Journal of the CONTENTS Reflections on 100 years of Rhodora. Janet R. Sullivan New England Botanical Club A review of the = _ A ms — Ann Wd Wo oo oO Triadenum fraseri 3 Lysimachia terrestris 2 38.5 Impatiens capensis 8 30.8 Galium trifidum 3 30.8 Chamaedaphne calyculata 6 231 Scutellaria galericulata 5 23.1 Carex canescens 8 15.4 Carex oligos, 13 15.4 Viola macloskeyi 6 15.4 well as the mosses Climacium dendroides (Hedw.) Web. & Mohr. peatlands may have a fringing Jris association less than a meter wide (Gates 1942). At Little Dollar Lake, I. versicolor was a 1999] Hellquist and Crow—Little Dollar Lake Peatland 63 prominent component of the C. canadensis CT with a distribution several meters in width, especially in the lagg areas of the south- ern-oriented peatland mats. Within the Jris versicolor—Lycopus uniflorus PHS there was one locality that had an entirely unique species composition com- pared to the rest of the peatland. This area was situated within the peNe i of the southwestern mat (Figure 1) in an area that ap- ently was influenced by runoff from a seasonal, dirt truck trail (Heliquist 1996). In this area, Typha latifolia L. was well estab- lished. Futyma (1982) noted the presence of Typha in the basin in the early 1980s, but made no reference to the location of the colony. Other species essentially limited to this distinct lagg hab- itat were Carex stipata Muhl., Epilobium ciliatum Raf., Polygo- num cilinode Michx., Rubus canadensis L., Scutellaria laterifiora L., and Rumex obtusifolius L. 1B. Sphagnum cuspidatum—Dulichium arundinaceum Phase The Sphagnum cuspidatum—Dulichium arundinaceum PHS oc- curred in two distinct areas in the southeastern and western lagg areas of the southern mat extension (Figure 3). This association was apparent in the wettest areas of lagg that contained exposed, mucky peat. In 1996, this area was covered by standing water roughly 0.25 to 0.50 m deep. The most frequent vascular species of this phase included Calamagrostis canadensis, Carex ad carpa, Chamaedaphne calyculata, and Potentilla palustris. Thirtee species with greater than 10% frequency were present Sona Glyceria canadensis (Michx.) Trin., Carex utriculata F Boott, Equisetum fluviatile L., and Salix pedicellaris Pursh (Table 4). aie dominant peat mosses of this phase were Sphagnum re- curvum and S. cuspidatum (Table 4). Other important bryophyte species included members of the Amblystegiaceae such as Warns- torfia fluitans, W. exannulata (Schimp. in B. S.G.) Loeske [Dre- panocladus exannulatus (Schimp. in B.S.G.) Warnst.], Calliergon cordifolium, and C. stramineum. Despite its infrequency in the quantitative sampling of this phase, W. exannulata formed an almost homogeneous bryophyte cover on exposed, mucky peat in the southeastern lagg of the southern mat extension. In 1996, this same population of W. exannulata was still vigorous despite being submerged in standing water. Warnstorfia exannulata is considered a dominant species of poor fens in Alberta along with Sphagnum angustifolium (C. Jens. ex Russ.) C. Jens., S. majus 64 Rhodora [Vol. 101 Table 4. Mean percent cover and percent frequency of dominant and as- sociated species in the Sphagnum cuspidatum—Dulichium arundinaceum phase of the Calamagrostis canadensis cover type (14 quadrats). Species having less than 10% frequency are not inched in age table. *TWINSPAN indicator species. Mean % Cover % Frequency BRYOPHYTES Sphagnum recurvum 38 21.4 Sphagnum cuspidatum* 27 S71 Warnstorfia fluitans 8 14.3 Sphagnum maju | 14.3 Calliergon stramineum 30 14.3 Calliergon cordifolium 27 jf By | VASCULAR PLANTS Calamagrostis canadensis* 14 78.6 Carex lasiocarpa* 43 78.6 Chamaedaphne calyculata 24 50.0 Potentilla palustris S, 50.0 Lysimachia thyrsiflora* 1 me | Triadenum fraseri <1 35:1 Glyceria canade 32 28.6 Dulichium arundinaceum* 20 28.6 Acer rubrum ca | 21.4 Carex utriculata 8 14.3 Vaccinium macrocarpon 3 14.3 Equisetum fluviatile 2 14.3 Salix pedicellaris 23 14.3 (Russ.) C. Jens., and S. jensenii H. Lindb. (Vitt and Chee 1990). Although members of the Amblystegiaceae tend to prefer more minerotrophic microhabitats, W. fluitans and C. stramineum seem to occur in acid to intermediate acid habitats (ca. pH 3.7-6.0; Gorham and Janssens 1992b). At Little Dollar Lake, several emergent vascular species col- onized this rich muck including Carex lasiocarpa, Dulichium arundinaceum (L.) Britton, Glyceria borealis (Nash) Batchelder, Juncus alpinus Vill., Potentilla palustris, and Puccinellia pallida (Torr.) R. T. Clausen. The only locality of Carex chordorrhiza L. f. was in the exposed peat of the southeastern lagg. Carex chor- dorrhiza is considered a species indicative of poor fens in Min- nesota (Wheeler et al. 1983) and rich fen conditions in Alberta (Vitt and Chee 1990). 1999] Hellquist and Crow—Little Dollar Lake Peatland 65 Table 5. Mean percent cover and percent frequency of dominant and as- sociated species in the Chamaedaphne calyculata cover type (220 quadrats). Species having less than 10% frequency are not included in the table. *TWINSPAN indicator species. Mean % Cover % Frequency BRYOPHYTES Sphagnum recurvum 74 83.2 Sphagnum majus 50 36.3 Sphagnum magellanicum* 14 26.5 Aulocomnium palustre 6 1S Sphagnum capillifolium* 18 10.6 VASCULAR SPECIES Chamaedaphne calyculata 46 99.1 Carex oligospe 26 94.7 Kalmia polifoli 15 62.8 Andromeda glaucophylla 11 36.3 cer rubrum 30 L. orientalis 86 4 7 yw 3 Vv AA EH L. styraciflua «JTS x -- matK 0 -- trnL intron --GBSS___OGS -- outgroups Figure 1. The single most parsimonious tree of 535 steps of Liquidambar based on sequences of ITS, matK, GBSS, and trnL intron. CI = 0.98. Num- bers above and below the branches are bootstrap percentages and decay index values, respectively. Symbols represent unambiguous, potentially informative changes of each data set along the branches. The trnL intron data set had 21 variable sites, two of which were phylogenetically informative. The sequence divergences be- tween Liquidambar species were from 0—0.8%. Liquidambar acalycina and L. formosana had identical trnL intron sequences. The parsimony analysis generated two equally short trees, one of which showed the tree topology produced by the ITS data, while the other tree did not resolve the relationships of L. formosana, L. acalycina, and the clade of L. orientalis and L. styraciflua. The consistency index was 1.0. There were 105 variable sites in the GBSS data set, 15 of which were informative. Parsimony analysis resulted in one single short- est tree of 111 steps, with a consistency index of 0.96. In the 90 Rhodora [Vol. 101 phylogenetic tree, eight and seven informative sites supported the clade of Liguidambar orientalis—L. styraciflua and L. acalycina— L. formosana, respectively. Bootstrap values for the two clades were 78% and 82%, respectively. The four data sets were congruent, including matK, ITS, trnL intron, and GBSS, and the combination of them created a data set 164 characters. The parsimony analysis, using both Zxbuck- landia and Mytilaria as outgroups with Mytilaria GBSS sequenc- es coded as missing data, produced a single most parsimonious tree with a consistency index of 0.98. The gE tree showed the same species relationships as described in al (1997a). Both bootstrap percentages and decay values were high, 100% and 18 steps for the L. acalycina—L. formosana clade, and 86% and three steps for the clade of L. orientalis and L. styra- ciflua. Figure 1 shows the number of unambiguous changes from each of the four data sets that support the two clades. This follow-up study strongly substantiates the previous hy- pothesis that the western Asian species Liguidambar orientalis is more closely related to the New World species L. styraciflua than to the southeast Asian species. Additionally, we conclude that sequences of the GBSS gene, especially the introns, provide an- other informative nuclear marker (besides nrDNA ITS) in resolv- ing phylogenetic relationships among closely related species. ACKNOWLEDGMENTS. We thank Robie Mason-Gamer and Toby Kellogg for assistance in designing GBSS gene primers. This study was partially supported by a Putnam Postdoctoral Fellow- ship through the Arnold Arboretum of Harvard University to JL. LITERATURE CITED BaALDwiINn, B. G., M. J. SANDERSON, J. M. Porter, M. FE WOIJCIECHOWSKI, AND - J. DONOGHUE. 1995. The ITS region of nuclear ribosomal DNA: A valuable apt > evidence on angiosperm phylogeny. Ann. Missouri Bot. Gard. 82: ce BocLe, A. L. 1986. pee floral morphology and vascular anatomy of the Ha- mamelidaceae: Subfamily Liquidambaroideae. Ann. Missouri Bot. Gard. 73: 325-347. Dat, W., W. DENG, W. Cul, S. ZHAO, AND X. WANG. 1996. Molecular cloning and sequence of potato granule-bound starch synthase gene. Acta Bot. Sin. 38: 777-784. DonoGuHuE, M. J., R. G. OLMsTEAD, J. E Smitu, and J. D. PALMER. 1992. 1999] Note 91 Phylogenetic relationships of Dipsacales based on rbcL sequences. Ann. Missouri Bot. Gard. 79: 333-345 FELSENSTEIN, J. 1985. Confidence limits on phylogeny: An approach using the bootstrap. Evolution hy 783-791. Hoey, M. T. AND C. R. Parks. 1991. Isozyme divergence between eastern Asian, North American, Cid Turkish species of Liguidambar (Hama- ae Amer. J. Pos 78: 938-947. Li, J., A. L. BOGLE, AND A. S. KLEIN. 1997a. Interspecific SRT and genetic divergence of the disjunct genus Liqguidambar (Hamamelidaceae) inferred from DNA sequences of plastid gene matK. Shindinn’ 99: 229- 0 ——-, AND 997b. Phylogenetic relationships in the Cor- ylopss —— (Hamamelidaceae): Evidence from sequences of the in- bed spacers of nuclear ribosomal DNA and morphology. Rhodora 99. 302 318 AND K. PAN. 1997c. Close relationship between Shandpdeniivon and Parone (Hamamelidaceae), evidence from ITS se- quences of nuclear ribosomal DNA. Acta Phytotax. Sin. 35: 481—493. SworFForD, D. L. 1993. PAUP: Phylogenetic Analysis Using Parsimony, ver- sion 3.1.1. AOE ts of Molecular Systematics, Smithsonian Institu- tion, Washington, Peer, P, L. GrELty, fi PauTOu, AND J. Bouvet. 1991. Universal primers for amplification of three non-coding regions of chloroplast DNA. PI. Molec. Biol. 17: 1105—1109. RHODORA, Vol. 101, No. 905, pp. 92-94, 1999 BOOK REVIEW The Savage Garden: Cultivating Carnivorous Plants by Peter "Amato. 1998. xxii + 314 pp. more than 200 photographs and illustrations, most in color. ISBN 0-89815-915-6 $19.95 (paperback). Ten Speed Press, Berkeley, CA. Carnivorous plants, a group of about 500 species and numerous cultivars, have captivated botanists, horticulturalists, and students of all ages since Darwin (1875) wrote one of the earliest books on the group. The more easily grown species and those locally abundant in boggy areas have been widely employed by teachers to stimulate interest in young students in plant biology and ecol- ogy. Unfortunately, until recently, little information has been available in book form in public and college libraries for teachers and students who want to know more about growing these plants. In recent years, three books with a worldwide scope have been published and widely distributed: Pietropaolo and Pietropaolo (1986), Lecoufle (1989, in translation), and Cheers (1992). These three works cover some of the same ground as the earlier and excellent books by Slack (1980, 1988) which are difficult to ob- tain. Peter D’Amato’s book surpasses these earlier works in the sheer volume of cultivation information and lore provided for individual species and cultivars. It is based on Peter’s many years of experience with growing these plants; of selling them through his greenhouse, mail order and Internet nursery, California Car- nivores; and of making numerous presentations at schools, hob- byist meetings, and on television. His goal, as he described it during a talk at the first meeting of the International Carnivorous Plant Society in Atlanta in 1997, was to popularize the plants and show how they can be grown in a wide variety of ways indoors and out. The range of creative, and sometimes whimsical, ways to grow these plants, well illustrated and described in the book, is its strongest feature. Since the book covers species worldwide, it also provides a starting point for exploring plant diversity and plant geography. The amazing radiation of Drosera species in Australia, and the narrowly restricted endemic genera Darling- tonia, Heliamphora, and Cephalotus which use trapping mecha- nisms similar to Sarracenia and Nepenthes are good examples. 92 1999] Book Review 93 A few minor faults should be noted. Many of the color pho- tographs are small, but this allows for more text to flow around the figures creating a tighter integration of the two. On the other hand, this design foregoes including any large in situ shots such as those in Schnell’s (1976) Carnivorous Plants of the United States and Canada or Clarke’s (1997) truly stunning book on the Nepenthes of Borneo. As noted in the review published in the September (1998) issue of the Carnivorous Plant Newsletter, there are some spelling errors and some of the cultivars listed are not well documented in the literature. Readers interested in an overview of the unique physiological and ultrastructural features of carnivorous plants may wish to supplement this book with the earlier work by Juniper et al. (1989). Some of the guidelines presented for growing individual species may be less effective outside the northern California climate where the author lives. Apparently, no hardcover edition is available. Overall, the well named Savage Garden is a bargain for the wealth of information it contains. As a CP enthusiast, I use it often. It may well achieve “‘bible’’ status for growers of these plants, especially as tissue culture (described in the first Appendix by expert Rob Gagliardo) makes more species available to the general public. I would recommend this book for all libraries, and to botanists and horticulturalists as a one-book gateway to know- ing more about these fascinating plants. LITERATURE CITED Cueers, G. 1992. A Guide to the ‘nail Plants of the World. HarperCollins Publishers, New Yor CLARKE, C. 1997. Nepenthes of a Natural History Publications, Kota Kinabulu, Sabah, Malaysia Darwin, C. 1875. fanectiocninns Plants. John Murray Publishers, London. JuNIPER, B. E., R. J. RoBins, AND D. M. JoEL. 1989. The Carnivorous Plants. Academic Press, London. LECOUFLE, M. 1989. Carnivorous Plants: Care and Cultivation. Cassell Vil- liers House, London. (English translation.) MEyYERS-RICE, B. 1998. Book review. Carniv. Pl. Newslett. 27: 72-7 rete J. AND P. eas 1986. ease etsivit Plants of the World. mber Press, Portland, O Papers D. E. 1976. hae Plants of the United States and Canada. John E Blair Publisher, Winston-Salem, NC. 94 Rhodora [Vol. 101 SLack, A. 1980. Carnivorous Plants. MIT Press, Cambridge, MA. —. 1988. Insect-Eating Plants and How to Grow Them. University of Washington Press, Seattle, WA. —DAvID LANE, Biological Sciences Library, Kendall Hall, Uni- versity of New Hampshire, 129 Main St., Durham, NH 03824- 3590. RHODORA, Vol. 101, No. 905, pp. 95-100, 1999 NEBC MEETING NEWS October 1998. Dr. Lisa A. Standley, the Club’s Vice President, spoke on the topic ‘‘Beyond the Brooks Range—Flora and Fauna of the Arctic National Wildlife Refuge.’”” Among Dr. Standley’s many activities, we learned, is participating in Sierra Club out- ings. Twice in recent years, she has enjoyed ten-day backpacking trips to the Arctic National Wildlife Refuge in northeastern Alas- ka. Both trips were in mid-June, and started with a flight into Fairbanks and transfer to a smaller plane that flew through passes in the Brooks Range to the Romanzof Mountains and the coastal plain of the Beaufort Sea, a couple of hundred miles north of the Arctic Circle. The area hiked was between two rivers, the Jago and the Aichilik, which flow northward from the mountains, crossing the coastal plain to the sea. The Refuge is contiguous with Indian lands and National Parks, which add to the wilderness landscape. It is home to the caribou’s Porcupine Herd calving grounds and their migration routes to the mountains. Fortunately for us, Lisa was armed with a good camera and the ability to use it well. We were treated to excellent images of the region’s plant life, interspersed with those of the often present and possibly curious caribou. Also, landscape shots illustrated some of the Ref- uge’s varied habitats and unadulterated beauty. The hiking area ranged in elevation from near sea-level to around 5000 ft. Most of the landscape is devoid of tall trees, although some of the narrower valleys supported white and black spruce in sheltered areas. Low willows were the more typical woody vegetation. The narrow valleys generally run east-west while larger river valleys run north-south. Precipitation is surprisingly low there, with only about 10 inches per year, Standley said. The few glaciers seen while crossing the Brooks Range were relatively small and not growing, evidently remnants of earlier times with higher rates of precipitation. The slide images gave a good sampling of the dominant plant families in the Refuge and the Arctic region, in general. The Cyperaceae, a family Standley knows especially well because of her research on the genus Carex, is one of them. Sedges were well represented and tipsy tussocks of cottongrass were frequently underfoot. More frustrating for Lisa than the tipsy tussocks, per- haps, was that nearly all the sedges present in June were flowering rather than fruiting, making identification very challenging. An- 95 96 Rhodora [Vol. 101 other family well represented was the Salicaceae. A favorite for Standley was Salix minima which stood less than an inch tall with catkins of reddish flowers. The Saxifrage family was rep- resented by several species of Saxifraga including S. oppositifol- ia, a circumboreal species present in New England, and the very unusual S. eschscholtzii with its tiny cushion-like rosettes of suc- culent leaves only 2 mm across. Jumping to the Rose family, we saw Potentilla hypartica (or P. nana, in some books), a close relative of New England’s federally endangered P. robbinsiana. Also representing the Rosaceae were both species of Dryas. The hikers liked seeing Dryas, since it meant they would be walking on gravel substrate and not wobbly tussocks. Ericads were also represented in the tundra by white flowering Cassiope [or Har- rimanella in some references], pink flowered Rhododendron lap- ponicum described by Standley as weedy everywhere, and Loise- leuria seen at around 5000 ft. elevation. Also illustrated by Standley were: a Douglasia species (Pri- mulaceae) which is endemic to Alaska and the Yukon; a Hedy- sarum (Fabaceae) which has aromatic, edible roots eaten by griz- zly bears; yellow poppies, which trap heat and attract flies in cup- like flowers that tilt toward the sun, which in June’s solstice sky shines for 24 hours per day; nitrophilous, orange-colored lichens growing on rocks where birds perch; caribou trails made in the tundra in the 1940s; caribou skeletons used for drying wet socks; musk-oxen simulating “fringed sofas” swaying in the breeze; ae- rial views of river meanders revealing a hundred or more years of geomorphology; vertical Jurassic formations with marine fos- sils; cliffs with gyrfalcon nest sites: sloping bogs at 4000-5000 ft.; and a grizzly sow with cub. Standley recommended Pielou’s Arctic Naturalist and Birds of laska. 1999] NEBC Meeting News 97 nine sample sites within a continuous, unfragmented habitat of the Presidential Range in the White Mountains of New Hamp- shire. Previous studies on the effects of fragmentation on loss of genetic diversity, he said, have involved habitats with a relatively recent history of fragmentation (i.e., less than a few hundred years) and only one taxon. He thought, by examining high peak populations presumably separated for thousands of years, that the effects of time on genetic drift and genetic diversity might be more apparent. By examining three species, he hoped to reduce the possibility of any erroneous conclusions made by assuming that what is true for one is true for all. Lindwall identified the three key questions he wanted to answer in the study as: 1) Do fragmented plant populations in the Adirondack peaks have less diversity than the continuous population in the White Mountains? 2) Is there more gene flow in the Presidential Range than among the fragmented populations in the Adirondacks? 3) What effect does greater habitat area have on diversity in the White Moun- tains versus the smaller area for each of the isolated Adirondack sites? A fortuitous coincidence of Lindwall’s site design, he added, was that the overall land area and distances between sites for the two study areas were approximately the same. To quantify the relative abundance and frequency of each species, 6000 plots, each one m2, were examined. The genetic diversity was assessed using allozyme analysis. The three species studied were Minuar- tia groenlandica, which appears to be exploiting disturbed trail- edge habitat, Carex bigelowii, which forms large patches in the White Mountains, and Diapensia lapponica, a monotypic genus found in tundra. Three thousand tissue samples were taken and analyzed during the study. For each ines Lindwall’s three questions, the answers were ‘“‘ves,”” “no,” and “‘maybe.’’ Did the fragmented Adirondacks have lower genetic diversity? For Diapensia lapponica, the an- swer was a Statistically significant “tyes,” but for Minuartia groenlandica, he found higher diversity at all Adirondack peak sites than at the Presidential Range subsites. The results for Carex bigelowii were not as easy to interpret. The overall genetic var- iability was higher in the White Mountains, but because C. bi- gelowii is less abundant in the Adirondacks than the Presidential Range, the sample size was small and only one of four indices was higher, statistically. Thus, we have a “‘maybe.”” What about 98 Rhodora [Vol. 101 gene flow? Lindwall created dendrograms to illustrate degrees of similarity (or difference) in both genetics and geographic dis- tances among the populations. Comparing Nei’s index of genetic identity for each of the three species relative to the Adirondacks and Presidentials, the answers were again mixed. For C. bigelo- wii, there was a close relationship among all sites in New Hamp- shire but not so among the New York sites. Minuartia groenlan- dica, on the other hand, showed no particular pattern with gen- erally good gene flow across the board. However, the most ge- netically distant population in the Adirondacks was from the most distantly isolated peak, the Gothics. The story with D. lapponica also seemed to relate to distance between sites. In both areas there appeared to be good gene flow with near neighbors, such as among the four McIntyre Ridge peaks in the Adirondacks, but less so when distance was greater between sites. What role does habitat area play? With C. bigelowii, there was a clear relation- ship: bigger places had more variability. Just the Opposite was true for Minuartia: the smaller sites in the Adirondacks had sta- tistically higher variability than the continuous population in the Presidentials. For Diapensia, size appeared to have no effect, and thus we have a “‘maybe”’ answer. There was one general conclusion that fit all three species, Lindwall said in summary: The greatest amount of genetic di- versity occurs where each species is the most abundant. He also concluded that we should neither assume that species will behave the same despite similar histories, nor for conservation planning purposes assume that the largest habitat area will support the most diverse population of a given species. December 1998. The program, entitled “Verdant Venues and Ventures: Visible and Verbal Visions” represented the annual event where Club members are invited to make short presenta- tions on their explorations over the year. Keith Williams led off with images from a South America vacation trip with his wife in May. It was the middle of the dry season in Brazil, their first destination, but they still saw lots of water because much of their time was spent on the coast and in the Pantanal, a huge wetland that extends into two other South American countries. Plants fea- tured in the slide images were Tabebuia alba, an endangered tree species in Brazil; Cuphea melvilla, a prolific shrub along the Pan- tanal waterways; and Ludwigia inclinata and Cabomba furcata 1999] NEBC Meeting News 99 growing in sloughs and shallows of the Pantanal. He ended with shots from Peru’s Inca Trail to Machu Picchu and an image of an Equisetum growing from the mortar of stone ruins. Marsha Salett followed Keith with a brief introduction to her Master’s degree project at the University of Massachusetts—Bos- ton which is to create a CD-ROM version of a natural history guide to bogs of southern New England. She showed images of several bogs with public access that she might feature in the guide, as well as a few that lack easy access or boardwalks that she may omit. Her intent is to present explanations and illustra- tions of bog types and common species such as Kalmia angus- tifolia and Ledum groenlandicum. Dichotomous keys and images of plants in flower and fruit will be provided to help with iden- tifications. Lois Somers then took us back to the tropics with images of a trip with husband Paul to Costa Rica. Being a registered nurse, not a botanist, she used a few wildlife images to illustrate some of the critters botanists need to be on the watch for while probing the greenery. The images included an orange-kneed tarantula seen in the Monteverde cloud forest and an eyelash viper seen at Brau- lio Carrillo National Park. Aquatic critters to be aware of included caiman seen on the Cano Negro River near the Nicaraguan border and the much larger and fiercer crocodiles of the Palo Verde re- gion. Joanne Sharpe’s slides started in Costa Rica with an image of Danaea wendlandii, one of the fern species she studied there for six years. She then took us to Puerto Rico for a look at distur- bance studies of ferns in a palm forest before and after Hurricane Georges, and in mangrove swamps, where the 14 ft. tall leather fern, Acrostichum danaeifolium, was regenerating following four years of hydrologic disturbance from dike construction. Her last stop was Maine with images from the Coastal Maine Botanical Gardens in Boothbay, where Nyssa sylvatica can be found at or near its northern limit. David Hunt continued the regional theme with images from New York where he has been helping to refine the state’s plant community classification, particularly in the Northern Appala- chian Ecoregion. His images included riverside ice meadows with Prunus pumila and Andropogon gerardii and pine-dominated rocky summit communities with either pitch or red pine and as- sociates such as Vaccinium myrtilloides, Amelanchier bartrami- 100 Rhodora [Vol. 101 ana, and Oryzopsis pungens. He then took us underwater at Lake George where he has been doing underwater vegetation sampling at depths up to 40 ft. In shallow bays he found Potamogeton amplifolius—Vallisneria americana and Eriocaulon aquaticum— Elatine americana to be common community types, whereas sandy deltas had associations of Lobelia dortmanna and Myrio- phyllum pinnatum. In deeper waters he found associations of Na- jas flexilis, Potamogeton gramineus, and P. perfoliatus. At 30 ft., he found beds of Jsoetes macrospora and Potamogeton robbinsii, and at 40 ft., a dense cover of Nitella flexilis. With this success, he’s now tackling marine eelgrass environments of Long Island. The next three presenters came as a team representing the new- formed Botanical Club of Cape Cod and the Islands. Don Schall spoke about the group’s search for and likely rediscovery of an extant population of Asclepias purpurascens on the Cape and the discovery of water hyacinth, Eichhornia crassipes, thriv- ing in a spring upwelling near a Barnstable cranberry bog. Mario DiGregorio discussed, with a vial sample in hand, the group’s discovery of a county record for Wolffia papulifera from a fresh- water pond in Barnstable and showed images of sandplain grass- land rarities: Liatris scariosa var. novae-angliae being visited by a monarch butterfly, Aster concolor at the northern limit of its range, Aristida purpurascens, New England’s only perennial awn- grass, and Prenanthes serpentaria from Nantucket. Pamela Pol- loni continued with the discussion of P. serpentaria by pointing out its hairy calyx, which distinguishes it from P. trifoliata, and other aspects of its life history such as pollination by Bombus bees and how to recognize the juvenile plants. —PauL Somers, Recording Secretary. INFORMATION FOR CONTRIBUTORS TO RHODORA Submission of a manuscript implies it is not being considered for publication simultaneously elsewhere, either in whole or in part. GENERAL: Manuscripts should be submitted in triplicate. The text must be double-spaced throughout, including tables, figure legends, and literature citations. Use a non-proportional font throughout and do not justify the right margin. Do not indicate the style of type through the use of capitals, underscoring, or bold, except for names of genera and species which should be in italics or underscored throughout. Do not underline punctuation. All pages should be num- bered in the upper right-hand corner. For guidance in matters not addressed here, consult the editorial office by phone at (603) 862- 3205, FAX (603) 862-4757, or e-mail: janets@christa.unh.edu. Brev- ity is urged for all submissions. Submit manuscripts to the Editor-in- Chief. TITLE, AUTHOR(S), AND ADDRESS(ES): Center title, in capital letters. Omit authors of scientific names. Below title, include au- thor(s) name(s), affiliation(s), and address(es). If “‘current address” is different, it should follow immediately below, not as a footnote. 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Cross-check spelling of author(s) name(s) and dates of publication. Literature citations in the text should be as follows: Hill (1982) or (Hill 1982). For two or more authors, cite as follows: Angelo and Boufford (1996) or (Angelo and Boufford 1996). Cite several refer- ences alphabetically by first author, rather than chronologically. With- in parentheses, use a semicolon to separate different types of citations (Hill 1982; Angelo and Boufford 1996) or (Figure 4; Table 2). FLORAS AND TAXONOMIC TREATMENTS: Specimen citation should be selected critically, especially for common species of broad 101 102 INFORMATION FOR CONTRIBUTORS distribution. Keys and synonymy for systematic revisions should be prepared in the style of ‘““A Monograph of the Genus Malvastrum,” S. R. Hill, RHODORA 84: 159-264, 1982. Designation of a new taxon should carry a Latin diagnosis (rather than a full Latin descrip- tion), which sets forth succinctly how the new taxon differs from its congeners. 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THE NEW ENGLAND BOTANICAL CLUB 22 Divinity Avenue Cambridge, MA 02138 The New England Botanical Club is a nonprofit organization that promotes the study of plants of North America, especially the flora of New England and adjacent areas. The Club holds regular meetings, and has a large herbarium of New England plants and a library. It publishes a quarterly journal, RHO- DORA, which is now in its 101st year and contains about 400 pages per volume. Visit our web site at http://www.herbaria. harvard.edu/nebc/ Membership is open to all persons interested in systematics and field botany. Annual dues are $35.00, including a subscrip- tion to RHODORA. Members living within about 200 miles of Boston receive notices of the Club meetings. To join, please fill out this membership application and send with enclosed dues to the above address. Regular Member $35.00 Family Rate $45.00 Student Member $25.00 For this calendar year For the next calendar year ee Name Address City & State Zip Phone FAX email Special interests (optional): Ann a ing 3 1 753 00364 9 THE NEW ENGLAND BOTANICAL CLUB Elected Officers and Council Members for 1998-1999: President: David S. Conant, Department of Natural Sciences, yndon State College, Lyndonville, VT 05851 Vice-President (and Program Chair): Lisa A. Standley, Vanasse Hangen Brustlin, Inc., 101 Walnut St., PO. Box 9151, Wa- tertown, MA 02272 Corresponding Secretary: Nancy M. Eyster-Smith, Department of Natural Sciences, Bentley College, Waltham, MA 02154- 4705 Treasurer: Harold G. Brotzman, Box 9092, Department of Bi- ology, Massachusetts College of Liberal Arts, North Adams, MA 01247-4100 Recording Secretary: Paul Somers Curator of Vascular Plants: Raymond Angelo Assistant Curator of Vascular Plants: Pamela B. Weatherbee Curator of Nonvascular Plants: Anna M. Reid Librarian: Leslie J. Mehrhoff Councillors: W. Donald Hudson, Jr. (Past President) Michael J. Donoghue 1999 Arthur V. Gilman 2000 Karen B. Searcy 2001 Matthew Hickler (Graduate Student Member) 1999 Appointed Councillors: David E. Boufford, Associate Curator Janet R. Sullivan, Editor-in-Chief, Rhodora Journal of the New England Botanical Club CONTENTS Inventory and vegetation classification of floodplain forest communities in M chusetts. Jennifer B. Kearsley 105 Jaltomata lojae (Solanaceae): Description and floral — of a new An- ean species. Thomas Mione and Luis A. Seraz 136 The reproductive biology of Magnolia evn Larry K. Allain, Mi- Zavada, and Douglas G. Matth 143 The taxonomy of Carex section Scirpinae (Cyperaceae). Debra A. Dunlop and Garrett E. Crow 163 NEW ENGLAND NOTE Rare and non-native plants of Massachusetts’ floodplain forests. Jennifer Kearsley 200 BOOK REVIEW Wild Orchids Across North America: A Botanical Travelogue .......... 