nuclear ribosomal DNA ITS sequences. 33 mai enna aa 295 ycal extinction 280— se of native a a from Worcester, Massachusetts. 325— 349 Shix- Dono- ei Louisiana 429—432 Lycopodiaceae 296-298 Lycopodiella Xrobusta, comb. et stat nov. 296-298 51-160: distri- Lawrence Lycopus europaeus bution map along St. River 154 Maine 77-82, 205-207, 422—428 Manatee Springs State Park, Florida 42-76; plant community map 48; checklist of vascular plants 60—76 Massachusetts 201-204, 219-252, 299-301, 302-303, 325-349, 422-423 McCombs, Martha R. 350—372 McDevit, Daniel C. and Craig W. Schneider. The survival of Vauch- eria (Waucheriaceae) propagules in New England riparian sediments alter repeated freeze/thaw cycles. 161—169 Mejia-Saulés, Teresa, Gonzalo Cas- tillo-Campos, and Sergio Avenda- no Reyes. New reports of Poaceae 2002| in the rocky substratum of Munic- ae of Perote, Veracruz, Mexi- co. 304—308 (Note Te 304—308 Miller, Norton G. Aneura maxima (Hepaticae: Aneuraceae) in Maine, U.S.A. 77-82 (New) England Morales, J. Francisco. Studies in neotropical Apocynaceae I: A re- vision of the genus Laubertia. Morales, J. Francisco. Studies in neotropical Apocynaceae II: A re- view of the genus Fernaldia. 186— 200 Morella caroliniensis, M. cerifera 31-41 Morphology, Aneura maxima and A. pinguis 77-82 Muhlenbergia glabrata 304-30 Myrica caroliniensis, identity history of 31-41 Myrica caroliniensis, M. cerifera, M. cers M. pensylvanica 3-4 himieace pa anne in ane 396-421: M ee — 400; M. aqua- ticum 400— . dicoccum 401403: M. Teter 4 403— 404; M. oguraense —405; M sibiricum 405-407; M. a I( tetrandrum M. eniereares 411—412; ussuriense 412-414; M. verti- ees 414-416; i to species so oe in China 397— ; distribution maps 416-418 a llum pinnatum 373-395 and spicatum 410— Naiads 373-395 Najas grac pete 373-395 Native species 325-349, 350-372 Naturalized species, Massachusetts 302-303 eee Meeting News 101-105, 215- , 317-322, 434-438 scnc Membership Form [14 Index to Volume 104 Neotropics 170-185, 186—200 New Books 100, 316, 433 New combination in) Lycopodiella (Lycopodiaceae). A, 296-298 New England ee New England 422—428, New native plant for 7s Carex backii (Cyperaceaec). A, 201-204 (New England Note) New records: Canada 83-85; Indiana 373-395; Maine 7 28: ene 201-204, 299- 301, 422-428; Me pee North C aeoling 86- ol: Ohi 350-372 New records for Cea. foggil in New England. 422—428 (New England Note) New records of vascular plants for Ohio and Cuyahoga County, Ohio. 350-372 New records with ecological notes for rare aquatic vascular plants in Indiana. Part 1. 373-395 New reports of Poaceae in the rocky substratum of Municipality of Per- ote, Veracruz, Mexico. 304—308 (Note) New variety 271-279 Newcomer, y W. 422-428 Bets North Carolina 86-9] Nova Scotia 280—2 Nuclear ribosomal 117-133 5S DNA (nrDNA) Occurrence of Scirpus (Cyperaceae) in Maine. 205—2 New England Note) Ohio 350—372 Ondricek, Robin on Orchidaceae 3 Orchids 14—30, eT Order Form, Index to Volumes 76— 100. 115, 324 georgianus O7 a7 Parker, Bruce C. 309-311 Penikese Island 219—252 Phylogenetic relationships and_ bio- geography of Stewartia (Camel- lioideae, Theaceae) inferred from 444 peep DNA ITS se- 117-13 Phylogeny 117-1 3 Platanthera leuc ili as Plumbaginaceae 280— Poaceae 304—308 nuclear quences. Pollination 14—30 Pollomi, Pamela . ae 252 Pondweeds 373— Potamogeton a a P. bic siege tus 385; P. epihydrus 380-387; pulcher 387-389, P. vaseyi . 390 Prairie Peninsula 373-395 Prescribed burns 219—252 Propagules 161—169 Quebec 83-85, 151-160 Quercus 134-150; VIrginiana 136; Q. pees 136-137: QO. in- cana ie elliottti, sp. nov. 138- *O: mic paul 140-141; . marilandica 141-142: O. nigra 2705 ta 142-144; Q. laevis 8 oe alba 145: QO. lyrata 45: 2 sinuata 145-147; O. stel- ae Rare and endangered macrophytes 373-395 Rare species: Indiana 373—395; Maine 205-207; New England 253-2 Penikese Island 219-252; Piciaith: ~ —_ era leucophaea |4—3¢ Raveill, . Allozyme evidence for the hybrid origin of Desmo- dium humifusum (Fabaceae). 253— 270 Recent plant collections—Massachu- etts. 299-301 (New. England ae inva- water-hore- hound, Lycopus europaeus (Labia- tac), me - ] Lawrence River, Québec. Re orien In sea ee 280-295 Red cedar 219-252 Reekie, Edward G, Reconstructing the biological sion of European 280-295 Rhodora [Vol. 104 Reid, Brian L. Review of the genus Fernaldia 186— 200 Reviewers of Manuscripts 439 Revision of the 170-185 apa of the genus Myriophyllum Haloragaceae) in China 396—421 Re ne Sergio Avendano 304-308 Richard J. LeBlond and Alan’ S. Weakley. Schizaea pusilla in N Carolina. 