206 NEBC MEETING NEWS 208 Information for Contributors 213 NEBC Membership Form 215 NEBC Officers and Council Members inside back cover MISSOURI BOTANICAL JUL 1 3 1999 GARDEN LIBRARY Vol. 101 Spring, 1999 No. 906 Issued: June 29, 1999 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, NH 03824 ANTOINETTE P. HARTGERINK, Managing Editor Department of Plant Biology, University of New Hampshire, Durham, NH 03824 Associate Editors HAROLD G. BROTZMAN STEVEN R. HILL DAVID S. CONANT THOMAS D. LEE GARRETT E. CROW THOMAS MIONE K. N. 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., 1041 New Hampshire St., Lawrence, KS 66044-0368. Periodicals postage paid at Lawrence, KS. POSTMASTER: Send address changes to RHODORA, PO. 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. ACCREDITED with the International Association for Plant Taxonomy for the purpose of registration of new names of vascular plants (ex- cluding fossils). SUBSCRIPTIONS: $75 per calendar year, net, postpaid, in funds paya- ble at par in United States currency. Remittances payable to RHO- DORA. Send to RHODORA, PO. Box 1897, Lawrence, KS 66044- 8897. MEMBERSHIPS: Regular $35; Family $45; Student $25. Application form printed herein BACK ISSUES: Questions on availability of back issues should be ad- dressed to Dr. Cathy A. Paris, Department of Botany, University of Vermont, Burlington, VT 05405-0086. E-mail: cparis @ moose. uvm.edu. ADDRESS CHANGES: In order to receive the next number 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. 101, No. 906, pp. 105-135, 1999 INVENTORY AND VEGETATION CLASSIFICATION OF FLOODPLAIN FOREST COMMUNITIES IN Missoury BOTANICAL MASSACHUSETTS JENNIFER B. KEARSLEY JUL 0 . 1999 Massachusetts Natural Heritage & Endangered Species Program, Massachusetts Division of Fisheries & aa Route 13.BARDEN LIBRARY Westborough, MA 015 ABSTRACT. Floodplain forests on eleven rivers in Massachusetts were sur- veyed to determine the variation in vegetation and soils across a range of hydrologic, physiographic, and climatic conditions. Quantitative vegetation data collected from 124 plots at 43 sites were analyzed using SPAN and DECORANA (DCA), and six community types were identified. The six pes were: Type I—Riverine island floodplain forests (Acer saccharinum— Populus deltoides-Acer negundo—Matteuccia struthiopteris association); Type II—Major-river floodplain forests (A. saccharinum—P. deltoides—Lapor- tea canadensis association); Type I1I—Transitional floodplain forests (A. sac- charinum—Arisaema dracontium association); Type [V—Small-river flood- plain forests (A. saccharinum—Fraxinus pennsylvanica—Quercus palustris as- sociation); Type V—Alluvial swamp forests (Acer rubrum—A. saccharinum— Q. bicolor association); and 1 VI—Alluvial terrace forests (A. rubrum— densis, Boehmeria cylindrica, and Onoclea sensibilis. Results of the classi- rang: showed variation in floodplain forest vegetation composition among ers in Massachusetts corresponding to significant differences in soil mot- da soil texture, presence/absence of a surface organic layer, and soil pH. Key Words: Acer saccharinum, community classification, DECORANA, floodplain forest, Massachusetts, ordination, TWINSPAN Floodplain forests, which develop on alluvial mineral soils within the zone of active flooding of rivers and streams, are con- sidered to be among the most threatened, globally significant wet- land community types in New England. Due to their high soil fertility and scenic qualities, floodplain forests have largely been converted to agriculture or lost to housing and industrial devel- opment. While several studies have addressed the relationship between floodplain forest vegetation and environmental variables within a single site or river basin in New England (Metzler and amman 1985; Veneman and Tiner 1990), this study addresses the variability in floodplain forest vegetation and environments across river basins and physiographic regions. The objectives of 105 106 Rhodora [Vol. 101 the current study were to conduct a statewide vegetation classi- fication of floodplain forest communities, to determine the distri- bution of defined community types across drainage basins and rivers, and to assess differences in environmental parameters among the identified floodplain forest community types. The Massachusetts inventory and classification work is part of a regional effort to classify floodplain forests by state Natural Heritage Programs and The Nature Conservancy. Results of these projects will provide the baseline community data necessary for future in-depth studies of floodplain forest communities, and for land protection and conservation of these ecologically significant wetland communities. MATERIALS AND METHODS Site selection. Potential floodplain forest sites were identified using USGS topographic quadrangles, Natural Resource Conser- vation Service soil surveys, and color-infrared (CIR) aerial pho- tography. A combination of 1:25,000 scale, leaf-on CIR aerial photography from an unpublished community inventory of the Connecticut River Valley (Motzkin 1993), and 1:12,000 scale, leaf-off CIR aerial photography obtained from the Massachusetts Department of Environmental Protection Wetlands Conservancy ogram were used. Potential floodplain forest sites were identi- fied using the following criteria: (1) low, forested sections of greater than 3 ha occurring within 1—2 contour intervals (10—20 ft. elevation) of river’s edge; (2) presence of alluvial soils; and (3) evidence of spring flooding and forest vegetation on aerial photography. The Massachusetts Natural Heritage and Endan- gered Species Program Biological and Conservation Database was also used to locate potential floodplain forest sites by iden- tifying localities of tracked, state-protected rare species known to occur in floodplain forest habitats. Using the resources and criteria listed above, 144 potential floodplain forest sites were identified in the state. Based on pre- a Figure 1. Massachusetts’ rivers and sub-ecoregions with sites surveyed for floodplain forest vegetation classification. Sub-ecoregions containing sur- vey sites are shaded in grey. SELECTED SUB-ECOREGIONS OF MASSACHUSETTS A - Western New England Marble Valleys B - Connecticut Valle C - Southern New England Coastal Plains D - Narragansett & Bristol Lowland -_— — (5 9 «/ N River A Ecoregion Boundary /V Sub-ecoregion Boundary Or ley Lo . “ Po) ' ri rn rn rs ta ym~ wa ee mt m s © 1 8 9 40 50 kilometers gl ‘ “s Pa AN "Shececaaca™ [6661 SONMUNUIUIOD 1S910.J UTe[dpoo],j—Aoz]sieay 108 Rhodora [Vol. 101 Table 1. Drainage basins and river sections with sampled floodplain forest communities. Tributaries refer to third order or smaller streams. 50% Ex- ceedance values indicate the discharge of 50% of flows annually, averaged over the period of record (Socolow et al. 1995). Exceedance values are given in cubic feet per second (cfs). Drainage 50% Num- Basin Exceed- Num-_ ber Area (sq. ance berof of Basin River miles) (cfs) Sites Plots Blackstone Blackstone 25.6 422 1 1 Connecticut Connecticut at 9,660 11,000 17 31 Connecticut tributaries — — 7 24 Deerfield 557 950 7 11 Housatonic Housatonic at 465 456 5 13 Ashley Falls, MA Ipswich Ipswich 44.5 37 1 4 Merrimack Assabet 116 125 1 4 Concord 400 481 1 4 Merrimack 4,635 5,110 4 4 Nashua 435 365 2 43 Nashua tips — — 1 3 Shawshee 36.5 38 2, 6 Taunton SERIE 84.3 113 1 6 Totals 43 124 liminary field checks of potential sites, 55 were found to be semi-natural forested floodplain sites with evidence of periodic flooding (e.g. floodlines on trees, flood debris, or scoured sur- faces) and a relative lack of evidence of human disturbance (e.g. limited clearings or non-native plant species). Quantitative veg- etation and environmental data were collected at 43 of the semi- natural forested floodplain sites that were distributed across eleven rivers and four physiographic provinces, or sub-ecore- gions (Figure 1). e eleven rivers ranged in drainage basin area from 25— 10,000 square miles, and in mean 50% exceedance values from 30—11,000 cubic feet per second (cfs; Table 1). Fifty percent ex- ceedance values are used as indicators of average river discharge; they indicate the minimum discharge in cfs that 50% of all flows exceed annually, averaged over the period of record (Socolow et al. 1995). Identified floodplain forest sites ranged in size from 1 1999] Kearsley—Floodplain Forest Communities 109 to 30 ha. Five sites less than the minimum size criterion of 3 ha were included because they either occurred on state-owned land with easy access (3 sites) or occurred on the Merrimack River 2 sites) where potential sampling sites were limited. Study area. The eleven rivers sampled in this study are lo- cated within four subregions of the two ecological regions, or ecoregions, occurring in Massachusetts: the Northern Highlands Ecoregion and the Northeastern Coastal Zone (Figure 1; Griffith et al. 1994). These ecoregions are defined as areas with distinct geology, landforms, soils, vegetation, climate, wildlife, water, and human influences (Griffith et al. 1994). The Northern Highlands Ecoregion includes all of Massachu- setts west of the Connecticut River Valley and the Worcester Pla- teau in north-central Massachusetts as well as most of northern New England and the Adirondack Mountains in New York (Grif- fith et al. 1994). It roughly corresponds to the Adirondack-New England mixed forest-coniferous forest—alpine meadow province described by Bailey (1995). The Northeastern Coastal Zone in- cludes eastern and coastal Massachusetts, most of southern New England, and coastal regions of New Hampshire and southern Maine (Griffith et al. 1994). It falls within the central Appalachian broadleaf forest—-coniferous forest-meadow province described by Bailey (1995). The lower Housatonic River runs through the Western New England Marble Valleys subregion of the Northern Highlands Ecoregion (Figure 1). Bedrock in this region, also known as the Berkshire Valley, consists of calcitic and dolomitic marbles and limestones; surface water alkalinity values in the area are high 1000 yeq/L; Griffith et al. 1994). The Connecticut and Deer- field Rivers and the lower reaches of their tributaries are included in the Connecticut Valley subregion of the Northeastern Coastal Zone (Figure 1). The Connecticut Valley is characterized by thick outwash, alluvial, and lake bottom deposits overlaying sedimen- tary bedrock. Surface water alkalinity values are generally above 500 wed/L. The Blackstone, Concord, Assabet, Merrimack, Shawsheen, Ipswich, and Nashua Rivers occur within the Southern New Eng- land Coastal Plains and Hills subregion (Figure 1). This is the largest subregion in southern New England and is variable in its topography and bedrock. Bedrock types in the subregion are 110 Rhodora [Vol. 101 mostly granites, schist, and gneiss, and surface water alkalinity values are generally lower than in the Connecticut Valley, ranging from less than 50 to 500 peq/L. The Threemile River occurs in the Narragansett Bristol Lowland subregion (Figure 1). The Nar- ragansett Basin is similar to the Coastal Plains and Hills subre- gion, but bedrock outcrops are not common, and thick glacial till and outwash deposits cover the area. Surface water alkalinity val- ues are generally between 100 to 300 wedq/L, but several areas have values less than 50 peq/L (Griffith et al. 1994). Field methods. Vegetation was sampled in 10 m X 20 m (0.02 ha) rectangular plots placed along transects perpendicular to the river. At most sites, two or more transects were placed at least 50 m apart. Each transect was walked and changes in to- aphy and vegetation were described. A plot was placed with- in each identified topographic or vegetation unit. In small flood- plain forests (=3 ha), one or two plots were subjectively placed within the “‘typical’’ vegetation type(s) and not along transects. The number of plots per site ranged from 1 at small sites to 8 at large sites. Plots were placed with their long axis parallel to the river. Percent cover of trees (stems >10 cm DBH), shrubs (stems <10 cm DBH), saplings, and vines was visually estimated within each 0.02 ha plot, and percent cover of herbs and seedlings was vi- sually estimated within two 0.0004 ha (2 m X 2 m) square sub- plots. Herbaceous taxa (<1 m tall) occurring within the 0.02 ha plot, but not within the subplots, were also recorded. Nomencla- ture follows Kartesz (1994). Percent cover for all taxa was esti- mated using a modified Braun-Blanquet cover scale with the fol- lowing cover classes: r (single occurrence), <1%, 1-5%, 6-10%, 11-20%, 21-25%, 26-35%, 36-45%, 46-50%, 51-55%, 56— 65%, 66-75%, 76-85%, 86-95%, and 96-100%. The average height and average percent cover of each vegetation stratum were also visually estimated and recorded. Vegetation data from 124 plots were included in the vegetation classification (Table 1). Eighty-nine plots were surveyed between July and September, 1997, using the methods described above. Existing data from 35 plots collected with equivalent methodol- ogies by other sources were included in the vegetation classifi- cation. Those data were: 10 plots from the Deerfield River (Thompson and Jenkins 1992), 16 plots from the Nashua River 1999] Kearsley—Floodplain Forest Communities 111 (Searcy et al. 1993), 7 plots from the Connecticut River and its tributaries (Motzkin 1993, 1995; Massachusetts Audubon Society, unpubl. data), and 2 plots from the Ipswich River (Massachusetts Audubon Society, unpubl. data). Environmental data were collected from the 89 plots sampled in 1997. At each plot, one 60 cm deep soil pit was dug and the following soil characteristics were described: depth, soil texture, and color of horizons; depth to mottling; color of mottles; depth of root penetration; and average pH of the mineral soil. The fol- lowing environmental data were also collected for each plot: to- pographic position (terrace, levee, level floodplain, depression), height of floodlines, and the number of stumps, and uprooted and snapped trees. Any evidence of disturbance or land use was also noted. Data analysis. Vegetation cover data were analyzed using two-way indicator species analysis (TWINSPAN) and ordination techniques (DCA) contained in the PC-ORD Version 3.0 statis- tical package (McCune and Mefford 1997). TWINSPAN (Hill 1979a) was used to identify floodplain forest types, and DCA (Hill 1979b) was used to illustrate the relationship between types. Default settings were used with the following exceptions: Braun- Blanquet cut-levels (0, 5, 26, 51, and 76) were used in the TWIN- SPAN analysis, and downweighting of rare species was used in DCA. Community types were based on both TWINSPAN and DCA results. Species indicator values for the community types were calculated using the Indicator Species Method of Dufréne and Legendre (1997) in the PC-ORD Version 3.0 statistical pack- age (McCune and Mefford 1997). Indicator species defined by the Indicator Species Method were used instead of TWINSPAN indicator species to describe community types because: (1) final community types were based on both TWINSPAN and DCA re- sults, and (2) the Indicator Species Method defines indicator spe- cies as those species present in the majority of sites belonging to a group, while TWINSPAN defines them as those species that are found mostly in a single group, but not necessarily in the majority of that group’s sites (Dufréne and Legendre 1997). A Monte Carlo technique was used to test for the statistical signif- icance of indicator values. Species richness, evenness (E), and Shannon diversity index (H’) values were calculated for each plot using the method out- 112 Rhodora [Vol. 101 lined in the PC-ORD statistical package (McCune and Mefford 1997). Single factor analysis of variance (ANOVA) was used to test if the indices were significantly different among community types. A multi-response permutation procedure (MRPP; Zimmer- man et al. 1985) was used to test if defined community types differed significantly in height and total percent cover of the fol- lowing six strata: emergent canopy, tree canopy, tree sub-canopy, tall shrubs, short shrubs, and herbs. In order to test for differences in soil characteristics among the defined community types, an MRPP was run on the following variables: presence/absence of soil mottling, depth to mottling (cm), soil texture at 10 cm intervals (sand, loamy sand, sandy loam, and silt loam), presence/absence of a surface organic layer, depth of organic layer, pH, and presence/absence of hydric soil. Hydric soil determination was based on the criteria outlined by the Natural Resource Conservation Service (1994). Individual ANOVA and chi-square tests were used to determine which soil variables were significantly different (at p < 0.01) among defined community types. Chi-square tests were used to test for signifi- cant differences in presence/absence of floodlines and topograph- ic position. RESULTS Vegetation classification. Six vegetation community types were recognized based on the TWINSPAN (Figure 2) and DCA (Figure 3) results. There was general agreement between TWIN- SPAN groups and DCA output (Figure 3). The six community types described below are primarily based on the TWINSPAN output (Figure 2) with two exceptions. First, twelve plots classi- fied as one type by the TWINSPAN analysis were moved to a different community type based on the DCA results. For example, TWINSPAN classified ten plots as Type I (Figure 2), but the DCA results showed that two plots classified as Type II by TWIN were more closely related to plots classified as Type I. Therefore, Type I is described below as containing twelve plots, and twelve plots were included in the environmental analyses. The second exception is that four plots classified by TWINSPAN were eliminated from the final community types described below and from the environmental analyses because they were found to be distinct species assemblages, unlike all other plots. 1999] Kearsley—Floodplain Forest Communities 113 MAST (3) ARTR (1) ULRU (1) TYPE I MAST (1) 4 LACA (1) (N=10) PODE (1) TYPE II ACSA (4) (N=36) LACA (1) TYPE Il (N=18) TYPE IV (N=28) * BOCY (1) Ae! All OSRE (1) peeee (N=17) VIDE (1) ACRU (1) CAOV (1) ht), TYPE VI (N=15) Figure 2. TWINSPAN dendrogram of floodplain forest community types with TWINSPAN indicator species listed at divisions. Species codes follow those listed in Table 2. Numbers in parentheses next to each species indicate the minimum percent cover class for that species in all plots within the com- munity type: (1) 0-4%, (2) 5—25%, (3) 26-50%, (4) 51-75%. Sample sizes refer to the number of plots classified as each type in the TWINSPAN anal- ysis. The primary division of plots in TWINSPAN was made based on the occurrence of Acer saccharinum and A. rubrum with Types I, II, II, and IV having A. saccharinum dominant in the overstory and Types V and VI having A. rubrum dominant (Figure 2). Var- iation in species composition across DCA ordination Axis 1 (R? = 0.227) was also associated with a decrease in A. saccharinum and increase in A. rubrum. Variation in species composition across DCA Axis 2 (R? = 0.127) was associated with an increase in Ulmus rubra and Impatiens pallida and a decrease in Populus deltoides and Laportea canadensis. Axis 3 did not characterize much additional variation in species composition (R2 = 0.038). Plots of Types II and III were closely related in TWINSPAN 114 Rhodora [Vol. 101 400 5 Oy 300 4 key oO O Vv re es - & : 5 200 - > uv + 9 A A Oo V, Bogen tries ¢ Boo eae: a es de yan A s. 100 ee a! a. © m ore) & A O 0 5 A 0 100 200 300 400 500 Axis 1 Figure 3. First two axes of the floodplain forest DCA output with plots coded by TWINSPAN group. Squares = Type I, circles = Type II, down- facing triangles = Type III, pluses = Type IV; up-facing triangles = Type V, and diamonds = Type VI. clustering where they divided at the fourth division (Figure 2), but they were well separated in the DCA analysis along both Axes 1 and 2 (Figure 3). Two hundred and fourteen vascular plant species were identi- fied in the floodplain forest sampling. A subset of those species with indicator values that were significant (p < 0.05) and/or greater than 10% are listed in Table 2. Indicator values represent a combination of species relative abundance and relative fre- quency of occurrence in the identified community types (Dufréne and Legendre 1997). For example, an indicator value of 10% assumes that the species was present in at least 33% of sites in a community type, and that the relative abundance of the species was at least 33% in one of the community types. If one of the two measures was 100% then the other was at least 10% (Dufréne and Legendre 1997). Thirty-four taxa with indicator values great- er than 20% were plotted according to their DCA axis loading scores for Axis 1 and Axis 2 to illustrate the relationship between species abundance and community types (Figure 4). 1999] Kearsley—Floodplain Forest Communities 115 RHTY 600 fe) 500 URSE 4 ° ULRU 400 a IMP/4O 1 MAST ois 300 - vin ye GECA = ARTR < TO LYUN BOCY CAOV 200 4 PODE © dee © UVSE, ° 2 fe) — Aku 7 © OSCI CIAR MACA~ QBSA ° fo) 100 5 in QUBI IVSA LYSU OSRE GvDGR zt 9° fe) ° LEVI © L 0 LACA ° OAM re} °o -100 T T T T T T ce fo ae -100 0 100 200 ©6300 400 500 600 Axis 1 Figure 4. Floodplain forest species with indicator values that are >20% and significant (p < 0.05) plotted by their axis loading scores for DCA Axis Acer saccharinum was the most common tree species encoun- tered and therefore had a low maximum indicator value (22%) for any single community type (Table 2). Acer saccharinum at- tained its highest indicator values in Types II (20%), III (20%), and IV (22%). Ulmus rubra was a strong indicator species (46%) for Type I, Populus deltoides for Types I and II (24% and 28%, respectively), and A. rubrum and Carya ovata for Type VI (60% and 49%, respectively; Table 2). Shrubs and saplings were im- portant components of Types I, V, and VI: Berberis thunbergii, Rhus typhina, and Rosa multiflora had high indicator values in Type I floodplain forests; Cornus amomum and Rhamnus fran- gula in Type V; and Ilex verticillata, Viburnum dentatum, and Quercus bicolor seedlings in Type VI (Table 2: Figure 4). Table 2. species Indicator values and associated p values for floodplain forest taxa listed by community type. * indicates non-native Indicator Values by Community Type Species Name CODE I II Il IV Vv VI p value Acer saccharinum ACSA 10 20 20 22 14 1 0.035 TYPE I Matteuccia struthiopteris MAST 61 10 1B 0 0 0 0 Imus rubra ULRU 46 3 0 0 0 0 0 Impatiens pallida IMPA 38 2 0 0 0 0 0 Arisaema triphyllum ARTR Sf 6 11 2 0 4 0 U. rubra seedling URSE 31 0 0 0 0 0 0 A. negundo sapling ANSA 25 1 0 0 0 0 0 Rhus - e ina RHTY 25 0 0 0 0 0 0 Vitis ri VIRI 25 1 7 0 0 0 0.017 eBarberis finbeet BETH 24 0 0 0 0 0 0.006 negundo seedlings ANSE 22 0 0 0 0) 0 0.001 *Rosa multifl ROMA 20 2 0 0 0 0 0.027 Geum canadense GECA 18 3 8 0 0 0 0.031 Acer negund ACNE 16 3 0 0 0 0 0.042 *Celastrus orbiculata CEOR 10 3 1 3 0 0 0.26 Eupatorium rugosum EURU 10 6 9 0 0 0 0.437 TYPE I Laportea canadensis LACA 0 84 3 0 0 0 Leersia virginica LEVI 0 29 19 2 2 0 0.058 Populus deltoides PODE 24 28 1 0 0 0 0.001 Helianthus tuberosus HETU 0 15 0 0 0 0) 0.046 ee — On eIOopoyy IOI TOA] Table 2. Continued. Indicator Values by Community Type Species Name CODE I II Ii IV Vv VI p value *Glechoma hederacea GLHE i 14 1 0 0 0 0.214 *Chelindonium majus CHMA 0 13 0) 0 0 0 0.073 TYPE Il Boehmeria cylindrica BOCY 0 l 32 24 20 0 0.013 Cinna arundi CIAR 0 2 a2 1 11 0 0.015 Onoclea sensibilis ONSE 1 0) PM 25 13 7, 0.013 Toxicodendron radicans TORA 0 1 25 9 18 Is 0.127 Amphicarpaea bracteata AMBR 0 Zz 24 l 0 0 0.002 ricana sapling UASA 0 zZ 20 0 0) 1 0.054 *Alliaria petiolata ALOF 0 2 18 0 0 0 0.054 Arisaema dracontium ARDR 0 0 16 0 0 0 0.011 Polygonum virginianum TOVI 1 2 16 0 0 0 0.026 F, americana sapling FASA 0 0 14 0 0 0 0.045 . Saccharinum sapling ASSA 2 4 13 3 1 0 0.21 Plena occidentalis PLOC 2 0 12 0 0 0 0.05 TYPE IV A. saccharinum seedlings ASSE 0 I 0 39 8 0 0.001 Sium s SISU 0 0 0 29 0 0 *Lysimachia nummularia LYNU 0 0 0 2 0 0 0.026 F. pennsylvanica seedlings FPSE 9 0 0 20 0 0 0.016 Cephalanthus occidentalis CEOC 0 0 0 19 1 0 0.01 [6661 SOnIUNUWIWOD 3se10,j ule[dpoo,j—Aoysieay Table 2. Continued. Indicator Values by Community Type Species Name CODE I Il Il IV Vv VI p value Fraxinus pennsylvanica FRPE 0 0 1 19 6 1 0.039 Lysimachia terrestris EYTE 0 0 0 18 0 2 0.012 Quercus palustris QUPA 0 0 0 18 0 0 0.007 *Myosotis scorpioides MYSC 0 0 0 17 0 0 0.043 Cicuta maculata CIMA 0 0 0) 14 0) 0 0.035 Leersia oryzoides LEOR 0 0 0 14 0 0 0.029 Q. palustris seedlings QPSE 0 0 0 14 0 0 0.055 Phalaris arundinacea PHAR 0 0 0 i 0 0 0.035 *Polygonum persicaria PLRS 0 0 1 13 0) 0 0.139 Carex — CALU 0 0 0 11 0 0 0.065 Carex typ CATY 0 0 0 11 0 0 0.079 Lobelia surdiedlis LOCA 0 0 0) 11 0 0) 0.078 TYPE V Osmunda regalis OSRE 0 0 0 0 53 0 0 Quercus bicolor QUBI 0 0 0 0) 39 4 0 Vitis labrusc VILA 0 0 0 0 31 0 0 *Lythrum aon LYSA 0 0) 0 6 25 0 0.003 Cornus amomu COAM 0 0 0 6 24 3 0.013 Lycopus si riak LYUN 0 0 0 0 20 0 0.003 Carex crinita CACR 0 0 0 9 15 0 0.095 reanmend — RH 0 0 0 3 ibs: 6 0.047 Betula BENI 0 0 0 4 13 0) 0.042 GC. “att cacti. QBSE 0 0 0 0 13 8 0.067 SII vIopouy IOI ISA] Table 2. Continued. Indicator Values by Community Type Species Name CODE I Il iW IV Vv VI p value TYPE VI Acer rubrum ACRU 0 10) 0) 0 13 60 0 Viburnum ae VIDE 0 0 0 0 16 54 0 CAOV 0 0 0 0 0 49 0 Ilex Cepia: ILVE 0 0 0 1 0 47 0 Athyrium felix-femina ATFI 0 0 0 0) 1 37 0 Mai emum canadense MACA 0 0) 0 0 4 a7 0 Uvularia sessilifolia UVSE 0 0 0 0 0 35 0 smunda cinnamomea OSCI 0) 0 0 0 0) 29 0) Q. bicolor saplings QBSA 0 0) 0 0 0) P| 0.003 Tilia american TIAM 1 0 0 0 0 19 0.004 P. serotina seedlings PSSE 0 0 0 0 0 14 0.017 Ulmus american ULAM 2 2 2 1 6 12 0.308 Circaea lutetiana ssp. CIQU + 1 1 0 1 11 0.206 canadensis gercmiais quinquefolia PAQU a 4 0 3 0 11 0.825 A. rubrum seedlings ARSE 0 @) 0 1 0 10 0.068 Prunus een PRSE 4 0 0 0 5 6 0.511 [6661 SeUNUIWOD 3seI0,j ure[|dpool,.j—Ae|sieoy 120 Rhodora [Vol. 101 Variation in abundance of woody vines along DCA Axis 1 was associated with an increase in Vitis labrusca and a decrease in V. riparia (Figure 4); V. labrusca attained its highest indicator value in Type V, while V. riparia was most abundant in Type I (Table 2). Toxicodendron radicans occurred across all plots, but was most abundant in Types III, IV, V, and VI (Table 2). Variation in herbaceous species composition across DCA Axis 1 was associ- ated with a decrease in Matteuccia struthiopteris and Laportea canadensis and an increase in Osmunda regalis and O. cinna- momea (Figure 4). Boehmeria cylindrica and Onoclea sensibilis had intermediate Axis | loading scores (Figure 4) and high in- dicator values for Types III, IV, and V (Table 2). Variation in herbaceous species composition across Axis 2 was associated with decreasing L. canadensis and increasing Impatiens pallida and M. struthiopteris (Figure 4). Community structure was similar among floodplain forest types, and the six community types did not differ significantly in the height and total percent cover of vegetation strata (MRPP p = 0.45). All types were characterized by a dense, tall tree canopy (20 m mean height, 70% mean cover) above a diffuse subcanopy (7 m mean height, 19% mean cover) and very limited to absent shrub layer (1.5 m mean height, 9% mean cover). The herbaceous cover was dense (80% mean cover) in most plots, and tall (1-2 m) when Laportea canadensis or Impatiens spp. were dominant (Types I and II; Table 2). Species richness was not significantly different among identified types (p = 0.217), but types did differ significantly in species diversity (p = 0.029) and species evenness (p = 0.0004) with Type VI forests having the highest values for both (H’ = 1.75, E = 0.71). Environmental parameters. Soil profiles of Types II, IV, V, and VI were typically hydric silt loams with soil mottling, while soil profiles of Types I and II were nonhydric, sandy loams, loamy sands, or sands without soil mottling (Table 3). Soil pro- files of Types V and VI usually had a surface organic layer, while soil profiles of Types I, II, and III usually lacked a surface organic layer (Table 3). Soil pH was least acidic in Types I, II, and III (Table 3). Results of the MRPP of soil variables showed that the observed differences in soil profiles among the six floodplain forest com- munity types were statistically significant (p = 0.001). Soil tex- 1999] Kearsley—Floodplain Forest Communities 121 ture, pH, and presence/absence of soil mottles, surface organic layer, and hydric soils were all significantly different (p < 0.001) among community types, while depth to mottling and depth of organic layer were not significantly different (Table 3). Presence/absence of floodlines and topographic position were also significantly different among defined types (p < 0.001). Most floodplain plots occurred on level floodplains, except for Type I communities which typically occurred on elevated sections of riv- erine islands and Type VI communities which occurred on ridges or high terraces (Table 3). Floodlines were visible on tree trunks in 41% of all plots, and they were most common in communit Types III and IV (Table 3). Snapped trunks were observed in 34% of all plots, uprooted trunks in 7%, and stumps (cut or bea- ver-cut) in 16%. When present, there were usually only one or two downed trees per 0.02 ha plot, and downed trees did not appear to be abundant overall in the floodplain forests invento- ried. COMMUNITY TYPE DESCRIPTIONS Type I—Riverine island floodplain forests (Acer sacchar- inum—Populus deltoides—Acer negundo—Matteuccia struthiop- teris association). Type I communities (12 plots at 8 sites) were open-canopy floodplain forests occurring on elevated sections of riverine islands and riverbanks of major rivers with high levels of disturbance. The community type was limited to the Connect- icut, Deerfield, and Housatonic Rivers in Massachusetts (Table 4). Plots classified as Type I were most likely to occur at sites where the vegetation in all plots at the site was classified as Type I. Table 5 shows that of the eight sites with plots classified as Type I, five had all plots classified as Type I, three had plots classified as both Types I and II, and one had plots classified as Types I, II, and III. Type I communities were never associated at sites with vegetation classified as Types IV, V, or VI (Table 5). Soils of Type I communities were typically nonhydric, sandy loams without soil mottles and without a surface organic layer. Soil pH ranged from 5.5 on the Connecticut and Deerfield River plots to 8.0 on the Housatonic River (Table 3). The overstory of Type I communities was a mixture of Acer saccharinum and Populus deltoides. Platanus occidentalis and Table 3. Environmental data for floodplain forest community types. *Soil texture: S = San Loam, ST = Silt Loam. **Topographical position of plots within floodplain DP = Depression. d, LS = Loamy Sand, SL = Sandy : TR = Terrace, LF = Level Floodplain, LE = Levee, % of Plots with Soil % of Plots Organic ith Soil Layer % of % of Plots Pe IONE 4 of Plots wih eck © ea wes % of Plots in each ber (me *Soil Texture at 10 cm depth wit — Fipodlines **Topographical Position of depth of, in Mean pH___ Hydric (height range peerer Type Plots to, in cm) s LS SL Si cm) (range) Soil in cm) TR LF LE. “DP I 11 0 0 0 2°94 0 6.6 (5.5-8.0) O 36 (43-135) 64 27 9 0 I ae AZAD 5 14 50 ai 4 (1.0) 6.3 (4.5-8.0) 0 13 (93-220) 13 S61 26 0 il 16 75 (18.3) 0 G i= 3] 63 0 6.2 (4.5-8.0) 38 62 (60-303) 19p 75 0 6 IV 16 88 (14.0) 6 6 2B 76 = =35. (5.0) 5.2 (4.5-6.0) 59 69 (44-265) 0 94 6 0 Vv 17 67 (26.1) 0 6 > 27 67 67 (5.0) 4.8 (4.5-5.5) 40 47 (45-122) 0 80 7 13 VI se “60(173) —0 0 40 60 60(5.3) 5.0 (4.5-6.0) 40 0 60 40 0 0) ALL ar SSG Ss 2 5 i 42 S51 23 (4.8) 5.7 (4.5-8.0) 30 41 (43-303) 19e- 73 10 3 col elopoyy TOT TOA] 1999] Kearsley—Floodplain Forest Communities 123 Table 4. oe of plots by each river in the defined floodplain forest community ty Community Type Basin River I | Gakias |5 Wieaire 5 Vooryy Blackstone Blackstone 1 Connecticut Connecticut 5. 20 1 Connecticut tributaries 1 er 12 2 Deerfield 2 8 1 Housatonic Housatonic 1 3 8 Ipswic Ipswich Merrimack Assabet 1 2 1 Concord 4 Merrimack 2 Nashua 1 5 9 Nashua tributary 3 Shawsheen 2 4 Taunton Threemile 3 2 1 Totals Boy 2 bo eR aS 14 Fraxinus americana were occasional canopy associates. Ulmus rubra, A. negundo, and Celtis occidentalis (on the Housatonic River) were common in the subcanopy (Table 2). The shrub/sap- ling layer was patchy and composed of taxa typical of disturbed areas, including Rhus typhina, Rosa multiflora, Berberis thunber- gui, and Celastrus orbiculata. Berberis thunbergii was observed n this floodplain forest community type more frequently than in any other (Table 2). The herb layer was dominated by Matteuccia struthiopteris (most plots had greater than 40% cover) or by a Table 5. Number of sites with plots of one defined floodplain forest com- munity type that also have plots of other community types. Total # of Sites with Plots of Type I Type II Type III Type IV Type V Type VI Type Type I 5 3 1 8 Type I 3 7 7 I? Type Il 1 7 3 1 11 Type IV 1 ii 4 a 2 Type V ~ 2 3 6 Type VI S 3 2 = 124 Rhodora [Vol. 101 dense, tall layer of Jmpatiens pallida over M. struthiopteris. La- portea canadensis occurred in low amounts but was never abun- dant. Other common herbaceous taxa were Eupatorium rugosum, Arisaema triphyllum, and Geum canadense. Vitis riparia was a strong indicator vine (Table 2), and Parthenocissus quinquefolia was also common. Type Il—Major-river floodplain forests (Acer saccharin- um—Populus deltoides—Laportea canadensis association). Type II communities (32 plots at 17 sites) occurred on mainstem sec- tions of the Connecticut, Deerfield, and Housatonic Rivers (Table 4). Plots classified as Type II were most likely to occur at sites with other plots classified as Type II or at sites with plots clas- sified as Type III (Table 5). Type II communities sometimes oc- curred associated with vegetation classified as Type I, but never with Types IV, V, or VI (Table 5). Soils were predominantly sandy loams without soil mottles (13% of plots had mottles) and without a surface organic layer (only 4% of plots had an organic layer). Soil pH ranged from 4.5 on the Connecticut and Deerfield Rivers to 8.0 on the Housatonic (Table 3). Acer saccharinum was strongly dominant in the overstory (>60% cover in most plots) mixed with lesser amounts of Po- pulus deltoides. Ulmus americana and/or U. rubra occurred in the subcanopy. Shrubs were generally lacking. The herbaceous layer was usually dominated by a 1-2 m tall, dense cover of Laportea canadensis, and Matteuccia struthiopteris was some- times abundant. Leersia virginica was consistently represented, but in low amounts (typically <5% cover). Other common as- sociates were Cinna arundinacea, Impatiens sp., Boehmeria cy- lindrica, and Arisaema triphyllum. Non-native plant species were usually less abundant than in Type I communities, but Polygonum cuspidatum often formed large patches along heavily scoured le- vees or in areas where the canopy was open. Other common non- native taxa were Glechoma hederacea and Alliaria petiolata. Type II—Transitional floodplain forests (Acer saccharin- um—Arisaema dracontium association). Type III communities (19 plots at 11 sites) occurred on third-order or smaller tributaries of the Connecticut River, on the Housatonic River, and in de- pressions within Major-river floodplain forests (Types I and II) of the Connecticut and Deerfield Rivers (Table 4). Plots classified 1999] Kearsley—Floodplain Forest Communities 125 as Type III were found associated at sites with plots classified as Types I, II, and IV (Table 5). Type III communities were inter- mediate in soil texture and drainage between the sandy, well- drained soils of Types I and II and the highly mottled, poorly drained silt loams of Type IV. Soil texture was silt loam or very fine sandy loam. Soils were poorly drained, and 75% of plots had soil mottling. None of the plots had a surface organic layer. The pH ranged from 4.5 to 8.0 (Table 3). The Type III vegetation association was transitional between Major-river (Types I, II) and Small-river (Type IV) floodplain forest vegetation, and shared taxa with Types I, II and IV (Table 2; Figure 3). In Type III communities, Acer saccharinum was dominant in the canopy, but unlike Types I and II, Populus del- toides was typically absent. As in Type IV plots, Fraxinus penn- sylvanica and Ulmus americana were present. A shrub layer was lacking; however, saplings of overstory trees were common. Vines were abundant with Amphicarpaea bracteata most com- mon. In contrast to Type II plots, Laportea canadensis was not dominant, but it was present in low amounts in all plots (5—15% cover). The herbaceous layer was typically an even mixture of L. canadensis, Matteuccia struthiopteris, Onoclea sensibilis, and Boehmeria cylindrica. Common associates were Leersia virgini- ca, Arisaema triphyllum, Bidens frondosa, Cinna arundinacea, and Impatiens sp. Arisaema dracontium (a state-protected rare species) was associated with this floodplain forest community type and serves as a good indicator species of the type (Indicator value = 16; Table 2; Kearsley 1999). Type IV—Small-river floodplain forests (Acer saccharin- um-—Fraxinus pennsylvanica—Quercus palustris associa- tion). Type IV communities (28 plots at 12 sites) occurred on third order or smaller tributaries of the Connecticut and Nashua Rivers, on smaller rivers of eastern Massachusetts where banks are low and overbank flooding occurs (Ipswich, Assabet, Shaw- sheen, and Threemile), and on edges of riverine islands of the Merrimack River (Table 4). Vegetation classified as Type IV was sometimes associated at sites with vegetation classified as Types Ill, V, and VI (Table 5). Soils were a mixture of silt loams and fine sandy loams. Fifty-nine percent of soil profiles were classi- fied as hydric; 88% had soil mottles and 35% had a surface or- ganic layer. The pH ranged from 4.5 to 6.0 (Table 3). 126 Rhodora [Vol. 101 As in Types I, II, and Ill, Acer saccharinum was dominant in the overstory of Type IV communities, but the understory of Type IV communities more closely resembled that of A. rubrum A\l- luvial swamp forests (Type V) and Alluvial terrace forests (Type VI). Populus deltoides and A. rubrum were both absent in the canopy of Type IV communities. Quercus palustris was a com- mon associate in the Connecticut River basin, and Betula nigra in the Merrimack River basin. Type IV floodplain forest plots had a more substantial shrub layer than both Major-river (Types I and II) and Transitional (Type III) types, but less than both Types V and VI. The shrub layer of Type IV communities consisted main- ly of Cornus amomum and Cephalanthus occidentalis. Fraxinus pennsylvanica saplings were present in most plots ere was greater herbaceous diversity in Small-river flood- plain forests than in floodplain forest Types I, II, and III. Onoclea sensibilis and Boehmeria cylindrica were most common, but as- sociates included Acer saccharinum seedlings, Cicuta maculata, Lysimachia terrestris, Sium suave, and non-native taxa, such as L. nummularia, Myosotis scorpioides, Rhamnus frangula, and Lythrum salicaria. Four state-protected rare plant species were associated with this community type: Mimulus alatus (State En- dangered), Carex typhina (State Threatened), C. grayi (State Threatened), and Rumex verticillatus (State Threatened; Kearsley 1999). Type V—Alluvial swamp forests (Acer rubrum—A. sacchar- inum-—Quercus bicolor association). Type V plots (15 plots at 6 sites) occurred along mainstem sections of smaller rivers in east- ern Massachusetts (Assabet, Concord, Nashua, Shawsheen, and Threemile; Table 4). Plots classified as Type V were often asso- ciated at sites with plots classified as Types IV and VI (Table 5). This type appeared to be wetter and more seasonally inundated than the four Acer saccharinum dominated types (Types I-IV). Soils were typically silt loams; 67% had soil mottles and 67% had a surface organic layer. The pH ranged from 4.5 to 5.5 (Table 3). Vegetation of plots classified as Type V was variable, as in- dicated by the lack of strong clustering in the DCA analysis (Fig- ure 3). In general, the overstory of Type V plots was character- ized by a mixture of Acer saccharinum and A. rubrum with lesser amounts of Fraxinus pennsylvanica and/or Quercus bicolor. Un- like Types I-IV, Type V communities had a well-developed shrub 1999] Kearsley—Floodplain Forest Communities F27 layer dominated by Viburnum dentatum, Cornus amomum, and the non-native plant Rhamnus frangula. As in Type IV commu- nities, the herbaceous layer of Type V communities was charac- terized by a mixture of Onoclea sensibilis and Boehmeria cylin- drica; however, the herbaceous layer differed in also having Q. bicolor seedlings, Osmunda regalis, and Carex crinita. Type VI—Alluvial terrace forests (Acer rubrum—Carya ova- ta—Prunus serotina association). Type VI plots (14 plots at 5 sites) occupied upland ridges within Alluvial swamp forests (Type V) and high terraces above the active flood zone. Plots of this type occurred on the Nashua, Assabet, Blackstone, and Threemile Riv- ers and on high terraces in the Connecticut River basin (Table 4). Vegetation classified as Type VI was often associated with Types and V (Table 5). Type VI forests were river influenced and mesic, but they did not appear to experience regular flooding as indicated by the presence of a distinct soil organic layer. Soils were typically silt loams; 60% had soil mottles and 60% had a surface organic layer. The pH ranged from 4.5 to 6.0 (Table 3). This community type showed the greatest within-group vari- ability in plot composition, as indicated by the point spread in the DCA output (Figure 3). Although the plots were not highly clus- tered, they were well-differentiated from the other five floodplain forest community types. Acer rubrum was dominant in the canopy, and typically mixed with varying amounts of mesic hardwoods including Carya ovata, Prunus serotina, Ulmus americana, and Tilia americana, As in Type V communities, the shrub layer was well-developed, and Viburnum dentatum and Ilex verticillata were most common. The herbaceous layer was a species-rich mixture of Onoclea sensibilis, Maianthemum canadense, Athyrium felix- femina, Osmunda cinnamomea, and Uvularia sessilifolia. DISCUSSION The greatest differences in vegetation composition and envi- ronmental characteristics among the floodplain forest community types occurred between Types I-III and Types IV—VI. These two groups were well-separated floristically, environmentally, and spatially. Types I-III occurred at sites on the Connecticut, Deer- field, and Housatonic Rivers, and Types IV and V occurred on small tributaries of the Connecticut River or on rivers in eastern 128 Rhodora [Vol. 101 Massachusetts. Type VI occurred on elevated ridges and high terraces across the state, but more data are needed to clarify the distribution of and variation in high-terrace forests statewide. Vegetation classified as Types I, II, and III often occurred to- gether at a site. Sites with vegetation that was primarily classified as Type II had patches, usually on elevated sections, where Mat- teuccia struthiopteris was dominant (Type I), and depressions where Boehmeria cylindrica and Onoclea sensibilis were domi- nant (Type III). Similarly, sites with vegetation primarily classi- fied as Type III had elevated sections dominated by Laportea canadensis (Type II). Overlap between community types within a single site also occurred among Types IV, V, and VI. Type IV forests in eastern Massachusetts had Quercus bicolor dominant (Type V) in low-lying, wet depressions and mesic, mixed-decid- uous patches (Type VI) in elevated areas. Although Types I-III and Types IV—VI were generally well- differentiated, overlap among defined community types did occur, particularly among Types III, IV, and V. These three ty e all characterized by Boehmeria cylindrica and Onoclea sensibilis in the herbaceous layer, and differences in vegetation composition among the three types was subtle; however, differences in soil profiles 3g location supported the three types as recognizable as- sociation Types 1 and II were also similar, and the differences observed in vegetation composition appeared to be primarily related to greater disturbance in areas classified as Type I. Riverine islands of the Connecticut and Deerfield Rivers where Type I vegetation assemblages were found had many canopy openings that were created by campers. Abandoned campsites were filling in with Rhus typhina, Polygonum cuspidatum, and a mixture of vines, including Celastrus orbiculata and Vitis riparia. The abundance of Acer negundo, which was associated with open, disturbed areas of Ohio floodplain forests (Hardin et al. 1989), was also indicative of greater disturbance in Type I forest plots. Comparison to other floodplain forest classifications. Type I and II Major-river floodplain forests are similar in vegetation composition to floodplain forests of larger rivers in other New England states. Type I forests correspond to the Acer negundo— Matteuccia struthiopteris association described in Vermont, which occurs on sandy loams or open cobbles within the active 1999] Kearsley—Floodplain Forest Communities 129 floodplain of larger rivers. Type II forests correspond to Ver- mont’s A. saccharinum—M. struthiopteris association, which oc- curs on coarse soils of levees of larger rivers (Sorenson et al. 1998). Elsewhere in New England a distinction between the two community types has not been made; however, Types I and II hs correspond to the A. saccharinum—M. struthiopteris—La- rtea ees association in New Hampshire (Bechtel and shes 1998), . Saccharinum—Eupatorium rugosum asso- ciation in pe omealrsiee (Metzler and Damman 1985), and to the A. saccharinum temporarily flooded forest alliance described for the eastern United States (Sneddon et al. 1998). Type I also con- tains elements of the A. negundo temporarily flooded forest alli- ance described for the Southeast which includes early succes- sional vegetation of active floodplains and sandbars with a heavy vine component (Sneddon et al. 1998). ype Ill Acer saccharinum—Fraxinus pennsylvanica—Ulmus americana floodplain forests correspond to A. saccharinum/On- oclea sensibilis floodplain forests described for Connecticut (Metzler and Damman 1985). Similar to Type III forests, Mat- teuccia struthiopteris was the dominant herbaceous taxon on the highest ridges within the community type in Connecticut (Metzler and Damman 1985). Type III forests are also closely affiliated with A. saccharinum—O. sensibilis-Boehmeria cylindrica com- munities occurring on silty soils of lower watersheds and lake- shores in Vermont (Sorenson et al. 1998) and A. saccharinum— Carex crinita—O. sensibilis associations in New Hampshire (Be- chtel and Sperduto 1998). Type III forests correspond to the A. saccharinum-U. americana—O. sensibilis temporarily flooded for- est community described for the Northeast (Sneddon et al. 1998). oodplain forest associations similar in composition to Types I, Il, and III also occur on large rivers throughout the north- central United States. Acer saccharinum dominated forests have been described in detail in southeastern Wisconsin (Dunn and Stearns 1987; Menges 1986; Menges and Waller 1983), Ohio (Hardin et al. 1989; Hardin and Wistendahl 1983), northern Mis- souri (Dollar et al. 1992), New Jersey (Buell and Wistendahl 1955; Frye and Quinn 1979) and central Illinois (Brown and Pe- terson 1983; Peterson and Rolfe 1982). In all the forests de- scribed, A. saccharinum was mixed with Fraxinus pennsylvanica and Ulmus americana in the canopy, and Laportea canadensis was a major component of the understory. 130 Rhodora [Vol. 101 Type IV forests are similar in vegetation composition and structure to Quercus palustris—Fraxinus pennsylvanica forests de- scribed in Connecticut (Metzler and Damman 1985), Acer rub- rum/Onoclea—Boehmeria alluvial forests in Rhode Island (Barrett and Enser 1997), and Q. palustris—A. rubrum—Carex grayi-Geum canadense temporarily flooded forests described for New England (Sneddon et al. 1998). Type IV forests as described here differed from the above community types in having A. saccharinum rather than A. rubrum codominant with Q. palustris in the overstory. Floodplain forest plots classified as Type V appeared to be seasonally saturated as indicated by prominent soil mottling close to the soil surface. Type V forests are similar in composition to Acer rubrum—Onoclea sensibilis forested wetlands found on poor- ly drained glacial lake sediments in Connecticut (Metzler and Tiner 1992) and to A. rubrum alluvial forests described for Mas- sachusetts oxbows (Holland and Burk 1984). They correspond to the A. rubrum—Fraxinus pennsylvanica seasonally flooded forest alliance of low terraces and bottomlands occurring throughout the Eastern United States (Sneddon et al. 1998), and they are prob- ably common in Massachusetts. Acer rubrum alluvial swamp for- ests similar to Type V forests have been described as a species- rich variant of A. rubrum swamps that are abundant throughout southern New England (Golet et al. 1993). Type VI includes high floodplains that are flooded very infre- quently, perhaps no more than several times per century (Jahns 1947). Type VI forests are similar in composition to high-terrace Acer saccharum-Tilia americana—Matteuccia struthiopteris for- ests in Vermont (Sorenson et al. 1998) and to high-terrace A. saccharum—A. saccharinum—Fraxinus americana forests in New Hampshire (Bechtel and Sperduto 1998), except that A. sacchar- um was uncommon in high-terrace plots in Massachusetts. Type VI forests are most closely related to A. rubrum—Prunus serotina/ Athyrium felix-femina forests in New Hampshire which also had Maianthemum canadense and Uvularia sessilifolia as common associates (Bechtel and Sperduto 1998). Vegetation patterns in relation to environmental parame- ers. The relationships between vegetation patterns and soil characteristics shown here are similar to those found across flood- plain forest communities of the north-central United States. In New York, Acer saccharinum—Fraxinus Sp. associations similar 1999] Kearsley—Floodplain Forest Communities 131 to Types III and IV occurred on fine-textured soils, while Populus deltoides—Platanus occidentalis—Ulmus sp. associations similar to ype I occurred on coarser-textured, alluvial sands and gravels (Huenneke 1982). In southern Illinois, Robertson and Weaver (1978) found that Fraxinus pennsylvanica attained its highest im- portance under prolonged and deep flooding, and occurred only at sites that had mottling near the surface. In Massachusetts, F. pennsylvanica attained its highest indicator values of 19 and 6 in community Types IV and V, respectively (Table 2). Results of the current study support the conclusions of Veneman and Tiner (1990) that Boehmeria cylindrica is restricted to hydric floodplain forest soils, while Laportea canadensis and Matteuccia struthiop- teris are restricted to nonhydric soils. Boehmeria cylindrica was an indicator of floodplain forest community Types III, IV, and V, which occurred mainly on hydric silt loams. Matteuccia stru- thiopteris and L. canadensis were strong indicators of Type I and ype Il, respectively, which both had predominantly coarse-tex- tured, nonhydric soil profiles. Soil pH was significantly different among floodplain forest community types in Massachusetts because there was a strong geographical distributional pattern among types, and soil pH ap- peared to be correlated with geographic location. Soils on the Connecticut, Deerfield, and Housatonic Rivers all had higher pH than those along rivers in eastern Massachusetts (Tables 3 and 4). That difference may be, in part, related to differences in flooding frequency among rivers. Dollar et al. (1992) found soil pH to be the closest correlate with variation in vegetation in northern Mis- souri floodplain forests and suggested that sites with higher flood- ing frequency and input of fresh alluvium had higher pH. This is supported in Massachusetts, where Types I, II, and IL, which were located on large rivers with broad alluvial deposits (the Con- necticut, Deerfield, and sections of the Housatonic Rivers), had the highest mean pH values. High pH values on the Housatonic River (pH 7-8) are also related to the influence of carbonate rich edrock in western Massachusetts. CONSERVATION IMPLICATIONS Thirty-eight natural or semi-natural true floodplain forest sites (Types I-IV) ranging in size from 1 to 30 ha were identified in Massachusetts: 20 were primarily Major-river floodplain forest 132 Rhodora [Vol. 101 communities (Types I and II), 6 were Transitional (Type III), and 12 were Small-river types (Type IV). Of the 38 sites identified, only 10 were found to be high-quality examples based on their condition, size, and landscape context. The ten high-quality flood- plain forests included five Major-river sites (four on the Con- necticut River and one on the Housatonic River), one Transitional site on the Mill River in Hatfield, and four Small-river sites (three in the Connecticut River basin and one on the Threemile River). With the exception of the one site on the Housatonic River, all of the high-quality examples of Major-river floodplain forests oc- curred on either public land or privately owned conservation land. Transitional and Small-river floodplain forests are less well-pro- tected in Massachusetts. Due to their limited occurrence in the state and the habitat that they provide for five state-protected rare plant species, Transitional and Small-river floodplain forest com- munities warrant active protection efforts. Although land acquisition and conservation restrictions are im- their distribution, vegetation composition, environmental charac- teristics, and quality in Massachusetts. Type V Alluvial swamp forests are probably widespread, but high-quality examples may be limited. An in-depth inventory and classification of those com- munities is warranted with emphasis on identifying remaining high-quality examples. Type VI High-terrace floodplain forests are very limited in Massachusetts because most river terraces have been cleared and ine a 1999] Kearsley—Floodplain Forest Communities 133 converted to agriculture. Some high-terrace forests occur on the Millers, Westfield, and Green Rivers and parts of the Connecticut River, but more detailed inventories and vegetation and environ- mental analyses are warranted. ACKNOWLEDGMENTS. I am indebted to the many people who helped make this project possible. I would especially like to thank Pat Swain for her assistance in the field and for her insights and support throughout the project. Rebecca Anderson provided dedi- cated and invaluable volunteer field assistance during the field sea- son. Darren Singer, Karen Searcy, Pam Weatherbee, Matt Hickler, Betty Anderson also assisted with field data collection. Jeanne An- derson and the Massachusetts Audubon Society (MAS) contributed site information and data from MAS properties. Janice Stone and Donna Peterson at the University of Massachusetts Resource Map- ping Center helped with air photo interpretation. Peter Fletcher, Al Averill, and Peter Veneman provided help with soil samplin methodology. Study design and data interpretations benefited great- ly from discussions with Glenn Motzkin, Pat Swain, David Foster, Doug Bechtel, Eric Sorenson, Dan Sperduto, Julie Lundgren, and Mark Anderson. Figure 1 was prepared by Dorothy Graaskamp at the Massachusetts Department of Fisheries, Wildlife and Environ- mental Law Enforcement’s Geographical Information Systems Pro- gram. I also thank Henry Woolsey and the Massachusetts Natural Heritage Program staff for their support. This project was funded in part by a State Wetlands Protection Development Grant from the U.S. Environmental Protection Agency. LITERATURE CITED BaiLey, R. G. 1995. Description of the Ecoregions of the United States, 2nd . Misc. aay No. 1391 (rev.), U.S.D.A. Forest Service, Washington, DC. BARRETT, N. AND R. ENSER. 1997, Alluvial plant communities within the span Rasta major basin, Rhode Island. Unpubl. report, Rhode Is- land Natural Heritage Program, Providence, RI BECHTEL, D. A. AND D. D. SperpuTo. 1998. Floodplain forest natural com- munities along major rivers in New Hampshire. Unpubl. report submitted to the U.S. Environmental Protection Agency, New Hampshire Natural Heritage Inventory Program, Concord, NH. Brown, S. AND D. L. PETERSON. 1983. Structural characteristics and biomass 134 Rhodora [Vol. 101 production of two Illinois bottomland forests. Amer. Mid]. Naturalist 110: 107-117. BuELL, M. EF AND W. A. WISTENDAHL. 1955. Floodplain forests of the Raritan River. Bull. Torrey Bot. Club 82: 463-472. Do.iar, K. E., S. G. PALLARDY, AND H. G. Garret. 1992. Composition and environment of floodplain forests of northern Missouri. Canad. J. Forest. Res. 22: 1343-1350. DuFRENE, M. AND P. LEGENDRE. 1997. Species assemblages and indicator spe- cies: The need for a flexible asymmetrical approach. Ecol. Monogr. 67: 345-366. Dunn, C. P. AND FE STEARNS. 1987. A comparison of vegetation and soils in floodplain and basin forested wetlands of southeastern Wisconsin. Amer. Midl. Naturalist 118: 374— Frye, R. J., II AnD J. A. QUINN. 1979. Forest development in pate to topography and soils on a floodplain of the Raritan River, New Jerse Bull. Torrey Bot. Club 106: 334-345. GoLeET, F C., A. J. K. CaLHoun, W. R. DERAGON, D. J. Lowry, AND A. J. GoLD. 1993. Ecology of Red Maple Swamps in the Glaciated Northeast: A hae Profile. Biological Report 12, U.S. Dept. of Interior, Fish and Wildlife Service, Washington, DC GrirFiTH, G. E., J. M. OMERNIK, S. M. PIERSON, AND C. W. KIILSGAARD, 1994, The Massachusetts Ecological Regions Project. Publication No. 17587- 74-6/94-DEP, U.S. Environmental Protection Agency, Washington, DC. Haro, E. D., K. P. Lewis, AND W. A. WISTENDAHL. 1989. Gradient analysis of floodplain forests along three rivers in unglaciated Ohio. Bull. Torre ey Bot. Club 116: 258-264 D W. A. WISTENDAHL. 1983. The effects of floodplain trees on Bienen vegetation patterns, microtopography and litter. Bull. Torrey Bot. Club 110: 23-30. HILL, M. O. 1979a. TWINSPAN—a FORTRAN program for arranging mul- tivariate data in an ordered two-way table by Semaine of the indi- viduals and attributes. Cornell University, Ithaca, ——— 1979b. DECORANA—a FORTRAN p rogram as detrended corre- spondence analysis and reciprocal averaging. ‘Co mell University, Ithac a M. M. ann C. J. Burk. 1984. The herb — of three ait oxbow swamp forests. Rhodora 86: 397-415. eeranici L. E 1982. Wetland forests of Tompkins County, New York. Bull. Torrey Bot. Club 109: 51-63. JAHNS, R. H. 1947. Geologic Features of the Connecticut Valley, Massachu- setts, as Related to Recent Floods. Water-Supply paper 996, U.S. Dept. of Interior, Washington, DC. Kartesz, J. T. 1994. A A i ia of the Vascular es of the United States, Canada, and Greenland. 2nd ed. Timber Press, Portland, OR. KEARSLEY, J. B. 1999. Rare te non-native plants of eae aaENr flood- plain forests. Rhodora 101:200—205. McCune, B. AND M. J. MEFFORD. 1997. PC-ORD. Multivariate Analysis of Ecological Data, Version 3.0. MJM Software Design, Gleneden Beach, OR. MENGEs, E. S. 1986. Environmental correlates of herb species composition 1999] Kearsley—Floodplain Forest Communities 135 in ast hci Wisconsin floodplain forests. Amer. Midl. Naturalist hg DAR. i WALLER. 1983. Plant strategies in relation to elevation sai light in floodplain herbs. Amer. Naturalist 122: 454-473. METZLER, K. J. AND A. W. H. DAMMAN. 1985. Vegetation patterns in the Connecticut River flood plain in bee to frequency and duration of oe Naturalist Canad. 112: —547. DR. W. Tiner. 1992. ct of Connecticut. Report invest. 13, eataiciens Geological and Natural History Survey, Hartford, CT. MorzkIn, G. 1993. Uncommon plant communities of the Connecticut Valley, Massachusetts. Unpubl. report, Massachusetts Natural Heritage and En- dangered Species Program, Westborough, MA. . 1995. Inventory of uncommon plant communities of western Mas- sachusetts: 1993-1994. Unpubl. ee niacarie ni Natural Heritage and Endangered Species Program, Westborough, M NATURAL RESOURCE CONSERVATION SERVICE. {oe Field me for identifying and interpreting soil condition (Draft). Unpubl. draft, Natural Resource Conservation Service, Washington, DC. PETERSON, D. L. AND G. L. ROLFE. 1982. Nutrient dynamics of herbaceous vegetation in upland and floodplain forest communities. Amer. Midl. Naturalist 107: 325-339. ee a A. AND G. T. WEAVER. 1978. Vegetation and tree species pat- terns near the northern terminus of the southern floodplain forest. Ecol. SEARCY, K. B., B. LINDWALL, AND T. Enz. 1993. Plant community and veg- etation analysis of floodplain areas along the Nashua River, Fort Devens, with particular emphasis on the floodplain and adjacent slopes of the Massachusetts Natural Heritage and Endangered Species Program, West- borough, : SNEDDON, L., M. ANDERSON, AND K. METZLER, eds. 1998. International clas- sification of ecological communities: Terrestrial vegetation of the Eastern Region (draft 2/98). Unpubl. report. The Nature Conservancy, Eastern Regional Sie Boston, MA SocoLow, R. S., L. ¥. Comeau, R. G. CASEY, AND L. R. RAMSBEY. 1995. Water a Data: Massachusetts and Rhode Island Water Year 1995. Water-Data report MA-RI-95-1, U.S. Geological Survey, Wash- DC ington } Gan E., M. Lapin, B. ENGSTROM, AND R. Popp. 1998. Floodplain forests of Vermont: Some sites of ecological significance. Unpubl. report s — to the U.S. Environmental Protection Agency, Vermont Nongame Natural Heritage Program, Waterbury, VT. Tuostson, E. AND J. JENKINS. 1992. Natural communities of the Deerfield VENEMAN, P. L. M. AND R. W. TINER. ren So oil-vegetation correlations in the Connecticut River floodplain of western Massachusetts. Biological Report 90(6), U.S. Fish and Wildlife ie. shington, DC. ZIMMERMAN, G. M.., OETZ, AND . MIELKE, JR. 1985. Use of an im reiki ‘statistical method for group ce a study effects of prai- e fire. Ecology 66: 606-611 RHODORA, Vol. 101, No. 906, pp. 136-142, 1999 JALTOMATA LOJAE (SOLANACEAE): DESCRIPTION AND FLORAL BIOLOGY OF A NEW ANDEAN SPECIES THOMAS MIONE AND Luis A. SERAZO Department of Biological Sciences, Central Connecticut State University, New Britain, CT 06050-4010 ern Peru is described. This species is distinguished by the following features: ible, herkogamous, and sometimes protogynous. Microscopic, densely stain- ing, multicellular glands that may be osmophores are located on the perianth. erries are eaten by humans. Key Words: edible fruit, floral biology, herkogamy, Jaltomata, osmophore, olanaceae Jaltomata includes about 40 neotropical herbaceous and shrub- by species distributed from Arizona, U.S.A., to Bolivia, as well as on the Galapagos Islands and the Greater Antilles (Mione et al. 1993). The fruits of most Species are eaten uncooked (Davis and Bye 1982; Mione 1992). This description of a new species is a contribution to ongoing systematic studies of this genus (Mi- one 1999; Mione and Bye 1996; Mione and Leiva 1997). MATERIALS AND METHODS 136 1999] Mione and Serazo—Jaltomata lojae 137 the nose. To examine seed set, flowers were manually pollinated during the pistillate and hermaphroditic phases. Undehisced an- thers were removed at the time of pollination. TAXONOMIC TREATMENT AND DISTRIBUTION Jaltomata lojae Mione, sp. nov. TyPE: ECUADOR. Prov. Loja: by guard station on the E side of Celica, 1950 m, growing along roadside, 2 May 1991, D. M. Spooner, R. Castillo, and L. Lépez 5037 (HOLOTYPE: CONN). Figures 1 and 2. A J. cajamarca Mione atque J. sagasteguii Mione foliis pilis dactyliformibus, corolla sub anthesi rotata et ovarii disco lato dif- ert. Perennial shrub; branches and leaves densely villous, bearing finger hairs, these sometimes gland-tipped, 0.15—6 mm long, be- coming less villous with age. Leaves alternate, often geminate, ovate, to 15 cm long X 6 cm wide, the margin entire to sinuate- ae Deeper the petiole to 4.5 cm long. Inflorescence um- , to 9-flowered. Peduncle 5—9 mm long (at flowering), sedienlé 8-11 mm long (at flowering), both having a dense cov- ering of erect finger hairs to 1.5 mm long, some hairs gland- tipped. Calyx (at flowering) light green, 9.5 mm in diameter, strongly reflexed, abaxially villous with gland-tipped finger hairs, forked hairs, and dendritic hairs all 1-2 mm long, and abaxially and adaxially with stalked multicellular glands 62-80 ym long (Figure 2), at fruit maturity calyx diameter to 12 mm. Corolla infundibular when partially open, rotate when fully open, white, with two green proximally positioned maculae straddling the ra- dial vein to each corolla lobe (Figure 1), 5 prominent lobes al- ternating with 5 small lobules, 25—27 mm in diameter on plants we grew, 18 mm in diameter on the holotype, adaxially glabrous, abaxially with sparsely but evenly distributed stalked multicel- lular glands 62—80 ym long (Figure 2). Two classes of hairs ex- tend out from the corolla margin: marginal hairs to 110 ym long, and submarginal (attached abaxially) hairs to 0.5 mm long. Sta- mens 5, 5 mm long; filaments on living plants angling away from style and slightly curved outward, villous on basal 1/4 of the length with unpigmented finger hairs to 1.2 mm long; anthers 2.1—2.5 mm long prior to dehiscing, 1.6—-1.8 mm long after de- hiscing, when corolla is fully open exserted out of corolla 2 mm, 138 Rhodora [Vol. 101 Jaltomata lojae. ER on plant grown from seed of the Figure 1. Prt pit, Bac Units are millimete 1999] Figure 2. Perianth gland (75 ym long). Multicellular head stains deeply with neutral red, the stalk cell does not. Drawn by L. A. S. from plant grown from seed of the type collection. otherwise included. Pollen grains 32.5 1m average diameter (n = 22 grains). Ovarian nectar disk orange (visible by eye on living flowers). Style 9-11 mm long, slender and straight (Figure 1), exserted 3-8 mm beyond the anthers; stigma shallowly bilobed, 0.4 to 0.75 mm wide on pressed specimens, broader than the style (Figure 1), the papillae 30-60 ym long. Mature fruits orange. 