86-91 (Note) Riparian sediments 161—169 Rolla Milton Tryon Jr., Scientist, Teacher, and 92-95 (In Memoriam) Rosaceae 302-303 Rosen, David J. and Stanley D. Jones. The deletion of ln hermaphroditus (Cyperaceae: Te- tragoni) from the Louisiana flora. 429—432 (Note) 219-252 genus Laubertia a ~ ~ 1916-2001: Mentor. Rosenzweig, Michael S. and Bruce C. Parker. Turion production by Ruppia maritima in Chesapeake Bay. 309-311 (Note) Ruppia maritima 309-31 | 280-295 Schizaea pusilla in North Carolina 86-91 (Note) Schneider, Craig W. 161-169 Scirpus ANCISTVOC hac Tus (Cypera- >): First record in Canada. 83— Salt marsh Scirpus georgianus 205-207 Scribailo, Robin W. and Mitchell S. lix. New records with ecological notes for rare vascular plants in Indiana. Part 1. 373-395 Sea lavender 280—295 Searcy, Karen B. 201—204 Seed banks ees. as 295 Seed dispersal 280— Seed germination ia Seed size |—I- caine growth |-13 Seedling recruitment in sea lavender 280-295 aquatic 2002 | Setarta reverchonit subsp. ramiseta Marian and John Cawly. Ef- fect of achene morphology and mass on germination and seedling growth of Boltonia decurrens (As- pepeeue a threatened floodplain species. I-13 Somers, Paul 201-204, 219-— South Carolina 134-150 Species loss 325—349 St. Lawrence River 151—160 Statement of Ownership 116 Stewartia 117-133; seal map 118; phylogeny 124—12 Studies in neotropical ee I: A revision of the genus Laub- ertia. 170-185 eae in neotropical Apocynaceae ‘the genus /ernal- > | oe 186—200 Sustainable harvesting 280—295 Taxonomic revision of the genus My- riophyllum (Haloragaceae) in Chi- na. 396-421 Taxonomy 31-41, 134-150, 170- 85, 186 I —200, 271-279, 296-298, 396-42 | Regt Philip D. 280—295 survival of Vaucheria (Vauch- eriaceae) propagules in New Eng- land riparian sediments after re- peated freeze/thaw cycles. 161— 169 Theaceae 117-133 Thomas Walter’s oaks from_ the coastal region of South Carolina. 134-150 Threatened species: Boltonia decur- Index to Volume 104 445 rens 1—| - nee leuco- phaea \4— Hyon, Rolla en Jr. 92-95 (In Memoriam) Tucker, Gordon C. 83— Turion production by Ruppia mari- 1] tima in Chesapeake Bay. 309—; (Note) 1916-2001 Varieties of Astragalus pulsiferae (Leguminosae). 271-279 Vaucheria 16 : = : _ Virginia 309— Weakley, Alan S. 86-91 Welsh, Stanley L., Robin Ondricek. and Glenn Clifton. Varieties of As- tragalus as (Legumino- sae). 271- Wilbur, seta a The identity and history of Myrica caroliniensis Myricaceae). < Wilbur, Robert L. Thomas Walter's oaks rom the eer region of ilder, George J. ‘and a R, New records of vas- cular plants for Ohio and Cuya- hoga County, Ohio. gs Woody vegetation 219-252 Worcester, Massachusetts 325-349 < aS O 2 O Yang, Shixiong 117-133 Zebryk, Tad M. Recent plant collec- ttons—Massachusetts. 299-30] (New England Note) Zettler, Lawrence, ae 14-30 Zhen-yu, Li 396-421 Zika, Peter FE ees divarica- tus (Rosaceae) naturalized in Mas- sachusetts. 302-303 (New Eng- land Note) —~ ORDER FORM Index to Volumes 76—100 The cumulative Index to Volumes 76-100 of Rhodora is now available. The index is divided into two parts: an index to sci- entific names and an index to authors and subjects, created di- rectly from the journal issues. Copies of the Index to Volumes 1—SO and the Index to Volumes 51-75 are still available. For more information on the first two cumulative indices contact Dr. Cathy Paris, Back Issues Manager (cparis@zoo.uvm.edu) or visit the New England Botanical Club web site (http://www.huh.harvard.edu/nebc/). To order the Index to Volumes 76—100 mail this form and payment to: RHODORA, Allen Press, Inc., PO. Box 1897, Lawrence, KS 66044-8897. Name Address City State. Zp Member $25.00 Other $50.00 funds payable at par in United States currency THE NEW ENGLAND BOTANICAL CLUB Elected Officers and Council Members for 2002—2003: President: Paul Somers, Massachusetts Natural Heritage and Endangered Species Program, | Rabbit Hill Rd., Rt. 135, Westborough, MA 01581 Vice-President (and Program Chair): Arthur V. Gilman, Wm. D. Countryman Environmental Assessment and Planning, 868 Winch Hill Rd., Northfield, VT 05663 Corresponding Secretary: Nancy M. Eyster-Smith, Department of Natural Sciences, Bentley College, Waltham, MA 02154- 4705 Treasurer: Harold G. Brotzman, Box 9092, Department of Biology, Massachusetts College of Liberal Arts, North Adams, MA 02147-4100 Recording Secretary: Neal W. Anderson Curator of Vascular Plants: Raymond Angelo Assistant Curator of Vascular Plants: Erika Sonder Curator of Nonvascular Plants: Anna M. Reid Librarian: Leslie J. Mehrhoff Councillors: Lisa A. Standley (Past President) Judy Warnement 2003 Karen B. Searcy 2004 Kanchi Gandhi 2005 Jennifer Forman (Graduate Student Member) 2004 Appointed Councillors: David E. Boufford, Associate Curator Janet R. Sullivan, Editor-in-Chief, Rhodora