140 Rhodora [Vol. 101 Seeds numerous, ovate to reniform, 1.3—1.6 mm long. Chromo- some number n = 12 (nine counts). DISTRIBUTION, ECOLOGY, LOCAL NAMES, USES. This species is dis- tributed in southern Ecuador and northern Peru in disturbed hab- itats from 1900 to 2700 m elevation. Its local names are ‘“‘ubillos”’ (Ellemann 66799) and “uvilla’” (van den Eynden and Cueva 342). It has been used ‘“‘for sunburn, used with alcohol for bath’’ (Ellemann 66799) and the fruits are edible (van den Eynden and Cueva 342). OTHER SPECIMENS EXAMINED: specimens made from plants grown from seeds of the type collection Mione 560 (CONN, HAO, MO, NY, QCA). Ecuador. Prov. CHIMBORAZO: Cafion on the Rio Chanchan, directly above the village of Huigra, 29-31 May 1945, Camp E-3498 (Ny); Prov. LOJA: Lugma Huycu 12 km north of Saraguro, 19 Jan 1989, Ellemann 66799 (AAU not seen, NY); Cerro Sozoranga, Colaisaca-Utuana, km 0.5, 24 Apr 1994, Jorgensen et al. 567 (MO not seen, Ny); Sevillan, 26 Mar 1995, van den Eynden and Cueva 342 (LOJA not seen, NY). Peru. Dept. prura, Prov. Huancabamba: Abra de Porculla, entre Olmos y Jaén, ladera con monte bajo, 22 Apr 1964, Ferreyra 15667 (kK, us); Carretera entre Canchaque y Huancabamba, km del 16 al 25 desde Canchaque, 17 Apr 1987, Diaz and Baldeén 2395 (Mo, NY); Perculla, 2 May 1981, Llatas and Laes 631 (HUT not seen, MO, NY). FLORAL BIOLOGY To look for structures that may release scent, flowers were stained in dilute neutral red for several hours. The stigma (in- cluding papillae) stained darkly with neutral red but the style did not. Anthers and pollen stained darkly while filaments did not. Multicellular glands, located on the abaxial face of the corolla and both faces of the calyx, stained darkly, except for the stalk cell (Figure 2). Similar glands have been described on the leaves of Solanum (Seithe 1979) and Physalis (Seithe and Sullivan 1990). Neither the corolla margin hairs nor the hairs of the fila- ments absorbed stain. On the multicellular finger hairs only the glandular tip, when present, absorbed stain. Hand-sections of epi- ermal tissue of the ovary disk, stained and then observed with a compound microscope, revealed that the nectary disk absorbs Stain only in the immediate vicinity of the stomata. The guard cells stained deeply, and the cells surrounding the guard cells also absorbed stain but staining was less intense. Flowers produced a 1999] Mione and Serazo—Jaltomata lojae 141 subtle fragrance that was described as licorice-like, vanilla-like, or faintly sweet by the seven people polled and the authors. (Two of these people also detected a fragrance in the empty/control jar.) Flowers remained open 5 to 7 days (mean 5.7 days, n = 9 flowers) and closed each night. Within an inflorescence one to four flowers were open at a time. The corolla was pale green prior to anthesis but after the corolla opened for the first time it became white and remained white. The anthers of a flower either dehisced a few at a time over the course of several hours, or two or three of the anthers dehisced one day and the others dehisced the next day. Anthers dehisced prior to 8:30 am. Some flowers exhibited one day of protogyny, with the stigma protruding through the partially open corolla. Two out of three flowers man- ually pollinated during the pistillate phase set seed, as did many (percentage not calculated) of the flowers that were manually pol- linated during the hermaphroditic phase. In the greenhouse, plants did not set fruit unless hand-pollinated. This is likely due to her- kogamy: the stigma is located 3 to 8 mm from the anthers. Nectar drops at the base of the corolla (alternating with the stamens) were large enough to be observed by eye. The broad, orange ovary disk is concentric around the green ovary and increases the diameter of the ovary by 1 to 1.5 mm over what the ovary di- ameter would be without the disk. Nectar seemed to be secreted by the ovarian disk but may also be secreted by the base of the corolla. ACKNOWLEDGMENTS. We thank David M. Spooner for seeds and review of the manuscript, Clinton E. Morse for care of living plants, Kancheepuram N. Gandhi for Latin translation, and Janet R. Sullivan and Gregory J. Anderson for review. LITERATURE CITED Davis, T. IV AND R. A. BYE, JR. 1982. Ethnobotany and progressive domes- tication of Jaltomata (Solanaceae) in Mexico and Central America. con. Bot. 36: 225-241 Kearns, C. A. AND D. W. INouyE. 1993. Techniques for Pollination Biolo- gists. University Press of Colorado, Niwot, C Miong, T. 1992. The systematics and speinieien of Tiidonialen (Solanaceae). Ph.D. dissertation, Univ. Connecticut, Storrs, CT. 142 Rhodora [Vol. 101 . 1999. Jaltomata Il: New combinations for five South American spe- cies (Solanaceae). Brittonia 51: 31-33. , G. J. ANDERSON, AND M. NEE. 1993. Jaltomata I: Circumscription, description and new combinations for five South American species. Brit- tonia 45: 138-145. AND R. Bye, Jr. 1996. Jaltomata chihuahuensis (Solanaceae): A new combination and observations on ecology and ethnobotany. Novon 6: 8-81. AND S. LeIva G. 1997. A new Peruvian species of Jaltomata (Sola- naceae) with blood-red floral nectar. Rhodora 99: 283-286. SEITHE, A. 1979. Hair types as taxonomic characters in Solanum, pp. 307— 319. In: J. G. Hawkes, R. N. Lester, and A. D. Skelding, eds., The Biology and Taxonomy of the Solanaceae. Linnean Society Symposium Series No. 7. Academic Press, New York. DJ. R. SULLIVAN. 1990. Hair morphology and systematics of Phys- alis (Solanaceae). PI. Syst. Evol. 170: 190-204 RHODORA, Vol. 101, No. 906, pp. 143-162, 1999 THE REPRODUCTIVE BIOLOGY OF MAGNOLIA GRANDIFLORA LARRY K. ALLAIN National Wetlands Research Center, 700 Cajundome Blvd., Lafayette, LA 70504 MICHAEL S. ZAVADA! AND DouGLas G. MATTHEWS Department of Biology, Providence College, Providence, RI 02918 ACT. The reproductive Cape of Magnolia grandiflora was inves- ested at three localities in south Louisiana. Over the 3—4 day flowering period, the flowers of M. ine pore dec chaes in sex expression (protogyny), stigmatic receptivity (self- and cross-compatibility to self-incom- patibility), UV reflectance (strong reflectance of the stigmas to strong reflec- tance i carried pollen, bees (non-native Apis mellifera and indigenous Lasioglossum bruneri) were frequent floral visitors and were the only floral visitors whose behavior showed any correlation with the array of floral changes that occurred over the 3—4 day flowering period. Key Words: Magnolia grandiflora, reproductive biology, halictid bees, bees Magnolia is considered to be primitive among angiosperms. A widely accepted view of the evolution of the flower is the Euan- thial or Anthostrobilus theory. This hypothesis asserts that the primitive flower is large, actinomorphic, solitary, white (some- times pink or yellow), and borne terminally. The primitive flower may have a showy, undifferentiated perianth and numerous floral parts arranged spirally on an elongated axis. The stamens are broad, three veined, and each has four elongate microsporangia on its adaxial surface. The gynoecium is apocarpous, with con- duplicate carpels that enclose a few ovules (Arber and Parkin 1907; Bessey 1897, 1915; Canright 1952, 1960; Maneval 1914). This widely held view of the magnolian flower has been consid- ered a theoretical starting point for understanding angiosperm evolution (e.g., Cronquist 1981). However, recent phylogenetic analyses (Crane 1985; Donoghue and Doyle 1989; Loconte and Stevenson 1990, 1991; Nixon et al. 1994), and fossil evidence of ' Reprint requests should be addressed to MSZ. 143 144 Rhodora [Vol. 101 early angiosperms (Crane et al. 1994, 1995: Friis et al. 1994, 1995; Taylor and Hickey 1992) suggest that the morphological features of the archetypical angiosperm flower are unresolved. There is evidence to suggest that the diversification of angio- sperms is associated with the diversification of the Apidae (Mich- ener and Grimaldi 1988a, b; Crepet 1984, 1996; Crepet et al. 1991). The first occurrence of angiosperms, however, significantly predates the first occurrence of bees in the fossil record (Laban- deira and Sepkoski 1993). The importance of the Diptera as early angiosperm pollinators has recently attracted attention (Kearns 1992; Kearns and Inouye 1993: Ren 1998). The Diptera have their origin in the Late Triassic—Early Jurassic (Rohdendorf 1974), prior to the origin and the diversification of angiosperms. Recent studies have shown that the floral foraging behavior of partitus the bracts of the inflorescence produce a specialized high lipid tissue (50% by dry weight) that the beetles consume along with pollen (Beach 1982). Taxa that are unequivocally beetle pol- linated frequently have floral modifications that specifically influ- ence beetle behavior in the flower or inflorescence. Despite a diversity of insect groups reported to visit Magnolia, most investigators have focused on the occurrence of beetles in the flowers of Magnolia, and on their role in pollen transfer (Bak- with the floral behavior of the beetles. urposes of this study are as follows: a) to examine the floral characteristics that may function as insect attractants and/ 1999] Allain et al.—Reproductive Biology of Magnolia 145 or rewards, i.e., UV reflectance of floral parts, nectar secretion and composition, pollen availability, and the origin and longevity of the floral fragrance; b) to determine the self- and cross-com- patibility of the flowers during anthesis and how this may relate to floral characteristics that are pollinator attractants and rewards; and c) to determine the kinds, numbers, and pollen loads of the various floral visitors, their behavior in the flowers of Magnolia grandifiora, and how this behavior may be related to floral char- acteristics that function as pollinator attractants and rewards. MATERIALS AND METHODS The investigation was carried out at three localities in south Louisiana. Site #1 consisted of 59 cultivated trees located throughout the city of Lafayette, Louisiana, a suburban habitat. Site #2 was a single tree in the Louisiana State Arboretum, lo- cated in Evangeline Parish, Louisiana. The 121.41 hectare arbo- retum is a secondary growth forest that is 75-80 years old. The tree at Site #2 was used for observing pollinators in the forest canopy. Access to the 30—40 m forest canopy was achieved using mountain climbing techniques (Perry 1978). Site #3 consisted of a monoculture of over 30 trees of varying ages at the Louisiana Nursery, Opelousas, Louisiana. The nursery is located in a rural area and the trees were grown under horticultural conditions. Pollinator attractants and rewards. To determine UV re- flectance, flowers were photographed with a 35 mm camera using a Kodak Wratten UV Filter No. 18A. This filter transmits long wave UV radiation (320—400 nanometers). High speed (ASA 400) T-Max 400 professional black and white film was used to record the reflectance patterns. The origin of the floral fragrance was determined by dividing the flower into separate parts (gynoecium, androphore, upper pet- als, and lower petals) and placing the separated floral organs in sterilized glass jars for 20 minutes. Thirty volunteers rated each of the floral parts according to the intensity of the odor (0 being the least and 5 being the most fragrant). The flowers used for hand pollinations (see below) were mon- itored for the presence of a stigmatic exudate (nectar) over the four-day flowering period. The nectar used in the sugar analysis was collected from first-day flowers from a variety of plants at 146 Rhodora [Vol. 101 Site #1 by using capillary tubes. (Nectar production ceased after the first day of anthesis.) The nectar was immediately placed in a cooler with ice packs, transported to the lab, and refrigerated at 1°C. The nectar was run on 20 X 20 cm EM Reagent, Silica Gel 60 Type plates. The gel thickness was 0.25 mm. After the plates were spotted, the sugars were run over 15 cm with one or two ascents, against ten sugar standards of various dilutions of a 15% (w/v) stock solution. The standards included glucose, man- nose, fructose, galactose, xylose, ribose, rhamnose, sucrose, malt- ose, and mannitol. The solvent used was a 9:6:3:1 solution of n- butanol: acetic acid: chloroform: water. The plates were devel- oped at 100—110°C for 10-20 minutes after being sprayed with a 1:3 (v/v) solution of sulfuric acid: methanol. The presence or absence of protein in the nectar was determined by the application of 2% ninhydrin in acetone (w/v) to nectars spotted on a Thin Layer Chromotography (TLC) plate and then baked at 100°C. € mean number of pollen grains per flower was estimated by removing thirty stamens from three flowers collected from three different trees at Site #1. Each of the thirty stamens was agitated with 0.01% Tween 20 in a known volume of water. Pol- len grains were counted on a hemacytometer grid. The mean number of pollen grains per stamen and flower was calculated. In addition, the number of ovules per flower was recorded to derive an estimate of the pollen-ovule ratio (P/O). Determination of self- and cross-compatibility. At Site #1, flowers were hand pollinated to determine stigmatic receptivity and cross- and self-compatibility. The pollinations were divided among ten treatments, 14—39 plants per treatment. Four of the treatments consisted of cross-pollinations on days 1—4 of anthesis, four consisted of self-pollinations on days 1—4, one treatment was an unpollinated bagged control for days 1—4, and one treatment was an unbagged control for days 1—4, to determine the seed set under natural conditions. Prior to anthesis, the gynoecia of the hand pollinated flowers were bagged with tubular nylon hosiery. After the hand pollina- tions, the flowers were tagged. Pollen was collected for various pollinations in paper envelopes. The pollen was stored at —10°C with a desiccator (Williams 1980). To insure that all pollinations from the stored pollen had a similar level of Viability, the duration of pollen viability was tested. Flowers were collected from three 1999] Allain et al—Reproductive Biology of Magnolia 147 different trees during the first morning of anthesis, the petals were removed, and the floral column with the stamens was placed on a sheet of paper. Within 24 hr., the anthers dehisced, the floral column was removed, one aliquot of pollen was stored at room temperature (21°C), and one aliquot was stored at —10°C. Pollen viability was tested from each of the aliquots daily for five con- secutive days following anthesis using the method of Alexander (1969). The gynoecia were collected as they ripened and the number of stigmas, the number of carpels setting seed, and the number setting two seeds were recorded. The percentage of ovules fertil- ized was calculated by dividing the total seed set by the total number of ovules (2/carpel) and multiplying by 100. The per- centage of ovules fertilized for the ten treatments was compared using ANOVA (analysis of variance). A general linear model for unbalanced designs was chosen and a two-factor general linear model ANOVA was run on Minitab (Cruze and Weldon 1989). The model was fitted with two main effects, crossed and selfed treatments. The four days over which pollinations were made were treated as an interactive factor with four levels. F-ratios were compared at the 0.01 level of significance to detect differences among treatments. Significant differences between treatments were then tested using Tukey’s multiple-comparison (W) proce- dure with Alpha = 0.05 (Cruze and Weldon 1989; Ott 1988). Observations of insects. The types of insects and the number of visits by each taxon at Sites #1, #2, and #3 were recorded by direct observation or by videotape. Each of the three sites was monitored 14 times at equally spaced intervals between May 19 and July 29. Insect visitor data were recorded for an average of three hours per observation time, yielding a total of 42 hours of observations per site. Unfamiliar insects were collected for iden- tification and five individuals of each species were collected as voucher specimens. All insects except thysanopterans were placed in a kill jar containing ethyl acetate. Thysanopterans were fixed in alcohol-glycerine-acetic acid (AGA) killing solution. After two weeks in the AGA, the thysanopterans were dehydrated in an alcohol series and mounted on microscope slides for identification (Borror et al. 1989). For each site the number of visits by each type of insect was recorded. The duration of the visits was re- corded by direct observation and from time lapse and real-time 148 Rhodora [Vol. 101 videotaping of the flowers. The pollen load per individual of each insect species was calculated as the mean number of grains car- ried on the insect specimens collected. The pollen carried on the corbicula of Apis mellifera was not available for stigmatic de- position and so was not included in the pollen load of this species. However, the pollen carried on the legs of halictid bees dislodged easily and was available for stigmatic deposition; thus these pol- len grains were included in the pollen load for this species. The relative importance (RI) of various insect species as pol- linators was calculated using the following equation: ade 44°29'W, 28 Jul 1967, Hansen, Kliim-Nielsen & Ollgaard 67-535 (CAN); Ju- 178 Rhodora [Vol. 101 lianehaab, near Sgen, 30 Aug 1937, Grontved 2114 (DAO); Kangerdlugsuak, Knud Rasmussen Land, Skaergaerd Peninsula, 17 Aug 1936, Wager & Wager s.n. (DAO); Kangerssuneq Quigordleq, Anivia, 60°19'N, 44°07’W, 4 Jul 1966, Hansen 66-1047 (Mo); Kjerulf Fjord, 8 Aug 1937, Oosting 1014 (cas); Kong Oscars Fjord, 72°14’N, 23°55’W, 1 Jul 1956, Raup, Raup & Washborn 25 (CAN); Kangmiut, 60°00’N, 44°28’W, 2 Jul 1967, Hansen, Kliim-Nielsen & Allgarrd 67-924, 969 (NY); Praestefjeld, 66°55'N, 53°35’W, 5 Jul 1949, Gelt- ing s.n. (COLO); Quinqua, 60°21'N, 23 Jul 1925, Porsild & Porsild s.n. (us); Scoresbysund, 71°20'N, 24°40'W, 13 Aug 1937, Sorensen 259 (MT); Skeldal, 72°15'N, 24°W, 16 Jul 1963, Spearing et al. 171 (mic); Séudre Strémfjord, 7 Aug 1927, Erlanson 2584 (MICH, NY); Tasissarssik Fjord, 66°05’N, 37°00'W, 14 Jul 1963, Gribbon 28 (CAN); Tasiusak, 61°45'N, 25 Jul 1889, Hartz s.n. 1962, Hansen, Hansen & Petersen 2235 (DAO); Ymer Island, Botanikerbug- ten, 73°08'N, 25°10’W, 18 Aug 1932, Sérensen 3116 (CAN). Norway. Norland Province, Junkerdalen, Salten, Mt. Solvagtind, 66°48’N, 15°35’E, 9 Aug 1859, Behm s.n. (uc), 8 Aug 1859, Schlyter & Behm s.n. (F, Ny) 18 Aug 1883, Nessen 805 (F), 9 Jul 1948, Jordal 1202 (F, MICH DAO, UBC); Madadam Oblast, Chukotskiy Peninsula, (NE coast) near mouth of Chegitum River, 12 Aug 1971, Sekretareka, Sitin & Yurtsev (ALA); middle branch of Erguveem River (left bank) near mouth of Vatamkaivan River, Pepenveem River, | Aug 1970, Nechaeva (ALA); middle branch of Utaveem Mountains, middle branch of southern Pekul’nayevem River, 8 Aug 1979, Korobkov & Sekretareva (Ny); Anyuiskoye Upland, 15 Aug 1973, Petrovsky (ALA); southern spur of Teniah Mts., at source of the Loran River, 14 Aug 1972, Gorbukova, Makarova & Plieva (ALA); SW coast of Chukotskiy Pen- insula, near Nunligran settlement, 28 Aug 1970, Afonina, Korobkov, Plieva & Khrenov (ALA); Anyuiskoye Upland, Pogingen River, 8 Aug 1976, Pe- trovsky & Korobkov (ALA). United States. ALaska: Alaktak, Half Moon, 70°45'N, 155°00’W, 1 Aug 1949, Spetzman 2439 (CAN, US); Anaktuvuk Pass, 12-15 Aug 1960, Hultén DAO); Bonanza Creek, Eagle Summit, 16 Jul 1949, Scamman 5270 (GH); Brooks Range, airstrip at ‘Nolan’, 19 Jun 1949, Jordal 1839 (BH, MICH); 46 i of Arctic Village, 68°40'N, 146°30’W, 18 Aug 1973, Hettinger 814 (CAN); Cane Creek, 68°35'N, 144°50’W, 8 Aug 1972, Hettinger 159 1999] Dunlop and Crow—Taxonomy of Carex 179 (ALTA); Cape Beaufort, 3-7 Aug 1961, Hultén s.n. (DAO, GH); Chip River, 70°26'N, 154°50'W, 17 Jul 1956, Wiggins 13674 (ps, us); Chitina River head, 16 Jun 1925, Laing 20 (CAN); Chugach Mts., Anchorage, 29 Jun 1948, LePage 23355 (us); Circle Hot Springs, 138 mi. N of Fairbanks, 17—22 Jul 1936, Scamman 69 (Gu); Colville River, 150°45'W, 69°45'N, 10 Aug 1953, Cantlon et al. 649 (msc); Delta River, S of Donnely Inn, 10 Aug 1966, Foote 8075 (RM); Daipaious Creek, 21 Jul 1958, Packer s.n. (ALTA); Don- nely Dome, mile 250 Richardson Hwy., 63°47'N, 145°45'W, 2 Aug 1951, Cody 6284, 6286 (DAO); Farwell Lake, 62°33'N, 153°36'W, 3-4 Aug 1949, Drury 2463 (GH); Fairbanks, Miller House on Steese Hwy., 12—28 Jul 1940, Scamman 2009 (GH); Firth River, 2 mi. S of junction Firth & Mancha Creek, 11 Aug 1961, Stone (RM); Fish Creek, 70°19'N, 151°58'W, 26 Jul 1977, Murray & Johnson 6532 (CAN); Index Mountain 40 mi. ENE of Arctic Village, 68°15’N, 144°10’'W, 11 Jul 1973, Hettinger 235, 246 (CAN, ALTA); Jago Lake, 69°26’N, 143°47'W, 23 Jul 1957, Cantlon & Gillis 57-1295 (msc); King Lodge, Border, Dawson Road, (E of Chicken), 12 Jul 1963, Spetzman 4883 (CAN); Kodiak, 28 Jul 1904, Piper 4776 (us); Kogosuknuk River, 69°45'N, 151°45’W, 16 Jul 1953, Borman, Rebuck & Cantlon 35] (MSC); 69°46’N, 151°40’W, 16 Jul 1953, Borman, Rebuck & Cantlon 398, 403 (MSC); Kongekat River Hill, 138 mi. NNE of Arctic Village, 69°34'N, 141°50'W, 19 Jul 1973, Hettinger 347 (CAN, ALTA); Kaness River, 15 mi. SSE of Arctic Village, 67°46’N, 143°45'W, 3 Jul 1973, Hettinger & Boyce 175 & 176 (ALTA); Kotzebue Sound, 9-16 Aug 1945, Scamman 3967 (GH); Kotzebue, 12 Aug 1938, Anderson 4679 (cas); Big River, 61°55’N, 154°25’W, 10 Jul 1950, Drury 4211 (GH); Kuskokwim River, Swift River, 62°40'N, 152°30'W, 19 Jul 1961, Viereck 5067 (CAN); Kenai Peninsula, Steton Creek Valley, 3 Aug 1951, Calder 6444 (cas, DAO); Moose Pass, 60°32'N, 149°32'W, 31 Jul 1951, Calder 6388 (DAO); Knife Ridge, 2 mi. N of Knifeblade, 2 Aug 1951, Jones 717 (ws); Kokrines Mts., 65°17’N, 154°30'W, 6 Jul 1926, Porsild 659-60 (CAN, MT, US); Lake Noluk, 2 se 1950, Thompson 1337 (ps, us); Lake Schrader, 69°25’N, 145°00'W, 8 Jul 1948, Spetzman 529 (Ds, us); Lazy Mt., E of Palmer, 29 Jul 1965, Mitchell 729b6 (DAO); Livengood, 9 Jul 1944, Anderson 9019 (CAN, MSC); Lodiack, 21 Jul 1904, Piper 4776 (pom); Mt. McKinley National Park, 2 mi. N of N entrance, 29 Jul 1967, Hermann 21517 (micH, NY); Mt. Eielson, Coppers Mt., 11 Jul 1956, Viereck 1250A (MIN, RSA); Hines Creek, 6 Aug 1950, Bailey 5017 (uc); Savage River, 31 Jul 1932, Henderson 14790 (ORE); Tok- lat Cabin, 11 Jul 1939, Murie 35 (RM); Polychrome Pass, mile 43, 1-10 Jul 1964, Hultén s.n. (AA); 63°43'N, 149°15'W, 13-22 Jun 1937, Scamman 585 (GH); Meade River, ca. 15 km from Atkasuk, 70°28'N, 157°25'W, 30 Jun 1966, DeBenedictis 92, 534 (MICH); milepost 50, along Pipeline Haul Road, 6 Aug 1981, Allred, Welsh & White 1214 (rsa); Mt. Dustin, 21 mi. from Nome, 4 Jul 1938, Anderson 3768 (cas); Nabesna River, 7 Aug 1902, Schrader & Hartman 67 (us); Nelchina Caribou Range, Tyrone Creek, (us); Nome Quad, meadow below radio tower, Anvil Mt., 64°31'N, 165°30’W, 11 Jul 1982, Kelso 82-39 (COLO); Nome, 1900, Blaisdell 139 (uc); Ogotoruk Creek, Cape Thompson, 27 Jul 1966, DeBenedictis 421 (MICH); Okpilak Valley, 69°25'N, 144°02'W, 30 Jun 1958, Cantlon & Mal- 180 Rhodora [Vol. 101 com 58-0117 (msc); Old John Lake, 27 Jul 1950, Jordal 375] (BH, MICH, MT); Pastolik, 5 Jul 1928, Miller 88c (Us); Port San Juan, Evans Island, 10 Aug 1948, Eyerdam 7031 (MIN, OSC, RM, WS); Prince of Wales Island, Vir- ginia Mt., 7 mi. S of Pt. Baker, 19 Jul 1972, Jaques 1476 (osu); Rapids Lodge, 138 mi. S of Fairbanks on Richardson Hwy., 25-28 Aug 1937, Scamman 1046 (Gu); Seward Peninsula, 64°33'N, 163°45’W, 5-6 Aug 1926, Porsild & Porsild 1193-94 (CAN, GH); Sheenjek River, 68°22’N, 143°55’W, 19 Jun 1956, Schaller 54 (mT); Sheenjek River, 68°36’N, 143°45’'W, 11 Jun 1956, Schaller 163 (mT); Snow Camp, Sagavanirkton River, 1958, Korando & Shanks s.n. (NY); Sunset Pass, 69°40’N, 144°45'W, 13 Aug 1948, Spetz- 5433 (GH); Umiat Mt., 2 Jul 1953, Borman 3394 (mT); 40 mi. NW of Umiat, 29 Jul 1951, Jones 723 (ws); Umiat, 69°25'N, 152°10'W, 25 Jun 1953, Bormann, Rebuck & Cantlon 118 (msc); Upper Marshfork River, 46 mi. NNW of Arctic Village, 68°40'N, 146°30’W, 18 Aug 1973, Hettinger 814 (ALTA); White River Valley, 61°42’N, 141°39’W, 17 Aug 1968, Murray 2279 (CAN); Alaska-Yukon Boundary, Firth River and Mancha Creek, 11 Aug Hilltop Mine, 12 Jul 1967, Weber 13299 (coLo); Fairplay, Horseshoe Cirque, T10 R79W S12, 21 Jul 1985, Dunlop & Orlando 2025 (NHA); South Park, 1873, Wolfe 1002 (COLO, F, MICH, NY, US); IDAHO: County unknown, Soda Springs, 25 May 1934, Davis 83-34 (Ny); Upper Priest River, 20 Jul 1925, Epling 7513 (uc); MAINE: Aroostook Co., Aroostook River Basin, 15 Jun 1940, Chamberlain 1579 (MAINE, uc); Aroostook River, Fort Fairfield, 5 Jun 1901, Fernald s.n. (GH, MAINE); Piscataquis Co., Mt. Katahdin, 4 Jul 1856, Blake s.n. (MAINE, NHA); North Basin headwall below Hamlin Peak, 1 Aug 1929, Ewer 226 (mass); W end of North Basin, 26 Jul 1929, Ewer t: 2749 (US); MICHIGAN: Delta Co., Escanaba River ca. 1 mi. NE of Cornell, ca. 10 mi. NW of Gladstone, 23 Aug 1982, Voss 15553 (MICH); MINNESOTA: Gunsight Pass, 25 Aug 1919, Standley 18139 (ny, US); Pigeon Pass, 22 Jul 1958, Bamberg 92 (coLo); Reynolds Mt., 2 Aug 1960, Schofield s.n. (MON- 1999] Dunlop and Crow—Taxonomy of Carex 181 TU); St. Mary Lake, 6 Aug 1919, Standley 17150 (us); MacDonald Lake, 3 Aug 1895, Williams s.n. (NY, US); Altyn Peak, 13 Jul 1919, Standley 15596 (NY, Us); Divide Mountain, 9 Aug 1964, Harvey & Pemble 7175 (MONTU, WTU); Teton Co., Pine Butte Preserve, 23 Jun 1982, Lesica 2055 (MONTU); Antelope Butte, 22 Jul 1982, Lackschewitz & Ramsden 10049 (COLO, MON- TU); Duhr Fen, 16 Aug 1982, Lesica 2408 (MONTU, WTU); Mt. Patrick Pass, 30 Jul 1983, Lackschewitz 10609 (MONTU, NY); NEVADA: Elko Co. , Ruby Valley, Point Hot Springs, T27N RS8E S15, 20 Jun 1984, Tiehm, Atwood & Williams 8748 (NY, ORE); Ruby Mts., Seitz Lake, T32N RS58E $20, 16 Sep 1983, Goodrich, Smith & Tuhy 20183 (BRY); Ruby Mts., T29N R57E, NW of Harrison Pass, 15 Aug 1980, Atwood 7713 (sry); Lamoille Canyon, Thomas Canyon Camp, 15 Jun 1941, Holmgren 1130 (Ny, uc); W of Ruby Mts., along Rt. 229, T32N R60E S19, 4 Aug 1985, Dunlop & Orlando 2140 (NHA); White Pine Co., Monte Neva Hot Spring, 17 mi. N of McGill, 4 Aug 1985, Dunlop & Orlando 2130 (NHA); NEW HAMPSHIRE: Carroll Co., Albany, Mt. Chocorua, 4 Jul 1978, Storks 385 (NHA); Hart’s Location, Mt. Willard, Butterwort Flume, 23 Sep 1984, Dunlop & Orlando 1965-1968 (NHA); Coos Co., Mt. Washington, mete Geers 31 Jul 1977, Storks 147 (NHA); Grafton C4. Franconia, Cannon Mt., 4 Aug 1960, Hodgdon 11670 (NHA); alpine areas of Franconia ty. Oakes Bee (MASS); Mt. Lincoln, summit, 18 Jul 1915, Fernald & Smiley 11607 (cu, GH, NY, US); Mt. Lafay- ette, 23 Aug 1865, Blake (GH, NHA); Grafton Co., Lyme, on Winslow Ledge, 25 Jun 1984, Dunlop, Korpi & Hency 2394 (NHA); NEW YORK: Essex Co N end Indian Pass, 5 Aug 1948, Smith 4602 (NA); Wilmington, Whiteface Mt., 7 Jul 1986, Dunlop & Orlando 2414 (NHA); Avalanche Pass, Mt. Mar- : Dunn Co. Mts., east slope, 11 Aug 1951, Stevens 1293 (CAN, UC, US); Rolette Co., ie Rolette & Thorne, 3 Jun 1913, Lunell 767236 (MIN, US); OREGON: Wallowa Co., E side Lostine Canyon, 18 mi. above Lostine, 22 Jul 1933, Peck 17861 (DS, NY, WILLU); Hurricane Creek, 23 Jul 1944, Peck 22549 (uc WILLU); Ice Lake ecm T4S R44E S12, 11 Aug 1961, Mason 1902 (OSC); UTAH: Duchesne Co., Ashley National Forest, Four Lakes Basin, 22 Aug 1974, Gasieeh "3736 (BRY); Emery Co., Scad Valley, T15S R6E S27, 5 Aug 1984, Lewis & Lewis 7758 (BRY, UT); Garfield Co., Dixie National Forest, Pine Lake Campground, 31 Jul 1985, Dunlop & Orlando 2100 (NHA); Upper Henderson Canyon, T35S RIW S32, ca. 11 NE of Tropic, 4 Jul 1983, Tuhy 863 (RSA); Iron Co., Cedar Breaks, T36S ROW S24, ca. 13 mi. S of Parowan, near Brian Head, 20 Jul 1977, Welsh & Clark 15512 (BRY, NY); VERMONT: fhnaaite tbat Co., Mt. Equinox, Deer Knoll, 5 May 1985, D Dunlop, Brackley & Thompson 2003 (NHA); Lamoille Co., Smugglers Ca ' 1956, yates 2288 (vT); Orleans Co., Willoughby Cliffs, 19 Jul 1885, Deane s.n. (BH, GH, NY), 25 Jun 1949, Hodgdon 6046 (NHA);, Mt. Pisgah, Lake Willoughby, along Rt. 5A, 16 Aug aan Ahles 78931 (MASS); WYOM- ING: Johnson Co., Big Horn Mts., 33 mi. NW of Buffalo, TS53N R87W S27, 7 Aug 1979, Nelson 4705 (RM); Park Co., Clay Butte, 15 Aug 1979, Dorn 3377 (RM); Sheridan Co., Big Horn Mt., TS7N R9OW S19, 5 Aug 1979, 182 Rhodora [Vol. 101 Nelson 4681 (RM); Tongue River, Aug 1953, Beetle 6297 (RM); Uinta Co., 5 mi. SW of Hilliard, 25 Jun 1950, Beetle 11062 (RM). 1b. Carex scirpoidea Michx. ssp. pseudoscirpoidea (Rydberg) Dunlop, Novon 7: 355. 1997. Carex pseudoscirpoidea Ryd- berg, Mem. N.Y. Bot. Gard. 1: 78. 1900. Carex scirpoidea var. pseudoscirpoidea (Rydberg) Cronquist, Univ. Wash. Publ. Biol. 17(1): 325. 1969. Type: u.s.A. Montana: Spanish Basin, Jul 1896, Rydberg 3064 (LECTOTYPE: NY! designated by Mackenzie). Rhizomes elongate, with regularly spaced shoots, internodes 1—2 cm long. Culms usually 1—few per node, arising from shoots of the previous year and retaining the withered, persistent leaf bases of the previous year, scabrous towards apex. Pistillate culms 0.5-1.5 mm wide at top, 0.8-2 mm wide at the base, 5—31 cm tall. Staminate culms 0.9-1.3 mm wide at top, 1.2-2 mm wide at the base, 9-27 cm tall. Leaf sheaths of the rhizome and culm base red-brown to brown-black, glabrous, shiny, becoming fi- brous with age. Leaves of the flowering shoots 3—5, clustered, blades diverging from one region up to 20 mm above the culm base, adaxial surface glabrous, scabrous along the Margins; in pistillate plants 7-19 cm long, 1.2-3.5 mm wide; in staminate plants 9-12 cm long, 1.5—2.8 mm wide. Vegetative leaves 5-8 per shoot; in pistillate plants 7-21 cm long, 1.6-3 mm wide; in staminate plants 9-13 cm long, 1.5—2.5 mm wide; ligules semi- circular, 1—-1.4 mm in height, 1.7-3 mm wide. Inflorescences uni- sexual, unispicate (very rarely with a short sessile lateral spike), erect, linear to oblong, densely flowered: pistillate spikes 10—34 mm long, 3.5—5 mm wide; staminate spikes 10-20 mm long, 3.5— 4 mm wide. Involucral bracts often absent, when present folia- ceous, shorter than the inflorescence, 6-40 mm long, occasionally scale-like (less than 1 mm and similar to inflorescence scales), inserted on culms 10—47 mm below spike, base occasionally au- riculate. Pistillate scales ovate, 2—2.6 (3) mm long, 1.1-1.5 mm wide, longer and wider than the perigynia, apically obtuse, red- 1-1.6 mm wide, abruptly contracted to a beak, lacking a stipe, nerveless, white to light green, becoming red-brown to dark 1999] Dunlop and Crow—Taxonomy of Carex 183 brown towards the apex, hirsute with white to tan hairs; body tightly enveloping the achene; beak 0.1—0.3 mm long, red-brown, straight at maturity, orifice entire and oval. Achenes light brown, 1.5—-1.8 mm long, 0.9-1.2 mm wide, lacking a stipe, filling the perigynia or at least % the length and width. Rachilla absent. Anthers 3 mm long. DISTRIBUTION. Carex scirpoidea ssp. pseudoscirpoidea is widely distributed in the higher elevations of the western mountains (Fig- ure 2). It is found chiefly in the San Juan Mountains in Colorado; Uinta and La Salle Mountains in Utah; the Sierra Nevada Range in California; Steen Mountains in Oregon; the Sawtooth Range in Idaho; the Little Belts Range, Anaconda-Pintlar Range and Beartooth Plateau in Montana; and the Okanagan Range in east- ern Washington and southern British Columbia HABITAT. Subspecies pseudoscirpoidea occurs at elevations from 3300 to 3900 m, on dry ridge sites and alpine fellfields with gravelly and non-calcareous soils. Subspecies pseudoscirpoidea is distinct ecologically, occurring chiefly in high elevation sites in mountain ranges in the West. This taxon is distinguished by culms that arise from second year shoots, clothed at the base by the withered and persistent leaf bases of the previous year. Generally, a single culm arises from a node and internodes of the rhizome are elongated, typically 1— 2 cm. The leaves are clustered, diverging from the shoot axis at One point approximately 10-20 mm above the rhizome, in con- trast to other taxa in which the leaves diverge from the stem at scattered intervals along the shoot axis. The plants generally have shorter and wider leaves than those of ssp. scirpoidea. In lectotypifying Carex pseudoscirpoidea, Mackenzie (1935) chose Rydberg 3064 (NY) as the lectotype. Unfortunately, this is a staminate plant and does not possess some of the diagnostic features of the taxon. REPRESENTATIVE SPECIMENS: Canada. BRITISH COLUMBIA: Cathedral Ridge, ca. 4% mi. N of Monument 95, 28 Aug 1972, Douglas & Douglas 4629 (ALA); Cathedral Park, Lake Lady Slipper, 12 Jul 1975, Hainault 7728 (Dao), 14 Jul 1975, Hainault 7526 (DAO); Lakeview ridge, 28 Jul 1976, Hainault 7962 (DAO); Mt. Apex, 11 Aug 1964, McLean & Haupt 65-64 (DAO), McLean & Marchand 65-63 (DAO); Mt. Bomford, Cathedral Lakes, Ashnolda District, 49°N, 120°15'W, 11 Jul 1951, Taylor 1359 (uBc). 184 Bhidora [Vol. 101 United States. caLiForNiA: Alpine Co., Carson Pass, Round Top Lake to Fourth of July Lake, 29 Aug 1974, Taylor 4910 (DAV); Mono Co., Minarets Wilderness Area, Inyo National Forest, Dana Plateau, N of Mt. Dana, TIN R25E S28, 7 Aug 1985, Dunlop & Orlando 2158 (NHA); COLORADO: Chaffee Co., Monarch Pass, 20 mi. W of Salida, 22 Jun 1926, Erlanson 2020 (mIcH); Manassas Creek, 24 Jul 1919, Clokey 3337 (BH, Mo, Ny, RM, UC, US); Gun- 1999] Dunlop and Crow—Taxonomy of Carex 185 nison Co., North Pole Basin, 14 Jul 1955, Weber & Barclay 9193 (COLO, cs, DS, MT, NY, RM, RSA, UC); La Plata Co., San Juan N a Forest, Chicago Basin, ‘Ealona. 29 Jul 1962, Michener 724 (cOLo); Sea an Co., San Juan National Forest, Eldorado Lake, T40N ROW, 19 Jul 1971, instare s.n. (COLO); Lake, 15 Aug 1979, ele aaiay 9154 (MONTU); Beartooth Mountains near g state line, 25 mi. S ed Lodge, T9S R19E S32, 22 Jul 1955, Cronquist 7998 (CAN, CAS, CU, D DS, GH, MICH, MONTU, MT, NY, OSC, RM, A, UC, WTU); Hell Roaring Plateau, 25 Jul 1921, Simms & Zeh 640 (RM); R13W S30 & 31, 24 Aug 1985, Dunlop & Orlando 2280 (NHA); Anaconda Pintlar Wilderness, T3N R14W S36, SW of Mt. Tiny on Storm Lake trail to Goat Flats, 24 Aug 1985, Dunlop & Orlando 2285 (NHA); Madison Co., schewitz & Fageraas 1081 a&b (MONTU); OREGON: Harney Co., Steen Mts., bove Alberson, 5 Jul 1925, Peck 14272 (CAS, F, PH, WILLU); Big Indian Gorge, T33S R33E S35, 28 Aug 1980, Wright 1468 (osc); UTAH: Daggett ah Leidy Peak, 31 Jul 1929, Dremolski D-7 (RM); Duchesne Co., Atwood Bluebell Lake, T4N RSW S31 & 32, 30 Jul 1980, Neese & Welsh 214016 (Ny); Uinta Mts. N slope 1 mi. SE of Island Lake, 25 Sep 1983, Neely & 1980, Welsh & Neese 213930 (Ny); San Juan Co., La Sal Mountains, W end Dark Canyon, Mt. Peale, T27S R24E $13, 26 Jul 1985, Dunlop, Orlando ph sop i 2075 (NHA); Mt. Mellenthin, T27S R24E S12, 26 Jul 1984, Tuhy 1746 (Bry); Summit Co., Bald Mt. summit, 3 Sep 1953, Lewis 267 (BRY, CAS); near divide W of Fish Lake on N slopes of Uinta Mts., T2N R1IE S2, 3 Sep 1945, Harrison & Harrison 10971 (BRY, US); rocky washes ing Dollar Lake, Henry’s Forks Basin, 12 Aug 1936, Maguire, Hobson & Ma- streamlet bam Ashley National Forest, 22 mi Marsh Peak, 19 Aug 1982, Goodrich 1 Ati (BRY, UT); 2 mi. NW of Paradise Park Res., T3N R1IW S2, 5 Jul 1980, Goodrich 14253 (BRY); E side Leidy Peak, TIN R19E S31, 24 Jul 1986, Goodrich 22074 (BRY); East Fox Lake, Uinta River, 14 Aug 1953, Lewis 230 (BRY); WASHINGTON: Okanogan Co., 186 Rhodora [Vol. 101 Chopaka Mt. ca. 11.5 mi. NW of Loomis, 16 Jul 1972, Douglas & Douglas 3858 (DAO); Tiffany Lake Pass, 24 Jul 1931, Fiker 406 (wILLU, ws); Windy Peak, ca. 17 mi. NW of Loomis, 17 Aug 1971, Douglas 3120 (RM); WYOMING: Fremont Co., Shoshone National Forest, East Fork Wind River, 8 Aug 1962, Johnson 249, 250 (cs, RM); Wind River Range, Roaring Fork Mt., 2 mi. NW of Silas Lake, 20-26 Jul 1965, Scott 535 (CAN, GH, UC); Johnson Co., Big Horn Forest, Grasshopper Ridge, Elk Lake, 28 Aug 1961, Johnson 144 (RM); Park Co., Island Lake, 19 Jul 1948, Daubenmire 48330 (ws); Little Bear Beetle 16673 (RM); White Rock Mt., near Green River Lakes, 8 Aug 1925, Payson & Payson 4605 (MO, MSC, NY, PH, RM, WS). 1c. Carex scirpoidea ssp. convoluta (Kiikenthal) Dunlop, Novon 7: 355. 1997. C. scirpoidea var. convoluta Kiikenthal, in En- gler, Das Pflanzenreich 38 (IV:20): 81. 1909. Type: U.S.A. Michigan: Thunder Bay Island, 18 Aug 1895, Wheeler s.n. (HOLOTYPE: B destroyed; ISOTYPES: BH! CAN! GH! MICH! MIN! MSC! NY! POM! vT!). Rhizomes short. Culms one to several per node, arising from current year shoots (lacking any withered persistent leaf bases of the previous year), scabrous especially at the apex. Pistillate culms 0.4—0.7 mm wide at the top, 0.8-2.2 mm wide at the base, (9.2) 19.5—35 (38) cm tall. Staminate culms 0.5—0.9 mm at the top, 0.9—-1 mm wide at the base, 9-31 cm tall. Leaf sheaths of the rhizome and the culm base red-brown to brown-black, gla- brous, shiny, coriaceous with acute hard tips. Leaves of the flow- ering shoots 2—4, not clustered, adaxial surface glabrous, margins scabrous; in pistillate plants 7-18 cm long, (0.8) 1.1-1.5 (2) mm wide; in staminate plants 6-15 cm long, 1—-1.5 wide. Vegetative leaves 4—9 per shoot; in pistillate plants 10.5—23.4 cm long, (0.7) 1-2 mm wide; in staminate plants 11-16 cm long, 1-1.7 mm wide; ligules semicircular, 0.2—2 (3) mm in height, 1-2 mm wide. Inflorescences unisexual, unispicate (very rarely with a short lat- eral, sessile or subsessile spike), erect, linear, densely flowered; pistillate spikes slender, (10.6) 15-21 (30) mm long, 2.5-3.5 (4) wide; staminate spikes 13-30 mm long, 2-3.5 mm wide. Involucral bract usually absent, when present foliaceous, shorter an the inflorescence, S21 mm long, base inserted on culm S- 12 mm below spike, auriculate. Pistillate scales ovate to obovate, (1.5) 2—2.4 mm long, (0.9) 1-1.2 mm wide, equal to or shorter than perigynia, apex obtuse to acute, red-brown to dark brown with narrow to broad hyaline margins, central midrib narrow 1999] Dunlop and Crow—Taxonomy of Carex 187 green-tawny to dark brown, extending to scale apex; margins en- tire. Staminate scales similar to pistillate, 2.5-3.2 mm long, 0.7— 0.9 mm wide. Perigynia ovate to obovate, 1.5—2.6 mm long, (0.7) 1—1.2 mm wide, as wide as the subtending scale, abruptly con- tracted to a beak, lacking a stipe, nerves absent, basally light green, becoming tawny to red-brown towards apex, hirsute with white to tan hairs, body tightly enveloping the achene; beak 0.1 mm long, red-brown and hyaline at tip, straight at maturity, orifice entire and circular. Achenes dark brown, 1—1.5 mm long, 0.6—0.9 mm wide, sessile or with 0.25 mm stipe, filling the perigynia or at least %4 the length and width. Rachilla absent. Anthers 1.5—2 mm long. DISTRIBUTION. This is the most sie apr sais of the four subspecies (Figure 3), occurring only e shores of Lake Huron on the Bruce Peninsula, on aoe of - Manitoulin District, Ontario, and on Drummond Island and Thunder Bay Is- land, Michigan. HABITAT. Subspecies convoluta is associated with alvar com- munities, characterized by Catling and Brownell (1995) and Ste- phenson (1983), as areas with sparse vegetation and thin soil over flat limestone or marble substrate. These open alvar communities contain a number of plant species that are calcicolous, often drought resistant, and grow in cracks of outcropped limestone “‘pavements.” Subspecies convoluta is distinguished by narrow, convolute leaves of the vegetative and flowering shoots, a strongly cespitose habit, and a higher number of flowering culms per plant than other subspecies. These narrow-leaves plants were first described as a variety of Carex scirpoidea by Kiikenthal based on specimens collected by Wheeler in 1895 from Thunder Bay Island, Michigan. Unfortu- nately, Kiikenthal’s Carex herbarium, presumably including the holotype of C. scirpoidea vat. convoluta, was sent to Berlin (B) and was destroyed during World War II (Stafleu and Cowan 1979). Isotypes of Wheeler’s 1895, Thunder Bay Island, Michi- gan, collections are available. PRESENTATIVE SPECIMENS: Canada. ONTARIO: Algoma District, Great Cloche Island, 26 Jul 1956, Soper & Fleischmann 6633 (CAN); Bruce Co., 188 Rhodora [Vol. 101 Figure 3. Distribution of Carex scirpoidea ssp. convoluta. DAO, MO, MT); Howdenvale, 31 Jul 1936, Watson 2904 (CU, NY, US); Lion’s Head, 11 Jun 1932, Marie-Victorin, Rousseau & Prat 45-922 (DAO, GH, MT); Little Pine Tree Harbour, Zinker Island Cove, 30 Jun 1982, Webber 4552 (TRTE); Oliphant, 9 Aug 1971, Montgomery 3693 (WAT); Red Bay, 9 Jul 1941, Sargent 5 (GH); Sauble Beach, 20 Jun 1934, Taylor & Fernald s.n. (wis); Stokes Bay, 26 May 1934, Krotkov 8759 (ny, US), 8792 (GH); Barrie Island, May 1979, Hogg s.n. (CAN, WAT); Georgian Bay Island National Park, Cove 1999] Dunlop and Crow—Taxonomy of Carex 189 Island, Bass Bay, 22 Sep 1981, Bobbette 7403 (wat); Zinkan Island, 45°04'N, 81°29’W, 15 Jul 1975, Cuddy & Emalie 1815 (CAN); Manitoulin District, Barrie Island, Rozels Bay, 29 Jul 1985, Hellquist 15513 (Nasc); Green Island Harbour, 20 Jul 1976, Ringius & Wilson 327 (wat); Hensly Bay, 26 May 1978, Morton & Venn 11521 (wat); La Cloche Peninsula, 20 Aug 1932, Fassett 14899 (wis, GH), 11 Jul 1957, Pease & Bean 26203 (GH); Whitefish River W side Hwy. 68, 3 Jul 1976, Catling & McIntosh s.n. (DAO, wis); Misery Bay, 28 Jul 1972, MacDonald & White 3593 (CAN); Murphy Point, 14 Jul 1952, Senn 5974 (DAO, MT, NY, PAC, US, WS); Tamarack Cove, 20 Jul 1932, Koelz 4206 (MICH, wi: Tamarack Point, Grassl 4594 (MICH). United States. MICHIGAN: Alpena Co., Thunder Bay Island, 18 Aug 1895, Wheeler s.n. (BH, CAN, GH, MICH, MIN, MSC, NY, POM, VT); Chippewa Co., Drummond Island, Meade Island, 19 Jun 1979, Voss 15074 (MICH). ld. Carex scirpoidea Michx. ssp. stenochlaena (Holm) Live and Léve, Taxon 13: 202. 1964. C. scirpoidea Michx. var. stenochlaena Holm, Amer. J. Sci. TV 18: 20. 1904. C. sten- ochlaena (Holm) Mackenzie, Bull. Torrey Bot. Club 35: 269. 1908. TYPE: CANADA. British Columbia: Chilliwack Lake, by a rivulet, 4000 ft., 12 Jul 1901, Macoun 33728 (LECTOTYPE: CAN 21326! designated herein; ISOLECTOTYPES: CAN! MO! Msc! NY! us!). Rhizomes short. Culms several per node, arising from shoots of the current year, (lacking any withered persistent leaf bases of the previous year), scabrous toward the apex. Pistillate culms 0.6— 1 mm wide at the top, 1.4-2.1 mm wide at base, 24—34 cm tall. Staminate culms 0.5—0.8 mm wide at the top, 2.2-3 mm wide at the base, 14—26 cm tall. Leaf sheaths of the rhizome and the culm base red-brown to brown-black, glabrous, shiny, coriaceous. Leaves of the flowering shoot 3—5, not clustered, adaxial surface glabrous, margins scabrous; in pistillate plants 12.5—25 cm long, 1.42.1 mm wide; in staminate plants 14—20 cm long, 1.5-2.4 mm wide. Vegetative leaves 5—6 per shoot; in pistillate plants 10-28 cm long, 1—2.5 mm wide, in staminate plants 19-24 cm long, 1.4-2.5 mm wide; ligules semicircular, 0.8— 1.5 mm in height, 1-2.5 mm wide. Inflorescences unisexual, unispicate (oc- casionally with a single, short lateral spike), drooping on lax culms, mostly clavate, loosely-flowered especially at base; pistil- late spikes 25-30 mm long, 3.5—6.5 mm wide; staminate spikes (few seen) 18-25 mm long, 4—5 mm wide. Involucral bract sin- gle, foliaceous, shorter than the inflorescence, 4-40 mm long, base inserted on culms 10—33 mm below spike, auriculate. Pis- tillate scales oblong-lanceolate, 2.4—3.5 mm long, 1—-1.5 mm 190 Rhodora [Vol. 101 wide, equal to or shorter than perigynia, apex subacute to acute, red-brown to black, without hyaline margins; central midrib ob- scure, only slightly raised, red-brown, but lighter in color than rest of scale, extending to the apex; margins ciliate. Staminate scales similar to pistillate, (3.5) 4.5-5 (6) mm long, (0.7) 1—1.4 mm wide. Perigynia lanceolate to oblanceolate, (2.8) 3—4 (5) mm long, 0.9—1.4 (1.6) mm wide, as wide as subtending scale, taper- ing gradually to beak, with short basal stipe, adaxial surface nerveless or with few short obscure basal nerves, red-brown to black, rarely tan, hirsute with tan to brown hairs, loosely envel- oping the achene in upper %; beak 0.3-0.5 mm long, dark brown and not hyaline, reflexed at maturity, orifice entire and abaxially oblique. Achenes light brown, 1.2-2 mm long, 0.8—1 mm wide, with 0.25—0.50 mm stipe, filling % to % the length of the peri- gynia. Rachilla absent. Anthers 3—3.5 mm long. DISTRIBUTION. This subspecies occurs in the Cascade Mountains in Washington, the Bitterroot Mountains in Montana, the coastal ranges in southern British Columbia, and at a few localities in Alaska and the Yukon (Figure 4). HABITAT. In the Bitterroot Mountains of Ravalli County, Mon- tana, this subspecies grows on bedrock terraces between 1615— 2620 m elevation, especially between 2130-2440 m. In Wash- ington it grows mostly between 1460-2010 m. This taxon is associated with weakly acidic soils which have high levels of magnesium and low levels of calcium. Edaphic requirements may be an important isolating factor for this sub- species. More detailed studies of habitat requirements and micro- environments are required before one can speculate on the history of this taxon and how it is related to the other subspecies. Subspecies stenochlaena is distinguished by lanceolate peri- gynia which are greater than 3 mm long, taper gradually to a beak, and are over 2.5 times as long as wide. The pistillate spikes are clavate, loosely flowered at the base, and borne on slender, lax culms which cause the spikes to droop. The pistillate scales are over 3 mm long, subtending hirsute perigynia with tan, yel- low, or golden-brown hairs. Both perigynia and pistillate scales are dark brown to black. Beaks are dark and reflexed at maturity, with an oblique mouth. Figure 4. Distribution of Carex scirpoidea ssp. stenochlaena. A lectotype, designated herein, has been selected from Ma- coun’s specimens since no specimen was chosen by Holm. The lectotype (CAN 21326) is a Macoun specimen bearing a “Geologic Survey of Canada” label, while others bear “Ex. Herb. Geologic 192 Rhodora [Vol. 101 Survey of Canada” labels. The latter are isolectotypes bearin the same number as the lectotype, but having a slightly different wording of the locality data. Some geographically based variation is present within this sub- species. Specimens from Washington and the Bitterroot Range in Ravalli County, Montana have the longest perigynia, often reach- ing 4.3 mm, and are the most distinct from ssp. scirpoidea. Spec- imens from Alaska, northern British Columbia and the Yukon ex- hibit tendencies towards ssp. scirpoidea. Fifteen percent of these plants have perigynia just under 3 mm long (high end of ssp. scirpoidea range) and perigynia length-to-width ratios over 2.5 (unlike those of ssp. scirpoidea which are less than 2.5 times as long as wide); they lack the clavate, loosely flowered spikes char- acteristic of plants of ssp. stenochlaena in Washington and Ravalli County, Montana. The intergradation observed in plants from Brit- ish Columbia and elsewhere might be the result of hybridization between ssp. stenochlaena and ssp. scirpoidea at localities where their ranges overlap. For this reason, subspecies stenochlaena is recognized at the subspecific level, rather than the specific level. REPRESENTATIVE SPECIMENS: Canada. BRITISH COLUMBIA: Bluster Mt., eee ble Mts., 14 Jul 1938, Thompson & Thompson 455 (CAS, DAO, F, GH, MO, US, WTU); Chilliwack Valley, 49°10’N, 121°-122°25'W, 12 Jul 1901, pias 33728 (CAN, MO, MSC, NY, US); Katherine Lake, 57°26'N, 126°48’W, 25 Jul Hollister 43 (NY, US); Mt. Assiniboine Park, Lake Magog, 16 Jul 1937, Rose 37550 (CAS, UC); Mt. Chelam, 15 Aug 1901, Anderson s.n. (MO); Noaxe Lake, 4 Aug 1957, Brink s.n. (DAO, OsU, UBC); Ellis Point & Mercer Point, W of Jul 1964, Hett & Armstrong 399 (DAO); Mt. Klitsa, 22 Jul 1971, Pojar 1 7} (UBC); YUKON TERRITORY: Alsek Valley, ca. 8 mi. W of Mackintosh, 5 Jul & Breitung 1198] (CAN); Canol Road, Mt. Sheldon, 3 Aug 1944, Porsild & Breitung 11708 (can), 11703 (GH); Kluane Lake, near Rusty Glacier, W of B Landing, 61°16'N, 140°15’W, 9 Jul 1968, Murray 1671 (can); Mackintosh, mile 1002, Alaska Hwy., Alsek Valley Road, 4—5 Jul 1957, Schofield & Crum fs 7560, 7561 (CAN). United States. ALaska: Charlie River, 64°50’N, 143°40’W, 30 Aug 1956, Argus 872 (RM, SASK); pentane Range, Tazalina Glacier, 61°45’N, 146°30'W, 1999] Dunlop and Crow—Taxonomy of Carex 193 19 Jul 1957, Viereck 2194 (CAN); Juneau, 10 Jul 1917, Anderson 365 (NY); Mt. McKinley National Park, 2 mi. N of N entrance, 29 Jul 1967, Hermann 215] a (MICH, NY); Wonder Lake, Argus 660 (SASK); Mt. Hayes, Palmer 606 (us); M Roberts, Juneau, 26 Jun 1925, Anderson 2A233 (GH); Yes Bay, 16 Jul (98) Howell 1705 (CAS, MSC, NY, US); MONTANA: Ravalli Co., Bitterroot National Forest, Bailey Lake, 22 Aug 1985, Dunlop & Orlando 2272 (NHA); Bitterroot- Selway Divide above Baily Lake, 12 Jul 1969, Lackschewitz 1344 (NY, RM); affin Lake Basin, Chaffin Peak, 30 Aug 1971, Lackschewitz 3397 (MONTU); Sheafman Lake, 17 Aug 1979, Lackschewitz 2317 (MONTU); St. Joseph’s Peak, 31 Jul 1969, Lackschewitz & Fageraas 1631 (MONTU); Tin Cup Lake, 19 Jun 1971, Lackschewitz 2727 (MONTU); Trappers Peak, 14 Aug 1946, Hitchcock & Muhlick 15381 (CAS, NY, WTU); White Mt., 11 Aug 1970, Lackschewitz & Smith 2277 (MONTU); WASHINGTON: Chelan Co., Crown Point, Holden-Lyman Lake Trail, 20 Aug 1956, Raven 10176 (cas); Ingalls Peak, 20 Jul 1925, St. John & Thayer 7239 (ws); Mt. Stuart, 23 Jul 1933, Thompson 9580 (DS, GH, MO, NY, Olympic Mt., Aug 1895, Piper 2243 (BH, GH, WS); Jefferson Co., Mt. Anderso 28 Jul 1936, Meyer 686 (Mo, ws); King Co., Denny Creek, 19 Aug 1936. prey e 13684 (CAS, MO, NY, PH, WS, WTU); Guy Peak, Snoqualmie Pass, 7 Aug 1933, Thompson 9690 (Ny, WTU); Kittitas Co., ‘Wecaly Creek Trail, T22N RISE ca. S1, 17 Aug 1985, Daendois & arcnen 224] (NHA); Fish Lake, 17 Jun 1934, Thompson 10663-4 (CAS, DS, GH, NY, POM, US, WTU, UWT); Okanogan Co., Hart’s Pass, ca. 20 mi. E of Diablo, 11 Jul 1971, Douglas 2866 (ALTA); Horse Shoe Basin, Sep 1897, Elmer 684 (MIN, MO, NY, POM, US, VT, WS); Snohomish Co., Mt. Pugh, 18 Aug 1938, Thompson 14340 (CAS, GH, WTU); Whatcom Co., Crater Mt., 20 Aug 1971, Naas 1240 (RM); Twin Lakes, Jackson Mt., 7 Sep 1927, St. John 8941 (ws, RM); Mt. Shuksan, ca. 22 mi. N of Rockport, 13 Jul 1969, Douglas 1417 (RM). 2. Carex curatorum Stacey, Leafl. W. Bot. 2: 13. 1937. C. scir- poidea var. curatorum (Stacey) Cronquist, Intermountain Flora 6: 113. 1977. TYPE: u.s.A. Arizona: Grand Canyon Na- tional Park, Kaibab Trail to Roaring Springs, 23 Jun 1933, Eastwood & Howell 1100 (LECTOTYPE: CAS 204972! desig- nated herein). Carex haysii Welsh, Mem. N. Y. Bot. Gard. 64: 124. 1990. TyPE: U.S.A. tah: Washington Co., Zion Canyon, Lower Emerald Pool, Spring- dale Sandstone, hanging garden, sandy bank and cliff face, ca. 4300 ft. T41S RLOW S9, 5 Jun 1989, Welsh, Clark & Hays 24335 (HO- LOTYPE: BRY; ISOTYPES: CAS, MICH!, NY!, POM, RM, UT). Rhizomes short and forming mats. Culms one to several per node, arising from current year shoots (lacking the withered per- sistent leaf bases of the previous year), strongly scabrous the full length of culm. Pistillate culms 0.4—1.2 mm wide at the top, 0.7— 5 mm wide at the base, (23) 35-91 cm tall. Staminate culms 194 Rhodora [Vol. 101 0.5—1 mm wide at the top, 0.9—-4.5 mm wide at the base, 20-74 cm tall. Leaf sheaths of the rhizome and culm bases light brown to purple-black, glabrous, dull, coriaceous. Leaves of the flow- ering shoots 2—6, arising from the base and not clustered, adaxial surface sparsely pilose especially along median adaxial groove and veins, margins strongly scabrous; in pistillate plants 12—57 cm long, 1.5—2.3 mm wide; in staminate plants 12-25 cm long and 1.2—-3.1 mm wide. Vegetative leaves 4-11 per shoot; in pis- tillate plants 14-55 (79) cm long, 1.1—2.3 mm wide; in staminate plants 18-55 cm long, 1.2-1.9 mm wide; ligules triangular, 1.4— 4.2 (5) mm in height, 1.1—2.3 mm wide. Inflorescences unisexual (rarely bisexual), not strictly unispicate, (occasionally with 1-2 short lateral spikes of the same sex in the axil of the involucral bract or subtending the terminal spike), erect or drooping on lax culms, linear, loosely to densely flowered; pistillate spikes 17—43 mm long, 2.5-5 mm wide, staminate spikes 13-37 mm long, 2— 5 mm wide. Involucral bract usually present, usually single, fo- liaceous or scale-like, shorter than or equal to length of the inflo- rescence, 6-11 cm long, base inserted 5-75 mm below spike, auriculate. Pistillate scales oblong-lanceolate, 2—3.5 mm long, 0.7-1.9 mm wide, half as long as to equaling the perigynia, api- cally acute, red-brown with narrow to broad hyaline margins; central midrib narrow, extending to apex, occasionally prolonged into a short awn; margins entire, sometimes ciliate. Staminate scales similar to pistillate, 3.5—-4.3 mm long, 1-1.3 mm wide. Perigynia obovate to ovate, 2-3 (4) mm long, 1.5-1.8 mm wide, wider than subtending scale, tapering gradually or abruptly con- tracted to a beak, with few obscure nerves on the adaxial surface over the achene, marginal nerves evident, pale green to tawny becoming red-brown towards apex, hirsute with white hairs; body not tightly enveloping the achene; beak 0.1-0.5 mm long, red- brown and hyaline at tip, straight at maturity, orifice entire and adaxially oblique. Achenes dark brown, 1.2-2 mm long, 0.8—1.2 mm wide, stipe 0.25 mm, filling only 4—% the length of the per- igynia and % the width such that the perigynia sides are com- pressed and contracted at the base. Rachilla often present. Anthers 1.7—2.4 mm long. DISTRIBUTION. Carex curatorum is limited to southern Utah and adjacent northern Arizona (Figure 5), especially along the Colo- rado and San Juan Rivers and their tributaries. 1999] Dunlop and Crow—-Taxonomy of Carex Figure 5. Distribution of Carex curatorum. 196 Rhodora [Vol. 101 HABITAT. This taxon grows in riparian or hanging garden com- munities, and may have been represented by more populations before the damming and flooding of the Colorado River. It occurs on Navajo Sandstone, described as a crossbed of fine-grained sandstone and gray limestone (Lohman 1975), and on the Kay- enta Formation, both of which are widespread substrata in south- ern Utah Carex curatorum is a distinct southwestern endemic. It is dis- tinguished by sparsely pilose adaxial leaf surfaces, most obvious with the scanning electron microscope but visible with a dissec- tion microscope. Achenes are not tightly enveloped by the peri- gynia and the pistillate scales are mostly shorter than the peri- gynia. The scales and perigynia are characteristically lighter in color than other taxa of section Scirpinae. Plants flower from April to May. Upon maturity, the perigynia and enclosed achenes disarticulate from the spikes, and are read- ily dispersed, unlike Carex scirpoidea which retains achenes and erigynia sometimes until the next growing season. Maximum culm height is greater than that of C. scirpoidea, and the achene Ils only a small portion of the perigynium. The unique achene micromorphology of C. curatorum and its sparsely pilose adaxial leaf surfaces segregate it from all other members of section Scir- pinae. In naming this species Stacey (1937) selected the epithet ‘‘cur- atorum”’ to honor Alice Eastwood and J. T. Howell, curators at the California Academy of Sciences, and the collectors of the type specimens. Stacey (1937) designated both a staminate and pistil- late plant, on separate sheets and with different collection num- bers, to serve as types. The pistillate specimen (Eastwood & Howell 1100 cas 204973), is selected herein to serve as the lec- totype, since it possesses the diagnostic features of this taxon. € staminate specimen (Eastwood & Howell 1101 cas 204973) other specimens (Eastwood & Howell 1045 cAs, 1089 cas) collected along with the type remain important original ma- terial from the type locality. Carex haysii Welsh, described in 1990, is treated here as a synonym since all the characters that are said to distinguish it overlap with C. curatorum except for perigynium length. How- ever, on many specimens, the longest perigynia of C. haysii lacked fully formed achenes. 1999] Dunlop and Crow—Taxonomy of Carex 197 REPRESENTATIVE SPECIMENS: United States. ARIZONA: Coconino Co., Gran Canyon National Park, Kaibab Trail to Roaring Springs, 22 Sep 1938, wood & Howell 7073 (Cas, F, MICH, MT, NY, POM, UC, US, WTU); Grand Canyon National Park, False President Harding Rapids, mile 43, 17 Mar 1974, Kar- piscak & Theroux 94] (ARIZ); Colorado River, Buck Farm Canyon, 40.75 mi. below Lees Ferry, % mi. above river, 29 Apr 1970, Holmgren, Holmgren & Ross 15481 (coLo); Mohave Co., Grand Canyon National Monument, aaa eap Pt., Saddle Horse Springs, 13 Jun 1941, Cottam 8652 (coLo, uid UTAH Kane Co., confluence of San Juan & Colorado Rivers, on San J ca. % mi. above second hanging garden on W in side-canyon, 9 Jun 1972. Aiwoed 40a (BRY); Glen Canyon National Recreation Area, Lake Powell, vicinity rth Escalante, ca. 3800 ft., T40S R9E S36, 28 May 1983, Welsh 22113 ra NY); Long Canyon, Waterpocket Fold, T39S R9E S1, 24 May 1984, Welsh 22850 (Bry); Coyote Creek Canyon, near Jacob Hamblin Arch, T38S R8E, 29 Jul 1985, Dunlop & Orlando 2087 (NHA); Glen Canyon Nat. Rec. Area, Cow Canyon, Waterpocket Fold, T38S R9E $36, 26 Jul 1983, Welsh, Welsh & Chatterley 22350 (BRY); San Juan Co., Lake Powell, Double Cove Garden, T40S R9E S25, 3800 ft., 1 Jul 1983, Welsh 22322 (Byu, UT); Lake Powell, Ribbon Canyon, canyon sides and hanging gardens, T41S RIOE SS, 25 Apr 1983, Welsh 21730 (BRY); Washington Co., Zion Canyon, Lower Emerald Pool, Springdale Sandstone, hanging garden, sandy bank and cliff face, ca. 4300 ft., T41S R1OW S9, 5 Jun 1989, Welsh, Clark & Hays 24335 (BRY, CAS, NY, POM, RM UT); Kayenta rhea: ca. 6000 ft., Kolob Section, hanging garden below Kolob Arch, ca. . SE of Kolob cea visitor center, T39S R12W S1, 25 May 1989, ha 18499 (BRY, CAS, NY UTC); Zion Canyon, Weeping Rock, T41S R1OW S82, ca. 4300 ft., 9 Tul ‘1988, Welsh, Clark & Charlesworth 24059b (BRyY); Zion Canyon, Lower Emerald Pool, Springdale Sandstone, hanging garden, ca. 4300 ft., T41S RI1OW S9, 2 May 1989, Welsh & Clark 24233 (BRY). EXCLUDED TAXA Carex gigas (Holm) Mackenzie, Bull. Torrey Bot. Club 35: 268. 1908. Carex scirpoidea var. gigas Holm, Amer. J. Sci. IV 18: 20. 1904. Type: U.S.A. California: Siskiyou County, Mt. Eddy (not known). Carex scabriuscula Mackenzie, Bull. Torrey Bot. Club 35: 268. 1908. TYPE: U.S.A. Wet meadow in the Cascade Mountains, 30 Jun 1902, Cusick 2849 (HOLOTYPE: NY!; ISOTYPES: CU!, Ds!, ORE!, osc!, POM!, uc!, ws!). The taxonomic status of Carex gigas and C. scabriuscula, ser- pentine endemics in California and Oregon, remains problematic. These two taxa are excluded from section Scirpinae because they are rarely unispicate, often not dioecious, and do not possess the pubescence of the perigynia characteristic of the section. Exclu- 198 Rhodora [Vol. 101 sion is further supported by evidence from chromosome numbers, leaf surface features, and ecology (Dunlop 1990). ACKNOWLEDGMENTS. We would like to thank the curators of the following herbaria for loans: AA, ARIZ, ALA, ALTA, ASC, BH, BYU, CAN, CAS, COLO, CS, DAO, DAV, DS, F, HSC, MAINE, MASS, MBG, MICH, MIN, MONTU, MSC, MT, NA, NY, OS, ORE, OSC, PAC, POM, RSA, RM, SASK, ble UBC, UC, US, UT, USAS, USFS, VT, WAT, WILLU, WIS, e thank A. L. Bogle, T. D. Lee, A. A. Reznicek, L. x pits J. R. Sullivan, M. Wirth, and E. G. Voss for advice and/or comments on this manuscript. This work was submitted to the University of New Hampshire by D.A.D. in partial fulfill- ment for the requirements of a doctorate degree. Research, field work, and/or publication preparation was supported by the fol- lowing: an Andrew Mellon Fellowship to the Naturalist-Ecologist Training Program at the University of Michigan, two grants from the University of New Hampshire Central University Research Fund, a Graduate School Dissertation Fellowship from the Uni- versity of New Hampshire, a graduate student travel award from the New England Botanical Club, Sigma Xi Grant-in-Aid of Re- search, a Gilmore Grant from New England College, and support from the New Hampshire Agricultural Experiment Station. This paper is Scientific Contribution Number 1974 from the New Hampshire Agricultural Experiment Station. LITERATURE CITED BAILEY, L. H. 1887. A ne synopsis of North American Carices. Proc. Amer. peer Arts 22: 1-157. pices bs M. AND V. R. nea: 1995. A review of the alvars of the hake region: Distribution, floristic eit biogeography sd protection Canad. Field-Naturalist 109: 143- Crins, W. J. AND P. W. BaLL. 1989. Taxonomy of te Het flava complex (Crveschnaiigls in North America and northern Eurasia. I. Taxonomic eatment. Canad. J. Bot. 67: 1048-1065. fines D. A. 1990. The biosystematics of Carex section Scirpinae (Cyper- aceae). Ph.D. dissertation, Univ. New Ham pshire, Durham, NH. , 7. Taxonomic changes in Carex (section Scirpinae, Cyperaceac). Novon 7: 355-356 HERMANN, F. J. 1957. New ome from the Canadian Rocky Mountains. Leafil. W. Bot. 8: 109- ok tues ed ise oO Cyperaceae. XXII. The Cyperaceae of the Chilliwack Valley, British Columbia. hail J. Bot: Ser. 4, 18: 12-22. Sicheemeas V. 1935. Carex. In: V. L. Komarov, ed., Flora of the USSR 1999] Dunlop and Crow—Taxonomy of Carex 199 3: 111-464. (English translation by N. Landau, Smithsonian Institute d the National Science Foundation, 1964. 3: 86-369). KUKENTHAL, G. 1909. Cyperaceae—Caricoideae. Das Pflanzenreich 4(20): 1- LOHMAN, S. W. 1975. The Geologic — eis Arches National Park. U.S. Dept. Interior, Geologic Survey Bulletin MACKENZIE, K. K. 1908. Notes on ra i Torrey Bot. Club 35: 266— 270 1935. Cyperaceae—Cariceae. N. Amer. Fl. 18: 1-478. eee A. 1803. Flora Boreali-Americana. Vol. II. Facsimile Edition, 1974. Hafner Press, New Yor NELMES, E. 1951. Facts and speculations on phylogeny in the tribe Cariceae of the pigs I. Sem. considerations. Kew Bull. 1951: 427—436. Snes H. AND M. Dum 1941. A new species of Carex and some notes this eae in pis ‘Coa Rhodora 43: 413-425. PAX, EA ea Bias Carex. In: A. Engler and K. A. Prantl, eds., Nat. Pflanzen- n 2(2). W. Englemann, Leipzig. petites ae A. 1990. Evolution in sedges (Carex, Cyperaceae). Canad. J. Bot. 68: 1409-1432. RypsBerG, P. A. 1900. Catalogue of the Flora of ee and the Yellowstone National Park. Mem. New York Bot. Gard. 1: 78. STACEY, J. W. 1937. Notes on Carex VII. Leafi. i Bot. 2: 13-15. STAFLEU, E A. AND R. S. Cowan. 1979. Taxonomic Literature, Vol. I. H— Le. Regnum Veg. Vol. 98. Bohn, Scheltema & Holkema, Utrecht. STANDLEY, L. A. 1985. Systematics of the Acutae group - och (Cypera- pane Rhodora [Vol. 101 Table 2. Comparison of staminate and androgynous inflorescence dimen- sions that have been used to distinguish Acalypha hederacea and A. mono- vs. A. monostachya). Lengths are shown as mean + standard error. Proba- bilities are based on two-tailed f-tests assuming unequal sample variances. A. Plant Sexuality Unisexual Plants Monoecious (n = 47) (n = 144) Peduncle length (cm) 1.85 + 0.065 1.95 + 0.051 £17627: 03242 i rtion length (cm) 2.06 + 0.085 2.14 + 0.053 0.845 0.400 Total inflorescence ngth (cm) 3.91. 0.133. 4.10 + 0.093 P1222 O265 B. Original Determination A. hederacea A. monostachya (n= 14 (n = 50) Peduncle length (cm) 1.89 + 0.046 2.04 + 0.092 1522-03132 Staminate portion length (cm) 212-* 0/052 2.14 + 0.093 0.235 0.815 Total inflorescence length (cm) 4.00 + 0.088 4.18 + 0.163 0.979 0.330 These results do not support the use of staminate spike length to distinguish Acalypha hederacea and A. monostachya. It is pos- sible that the complex has two species, differing in sexuality, with broadly overlapping geographic ranges. However, I found no oth- er morphological features that differentiate two groups. I suspect that Miller (1964) studied relatively few specimens (she gave no sample size) and that she measured the entire staminate spike, whereas McVaugh (1961) measured only the fertile staminate portion, hence the differences in their observations. When inter- preted this way, their measurements are consistent with mine. Because I can find no way to distinguish two s cies, I agree with McVaugh (1961) that only a single species is involved, which should be called A. monostachya. ACALYPHA PHLEOIDES CAV. AND A. LINDHEIMERI MULL. ARG. Acalypha phleoides is traditionally interpreted as a species of the arid highlands of eastern and central Mexico, ranging as far 1999] Levin—Acalypha in North America 223 north as Chihuahua and Coahuila. Overlapping with this in north- ern Mexico and extending into Arizona, New Mexico, and Texas, is what is usually called A. lindheimeri. According to Miiller (1866), A. lindheimeri has acuminate rather than acute leaves, the terminal tooth of the bracts subtending the pistillate flowers pro- longed rather than equal to the other teeth, and more slender style branches. I examined more than 400 specimens from throughout the range of these two taxa. Leaves on plants from Texas are mostly rhombic-ovate and acuminate, though lower leaves tend to be broader relative to their length and have acute apices. Sometimes the bracts subtending the pistillate flowers have elongate terminal teeth, as they do on two of the isosyntypes of Acalypha lindhei- meri I have seen (Lindheimer 520 [CAN!, GH!, MO!] and Lindhei- mer 688 [GH!, MO!]), but more often all the teeth are subequal, s they are on the third isosyntype I examined (Wright 1815 [GH!]). Leaves of plants from central Mexico south are ovate to suborbicular (especially the lower leaves) with consistently acute apices. The teeth of the bracts subtending the pistillate flowers are consistently subequal. There is also a tendency for the more southerly plants to have denser and coarser pubescence than the more northerly plants, especially those from Texas. Plants from Arizona, e.g., Blumer 1498 (ARIZ!, GH!, ISC!, NMC!; the type of A. lindheimeri var. major Pax & K. Hoffm.), and northern Mexico are intermediate between the Texan and central Mexican plants, with no obvious discontinuities. I could distinguish no differences in style branch thickness. The difficulty of quantifying leaf shape and its variation within individual plants makes statistic analysis extremely problematic. However, because leaf shape varies ap- parently continuously throughout the range of the group and bract toothing shows no consistent pattern, it appears that A. lindhei- meri was based on a few extreme specimens of A. phleoides. It therefore seems preferable to treat all these plants as a single species using the older name, A. phleoides. ACALYPHA VIRGINICA GROUP The Acalypha virginica group has received the greatest study of any North American members of the genus. Through much of the 18" and 19" centuries, there was considerable disagreement in interpretation of this group, but most authors recognized either 224 Rhodora [Vol. 101 one or two taxa. Miiller (1865, 1866), however, recognized four taxa in the group, all as varieties of A. virginica. In a series of papers, Weatherby (1927, 1937, 1940) attempted to sort out the variation in the group and to resolve a nagging typification prob- lem surrounding the name A. virginica. Ultimately he recognized three species, A. gracilens A. Gray with three varieties [var. gra- cilens, var. fraseri (Mill. Arg.) Weath., and var. monococca En- gelm. ex A. Gray], A. rhomboidea Raf. with two varieties [var. rhomboidea and the newly described var. deamii (Weath.) Weath.], and A. virginica. Unfortunately, the typification problem was not finally resolved until 1990 (Reveal et al. 1990), when conservation of the name and type of A. virginica established Weatherby’s (1937) treatment. Weatherby’s treatment (1937) continues to be widely used to- day. However, two additional treatments have also had some in- fluence. First, Miller (1964, 1969, 1970; Gandhi and Hatch 1988) recognized five species in the group, Acalypha deamii (Weath.) H. E. Ahles, A. gracilens, A. monococca (Engelm. ex A. Gray) Lill. W. Miller & Gandhi, A. rhomboidea, and A. virginica. In addition to segregating A. monococca as a separate species, she further realigned A. gracilens by treating var. fraseri as a syno- nym of A. gracilens var. gracilens and recognizing A. gracilens var. delzii Lill. W. Miller. The second treatment was by Cooper- rider (1984, 1995). Studying almost exclusively plants from Ohio and therefore not considering A. gracilens or A. monococca, he treated the remaining three taxa as varieties of A. virginica [thus A. virginica var. deamii Weath., A. virginica var. rhomboidea (Raf.) Cooperr., and A. virginica var. virginica]. In justifying his treatment, Cooperrider (1984) stated that he saw many intermediates between his varieties of Acalypha vir- ginica, particularly between var. rhomboidea and var. virginica, and further that “‘no single reliable diagnostic character or com- bination of characters’’ separates these taxa. Based on my obser- vations, I disagree. Instead, the apparent intergradation and lack of diagnostic characters reflect two problems. First, most of the characters used by Weatherby (1927, 1937) and Miller (1964, 1970) are not always reliable. Second, and clearly related to the first, misidentifications are rife in herbaria. On the approximately 4,000 specimens of A. rhomboidea and A. virginica I examined, about 18% of the annotations applied since 1940 (sufficiently after Weatherby’s publications to allow them to become widely 1999] Levin—Acalypha in North America phe used) were misidentified, about half of these bearing names of the other taxon. (In this calculation I excluded Miller’s annota- tions, with which I almost entirely concur.) Cooperrider’s own identifications are instructive. I have seen 96 specimens of the two species he annotated. We agree on almost all that he called A. virginica var. rhomboidea, but we would agree on only 44% (eight of 18) that he called A. virginica var. virginica. Perhaps this high frequency of misidentifications prevented him from see- ing the clear and reliable distinctions between these taxa. I found a similar rate of misidentifications among specimens of A. gra- cilens and A. virginica. (Another consequence of the high rate of misidentifications is that most published range maps are not re- liable.) Most regional floras that treat several taxa in the Acalypha virginica group (e.g., Cooperrider 1995; Gleason and Cronquist 1991; Miller 1970; Mohlenbrock 1982, 1986; Radford et al. 1968; Steyermark 1963) distinguish them using some combination of stem pubescence (long spreading and short incurved vs. just short incurved), petiole length (either absolute or relative to leaf length), leaf shape, shape and number of teeth or lobes on the bracts subtending the pistillate flowers, pubescence on these bracts, carpel number, and seed size. Table 3 summarizes these characteristics for the five species recognized by Miller (1964, 1970) and me. Some of these characters are unambiguous and serve to distinguish individual taxa. For example, A. virginica is unique in having long spreading eglandular trichomes on the ab- axial surface of the pistillate bracts, and mature plants always bear these (the lowermost bracts on the plant may lack the spread- ing hairs, however). Similarly, A. deamii always has gynoecia with two carpels and A. monococca has gynoecia with one carpel, whereas the remaining species have three carpels. Also A. deamii has seeds that are at least 2.2 mm long, whereas the other species have seeds no more than 2.0 mm long, with the exception of A. monococca, which occasionally has seeds to 2.4 mm long. Out- group comparison with A. alopecuroides Jacq., A. arvensis Poepp., A. australis L., A. brachystachya Hornem., A. indica L., and A. mexicana Miill. Arg. suggests that these unique charac- teristics are all apomorphies of the individual species (Table 3). Other characters may be useful in separating otherwise similar species. A sample of 100 specimens each of Acalypha virginica and A. rhomboidea showed that the number of lobes on the pis- Table 3. Summary of characters often used to separate species in the Acalypha virginica group, compiled from herbarium specimens from throughout the geographic ranges of the species. Values for quantitative characters are ranges. Sample sizes are n = 100 except for A. deamii, a rare species for which n = 35. Asterisks (*) indicate unambiguous unique species-level apomorphies based on outgroup comparison with A. alopecuroides, A. arvensis, A. australis, A. brachystachya, A. indica, and A. mexicana. A. mono- Character A. deamii_ A. gracilens cocca A. rhomboidea A. virginica Stems with long spreading hairs Never Never Never About 5% of plants About 90% of plants Petiole length (cm) 2.5-7.0 0.2-1.4 0.2-0.9 0.8-7.0 0.7-3.6 Petiole length/leaf blade length 0.42-0.94 0.09-0.30 0.08-0.20 0.34—0.89 0.23—0.66 Leaf blade length/width 1.5—2.0 2.3-8.0 2.8-8.7 1.4-3.2 2.1-4.5 Pistillate bracts with long spreading No No No No Yes (except sometimes eglandular hairs the lowermost on the plant)* Pistillate bracts with red glands No Yes Yes No No Pistillate bract tooth (lobe) length (mm 4.5-9.0 0.6—2.2 0.9-2.2 1.8-9.0 1.6—5.0 Pistillate bract tooth (lobe) length/pis- tillate bract length 0.44-0.75 0.08-0.28 0.10-0.25 0.30-0.75 0.21-0.50 Number of teeth/lobes on pistillate bracts (average of 3—S bracts) 5.38.0 9.0-13.3 9.0-13.7 5.7-8.7 10.0-—13.7 Carpel number . 1% 3 Seed length (mm) 2.2-3.1* 1.1-1.9 1.62.4 1.3-2.0 1.3-1.8 9C7 elopoyuy TOI T°A] 1999] Levin—Acalypha in North America 227 tillate bracts is a reliable character distinguishing these species. It is true that individual bracts of A. rhomboidea may have as many as nine lobes and bracts of A. virginica may have as few as nine lobes. However, averaging three to five bracts per plant gave no more than 8.7 lobes/bract for A. rhomboidea and no fewer than 10 lobes/bract for A. virginica (Table 3; Figure 1). The presence of red glands on the pistillate bracts, at least on the tooth apices and often scattered on the abaxial surface, distinguishes A. gracilens from very young plants of A. virginica that have not yet produced bracts with spreading trichomes. (Acalypha mono- cocca also produces these red glands.) A particularly interesting situation that has not been noted previously is that scattered throughout the range of A. rhomboidea, but more frequent in the southern states, are small plants with short petioles, small leaves, and small pistillate bracts with short teeth. These often appear in herbaria labeled A. gracilens, presumably because of the short petioles and bract teeth. However, tooth number is consistently nine or fewer, the petioles are at least 40% the length of the leaf blades, the bract teeth are more than 30% the length of the bracts, and the bracts do not bear red glands. These characteristics clearly demonstrate that these plants are simply small A. rhomboidea rather than A. gracilens (Table 3; Figure 1) Some characteristics that are frequently used to distinguish spe- cies in the Acalypha virginica group are not consistent and over- lap among species. Notably, these include leaf shape (described by blade length/width; all species in this group have the same general shape, so this ratio is an appropriate statistic) and both absolute and relative petiole lengths (Table 3; Figure 1). Although very different species may show no overlap, others, like A. rhom- boidea and A. bene show no reliable differences in these characters. Neither is the presence or absence of spreading tri- chomes on the =o reliable. Nearly 10% of the A. virginica specimens I sampled lacked spreading trichomes, whereas 5% of . rhomboidea specimens eae bore them. Spreading stem trichomes are absent on the remaining species. It is likely that use of these unreliable ie acd has contributed to the rel- atively frequent misidentifications and the taxonomic confusion in this group. The data summarized in Table 3 demonstrate that, in fact, in- termediates among the taxa in the Acalypha virginica group are exceedingly rare or absent. The taxa are often found growing 228 Petiole length/leaf length Rhodora [Vol. 101 1.0- 0.8 0.6 0.4+ ! Number of carpellate bract lobes 1 2 4 6 8 10 Leaf length/width 4 | | | 0.0 0.2 0.4 0.6 0.8 Carpellate bract lobe length/bract length 1999] Levin—Acalypha in North America 229 nearby or together, notably A. deamii with A. rhomboidea; A. gracilens with A. monococca, A. rhomboidea, or A. virginica; A. monococca with A. virginica; and, most frequently, A. rhomboi- dea with A. virginica (pers. obs.). Yet examination of over 6500 specimens revealed no clear evidence for hybridization, nor have I found evidence for hybridization during field work throughout much of the range of these taxa. The presence of unique apomorphies (Table 3) for Acalypha deamii (two carpels/flower, seeds at least 2.2 mm long), A. mon- ococca (one carpel/flower), and A. virginica (long spreading eglandular trichomes on the abaxial surface of the pistillate bracts), supports recognition of these taxa as species under both the phylogenetic (Davis and Nixon 1992; Nixon and Wheeler 1990) and genealogical (Baum and Donoghue 1995; de Queiroz and Donoghue 1988; Olmstead 1995) species concepts. Because they are distinguishable by nonoverlapping characters, A. graci- lens and A. rhomboidea would also be species under the phylo- genetic species concept, but because they lack unique apomor- phies (insofar as known), they would be metaspecies under the genealogical species concept. In either case, under both of these cladistically based species concepts, all the taxa in the A. virgin- ica group should be recognized at the rank of species. I do not recognize any infraspecific taxa in A. gracilens because variation in this species is clinal over broad geographic areas (Levin 1998). Key to the species of the Acalypha virginica group. I also include in the key the introduced Acalypha australis, which may be confused with members of this group. 1. Leaf blades linear to oblong-lanceolate; petioles rarely more than /, the length of the leaf blades; bracts of pistillate flowers with deltoid teeth Y,,—Y, length of the bract, with sparse to dense sessile red glands ................ (2) Figure 1. Plots showing the variation in the sie legigil virginica group along four characters. All species are represented by n 100 except A. deamii, a rare species with n = 35. Note that both axes in the upper graph are logarithmic. Key to symbols: A. deamii O, A. gracilens (1, A. monococca ©, A. rhomboidea A, and A. virginica ¥*. 230 Rhodora [Vol. 101 2. Pistil with 3 carpels, usually producing 3 seeds ..... es AG pees. A. woes oanu calypha gracilens 2. Pistil with 1 carpel, producing 1 seed .............. pad ONES 4 Hal RS EE Acalypha monococca 1. Leaf blades broadly lanceolate to ovate; petioles more than Y, (usually more than /,) the length of the leaf blades; bracts of pistillate flowers either with triangular to lanceolate lobes more than Y, the length of the bract or with rounded teeth less than ¥,, the length of the bract, lacking red ROUEN SW 2K JUG OTE OS. auth 6a (3) 3. Bracts of pistillate flowers hirsute with dense long spreading non-glandular hairs (also ciliate and often with stalked glands), and with (9—) 10-14 (-16) lobes more than Y, the length of the bract; stems usually hirsute with long spreading hairs ..... tcc gap nA, © bees oie Ae att ek Weer calypha virginica 3. Bracts of pistillate flowers without long spreading non- glandular hairs (may be ciliate or with stalked glands), and with either (S—) 7-9 (-11) lobes more than ¥, the length of the bract or 12-15 teeth less than “9 the length of the bract; stems very rarely u ( 4. Bracts of pistillate flowers with 12-15 rounded teeth less than '/,) the length of the bract ....... eee ed Ce Oke Feo als Acalypha australis 4. Bracts of pistillate flowers with (5—) 7-9 (-11) lan- ceolate lobes more than ¥, the length of the bract ( 5. Pistils with 3 carpels; seeds <2.1 mm We ss (eee owebaimlio Acalypha rhomboidea 5. Pistils with 2 carpels; seeds >2.1 mm long ..... aaa 5 tn'etcaly ghee SDL SDS | E Acalypha deamii Taxonomic treatment of the Acalypha virginica group. 1. Acalypha deamii (Weath.) H. E. Ahles in Jones & Fuller, Vasc. Pl. Illinois 301. 1955. Bitsieats i hip L. var. deamii Weath., Rhodora 29: 197. 1927. TyPE: NITED STATES. Indiana: Dearborn Co., road along White Water Be Py mi. northeast of Logan, Oct 20, 1924. C.C. Deam 41107 1999] Levin—Acalypha in North America 231 (HOLOTYPE: IND!; ISOTYPE: GH!). Acalypha rhomboidea Raf. var. deamii (Weath.) Weath., Rhodora 39: 16. 1937 2. Acalypha gracilens A. Gray, Manual 408. 1848. Type: UNITED STATES. Virginia. F.J.X. Rugel s.n. (LECTOTYPE—designated by G. A. Levin, Syst. Bot. 23:285. 1998[1999]: GH!; ISOLEC- TOTYPE: G). Acalypha virginica L. var. gracilens (A. Gray) Miill. Arg., Linnaea 34: 45. 1865 Acalypha = L. var. fraseri Miill. Arg., Linnaea 34: 45. 1865. D STATES. South Carolina. J. Fraser s.n. (HOLOTYPE: G- Dc, microfiche!). Acalypha gracilens A. Gray var. fraseri (Miill. Arg.) Weath., Rhodora 29: 202. 1927. whist earn A. Gray var. delzii Lill. W. Miller, Sida 3: 447. 1969. Bahia STATES. Arkansas: Hot Spring Co., Bismarck PO., Jul “i "1957 oD: Bonded 39432 (HOLOTYPE: SMU!; ISOTYPES: GH!, KANU!, OKL!). 3. Acalypha monococca (Engelm. ex A. Gray) Lill. W. Miller & Gandhi, Sida 13: 123. 1988. Acalypha gracilens A. Gray var. monococca Engelm. ex A. Gray, Man- 2, 390. 1856. TYPE: UNITED STATES. Missouri: St. Louis Co., limestone precipices on the banks of the Mississippi River ISOTYPE: WIS!). Acalypha gracilens A. Gray ssp. monococca (En- gelm. ex A. Gray) G. L. Webster, J. Arnold Arbor. 48: 373. 1967. 4. Acalypha rhomboidea Raf., New FI. 1: 45. 1836. Type: UNIT- ED STATES. South Carolina. C.S. Rafinesque s.n. (LECTOTYPE— designated by C. A. Weatherby, Rhodera 42: 96. 1940: G- DEL, photo at GH!). Acalypha virginica L. var. genuina Miill. g., Linnaea 34: 44. 1865. Acalypha virginica L. var. rhom- boidea (Raf.) Cooperr., Michigan Bot. 23: 165. 1984. 5. Acalypha virginica L., Sp. P1. 1003. 1753. Type (conserved): UNITED STATES. Virginia. J. Clayton 20] (HOLOTYPE: BM, drawing at GH!). Acalypha virginica L. var. intermedia Miill. Arg., Linnaea 34: 45. 1865. Acalypha digyneia Raf., Fl. Ludov. 112. 1817. Type: none located. ACKNOWLEDGMENTS. ‘This research was supported by a grant from the Campus Research Board of the University of Illinois, Urbana-Champaign. I thank the curators and staff of the following herbaria for allowing me to study the specimens in their care: A, ARIZ, ASU, BKL, BM, BRIT, BUT, CAN, CAS, CHRB, CLEMS, CM, DAO, Za2 Rhodora [Vol. 101 DAV, DES, DHL, DS, EIU, F, FARM, FLAS, FSU, FTG, GA, GH, GMUF, IA, ILLS, IND, ISC, KANU, KE, LL, LSU, LYN, MARY, MICH, MIN, MO, MT, MU, NCU, ND, NEB, NEBC, NMC, NY, NYS, OKL, OKLA, OS, PAUH, PENN, NM, Rick Phillippe, “hin Hill, Tom Mione, and two anonymous re- viewers provided helpful comments on earlier versions of the manuscript. LITERATURE CITED Baum, D. A. AND M. - DONOGHUE. 1995. Choosing among alternative “‘phy- logenetic sel concepts. Syst. Bot. 20: 560-573. CavANILLES, A. J. 1800. Descripcion del genero Bonplandia, y de otras plan- tas. Anales Hist. Nat. Madrid 2: 12-142. COoPERRIDER, T. S. 1984. 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Evolution in the Ac. alypha gracilens/monococca complex (Euphorbiaceae): Morphological analysis. Syst. Bot. 23: 269-— 288. ———. In press. Acalypha pr spe sng In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mex- ico, Vol. 12. Oxford Univ. Press, New York and Oxford. McVauau, R. 1961. ea amernan novae Novo-Galicianae. Brittonia 13: 145-205. , L. W. 1964. A taxonomic treatment of the species of Acalypha in the United States. Ph.D. dissertation, Purdue Uni niversity, West Lafayette, IN. . 1969. Acalypha gracilens uy var. delzii L. Miller, var. nov. Sida a : 1970. Acalypha, pp. 943-947. In: D. S. Correll and M. C. Johnston, 1999] Levin—Acalypha in North America 233 eds., Manual of the Vascular Plants of Texas. Texas Research Founda- tion, Renner, MOHLENBROCK, R. H. 1982. psieae Plants: Basswoods to Spurges. South- ois Univ. Press, Car ey Us. . 1986. Guide to the Veal: Flora of Illinois, revised and enlarged ed. Southern Illinois Univ. Press, Carbondale, IL. MULLER, J. 1865. Vorlaiifige Mitteilungen aus dem fiir De Candolle’s Prod- i i 26. ten Manu : . 1866. Euphorbiaceae, pp. 189-1286. In: A. De Candolle, ed., Prod- romus Systematis Naturalis Regni Vegetabilis, Vol. 15(2). Victor Mas- son, Paris. Nixon, K. C. AND Q. D. WHEELER. 1990. An amplification of the phylogenetic species concept. Cladistics 6: 211-223. OLmsTEAD, R. G. 1995. Species concepts and plesiomorphic species. Syst. Pax, F AND K. HOFFMANN. 1924. VE oe fea ha ce ee ae— Acalyphinae, Additamentum VII, 1-231. In: A. Engler, ed., Das Pflanzenreich, IV, 147. XVI (Heft 5). RADFORD, A. E., H. A. AHLES, AND C. R. BELL. 1968. Manual of the Vascular Flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill, NC. REVEAL, J. L., C. E. JARvis, AND FR. Barrie. 1990. Proposal to conserve the name and type of Acalypha virginica L. (Euphorbiaceae). Taxon 39: 361-363. TEYERMARK, J. A. 1963. Flora of Missouri. Iowa State Univ. Press, Ames, IA. baa weer C. A. 1927. The group of Acalypha virginica in eastern North ca. Rhodora 29: 193-204. : 1937, The typification of Acalypha virginica L. Rhodora 39: 14-16. - 1940. Rafinesque types in the group of Acalypha virginica. Rhodora 2:96. RHODORA, Vol. 101, No. 907, pp. 234-263, 1999 THE F TER SPECIES OF VAUCHERIA (TRIBOPHYCEAE, CHRYSOPHYTA) FROM CTICUT CRAIG W. SCHNEIDER, CHRISTOPHER E. LANE, AND ANNA NORLAND! Department of Biology, Trinity College, Hartford, CT 06106-3100 ABSTRACT. Having scoured freshwater habitats throughout Connecticut for more than two years at 151 collection sites, we have discovered the presence of nine species and two varieties of the genus Vaucheria, including four species not Fe known from the state: V. prona, V. taylorii, V. uncin- ata, and ndulata. In addition, V. compacta, normally found in local es- tuaries, is fara in freshwater here, and is distinguished from its local freshwater variety, V. compacta var. dulcis. Culture conditions in the labo- ratory stimulated production of gametangia, allowing for identification of mostly vegetative a tic ca samples. A key to the species and varieties is provide Key Words: Vaucheria, Vaucheriaceae, Tribophyceae, Connecticut Vaucheria is a yellow-green alga commonly encountered along freshwater stream and river banks, in mud surrounding small ponds and marshes, in drainage ditches alongside roadways, and in puddles and vernal pools worldwide. In habitats such as these, coenocytic siphons of Vaucheria act as ecologically important substratum stabilizers, often during continual environmental dis- ruption. Although most species in the genus live in freshwater, several species are found in brackish water, and a few are com- pletely restricted to marine environments (Blum 1972). Vaucheria species have been shown to be euryhaline, often tolerating great variation in salinity in very short periods of time (Schneider et al. 1993). The first recorded collections of freshwater Vaucheria from Connecticut were included in the early and important North merican exsiccata, Phycotheca Boreali-Americana (P.B.-A., Collins et al. 1895, 1898, 1905). This exsiccata included three freshwater Vaucheria specimens from the state: V. terrestris Lyngb. [= V. frigida (Roth) C. Agardh], Fasc. II (1895), no. 78a ' Current address: Harvard University Graduate School of Education, Ap- pian Way, Cambridge, MA 02138. 234 1999] Schneider et al—Vaucheria from Connecticut 235 (Setchell, 9.x.1892, Norwich); V. aversa Hassall, Fasc. X (1898), no. 475 (Holden, 3.xiii.1892, Mill R.., Fairfield); and V. geminata (Vaucher) DC with some V. geminata var. racemosa (Vaucher) alz [= V. racemosa (Vaucher) DC], Fasc. XXVI (1905), no. 1287, (Holden, 9.iv.1892, Ash Creek, Bridgeport). In his mono- graphic treatment of freshwater and marine “green” algae from North America, Collins (1909) included only five freshwater spe- cies of Vaucheria for all of North America, reflecting the limited knowledge of the genus at that time. Of these, only V. aversa and V. sessilis (Vaucher) DC [= V. bursata (O. FE Miill.) C. Agardh] were reported from Connecticut. Conn and Webster (1908) pub- lished the first report on freshwater algae specifically from the state, and recorded only one species, V. bursata (as V. sessilis). Hylander (1925, 1928) included five freshwater species of Vauch- eria in his “Algae of Connecticut,”’ based on limited reports from throughout the state. These included V. aversa from Fairfield; V. bursata (as V. sessilis) from Brookfield and Middletown; V. fri- gida (as V. terrestris) from Litchfield, Hartford, New Haven, and New London counties; V. geminata from Bridgeport, Barkham- sted, and New Haven; and V. racemosa (as V. geminata vat. ra- cemosa) from New Haven (Hylander, 1925). Each of these spe- cies was then recorded in the first monograph of Vaucheria for North America by Helen Brown (1929). John Blum’s (1972) later seminal monograph on the ‘‘Family Vaucheriaceae,” published as part of the New York Botanical Garden’s ‘‘North American Flora,” included 22 freshwater and 20 marine species, but none with specific reference to the state. Since 1928, only one species has been added to the freshwater Vaucheria flora of Connecticut, when V. compacta var. dulcis J. Simons was found in abundant populations near the junction of the Farmington and Connecticut Rivers (Schneider et al. 1996). Thus, in the more than a century since the first reports of Vaucheria from Connecticut in P.B.-A., only six freshwater spe- cies have been reported. During this time, collections were made from only a few localities, mostly in the western part of the state, and only a few sites were visited throughout the year. Considering that fourteen freshwater species have been reported for New Eng- land, New York, and New Jersey (Blum 1972), not including one presently known only from Connecticut within that range (V. compacta var. dulcis), the timing seemed right to conduct a basic field and laboratory study to assess the diversity of freshwater 236 Rhodora [Vol. 101 Vaucheria in the state, and to broadly survey the local species distributions. Only four marine/brackish species of Vaucheria from Connecticut were recorded prior to a similar coastal study of the genus, and five species and one variety were subsequently added to the local flora after extensive field and culture work (Schneider et al. 1993). MATERIALS AND METHODS Since the initiation of this study in 1994, we have visited 151 riparian and limnic habitats, muddy wetlands, drainage ditches, and roadside catchments, often revisiting the same sites in dif- ferent seasons. Crude samples were collected either as siphonous algal mats attached to substratum, or mud itself with no obvious signs of Vaucheria. Using a flat spatula, substrata were cut into approximate 100 cm? quadrats, varying in thickness from 2—3 cm. At the site, after temperature was recorded for proximal standing water, pH (Corning pH 103 meter) and salinity were measured (Reichert-Jung refractometer 11719-2). Field evaluation of the substratum type (sand, mud, etc.) was made. Codes identifying the collecting sites are given in the Appendix (bold face), and numbers at the end of codes under ‘“‘REPRESENTATIVE COLLEC- TIONS” represent the first, second, third, etc. visit/collection in the same location. Collection data are followed by the first date of gametangia appearance in culture, or noted if they were fertile in the field collection. The crude samples were collected in labeled, zippered, plastic bags and transported on ice to the laboratory where they were immediately subdivided into three labeled culture dishes. Vas- cular plants, sticks, and leaves were removed from the surface. Bold’s basal medium (Bischoff and Bold 1963) was added to each culture dish to a depth of approximately one-third the thickness of the crude sample. Samples were distributed among three pho- toperiods (8L:16D, 12L:12D, 16L:8D) in Hotpack incubators (#352642) set at 15°C, an optimum temperature for the growth of Vaucheria (Schneider et al. 1993). Microscopical observations on each crude culture were made every five to ten days. Drawings were made with the aid of a Zeiss camera lucida, and microscope slide vouchers (20% or 40% Karo® corn syrup, 1% aniline blue in IN HCI in a ratio of 20:1:1) are deposited in Herbarium/C. W. Schneider at Trinity College. The P.B.-A. set utilized in the pre- 1999] Schneider et al.—Vaucheria from Connecticut 237 sent study was originally purchased by Wellesley College (Fahey and Doty 1955) and is presently located in the Herbarium/C. W. Schneider. Standard forms of author names follow Brummitt aiid Powell (1992). Because other works (e.g., Blum 1972; Entwisle 1988a) include world distributions for the species found in Con- necticut and are often not specific for a particular place, citing ‘widespread in North America,”’ we have listed states and prov- inces in the northeastern part of North America with known re- ports under the designation “‘NORTHEAST DISTRIBUTION.” RESULTS AND DISCUSSION Few samples collected were fertile in the field. The remainder of the collections with obvious Vaucheria siphons later produced gametangia after 10-14 days in culture, on the average. About 30% of the mud samples never produced Vaucheria after incu- bation in culture media. In others without obvious Vaucheria, either siphons existed within the mud below grade, or dormant zygotes or asexual spores were coaxed into germinating under favorable culture/incubator conditions. Many cultures produced two to six biotic sympatric species over time in a single crude sample. Most species discovered in this study presented unre- markable features from those reported for other North American populations, so the reader is referred to Blum (1972) for full descriptions including pertinent measurements. Where our obser- vations have been at variance with Blum (1972), these are noted below. We have discovered great variability in our extensive col- lections, allowing for a greater understanding of the species and overlap among them. Thus, when working with the polymorphic Vaucheria, it is advisable to sample widely for species and to rely on measurements and morphological characteristics of many in- dividuals before making taxonomic determinations. Although we have sampled extensively throughout the state, we have the fewest Vaucheria species records (four) from the northwestern Litchfield Co. This is in part due to more limited sampling there (14 sites), but mostly because of the poor, often sandy, substrata found in this part of the state. New records for Connecticut are noted with asterisks (*). 238 Rhodora [Vol. 101 VAUCHERIACEAE Dumortier 1822, p. 71 Vaucheria aversa Hassal 1843, p. 429. Figures 1-3, 46. TYPE LOCALITY: vic. Chestnut, England. NORTHEAST DISTRIBUTION: Maine, N.H., Mass., Conn., N.Y., N.J. REPRESENTATIVE COLLECTIONS: Fairfield Co.—KP1, CEL, coll. 25.x.1998, gam. 1.xi.1998; NR2, CEL, coll. 28.xi.1997, gam. 7.ii.1998; PB1, CEL, coll. 28.xi.1997, gam. 6.i1.1998; PN1, CEL, coll. 31.v.1998, gam. 6.vi.1998; SNR1, CEL, coll. 25.x.1998, gam. 1.xi.1998. Hartford Co.—BBB2, CWS, coll. 10.xii.1997, gam. 6.i.1998; BRWB, CWS, coll. 12.xii.1997, gam. 23.v.1998; c fe z gam. 3.iii.1998; GMS1, CEL/CWS, coll. 5.vi.1998, gam. 14.vi.1998; RDD1, AN, coll. 22.ii.1998, gam. 22.iv.1998; JB1, AN, coll. 22.ii.1998, gam. 28.v.1998; LJB1, AN, coll. 22.ii.1998, gam. 28.v.1998; PGB1, CEL, coll. 10.x.1997, gam. 22.iv.1998; SBBB1, CEL, coll. 21.x.1997, gam. 29.xi.1997; SCR1, CEL/CWS, coll. 5.vi.1998, gam. 31.ix.1998. Middlesex Co.—SR2, CWS, coll. 28.ix.1997, gam. 23.xii.1997. New Haven Co.— DLF1, AN/CEL, coll. 15.xi.1997, gam. 13.1.1998; EMR1, AN/CEL, coll. 3.xii.1997, gam. 3.iii.1998. New London Co.—ARI1, CEL, coll. 23.v.1998, gam. 7.vii.1998; BBS1, CWS, coll. 29.xi.1997, gam. 3.ii.1998; LBM1, CWS, coll. 29.xi.1997, gam. 13.i1.1998: LEL1, CWS, coll. 29.xi.1997, gam. 10.11.1998; ST1, CEL, coll. 23.v.1998, gam. in field. Tolland Co. gam. 10.x.1998; SKR1, CEL, coll. 11.xi.1997, gam. 12.i1.1998. Windham Co.—NK1, AN/CWS, coll. 18.ix.1997, gam. 21.v.1998: NK7, MED/CWS, coll. 29.ix.1998, gam. 22.x.1998: QR1, CEL, coll. 20.viii.1998, gam. 31.1x.1998. throughout Connecticut (Figure 46) in river beds, drainage ditch- es, and woodland trails—this species appeared in 20% of our crude cultures. Often, samples containing this species did not ap- pear as fertile siphons until two or more months in culture. In a few instances, after being in culture for up to 3 months, V. aversa became the dominant species in the crude samples, excluding others. Vaucheria aversa is one of two species in Connecticut, along with V. bursata, with sessile gametangia. These two can be easily distinguished by the angle and appearance of oogonia and by antheridial morphology. Oogonia of V. bursata have completely- 1999] Schneider et al.—Vaucheria from Connecticut 239 filled cavities at maturity and have beaks that are directed at 70— 80° angles from the siphons towards the adjacent long, curled, cylindrical antheridia (Figures 4-8). The oogonia of V. aversa have peripheral cavities when mature and arise at 20—30° angles, with their beaks deflected downwards to the siphons in the vicin- ity of the adjacent short, straight, fusiform to cylindrical anther- idia (Figures 1-3). Neither of these two could be confused with any other freshwater Connecticut species. Vaucheria bursata (O. F. Miill.) C. Agardh 1811, p. 21. Figures 4-8, 47. BASIONYM: Conferva bursata O. FE Miill. 1788, p. 96. TYPE LOCALITY: Geneva, Switzerland. NORTHEAST DISTRIBUTION: Maine, Mass., R.I., Vt., Conn., N.J. REPRESENTATIVE COLLECTIONS: Fairfield Co.—CD1, CEL, coll. 31.v.1998, gam. 28.vi.1998; CPG1, CEL, coll. 31.v.1998, gam. 6.vi.1998; KP1, CEL, coll. 25.x.1998, gam. 28.x.1998; NB1, CEL, coll. 28.xi.1997, gam. 12.i.1998; NCMI1, CEL, coll. 25.x.1998, gam. in field; MB1, CEL, coll. 28.xi.1997, gam. 3.iii.1998; PN1, CEL, coll. 31.v.1998, gam. 6.vi.1998; WMR1, CEL, CWS/CEL, coll. 5.vi.1998, gam. in field; FTS1, CWS, coll. 21.ix.1994, gam. 24.iv.1996; GSB1, CWS, coll. 12.xii.1997, gam. 3.ii.1998; NH1, CEL, coll. 8.x.1997, gam. 23.x.1997; PGB1, CEL, coll. 10.x.1997, gam. 17.x.1997; RGB1, CEL/CWS, coll. 5.vi.1998, gam. 14.vi.1998; RP2, AN, coll. 13.xi.1997, gam. 29.xi.1997; TMR1, AN/CEL, coll. 1.x.1997, gam. 14.x.1997; WBB1, CWS, coll. 10.xii.1997, gam. 25.v.1998; WHR1, AN, coll. 13.xi.1997, gam. 2.xii.1997; WPF2, AN, coll. 7.x.1997, gam. 31.x.1997. Litchfield Co.—BBR1, CEL, coll. 30.vii.1998, gam. 3.ix.1998; BNB1, CEL, coll. 1.ix.1998, gam. 29.ix.1998; EA1, CEL, coll. 1.ix.1998, gam. in field; OR2, CEL, coll. 30.vii.1998, gam. 21.ix.1998; KFB1, CEL, coll. 30.vii.1998, gam. 12.xi.1998; NCWRI1, CEL, coll. 30.vii.1998, gam 23.ix.1998. New London Co.—LBMI1, CWS, coll. 29.xi.1997, gam. 24.vi.1998; ST1, CEL, coll. 23.v.1998, gam. in field; TR2, AN/CEL, coll. 22.x.1997, gam. 1.xi.1997. Tolland Co.—SKR2, CEL, coll. 23.v.1998, gam. 9.ix.1998. REMARKS. Vaucheria bursata was first reported in Connecticut as V. sessilis (Vaucher) DC (Lamarck and de Candolle 1805) by Hylander (1928) and Brown (1929). Vaucheria sessilis was shown by Christensen (1973) to be a junior synonym of V. bur- sata, based upon Miiller’s (1788) illustrations of Conferva bur- sata, as well as collections Christensen made at Bad Meinberg, southwest of Hanover, Germany. Blum (1972) maintained V. ses- 240 Rhodora [Vol. 101 silis and two similar species, V. repens Hassall and V. clavata sensu Klebs, differentiating each by siphon diameter and number of oogonia in association with each antheridium. Based upon our collections of V. bursata from a variety of habitats in Connecticut, we find these distinctions untenable, and follow Entwisle (1987, 1988a) in considering these as heterotypic synonyms of V. bur- sata. For example, we have found individual siphons with one and two oogonia associated with an antheridium. Our collections of this cosmopolitan freshwater species come from all but two counties in the state, Middlesex and Windham (Figure 47). Vaucheria bursata appeared in 20% of our crudes cultures, thus being one of the most commonly encountered species in the state. Vaucheria compacta vat. compacta (Collins) Collins ex W. R. Taylor 1937, p. 226. Figures 9-11, 48. BasIONYM: Vaucheria piloboloides var. compacta Collins 1900, pid. TYPE LOCALITY: Malden Mass., United States. NORTHEAST DISTRIBUTION (freshwater only): Conn. COLLECTIONS: Fairfield Co.—CD1, CEL, coll. 31.v.1998, gam. 6.vi.1998; BPG1, CEL, coll. 31.v.1998, gam. 6.vi.1998. V. undulata and V. bursata, two other freshwater species, were ound. These are the first low-salinity collections from the north- east other than the records of its freshwater variety, V. compacta var. dulcis (see below). Vaucheria compacta is dioecious. After a month in freshwater 1999} Schneider et al.— Vaucheria from Connecticut 241 culture, both samples produced male gametangia only in the 16L/ 8D photoperiod regime, with the samples in 8L/16D and 12L/ 12D remaining vegetative. The antheridia were mostly of the typ- ical “brackish-type,”” with two or more lateral discharge pores (Figures 9, 10), not to be confused with those produced by V. compacta var. dulcis that have one or no lateral discharge pores (Figures 12, 13) (Schneider et al. 1996; Simons 1974). In both Greenwich samples, the siphons produced male gametangia for less than two weeks. Thereafter the cultures became overgrown by other freshwater algal species including Vaucheria. Vaucheria compacta var. dulcis J. Simons 1974, p. 624. Figures 12, 13, 49. TYPE LOCALITY: The Netherlands. NORTHEAST DISTRIBUTION: Conn., Delaware. REPRESENTATIVE COLLECTIONS: Hartford Co.—RPC, CWS, coll. 8.x.1992, gam. 21.x.1992; FW, CWS, coll. 8.x.1992, gam. 26.x.1992; FP2, CEL/CWS, coll. 5.vi.1998, gam. 24.vi.1998. REMARKS. Despite our extensive collections of Vaucheria in the state during all seasons over two years, we have not located this freshwater variety of V. compacta in locations outside those in the original report for North America of dense year-round pop- ulations in the Connecticut and Farmington Rivers below the Rainbow Dam (Figure 49; Schneider et al. 1996). These popu- lations remain present and abundant only in areas of the Far- mington River where they receive daily “‘hydrotidal”’ flooding from the twice-daily hydroelectrical generation produced by the dam, as well as natural tidal effects from Long Island Sound in the Connecticut River. Schneider et al. (1996) speculated that the variety’s exclusive distribution in Connecticut would be only to areas receiving daily freshwater tidal flushing and this has been borne out by a lack of collections from sites other than those in the original report. We cannot speculate why this variety hasn’t found its way to the lower Housatonic and Thames River systems, as both are affected by tidal waters. In the Farmington River, V. compacta var. dulcis thrives as thick continuous mat-like bands, especially along the banks with the greatest flow of water, and unlike other Vaucheria species in the state, it seems to grow to the exclusion of other species (Schneider et al. 1996) Despite years of culture work and field collections, we have 242 Rhodora [Vol. 101 yet to find or produce oogonia on plants from these sites, though antheridia have been abundant (Schneider et al. 1996). Neverthe- less, differences in the antheridial morphology allowed us to dif- ferentiate this European variety from the nominate species, Vaucheria compacta. The freshwater variety maintains its dis- tinctive antheridial characteristics despite drastic salinity changes in vitro (Schneider et al. 1996). Vaucheria frigida (Roth) C. Agardh 1824, p. 173. Figures 14-19, 50. BASIONYM: Conferva frigida Roth 1797, p. 166. TYPE LOCALITY: London, England. NORTHEAST DISTRIBUTION: N.B., Maine, Mass., R.I., Conn., N.Y. REPRESENTATIVE COLLECTIONS: Fairfield Co.—CPG1, CEL, coll. 31.v.1998 gam. 12.xi.1998; MBI, CEL, coll. 28.xi.1997, gam. 3.ii.1998: NR2, CEL coll. 28.xi.1997, gam. 6.xii.1997; PB1, CEL, coll. 28.xi.1997, gam. 6.1.1998. Hartford Co.—BBB2, CWS, coll. 10.xii.1997, gam. 6.1.1998; BUR1, CWS/ CEL, coll. 5.vi.1998, gam. 2.vii.1998; FS1, CWS, coll. 5.1x.1995, gam. 10.x.1995; JB1, AN, coll. 22.11.1998, gam. 28.v.1998: RGB2, CEL/CWS, coll. 5.vi.1998, gam. 17.vii.1998; WP2, AN, coll. 7.x.1997, gam. 29.xi.1997. New London Co.—SDD1, CWS, coll. 29.xi.1997, gam. 3.1.1998. Tolland Co.— RRB1, CEL, coll. 6.xi.1997, gam. 25.v.1998. Windham Co.—LR1, CEL, coll. 20.viii.1998, gam. 12.xi.1998; NK3, AN/CWS, coll. 18.ix.1997, gam. 29.xi.1997. ? '. REMARKS. For much of its taxonomic history, Vaucheria frigida was confused with another species originally described from Eu- rope, V. terrestris (Vaucher) DC (Christensen 1968). Early North American collectors apparently followed the account of V. ter- restris by Gotz (1897), a taxon now considered distinct from true V. terrestris (Blum 1972; Christensen 1968). Thus, early collec- tions of V. frigida from Connecticut were reported as V. terrestris (Blum 1953; Brown 1929; Hylander 1925, 1928), as well as dis- tributed in the P.B.-A. exsiccata (Collins et al. 1895, no. 78a). Vaucheria frigida occurs throughout North America, as well as being cosmopolitan elsewhere, while V. terrestris is presently rec- ognized only from Europe (Blum 1972). We have collected V. frigida in all seven Connecticut counties, and it appeared in 13% of our cultures (Figure 50). In Vaucheria frigida, thick-walled oogonia are borne singly on fruiting branches distal to single circinate antheridia. Long axes of oogonia are projected either horizontally or directed back at 1999] Schneider et al.—Vaucheria from Connecticut 243 Figures 1-19. Connecticut freshwater Vaucheria species. 1-3. V. aversa; 4-8. V. bursata; 9-11. Antheridia of V. compacta var. compacta; 12-13. Antheridia of V. compacta var. dulcis; 14-19. V. frigida. All scale bars = 100 pm. 244 Rhodora [Vol. 101 the supporting siphons bearing fruiting branches (pedicels). The species can be confused with single oogonial forms of V. undu- lata, but there are obvious differences between the two, not the least of which are the spiraled siphons of the latter (see below). Blum (1972) found considerably larger oogonia in V. frigida (80-— 135 X 106-165 wm) than in V. undulata (71-113 X 96-127 um). Our collections as well yielded larger oogonia for V. frigida (81— 110 X 93-140 pm) than for V. undulata (50-80 * 60-100 pm), but these had somewhat smaller dimensions than those listed by Blum (1972). While V. undulata often bears reproductive branch- es with a single oogonium, rarely are there vegetative siphons without reproductive branches bearing two oogonia. As siphons of V. undulata mature and bear more gametangia, the pedicels usually proliferate from one another, a phenomenon only ob- served once in eighteen fertile collections of V. frigida, and not nearly to the extent of that seen in V. undulata (see below). Vaucheria geminata (Vaucher) DC in Lamarck et DC 1805, p. 62. Figures 20-22, 51. BaSIONYM: Ectosperma geminata Vaucher 1803, p29. TYPE LOCALITY: Between Geneva and Versoix, Switzerland. NORTHEAST DISTRIBUTION: Maine, N.H., Vt., Mass., Conn., N.J. REPRESENTATIVE COLLECTIONS: Fairfield Co.—BSP1, CEL, coll. 31.v.1998, gam. 6.vi.1998; BUR1, CEL/CWS, coll. 5.v.1998, gam. 28.vi.1998; NB1, CEL, coll. 28.xi.1997, gam. 25.v.1998; PB1, CEL, coll. 28.xi.1997, gam. 22.iv.1998. Hartford Co.—EGST, CWS, coll. 12.xii.1997, gam. 3.ii.1998; GMS1, CEL/CWS, coll. 5.vi.1998, gam. 14.vi.1998; LJB1, AN, coll. 22.ii1.1998, gam. 14.iv.1998; LV1, CEL, coll. 21.x.1997, gam. 29.xi.1997; NBBB1, CWS, coll. 8.xi.1997, gam. 3.i.1998: NH1, CEL, coll. 8.x.1997, gam. 23.x.1997; QP1, AN/CEL, coll. 1.x.1997, gam. 22.iv.1998; SCR1, CEL/CWS, coll. 5.vi.1998, gam. 14.vi.1998; WHR1, AN, coll. 13.xi.1997, gam. 2.xii.1997. Litchfield Co—HOR2, CEL, coll. 30.vii.1998, gam. 20.xi.1998. Middlesex Co.—SR2, CWS, coll. 28.ix.1997, gam. 11.x.1997. New London Co.—CCRI, CEL, coll. 23.v.1998, gam. 6.vi.1998; LBM1, CWS, coll. 29.ix.1997, gam. 16.ii.1998; SDD1, CWS, coll. 29.ix.1997, gam. 22.iv.1998; ST1, CEL, coll. 23.v.1998, gam. 14.vi.1998. Windham Co.—NK3, AN/CWS, coll. 18.ix.1997, gam. 15.x.1997. REMARKS. Collins et al. (1905) first reported Vaucheria geminata from Connecticut, followed by the reports of Hylander (1925, 1928), and in this study it was collected in 18% of our samples from throughout the state (Figure 51). Vaucheria geminata almost invariably produces two opposite 1999} Schneider et al.—Vaucheria from Connecticut 245 oogonia on short to slightly extended pegs off erect reproductive branches at abaxial angles of 50—70° (Figures 21, 22). In a few instances, we found reproductive branch proliferation (Figure 20), a phenomenon also noted by Christensen (1969; fig. 9837a, i) for V. geminata. The oogonia are ovoid to slightly reniform in shape (Figures 20-22) and range from 60-78 X 62-90 um. Circinate antheridia are borne singly and terminally in between and distal to the oogonia, but mostly not projecting beyond the height of the oogonia (Figures 20-22). Fertile V. geminata appear very similar to forms of V. taylorii that bear two oogonia on repro- ductive branches (see below), with the distinguishing characters of not-overly-swollen pedicels (often distinctly swollen in V. tay- lorii) and the height of antheridia (V. taylorii mostly has its an- theridia exceeding the tops of the oogonia on pedicels). Oogonia are often distinctly flattened on adaxial surfaces in V. taylorii (see below), a characteristic not seen in V. geminata. As discussed later in this report, the Connecticut specimens of V. geminata and its var. racemosa distributed in P.B.-A. (XXVI:1287), are now seen to be a polymorphic collection of V. taylorii. *Vaucheria prona T. A. Chr. 1970, p. 250. Figures 23-27, 52. TYPE LOCALITY: Kongelunden, Amager, Denmark. NORTHEAST DISTRIBUTION: Mass., Conn., REPRESENTATIVE COLLECTIONS: Fairfield Co.—BSP1, CEL, coll. 31.v.1998, gam. 6.vi.1998; NR2, CEL, coll. 28.ix.1997, gam. 6.xii.1997; PB1, CEL, coll. 28.ix.1997, gam. 22.iv.1998; SNR1, CEL, coll. 25.x.1998, pam. in field. Hart- aZai. 1998, gam. 3.xi.1998; GRDD1, AN, ‘coll. 224 ii. 1998, gam. 22.4 iv. ‘1998: coll. 22.ii.1998, gam. 14.iv.1998. New London Co.—BBS1, CWS, coll. 29.xi.1997, gam. 3.11.1998; RRD1, AN/CEL, coll. 22.x.1997, gam ey Ses AN/CEL, coll. 22.x.1997, gam. 28.x.1997. Tolland Co.— SKR2, , coll. 23.v.1998, gam. 28.vi.1998. Windham Co.—NK3, AN/ CEL, coll. > th ix.1997, gam. 29.xi.1997. REMARKS. Previously encompassed under Vaucheria terrestris (Vaucher) DC (=V. frigida), Christensen (1970) separated V. prona from it, as the former had only a single pendent oogonium on a fertile branch. He noted that his paired, pendent oogonial 246 Rhodora [Vol. 101 plants actually represented what was later called V. hamata sensu Gétz (1897). In order to avoid continued taxonomic confusion, V. prona was established as a new species to accomodate this paired oogonial alga rather than reintroducing new concepts for long-established taxa (Christensen 1970). This common alga was designated by early North American workers as V. hamata, and is now recognized from collections from throughout the world (Entwisle 1988a). Our isolates of V. prona represent the first re- cords of this species from Connecticut, and only the second from New England. This species was found in 16% of our samples, but not in Litchfield, New Haven, and Middlesex counties (Figure 52). Oogonia on Vaucheria prona, like those of V. frigida, are formed distally to single antheridia on fertile branches, and are pendent on downwardly curved pegs off curved fertile branches. In some cultures, gametangia were formed terminally on the si- phons themselves or on greatly extended fruiting branches. In V. prona there are usually two large oogonia (42-68 X 55-78 jum) found on each fruiting branch, and when fully mature have op- positional tips (Figures 24, 26). Occasionally, we have found fruiting branches proliferating from others, while retaining two oogonia on the basal fruiting branch. Such occurrences are not nearly as common as those in V. undulata, a species that normally retains only a single oogonium on the basal branch. At one site (HLM), we often found V. prona producing three or four oogonia per pedicel (Figure 23), and in rare instances, we have found siphons producing only a single oogonium on a fertile branch (Figure 25). The multiple oogonial form is easily distinguished from V. taylorii by its pendent oogonia, a phenonmenon also nicely illustrated in Australian populations of this species (En- twisle 1988a; figs 85, 90). However, the single oogonial form of V. prona is difficult to distinguish from the single forms of V. frigida and V. undulata. Careful observations of large populations ; as trademark spiraled siphons (Figure 39) and, at maturity, consistently produces pro- re one oogonium in an op- 1999] Schneider et al.—Vaucheria from Connecticut 247 posite pair would have formed on a reproductive branch (see below). Vaucheria prona can be distinguished from the other lo- cal species with opposite oogonia, V. geminata, as this species has erect oogonia on straight reproductive branches (Figures 20— 22; Entwisle 1988b). ee iota taylorii Blum 1971, p. 191. Figures 28-34, 53. TYPE LOCALITY: Beaver Is., Charlevoix Co., Mich., United a NORTHEAST DISTRIBUTION: Maine, Conn. COLLECTIONS: Fairfield Co—KP1, CEL, coll. 25.x.1998, gam. in field; NB1, CEL, coll. 28.xi.1997, gam. 12.i.1998. Hartford Co.— ir, CeEs CWS, coll. 5.v.1998, gam. 29.vii.1998; FTS1, CWS, coll. 21.ix.1994, gam. 5.x.1994; GSB2, CWS/CEL, coll. 5.vi.1998, gam. 11.ix.1998; NRS1, CWS/ CEL, coll. 5.x.1998, gam. 30.xi.1998; WPI, AN, coll. 7.x.1997, gam. 14.x.1997. Litchfield Co.—EA1, CEL, coll. 1.ix.1998, gam. 24.x.1998; HOR2, CEL, coll. 30.viii.1998, gam. 21.ix.1998; KFB1, CEL, coll. 30.viii.1998, gam. in field; NN1, CEL, coll. 1.ix.1998, gam. 21.ix.1998. New London Co.—ST1, CEL, coll. 23.v.1998, gam. 14.vi.1998 REMARKS. Although our recent collections represent the first report of this species from Connecticut, specimens of Vaucheria taylorii were actually distributed in P.B.-A. (Collins et al. 1905) as V. geminata (Ash Creek, Bridgeport, Conn., Holden 9.iv.1892, no. 287). Collins et al. (1905) noted that this dried specimen con- tained “mostly the type [of V. geminata], but with some var. racemosa,” this being the sole basis for Hylander’s (1928) report of V. geminata var. racemosa [=V. racemosa (Vaucher) DC], a species we have not found in our extensive collections here. We have examined this exsiccata and have established that specimen no. 1287 represents a collection of V. taylorii with 2-6 oogonia on fertile branches rather than the two species proposed in the exsiccata. Also, as will be shown below, the illustration of V. geminata in Hylander (1928; pl. XX, Fig. 8), could possibly rep- resent V. taylorii depending upon the other reproductive branches that may have been associated with it in its original population. The oogonia of V. taylorii are borne at erect angles on pegs from the fruiting branches curving inwardly, often with their tips point- ing towards terminal antheridia. Vaucheria racemosa can easily be distinguished by its downcurved, long pegs from fertile branches bearing pendent oogonia (Blum 1972). Vaucheria ra- cemosa is in fact more similar to multiple oogonial forms of V. [Vol. 101 Figures 20-34. Connecticut freshwater Vaucheria species. 20-22. V. geminata; 23-27. V. prona; 28-34. V. taylorii. All scale bars = 100 Lm. 1999] Schneider et al.—Vaucheria from Connecticut 249 prona, but that species has long fruiting branches while those in V. racemosa are distinctly smaller (Blum 1972) The antheridial pedicels of Vaucheria taylorii project at least to the same height as the tops of oogonia (Figures 28, 30), usually exceeding them and at times being significantly higher (Figures , 34), one of the characters used to distinguish it from V. gem- inata (Blum 1971). Vaucheria taylorii is further noted by its swollen pedicels (Blum 1971), yet this character is variable and between the various collections we and others have made (Figures 28-34). Still, unlike V. geminata, some of the re- productive branches of V. taylorii in a population will be signif- icantly swollen centrally where oogonial pegs are formed (Figures 28, 31, 32). Oogonia range from 61-91 X 64-92 ym, and show a trend from larger and distinctly flattened on adaxial surfaces when only two are formed on a reproductive branch (Figure 29), to smaller and ovate to reniform when present in whorls of three to six (Figures 28, 32). e only other species we have found with more than two oogonia per fertile branch, Vaucheria prona, is easily distin- guished by its markedly pendent oogonial pegs (Figure 23). Along with V. compacta, V. taylorii is one of the least prevalent freshwater species in Connecticut, appearing in only 8% of our collections (Figure 53). *Vaucheria uncinata Kiitz. 1856, p. 21. Figures 35-38, 54. TYPE LOCALITY: Freiburg, Germany. NORTHEAST DISTRIBUTION: Maine, N.H., Mass., Conn., N-Y. REPRESENTATIVE COLLECTIONS: Fairfield Co.—SNR1, CEL, coll. 25.x.1998, gam. in field; NB1, CEL, coll. 28.xi.1997, gam. 25.v.1998. Hartford Co.— BBB2, CWS, coll. 10.xii.1997, gam. 13.i.1998; BUR1, fe ae coll. 5.vi.1998, gam. 28.vi.1998; CM1, CEL, coll. 1.ix.1998, gam. 10.ix.1998; FS3, CWS/CEL, coll. 5.vi.1998, gam. 28.vi.1998; GMS2, "CWSICEL. oul: 5.vi.1998, gam. 28.vi.1998; GRDD1, AN, coll. 22.11.1998, gam. 28.v.1998 GSB2, CWS/CEL, coll. 5.vi.1998, gam. 28.vi.1998; RGB2, CWS/CEL, coll. 5.vi.1998, gam. 28.vi.1998; SCR1, CEL/CWS, coll. 5.vi.1998, gam. 14.vi.1998. Litchfield Co.—PTB1, CEL, coll. 1.ix.1998, gam. 1.x.1998. Mid- dlesex Co.—SR2, CWS, coll. 28.ix.1997, gam. 11.x.1997. New London Co.—AR1, CEL, coll. 23.v.1998, gam. 7.vii.1998; CCR1, CEL, coll. 23.v.1998, gam. 28.vi.1998; HVP1, CEL, coll. 23.v.1998, gam. 7.vii. 1998. LBM1, CWS, coll. 29.xi.1997, gam. 24.vi.1998; ST1, CEL, coll. 23.v.1998, gam. 17.vii.1998; TR1, AN/CEL, coll. 22.x.1997, gam. 28.x.1997. Tolland Co.-—CB1, CEL, coll. 11.ix.1997, gam. 3.i1.1998;; RRB, CEL, coll. 250 Rhodora [Vol. 101 6.ix.1997, gam. 22.iv.1998. Windham Co.—NK1, AN/CWS, coll. 18.ix.1997, gam. 30.ix.1997; NK7, MED/CWS, coll. 29.ix.1998, gam. in field. REMARKS. The complicated taxonomic and nomenclatural history of this species is summarized by Blum (1953). Our collections represent the first record of Vaucheria uncinata for Connecticut, although Blum (1972) reported its distribution throughout the continental United States. It is a somewhat common species in our collections, found in 15% of all sites sampled, occurring al- most exclusively in the Connecticut River Valley or to the east of the Connecticut River (Figure 54). We rarely found V. uncinata on mud saturated with water in the field and it only appeared in our cultures when the crude samples became somewhat dried. Vaucheria uncinata is easily distinguished from the other local species by its unusually large, transversely elongated, pendent oogonia (108-150 wm in greatest dimension) that often appear to be sessile on the siphons due to their terminal position on long, hooked, or gallows-shaped, reproductive branches (Figures 35— 38). In certain locations during the fall, we have found fertile V. uncinata growing as an extensive mat over mud in non-flowing intermittent streams. *Vaucheria undulata C. C. Jao 1936, p. 741. Figures 39—45, 55 TYPE LOCALITY: Fou-tu-Kuan, Szechwan, China. NORTHEAST DISTRIBUTION: Maine, Mass., Conn., RA. NY.) NJ. REPRESENTATIVE COLLECTIONS: Fairfield Co.—AC1, CEL, coll. 30.viii.1998, gam. 18.ix.1998; BSP1, CEL, coll. 31.v.1998, gam. 6.vi.1998: CD1, CEL, coll. 31.v.1998, gam. 14.vi.1998; CPG1, CEL, coll. 31.v.1998, gam. BRWB, CWS, coll. 12.xii.1997, gam. 23.v.1998; CP1, AN, coll. 22.11.1998, gam. 3.ix.1998; FMT1, CEL, coll. 25.x.1998, gam. 12.xi.1998: FWB1, CEL/ CWS, coll. 5.vi.1998, gam. 6.x.1998; GMS1, CEL/CWS, coll. 5.vi.1998, gam. 28.vi.1998; GRDD1, AN, coll. 22.11.1998, gam. 30.v.1998: MP1, CEL, coll. 8.x.1997, gam. 15.x.1997: RGB2, CEL/CWS, coll. 5.vi.1998, gam gam. in field; PTB1, CEL, coll. 1.ix.1998, gam. 23.ix.1998. Middlesex Co.— SR2, CWS, coll. 28.ix.1997, gam. 23.xii.1997. New Haven Co.—DL1, CEL/ 1999] Schneider et al.—Vaucheria from Connecticut 251 Figures 35-45. Connecticut freshwater eis al species. 35-38. V. un- cinata; 39—45. V. undulata. All scale bars = 100 p 252 Rhodora [Vol. 101 coll. 23.v.1998, gam. 17.vii.1998; LBM1, CWS, coll. 29.xi.1997, gam. 24.vi.1998; TAB1, CEL, coll. 23.v.1998, gam. 17.vii.1998; TR1, CEL/AN, RRB1, CEL, coll. 6.xi.1997, gam. 20.v.1998: TD1, CWS, coll. 5.x.1997, gam. 10.x.1997; US1, CWS, coll. 7.xii.1997, gam. 28.v.1998. Windham Co.—BK1, CEL, coll. 20.viii.1998, gam. 28.i.1999; LR1, CEL, coll. 20.viii.1998, gam. 12.xi.1998; NK1, AN/CWS, coll. 18.ix.1997, gam. 16.vi.1998; NK7, MED/ CWS, coll. 29.ix.1998, gam. in field; QR1, CEL, coll. 20.viii.1998, gam. 10.ix.1998. REMARKS. Since its description from Szechwan, Vaucheria un- dulata, with its trademark spiraled siphons, has been found in many New England locations (Blum 1972; Colt 1985), and here we add it to the flora of Connecticut. We have found this species to be widespread throughout the state (Figure 55), and it was the most prevalent species in our crude cultures (39%). In the past, V. undulata was undoubtedly misidentified as other species, col- lectors not recognizing it by the majority of siphons that develop spirally at the tips (Figure 39). However, if gametangia are found on a siphon lacking spiraled development, they could easily be confused with V. frigida or V. prona, due to similar gametangial morphology and position. All three species have pendent oogonia that are borne on fruiting branches distal to single circinate an- theridia, V. undulata and V. frigida mostly with one oogonium per fruiting branch, V. prona usually with two. Unlike the others, V. undulata oospores often turn light brown prior to release from the mother siphon. Occasionally, V. taylorii produces a few un- dulating (spiraled?) siphons in a population, but the reproductive branches of the two species are distinct. diameter axes were usually spiraled, while those of greater di- ameter were mostly undulate to subundulate. In our cultures, the filaments were found to be of a similar dimorphic range, with approximately 60% showing spiral development in both size clas- ses. Our small-diameter siphons were distinctly spiraled, while 1999] Schneider et al.—Vaucheria from Connecticut 253 the larger ones (mature?) were more swollen and appeared to be undulate. Furthermore, only the larger-sized siphons in our cul- tures produced gametangia. Oogonia in Connecticut populations anged from 50-80 x 60-100 um. The reproductive branches ans proliferated at maturity, a phenomenon seen only oc- casionally in other species such as V. frigida, V. prona, and V. geminata (Figure 20), but never to the extent of V. undulata. New es branches were produced at the site where an oppo- oogonium would have formed on the previous pedicel in the same system (Figures 40, 41, 43), often repeating this develop- ment sequentially (Figures 42, 44). In our crude cultures, V. un- dulata often grew to the exclusion of other Vaucheria species, the latter only appearing after the bloom of V. undulata had passed (5—6 weeks). KEY TO THE FRESHWATER SPECIES OF VAUCHERIA IN CONNECTICUT 1. Antheridia and oogonia on separate plants (dioecious) .... 2 1. Antheridia and oogonia on the same plants (monoecious) ... 3 2. Antheridia with 1 terminal and mostly 2 lateral discharge pores (Figures 9-11) .... V. compacta var. compacta 2. Antheridia with 1 terminal and 1 or 0 lateral discharge potes (Figures IZ. 13)". ...... V. compacta var. dulcis 3. Oogonia and antheridia sessile or short-stalked on siphons, not borne on special bisexual fruiting branches (pedicels) ... 4 3. Oogonia and antheridia borne on special bisexual fruiting 5 4. Oogonia formed singly or in pairs with long, circinate an- theridia between them; oospores completely filling oo- gonial cavities at maturity; oogonial beaks oriented to- ward the antheridia, either parallel with the siphons or erect at oblique angles (Figures 4-8) ..... V. bursata 4. Oogonia formed singly, never paired around a single an- theridium, usually with small, cylindrical antheridia on both sides; oospores with obvious distal and peripheral oogonial cavities at maturity; oogonia with deflected beaks directed towards the siphons (Figures 1-3) .... Lita SE GEO ae Fe PRR we Saeed ia ae ree V. aversa 5. Oogonia formed singly on fruiting branches ............. 6 5. Oogonia usually 2 or more on each fruiting branch ...... i 6. Oogonia subspherical to transversely elongate, lacking 254 Rhodora [Vol. 101 ~ ~l s© © aks, pendent on hooked or gallows-shaped pedicels Ciianies SSO inti ec ee t.. Peve ree V. uncinata 6. Oogonia subspherical to obovate and reniform with small distal beaks, borne terminally on curved pedicels (Fig- pies Pdi cis. oo See Sa es V. frigida Population rarely containing spiraled siphons; oogonia almost always 2 per fruiting branch (occasionally 1 or 3—4, but these oogonia pendent), borne laterally to terminal erect antheridia; if proliferating additional fruiting branches, usually 2 oogonia maintained on the basal fruiting branch (Figure 20) ed . Population often containing spiraled siphons; oogonia highly variable in number per fruiting branch; if proliferating ad- ditional fruiting branches, usually 1 oogonium maintained on the basal fruiting branch (Figures 40-42) ......... 8. Oogonia pendent on obvious downcurved pegs, occasion- ally numbering 3—4 (Figure 23); fruiting branches usu- ally curved (Figures 24-27) .............. V. prona 8. Oogonia erect on pegs and directed away from siphons, almost uniformly numbering 2; fruiting branches straight and erect (Figures 20-22) ..... V. geminata Siphons commonly spiralling, especially the thinner ones in a population (Figure 39); oogonia | or 2 per fruiting branch, with variation common on individual siphons; fruiting branches commonly proliferating in chain-like fashion from one another at maturity (Figures 40—44); fruiting branches not swollen at points of oogonial attachment at Soturity. (PTS Oren Se oe a me woe os V. undulata Siphons only occasionally spiralling; 2 to 6 oogonia per fruit- ing branch, with great variation common on individual siphons; fruiting branches distinctly swollen at points of oogonial attachment at maturity (Figures 28-34) ei oa poe! big a Sin es etamiereialiavags GAR a3) acaatwid ac V. taylorii ACKNOWLEDGMENTS. We thank Courtney Hadly, Megan Dun- phy, Megan Garretson, and Ginny Schneider for their assistance on this project both in the field and in the lab. We acknowledge a Trinity College student research grant from the Faculty Re- search Committee to CEL. We appreciate the constructive criti- cism of the manuscript provided by Dr. Arthur Mathieson (UNH). 255 Schneider et al.— Vaucheria from Connecticut V. aversa 1999] mea de co COAT weet eee aly gow ary 7 tS “a Los aa ae oe Oe P< dis te Sy Sy aa Saree — ae SPR FLAG SEN ek a) Se oN PRD een fats oe bee Thames R. * ’ ' 4 « 3 . a y \ , S ’ . (ge eceage Sa eg os -# ‘ Ane ee J dee te Connecticut R. ie ¥ Te ~ J Fat ba) 1+. *, =: bi ~ bf oy. q * mre Ae ey eg big oy 2 . ey s : aes Sem to : ; VX tie he mo * thal - i = : ae : i“ Re DEANS : FP’! ee es rk Lv . ‘ 46 V. bursata 47 Figures 46—47. Distribution maps of Connecticut freshwater Vaucheria species. Thames R. wae! ou SS ‘a fap ae ahs ot sae ot te 256 Rhodora [Vol. 101 V. compacta var. compacta Viet. es 7 ke 1a, uP PS fe ASG on rene “$s be ne ig gt er ay 4 7X i ts ae Are mile : r as fA aes c TY te ee - a Hy 49. Distribution maps of Connecticut freshwater Vaucheria Figures 48— pecies. 257 Schneider et al.—Vaucheria from Connecticut Vi 1999] ida sit =e "rh Lone. nh tre td pent Thames R. Connecticut R 50 ta ipl V. ea! Thames R. Distribution maps of Connecticut freshwater Vaucheria a4 5 ech o'r, = Pace) on = ee c 3) ie 1 eos a = phic Nv if agree Y bide eee 3 7 x = A eae aan xt Nees = re sek wie } 4 ed le oS ae ee By Eile cee ea ee “40 q ayo pbs 0 2425 * Housatonic R. Figures 50-51. species. [Vol. 101 Rhodora : ee > oN De gs eae YF vo nThn “A ae ay = ara aR, apes Tele ae ee ee aie elt 4 . - - sa ci o> ats IAD AX ot Oo 5 Thames R Distribution maps of Connecticut freshwater Vaucheria oy ‘ Thames R. 52 Connecticut R. oe hae ate a a ~ Housatonic R. Fie Laks 258 V. prona V. taylorii Figures 52-53. species. Schneider et al.—Vaucheria from Connecticut 259 1999] inata uncina er ss ra (‘AP coon ama | 7h ee war koe: Thames R. Connecticut R. ousatonic R 54 V. undulata tg Thames R. Connecticut R. 35 Distribution maps of Connecticut freshwater Vaucheria Figures 54-55. species. 260 Rhodora [Vol. 101 LITERATURE CITED AGARDH, C. A. 1811. Dispositio Algarum Sueciae. . . Part 2. Berling’s Print- ing Office, Lund. . 1824. Systema Algarum. Lund. BIsCHOFF, H. AND H. C. BoLp. 1963. Phycological studies. IV. Some soil algae from oe Rock and related algal species. Univ. Texas Publ., No. 6318 Buu, J. L. ioe i, racemose Vaucheriae with inclined or pendent oogonia. Bull. 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Salinity preference of twenty species of Vaucheria (Tribophy- ae). J. Mar. Biol. Assoc. U.K. 68: 531-545. Cou E S. 1900. eee lists of New England plants, —V. Marine ae. Rhodora 2: 41-5 9. The green ve of North America. Tufts Coll. Stud., Sci. Ser. Z: a 480. » I. HOLDEN, AND W. A. SETCHELL. 1895. Phycotheca Boreali-Ameri- cana ae Algae of North America. Fascicle II. No. 51— 100, Mal- den, MA —_, ——_, ———.. 1898. Phycotheca Boreali-Americana (Exsic- cata), Alene ae thon America. Fascicle X. No. 451-500, Malden, MA. ——_, - 1905. Phycotheca Boreali-Americana (Exsic- cata), ‘Alene of North Amecina Fascicle XXVI. No. 1251— 1300, Malden, Coirsi. ron Jr. 1985. Vaucheria undulata Jao again in New England. Rho- dora 87: 597-599. Conn, H. W. AND L. W. Wesster. 1908. The algae of the fresh waters of Connecticut. Connecticut State Geol. Nat. Hist. Surv. 10: 5-73. 1999] Schneider et al.—Vaucheria from Connecticut 261 DumortigER, B.-C. 1822. Commentationes botanicae. Observations bota- ques. Tournay. ENTWISLE, T. J. 1987. An evaluation of taxonomic characters in the subsection Sessiles, section Corniculatae, of Vaucheria (Vaucheriaceae, Chrysophy- ta). Phycologia 26: 297-321. 1988a. A monograph of — aes IN Ken al in southeastem mainland Australia. A Bot. 1: 1-77. . 1988b. An evaluation of seconde characteristics in the Vaucheria prona prong ni (Vaucheriaceae, eereek vite Phycologia 27: 183-200. FaHey, E. M. AND M. S. Dory. 1955. An alphabetical index to the Phycotheca plone Unpublished mimeograph, Woods Hole, MA Gotz, H. 1897. Zur systematik der gattung Vaucheria DC speciell der arten 134. HassaL, A. H. 1843. Descriptions of British freshwater confervae, mostly new, with observations on some of the genera. Ann. Mag. Nat. Hist. 11: 428-437. HyLanber, C. J. 1925. Vaucheria. pp. sage In: The algae of Connecticut. . dissertation, Yale Univ., New He : 192 28. The algae of Connecticut. psa State Geol. Nat. Hist. Surv. 42: 9-245. Jao, C.-C. 1936. Studies on the freshwater algae of China. II. Vaucheriaceae from Szechwan. Sinensia 7: 730-747. KUTzING, FE. T. 1856. Tabulae Phycologicae. .. Vol. 6. Nordhausen Lamarck, J.-B., P. A. DE CANDOLLE, AND A. P. DE CANDOLLE. 1805. Flora Francaise, troisiéme édition, Vol. 3. Paris. MULLER, O. F. 1788. Histoire de confervis palustribus oculo nudo invisibi- ibus. ose Acta Acad. Sci. Imp. Petrop. 3: 89-98. Pecora, R. A. 1977. Brackish water species of cin ag oe Vaucheriales) from Louisiana and Texa: oun eee 5-0. Rot, A. W. 1797. Catalecta Botanica. . ig LG. Ske. Leipzig. SCHNEIDER, C. W., L. A. MACDONALD, J. F. ae AND S. W. HEMINWAY. 1993. The marine and brackish water species of aowpeoy (Tribophy- ceae, et ceo from Connecticut. Rhodora 95: 97-11 : . Ritey, AND B. M. StTockTON. 1996. Stability - antheridial Cal eg in freshwater North American Vaucheria compacta var. dul- cis J. Simons (Tribophyceae, "neil grown under different salin- ities. Aquatic Bot. 52: 301-31 Simons, J. 1974. Vaucheria ee. A euryhaline estuarine algal species. Acta Bot. Neerl. 23: 613-626. TAYLor, W. R. 1937. Notes on North Atlantic marine algae. I. Pap. Michigan cad. Sci. 22: 225-233. VAUCHER, J. P. 1803. Histoire des Conferves d’Eau Douce. J. J. Paschoud, Geneva APPENDIX FRESHWATER COLLECTION SITE CODES. AB — Tolland Co.: Ash Brook, Coventry. AC — Fairfield Co.: Ash Creek, Bridgeport. Rhodora [Vol. 101 AR — New London Co.: Blissville Brook, Lisbon. BBB — Hartford Co.: Beamans Brook, Bloomfield. BBR — Litchfield Co.: Blackberry River, North Caanan. BBS — New London Co.: Big Brook, Salem. BK — Windham Co.: Blackwell Brook, Brooklyn. BNB — Litchfield Co.: Butternut Brook, Litchfield. BPG — Fairfield Co.: Bruce Park, Greenwich. BRWB — Hartford Co.: Tunxis Floodwater Retention Reservoir, Wash Brook, Bloomfield. CM — Hartford Co.: Copper Mine Brook, Bristol. CP — Hartford Co.: Covilli’s Pond, Canton CPG FW — Hartford Co.: Farmington River, Windsor. FWB — Hartford Co.: Freshwater Brook, Enfield. GMS — Hartford Co.: Mountain Brook, Great Marsh, Suffield. GRDD — Hartford Co.: Gracey Road drainage ditch, Canton. GSB — Hartford Co.: Salmon Brook, Granby. HB — New Haven Co.: Junction Route 5 and Interstate 691, Meriden. HLM — Hartford Co.: Hockanum River, Laurel Marsh, Manchester. HOR — Litchfield Co.: Housatonic River, Sharon. JB — Hartford Co.: Jim Brook, Canton. LBM — New London Co.: Latimer Brook, Montville. LEL — New London Co.: Latimer Brook, East Lyme. MP — Hartford Co.: Drainage catchment basin (pond), Interstate-84, Manchester. 1999] Schneider et al.—Vaucheria from Connecticut 263 NB — Fairfield Co.: North Brook, Newtow NBBB — Hartford Co.: North Branch Bini Brook, Burlington. NCM — Fairfield Co.: Mill Pond, New Caanan NCWR — Litchfield Co.: Witing River, North Caanan. NH — Hartford Co.: Drainage ditch, Route 30, South Windsor. NK — Windham Co.: Nipmuck Trail, Ashford. NN — Litchfield Co.: Nonnewaug River, Woodbury. NRS — Hartford Co.: Floodplain vic. Wells Pond, West Hartford. PB — Fairfield Co.: Patterson Brook, Easton. PBW — Hartford Co.: Phelps Brook, Windsor. PGB — Hartford Co.: Plum Gully Brook, South Windsor. RGB — Hartford Co.: Rocky Gutter Brook, deste RP — Hartford Co.: Reservoir pond, West Hart RPC — Hartford Co.: Riverside Park, Cones River Hartford. RRB — Tolland Co.: Valley Falls Park, Vi RRD — New London Co.: Drainage eh, ‘el Road, Norwich. SBBB — Hartford Co.: South Branch Bunnel Brook, Burlington. SCR — Hartford Co.: Scantic River, Enfield SDD — New London Co.: Route 11 drainage pond, junction Route 82, Salem SKR — Tolland Co.: Skungamaug River, Coventry. SNR — Fairfield Co.: South Norwalk Reservoir, Wilton. TMR — Hartford Co:: Lazy Lane Road, Southington TSB — New Haven Co.: Transylvania Brook, Southbury. US — Tolland Co.: Stream, Route 171, Union. WBB — Hartford Co.: Wash Brook, Bloomfield. WBSP — New Haven Co.: Wharton Brook State Park, outlet brook, Wallingford. WCP — Hartford Co.: Woodridge Circle Pond, Canton. WHR — Hartford Co.: West Hartford Reservoir, West Hartford. WMR — Fairfield Co.: Stream, Wire Mill Road, Stamford. WP — Hartford Co.: Wells Pond, West Hartford. WPF — Hartford Co.: Wells Pond feeder stream, West Hartford. RHODORA, Vol. 101, No. 907, pp. 264—273, 1999 DISTURBANCE AS A FACTOR IN THE DISTRIBUTION OF SUGAR MAPLE AND THE INVASION OF NORWAY MAPLE INTO A MODIFIED WOODLAND REBECCA ANDERSON! Tufts University, Biology Department, Medford, MA 02155 ABSTRACT. Disturbances have the potential to increase the success of bi- ological invasions. Norway maple (Acer platanoides), a common street tree & @ 5 - o 2) - a @ g ct. = By 5 5 E a 5 = = & n = a9 5 yd ® > ® ™ s 8 > : 3 — z. invasion of Norway maple and in the distribution of sugar maple. Disturbed areas on the path and nearby undisturbed areas were surveyed for both species along transects running perpendicular to a road. Norway maples were present in greater number closer to the road and on the path, while the number of sugar maples was not significantly associated with either the road or the path. These results suggest that human-caused disturbances have a role in facili- tating the establishment of an invasive species. Key Words: invasive plants, Norway maple, Acer platanoides, sugar maple, cer saccharum, disturbance Biological invasions happen when non-native species are in- troduced into new environments (Drake 1988). Although many of these species are absorbed into the community without influ- encing it very much (Begon et al. 1996), the detrimental effects of some invasives have been recognized as one of the major threats to biological diversity (Soule and Kohm 1989). These spe- cies may alter population dynamics, community structure (Elton 1958; Mooney and Drake 1986), and ecosystem structure and diversity (Vitousek 1990). ile natural disturbances often increase species richness and diversity (Hobbs and Hunneke 1992), they can also increase the likelihood of invasion. The potential for invasion is enhanced when a combination of disturbances is present (Hobbs 1991). For this study, disturbance refers to events that stress the community and influence resource availability and mortality rates. Examples include soil exposure, edge effects, nutrient addition (Hobbs 1991), and any event that removes plants, leaving a space open for colonization (Begon et al. 1996). ' Current address: Harvard Forest, P.O. Box 68, Petersham, MA 01366. 264 1999] Anderson—Invasion of Norway Maple 265 One common type of disturbance occurs when an edge of a community faces a more open area, such as a field, path, road, or body of water. An “edge effect’? occurs because the edges of a forest have a different microclimate than the forest interior. This results in a different plant species composition, including a higher number of exotics, and different community structure. In North Carolina, Fraver (1994) measured the percent cover of individual species and the relative cover of exotic species, and found edge effects extended 20-60 m into the interior of the forest. In ad- dition, exotics were found mainly on the edges and did not make up a significant portion of the forest interior. In Pennsylvania and Delaware, the edge effect significantly influenced light, temper- ature, litter moisture, humidity, and shrub cover up to 50 m from the edge (Matlack 1993). primary reason for the spread and 0 psotscg aca of invasive species is their introduction by humans as crops, ornamentals, or for forestry (Elton 1958). Norway maple (Acer as: bin) is an ornamental that has been described as an invasive exotic in urban forests in New Jersey (Webb and Kaunzinger 1993), Penn- sylvania (Kloeppel and Abrams 1995), and Great Britain (Nowak and Rowntree 1990). Norway maple is the widest ranging maple in its native European habitat. Introduced into the United States in 1756 by John Bartram of Philadelphia, its optimal range in the United States includes coastal northern New England, southern New England, and the Midwest (Nowak and Rowntree 1990). It may be able to transform the native woodland by outcompeting sugar maple (Acer saccharum Marshall; Webb and Kaunzinger 1993), which is native to the United States. Norway maple and sugar maple grow side by side as street trees and in urban forests and share many life-history character- istics. Both species produce seeds in the fall that require 2-3 months of stratification before spring germination (Hartmann et al. 1990) and both have been described as shade tolerant in their native ranges (Diekmann 1996; Sipe and Bazzaz 1995). Sugar maple grows best in small gaps (Runkle 1984). Although it grows faster in a gap than in the closed canopy (Canham 1988), sugar maple has a strong negative correlation with large gaps of 400m? (Runkle 1984). Norway maple has certain advantages that allow it to outcom- pete sugar maple and influence species composition when it in- vades urban woodlands. Since its introduction, Norway maple has 266 Rhodora [Vol. 101 become one of the most widely planted street trees due to its longevity, disease resistance, and ability to withstand poor soils and pollution. In urban areas, its wide tolerances and abundant seeds allow it to colonize woodlots and urban forests (Spongberg 1990). Norway maple also has a physiological advantage over sugar maple due to its higher rate of photosynthesis, longer re- tention of leaves in the fall, and faster sapling growth (Kloeppel and Abrams 1995). In addition, Norway maple can influence community structure. In a New Jersey sugar maple/beech/oak for- est that had been invaded by Norway maple, the understory spe- cies richness for each of the canopy trees was compared. Norway maple had a significantly lower species richness in its understory than the other tree species (Wyckoff and Webb 1996). se of this study was to determine if there is a cor- relation between disturbance and the establishment of Norway maple. STUDY SITE The Middlesex Fells Reservation in eastern Massachusetts was created in 1894. European colonists, arriving in the area in the 1600s, used the land primarily for farming but also for mills, firewood, cattle grazing, and mining (Levin and Mahlstedt 1990). Today the Fells is a mostly wooded, popular urban retreat. Only 10 km north of Boston, it is located in the towns of Winchester, Stoneham, Melrose, Malden, and Medford. Routes 23 and 93 run through the middle of the Reservation and divide it into eastern and western halves. Two-lane paved roads run along the perim- eter, with Norway maples planted as street trees in some areas. The trees range from 10-40 cm in diameter. No record could be found of exactly when they were planted, but all are large enough to produce seeds. The western side of the Fells, where this study was done, cov- ers about 400 ha. The landscape changes greatly over short dis- tances, due to high ridges with exposed ledges that run north to south with slopes running down to streams, ponds, and large man- made reservoirs (Drayton and Primack 1996). Foot, animal, and mountain bike traffic can be heavy on the extensive 3-5 m wide Carriage roads and the smaller footpaths. The first major carriage paths were likely due to the creation of the reservoirs from 1870— 1901. Later, in the 1930s, the Civilian Conservation Corps and 1999] Anderson—Invasion of Norway Maple 267 the Works Progress Administration also built many roads and paths (Levin and Mahlstedt 1990). MATERIALS AND METHODS Twenty-four transects were established perpendicular to the paved roads along the western side of the Middlesex Fells Reservation. At each of twelve sites two parallel transects were set up, one with a path as the transect line (disturbed), and another 20 m away in an area with no path (undisturbed). Along each transect, 10 X 10 m plots were established with the first plot at the road. On the transect that ran along the path, half of each plot was placed on either side of the path. The next two plots leading away from the road (numbers 2 and 3) had 50 m between them, with 100 m between remaining plots (Figure 1). Sampling was dis- continued when the plots had no Norway maples, and a visual inspection showed no more further on. In each plot, the number of Norway and sugar maples above 50 cm in height was counted. To determine the influence of the path, a Mann-Whitney test was performed on the number of Norway maples and sugar ma- ples on and off of the path. To determine the influence of the road, a Mann-Whitney test was performed comparing the number of Norway maples and sugar maples in plots 1 and 2. Plots 2 and 3 were also compared. Only the first three plots at each site, both on and off the path, were used in the analysis due to the lack of replication for plots 4—6. All statistical analysis was done on SPSS (SPSS, Version 6.1, Chicago, Illinois). RESULTS The path and distance from the road (using plot as a proxy for distance) correlated with the presence of Norway maples but not the presence of sugar maples. Results of the Mann-Whitney tests indicated there were significantly more Norway maples closer to the road and on the path (Figure 2; path: df=65, Z=—2.538, P=0.011; plot 1—plot 2: df=49, Z=—3.628, P<0.001; plot 2—plot 3: df=41, Z=—1.345, P=0.179). Results of the Mann-Whitney tests indicated that the number of sugar maples was not affected by the road or path (Figure 3; path: df=65, Z=—0.367, P=0.713; plot 1—plot 2: df=49, Z=—0.504, P=0.614; plot 2—plot 3: df=41, Z=—0.699, P=0.484). 268 Rhodora [Vol. 101 path Figure 1. Representation of placement of transects and plots at the Mid- dlesex Fells Reservation. DISCUSSION Two specific factors were hypothesized to aid Norway maple’s invasion. First, that the road, planted with Norway maples as street trees, created an edge effect and acted as a seed source. 1999] Anderson—Invasion of Norway Maple 269 Mean Number of Norway Maples [7] path / Z non-path number of trees PLOT 1 PLOT 2 Figure 2. Mean number of Norway — in plots 1-3 at the Middlesex Fells Reservation. Error bars indicate +1 SE Second, that the paths running perpendicular to the road allowed Norway maple to penetrate greater distances along the path than in nearby nondisturbed areas with no paths. As hypothesized, there were significantly more Norway maples by the road and on the path (Figure 2). The effect of the road seems not to have exceeded 50 m, as there was no significant difference in the number of Norway maple trees in plots 2 and 3. It is possible that there were more Norway maples closer to the road because there were abundant seeds from the nearby street trees, and not due to Norway maple’s tolerance for salt or other edge effects. Even if edge effects were not a significant factor, the path seems to have acted as a place for Norway maple to become established. Since sugar maple abundance has been shown to have a sig- nificant negative correlation with large gaps (Runkle 1984), its abundance was expected to increase with distance from the road. Contrary to expectations, the presence of sugar maples was not significantly affected by the road or the path (Figure 3). The edge effects may not have been strong enough to influence the sugar maple. These results suggest that Norway maple relies more on dis- turbed habitats for establishment than sugar maple does. Invasive plants can follow one of two patterns, either advancing in a front or as scattered small populations (Baker 1986). Species that de- 270 Rhodora [Vol. 101 Mean Number of Sugar Maples sa path non-path number of trees 1 PLOT 2 PLOT 3 3. Mean number of sugar maples in plots 1—3 at the Middlesex Fells Reservation. Error bars indicate +1 SE. pend on disturbance for successful invasion favor the second pat- tern. These species also share many characteristics of early-suc- cessional plants, such as high rates of population growth, pho- tosynthesis, respiration, transpiration, and growth (Bazzaz 1986). In this study, there were often large numbers of Norway maples near the road, but they penetrated further into the forest with large spaces in-between, not in a solid front. In comparison to sugar maple (Kloeppel and Abrams 1995), Norway maple has many of the characteristics listed by Bazzaz that would make it better adapted than sugar maple to exploit disturbance. Norway maple may be contributing to a proposed trend of invasion by exotics and loss of native species in the Middlesex Fells. A survey of species lost over one hundred years found three trends: the number of native species declined, the number of ex- otics increased in proportion to the total flora, and species were most affected in moist habitats (Drayton and Primack 1996). This suggests several effects Norway maple could have in the Fells. For example, in this study, I observed that Norway maple invaded furthest in low, moist areas, which may lead to its contributing to any loss of native species in the moist areas. As Norway maple also has a lower species diversity under its canopy than sugar maple (Wyckoff and Webb 1996), this may also influence the Middlesex Fells’ herbaceous layer in the moister areas. As a common street tree, Norway maple has great invasive 1999] Anderson—Invasion of Norway Maple 271 potential. The likelihood of successful invasion depends on the nature and extent of the disturbance, the number of exotic seeds deposited each year, and how long the community is subject to the propagules (Rejmanek 1989). Thus, communities near street- planted Norway maples are at high risk of invasion. First, the street is a large disturbance, producing an edge effect and intro- ducing pollution and road salt into the community. Second, the woodland is subjected to an annual deposition of seeds from ma- ture Norway maple trees. Since Norway maple is widely planted, many communities are exposed to its seeds. Where the habitat is not conducive to invasion, the threat may not materialize, but having it planted so widely increases the chance that a vulnerable community will be nearby. Since Norway maple is becoming established in the Fells, and becoming the dominant tree near the road at many sites, it would be worthwhile to investigate other species in the Fells that might be influenced by Norway maples’ dense stands. It may be rea- sonable to cut down the roadside trees that are the seed source, replacing them with a native tree such as sugar maple. It is es- pecially important to remove those trees in the Fells that are in sensitive areas. These results also support restrictions on further path construction for recreational purposes. ACKNOWLEDGMENTS. This study was done as part of a Senior Honors Thesis at Tufts University. I would like to thank my com- mittee: Dr. George Ellmore, Dr. Sara Lewis, Dr. Colin Orians, and Dr. Chris Swan. I appreciate Richard Anderson’s generosity with the use of his computer and his statistical advice, and Jen- nifer Kearsley teaching me field methods and giving advice in the early planning of the project. I’m also grateful to three anon- ymous reviewers for their comments. LITERATURE CITED Baker, H. G. 1986. Patterns of plant invasion in North America, pp. 43-57. In: H. A. Mooney and J. A. Drake, eds., Ecology of Biological Invasions of North America and Hawaii. Springer-Verlag, New York. Bazzaz, FE. A. 1986. Life histories of colonizing gee Some A A genetic, and physiological features, pp. 96—10 : H. A. Mooney and J. A. Drake, eds., Ecology of Biological ac oe North America and Hawaii. Springer-Verlag, New York. 212 Rhodora [Vol. 101 BEGON, M., J. L. HARPER, AND C. R. TOWNSEND. 1996. Ecology. Blackwell Science Ltd., Cambridge, M CaNHAM, C. D. 1988. Growth and Seeoy architecture of shade-tolerant trees: Response to canop gaps. Ecology 69: 786-795. DIEKMANN, M. 1996. Ecological behavior of deciduous hardwood trees in Boreo-nemoral Sweden +4 relation to light and soil conditions. Forest Ecol. Managem. 86: Drake, J. A. 1988. Biologia invasions into nature reserves. Trends Ecol. Evol. 3: 186-18 DRAYTON, B. AND ry B. PRIMACK. 1996. Plant species lost in an isolated conservation area in metropolitan Boston from 1894 to 1993. Conser- vation Biol. 10: 30-39. oe _ = 1958. The Ecology of Invasions by Animals and Plants. Red- Press Limited, London Ee S. 1994. Vegetation responses — edge-to-interior gradients in the mixed hardwood forests of the Roanoke River basin, North Carolina. Conservation Biol. 8: 822-832. HARTMANN, H. T., D. E. KESTER, AND E T. Davies, Jr. 1990. Plant Propa- Hosss, R. J. 1991. Disturbance a precursor to weed invasion in native veg- etation. Pl. Protect. Quart. 6: 99-104. : HUENNEKE. 1992. Disturbance, diversity, and invasion: Im- slicatisee for big h ge ihe Biol. 6: 324-337. KLOEpPEL, B. D. AND M. D. ABRAMs. 1995. Ecophysiological attributes of the native Acer nai and ae exotic Acer platanoides in urban oak forests in persion. USA. Tree ime 15: 739-746. LEVIN, E. AND T. MAHLSTEDT. 1990. Middlesex Fells Reservation Historic Land-use Study. Metropolitan District cost Reservations and Historical Sites Division, Boston, MA. MatTLack, G. R. 1993. Microenvironment variation within and among forest edge sites in Lepeve United States. Biol. Conservation 66: 185-194. Mooney, H. A. and J. A. DRAKE, eds. 1986. Ecology of Biological Invasions of North pate and Hawaii. Springer-Verlag, New Nowak, D. J. AND R. A. ROWNTREE. 1990. Hist story and range of Norway 291-296. maple REJMANEK, M. 1989. tap of plant communities, a 369-388. In: J. ake, H. A. Mooney, F di Castri, R. H. Gro s, E J. Kruger, M. Rejmanek, and M. Williamson, eds., Biological lavciiie A Global Per- spective. John Wiley and Sons, New Yor RUNKLE, J. R. 1984. Development of wand Rebctsiin in treefall gaps in a beech-sugar maple forest. Holarc. Ecol. 7: 157-164. Sipe, T. W. AND E A. Bazzaz. 1995. Gap partitioning among maples (Acer) in central diy England: Survival and growth. Ecology 76: 1587~—1602. - AND K. A. Kou. nites Research Priorities for Conservation Diclows tsdend Press, Washin ease: S. A. 1990. bri n, A Reunion at Trees. Harvard University Press, Cam- hii P M. 1990. Biological invasions and ecosystem processes: To- 1999] Anderson—Invasion of Norway Maple 273 wards an integration of population biology and ecosystem studies. Oikos 57: 7-13. WEBB, S. a AND C. K. KAUNZINGER. 1993. Biological invasion of the Drew University (New ri forest preserve by Norway maple. Bull. Torrey Bok? Club 120: 343-— Wyckorr, P. H. anp S. cas B. 1996. Understory influence of the invasive Norway maple (Acer platanoides). Bull. Torrey Bot. Club 123: 197-205. RHODORA, Vol. 101, No. 907, pp. 274-276, 1999 CARDAMINE GEORGIANA (BRASSICACEAE), A NEW NAME REPLACING DENTARIA MICROPHYLLA IHSAN A. AL-SHEHBAZ Missouri Botanical Garden, PO. Box 299, St. Louis, MO 63166-0299 SUZANNE I. WARWICK Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Central Experimental Farm, Ottawa, Ontario KIA 0C6, Canada ABSTRACT. Cardamine georgiana is proposed to replace the illegitimate name C. microphylla (Willd.) O. E. Schulz (1903; non Adams 1817). A dis- cussion on the generic limit of Cardamine including Dentaria is presented. Key Words: Cardamine, Dentaria, Georgia Both Cardamine and Dentaria were simultaneously published by Linnaeus (1753). When uniting the two genera, Crantz (1769) adopted Cardamine for the combined genus and, therefore, this name has priority (Greuter et al. 1994; Article 11.5 ex. 14). Al- though in most treatments (Akeroyd and Marhold 1993; Al-Sheh- baz 1988; Cullen 1965; Rollins 1993; Schulz 1903, 1936) Den- taria is reduced to a synonym of Cardamine, some North Amer- ican authors (Detling 1936; Harriman 1965: Turrill et al. 1994) and eastern European authors (Busch 1909, 1939; Czerepanov 1995; Grossheim 1950; Khintibidze 1979) maintain it as a distinct genus. The alleged morphological differences between Dentaria (larg- er flowers, fleshier and larger rhizomes, often petiolate cotyle- dons, and stems narrowest at base) and Cardamine (usually small- er flowers, often nonfleshy and smaller rhizomes, usually sessile cotyledons, and stems usually broadest at base) Clearly do not hold if the two genera are examined on a worldwide basis (Al- Shehbaz 1988). As indicated by Franzke et al. (1998) and Swee- ney and Price (pers. comm.), molecular data clearly show that Dentaria is polyphyletic and is nested within Cardamine. ing the process of compiling a worldwide checklist of chro- mosome numbers of the Brassicaceae (Cruciferae), it became ev- ident that the transfer by Schulz (1903) of Dentaria microphylla Willd. to Cardamine created a later homonym of C. microphylla 274 1999] Al-Shehbaz and Warwick—Cardamine georgiana 275 Adams and two other homonyms (see below). Therefore, C. geor- giana is proposed as a nom. nov. to replace the illegitimate name. Dentaria microphylla which is narrowly endemic to Georgia and adjacent easternmost Turkey, was recognized in Dentaria by Busch (1909, 1939), Czerepanov (1995), Grossheim (1950), and Khintibidze (1979) under this correct name. It was treated in Car- damine by Cullen (1965), Kolakovsky (1982), and Schulz (1903) under the illegitimate later homonym C. microphylla (Willd.) O. E. Schulz. By contrast, the earlier homonym C. microphylla Ad- ams is a species distributed in northwestern Alaska, the Russian Far East, and eastern Siberia (Berkutenko 1988; Czerepanov 1995; Khatri 1990; Rollins 1993). Cardamine georgiana Al-Shehbaz & Warwick, nom. nov. Re- placed name: Dentaria microphylla Willdenow, Sp. Pl. 3: 479. 1800. Syn. Cardamine microphylla (Willd.) O. E. Schulz, Bot. Jahrb. Syst. 32: 342. 1903, not C. microphylla Adams, Mém. Soc. Imp. Naturalistes Moscou 5: 111. 1817, not C. microphylla J. Presi, Delic. Prag. 1: 15, 1822. TYPE: [GEorGIA]. Collector and locality unknown (HOLOTYPE: B not seen). ACKNOWLEDGMENT. We thank Michael G. Gilbert for his crit- ical review of the manuscript. LITERATURE CITED AKEROYD, J. R. AND K. Maruo_p. 1993. Cardamine, pp. 346-351. In: T. G. tin et al., eds., Flora Europaea, Vol. 1, 2nd ed. Cambridge Univ. Press, Cambridge, Englan AL-SHEHBAZ, I. A. 1988. The genera of Arabideae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arbor. 69: 85-166 BERKUTENKO, A. N. 1988. Brassicaceae, pp. 38-115. In: S. S. Charkevicz, ed., Plantae Vascular Orientis Extremi Sovietici, Vol. 3. Academy of Sciences, Lenin Buscu, N. 1909. inca pp. 356-370. In: N. Kusnezow, N. Busch, and A. Fomin, eds., Flora Caucasica Critica, Vol. 3(4). Typographie K. Matte- cena, Urev - L9So: Dentiria, pp. 144-153. In: V. L. Komarov, ed., Flora of the USSR, Vol. 8. Academy of Sciences of USSR, Moscow & Leningrad. Crantz, H. J. N. 1769. Classis Cruciformium Emendata. Leipzig. CULLEN, J. 1965. Cardamine, pp. 438-444. In: P. H. Davis, ed., Flora Turkey, Vol. 1. Edinburgh Univ. Press, Edinburgh. 276 Rhodora [Vol. 101 CZEREPANOV, S. K. 1995. Vascular Plants of Russia and Adjacent States (the Former USSR). Cambridge Univ. Press, Cambridge, Englan eekartee L. E. 1936. The genus Dentaria in the Pacific States. yy tin J. Bot. 70-576. FRANZKE, A., K. POLLMAN, W. BLEEKER, R. KOHORT, AND H. HurKa. 1998. Molecular systematics of Cardamine and allied genera (Brassicaceae): ITS and non-coding — DNA. Folia Geobot. 33: 225-240. GreuTer, W., FE R. Barrie, H. M. BurDET, W. G. CHALONER, V. DEMOULIN, T : menclature (Tokyo Code). Koeltz Scientific Books, Kénigstein y. GrossHEIM, A. A. 1950. Flora Kavkaza, 2nd ed. Academy of Sciences of SSR, Moscow & Leningrad. HARRIMAN, N. A. 1965. The genus Dentaria j (Cruciferae) i in eastern North Sl. wis A. A. 1982. Flora Abkhazia, Vol. 2, 2nd ed. Metsniebera, Tbi- panes C. 1753. Species Plantarum, Vol. 2. The Ray Society, London. ROLLINS, R. C. 1993. Cruciferae of Continental North America. Stanford Univ. Press, Stanford. ScHUuLz, O. E. 1903. Monographie der Gattung Cardamine. Bot. Jahrb. Syst. 32: 280-623. - 1936. Cruciferae, pp. 227-658. In: A. Engler and H. Harms, eds., Nat, ce 17b, 2nd ed. W. Englemann, Leipzig. TURRILL, N. L., . EVANS, AND G. S. GILLIAM. 1994, fdeaenictlite of West Virginia sears of the Dentaria complex (D. diphyllu Michx., D. het- erophylla Nutt., and D. laciniata Muhl. ex Willd. oe using above-ground vegetative characters. Castanea 59: 22-30. RHODORA, Vol. 101, No. 907, pp. 277-297, 1999 COVARIANCE OF LICHEN AND VASCULAR PLANT FLORAS JAMES P. BENNETT Biological Resources Division, U. S$. Geological Survey, Institute for Environmental Studies, University of Wisconsin, 504 Walnut St., Madison, WI 53705 CLIFFORD M. WETMORE Department of Plant Biology, University of Minnesota, 1445 Gortner Ave., St. Paul, MN 55108 ABSTRACT. The € geographic oe among taxonomic groups are im- Lakes parks, between 28-30% of either the vascular plant or lichen species were singletons (occurring in only one park), but the parks that contained the fekens species (2%) than vascular plants (4%) occurred in Latitude appeared to explain some of the variation between the two groups: vascular plants decreased with increasing latitude, while lichens increased. Key Words: floras, lichens, vascular plants, species-area curves The number of lichen species in North America, north of Mex- ico, is thought to be approximately 3600 (Esslinger and Egan 1995), while the number of vascular plant species approaches 22,000 (Kartesz 1994), a ratio of about six vascular plant species to one lichen species. Although general, ratios like this have been used to make broad biodiversity estimates (e.g., Hawksworth 1991). Such estimates can be made more useful if the geographic pattern of covariance of higher taxa can be ascertained. In recent years studies of spatial covariance of higher taxa have been con- ducted in order to determine patterns of biodiversity hotspots, or ail 278 Rhodora [Vol. 101 the relationships among taxa, if any exist (Faith and Walker 1996; Gaston 1996). Many factors can affect spatial covariance, includ- ing scale, environmental factors, and biological properties of the taxa. Several authors have pointed out that lichen and vascular plant diversities may not track each other in general due to cli- mate (Huston 1994) and habitat diversity (Galloway 1992; Gilbert 1977). Another study found that lichen biodiversity decreased significantly with an increase in vascular plant cover (Pharo and Beattie 1997). In addition, we suspected that the effects of air pollutants may also be a factor affecting lichen and vascular plant floras differentially because of the greater air pollution sensitiv- ities of lichens. This paper explores all of these issues using the lichen and vascular plant floras of nine national parks in the Great Lakes region of the United States. The lichen floras of nine national parks in the Great Lakes region of the north central United States have been well studied in recent years and are considered to be fairly complete. The parks include Apostle Islands National Lakeshore in Wisconsin (APIS), Cuyahoga Valley National Recreation Area in Ohio (CUVA), Grand Portage National Monument in Minnesota (GRPO), Indiana Dunes National Lakeshore in Indiana (INDU), Isle Royale National Park (ISRO) and Pictured Rocks National Lakeshore in Michigan (PIRO), St. Croix National Scenic River in Minnesota and Wisconsin (SACN), Sleeping Bear Dunes Na- tional Lakeshore in Michigan (SLBE), and Voyageurs National Park in Minnesota (VOYA; Figure 1). It is logical to study these floras because they should show affinities with one another, being associated geographically within the homogeneous Great Lakes/ north central U. S. region. The nine parks span a region of about 1200 km longitudinally, and in the aggregate, cover a total area of 201,000 km?. The vascular plant floras of these parks have been studied and were segregated into two groups by multivariate analyses (Bennett 1996a, 1996b). This study was also undertaken to determine if the same groups of parks are segregated based on the lichen floras. The management of natural area preserves is sometimes fo- cused on a select group of organisms, e.g., mammals, trees, rare plants, birds, or butterflies. It is often implicit in the management of these groups that the management of one group will also sat- isfactorily manage another group by association. This is because it is assumed that the biological groups in an area are related. For 1999] Bennett and Wetmore—Covariance of Floras 279 Figure 1. Map of nine Great Lakes national park units for which floras were used in this study. Park codes are explained in the introduction. the purposes of this study, it was assumed that vascular plants and lichens are related floristically somehow, although ecologi- cally they occupy habitats at very different scales and their flo- ristic relationships may be obscure. From a management per- spective however, it may be assumed that if there are more vas- cular plants in an area there will also be more lichens. This study was undertaken in order to determine the validity of this as- sumption by attempting to prove the hypothesis that there is no relationship between vascular plant and lichen floras. MATERIALS AND METHODS Lichen floristic field work in the nine parks was conducted during the summer in the years from 1978 to 1995, although collecting was not done in these particular parks every year (Wet- 280 Rhodora [Vol. 101 a . Numbers of collections, collecting days, and collection localities for lichens in nine Great Lakes national parks. See introduction for park names. Number of Collecting Collection Park Collections Days Localities APIS 1497 21 28 CUVA 3 11 31 GRPO 373 3 6 INDU 371 14 24 ISRO 5246 53 2 PIRO 1231 11 25 SACN 2327 88 77 SLBE 847 10 27 VOYA 8028 80 128 park were selected to include all habitats and vegetation types. The collection localities were distributed over the entire park, and vegetation types were studied multiple times. At each locality, all groups of lichens were collected (fruticose, foliose, squamulose, and crustose) and sufficient time was spent examining all sub- strates. At each locality all species found were collected to pro- vide relative abundance estimates, even though the same species might have been found at previous localities. A summary of the collecting efforts is shown in Table 1. Vascular plant floras and methods were described previously (Bennett 1996a). The nomenclature of each lichen flora was updated to the fifth checklist (Egan 1987) so as to standardize all the names. The names were then entered into a computer spreadsheet program (MICROSOFT EXCEL, Microsoft Corp., Seattle, WA) and the presence/absence of each species recorded in a field for each park. arious tallies and sorts were performed in order to perform qual- ity control procedures and to rank species by frequencies. This file was also used to analyze the data Statistically, using MINI- TAB (Minitab, State College, PA) and SYSTAT (SPSS, Chicago, IL). Relationships between variables were tested with Pearson’s linear correlation coefficients. Similarities between park floras were calculated using Jaccard’s 1999] Bennett and Wetmore—Covariance of Floras 281 Table 2. Area, numbers of lichen and vascular plant apace average Jac- card similarity based on lichens and vascular plants, and average distance from the other parks for nine Great Lakes national parks. See introduction for park names erat eee of Average Similarity (decimal ee Vascular Average Area degrees Vascular Lichen Plant Distance Park (km?) N) Lichen Plant (%) (%) (km) APIS 6605 47.000 285 509 39 35 388 CUVA 3136. =“ 4 4-283 66 855 18 26 842 GRPO 287 47.967 182 279 30 23 383 INDU 5203 41.625 62 1399 19 28 638 ISRO 54,140 48.000 554 698 31 OT 400 PIRO 25,545 46.567 245 123 36 39 415 SACN 26,459 46.000 282 1165 34 33 443 SLBE 23,663 44.875 182 928 33 oT 426 VOYA 55,955 48.500 458 603 35 35 566 similarity index, which measures the proportion of park pairs where species are present in both parks in the pair. Details are given in Bennett (1996b). The correlation of two similarity ma- trices was tested with Mantel’s test with PC-ORD (McCune and Mefford 1997). Park distances and latitudes were measured using STREET AT- LAS (De Lorme, Freeport, ME), a desktop computer mapping program. Straight line distances and latitudes were calculated us- ing the approximate park centroids for park locations. Distances were rounded to the nearest five kilometers. RESULTS A floristic summary of each park is shown in Table 2. Isle Royale had the greatest number of lichen species, while Indiana Dunes had the lowest. The greatest and lowest numbers of vas- cular plant species, however, were at Indiana Dunes and Grand Portage, respectively. Based on the lichen floras, seven of the parks were, on average, between 30 and 40% similar to all the other parks, except for Indiana Dunes and Cuyahoga Valley, which were almost 20% similar to all other parks. This is not too surprising, given that on average they are farther away from all the other parks. Overall similarity based on vascular plant floras 282 Rhodora [Vol. 101 followed the same pattern, except for Grand Portage, which had the lowest overall similarity, lower than Cuyahoga Valley and Indiana Dunes. Some of the variables were significantly correlated (Table 3): lichen numbers and park area (Figure 2), and average lichen sim- ilarity and average distance (Figure 3) or latitude were the stron- gest relationships. Number of lichens increased with latitude, while vascular plant numbers decreased (Figure 4). Numbers of lichens and vascular plants appeared to be negatively correlated, but the probability of the correlation occurring by chance was high (0.33). Park similarities based on lichens and vascular plants were comparable (P = 0.0517). Vascular plant similarity signif- icantly increased with area and did not appear related to latitude, while park lichen similarity increased with latitude and was not related to area. The lichen flora Jaccard similarity matrix for all possible 36 pairs of the nine parks is shown in Table 4. Two park pairs, Isle Royale and Voyageurs, and Apostle Islands and Pictured Rocks were greater than 50% similar, while the overall average was 31%. The comparable distance matrix (Table 5) shows that Cuy- ahoga Valley and Voyageurs are just over 1200 km apart, while Grand Portage and Isle Royale are the closest pair at only 60 km apart. The similarities for these same pairs were 11% and 30%, respectively (Table 4). When all 36 pairs of similarities and dis- tances were plotted against each other (Figure 5), there was a clear negative relationship, even though the maxima and minima did not all correspond. The overall average of vascular plant similarities was 32% (see Table 1 in Bennett 1996b), and only one park pair, Isle Royale and Pictured Rocks, was more than 50% similar. The similarities of Cuyahoga Valley and Voyageurs, and Grand Portage and Isle Royale were 23% and 30%, respectively. The overall vascular plant similarity matrix was significantly positively related to the lichen similarity matrix (Mantel r = 0.9151, t = 6.3718, P = 0.0000), suggesting that these parks show comparable degrees and patterns of similarity based on both vascular plant and lichen oras. A cluster analysis of the lichen flora Jaccard similarity matrix did not reveal any significant groupings because the similarities were not very diverse. However, a cluster analysis of the pres- ence/absence matrix of all species in the nine parks did reveal ble 3. Pearson’s linear correlation wapaie ator for variables given in a 2. Significance levels of coefficients are indicated Ta by preceding asterisks: * (0.05), ** (0.01), and * * (0.001). ' Number of speci Average Vascular Average Lichen Vascular Plant Area Latitude Lichens' Plants! Similarit Similarity Latitu , Lichens' ***0.9124 **().7968 Vascular plan —0.1671 *—0.6985 —0.3715 Average vee similarity 0.5214 **0.8379 0.6210 —0.4850 Average vascular plant similarity *0.7054 0.3882 0.5783 0.0440 6628 Average distance —0.2143 *—0.7685 —0.4796 0.4002 **—(0.8083 —0.4540 [6661 selOLJ JO duRLRAOD—oI0UNE AM pur youUDg €87 284 Rhodora [Vol. 101 500 Lichens = 0.7817 area + 54.262 s R? = 0.8325 : 3 400 ‘ ® > 300 | 2 200 | = 2 100 | 0 ' T T T T 0 > 200 S00 400. 500. 800 Area (km?) Figure 2. Relationship between number of lichen species and area for nine Great Lakes national parks, and fitted linear regression line. some interesting groups (Figure 6). At the highest similarity level, abo %, four parks (Apostle Islands and Pictured Rocks, and Cuyahoga Valley and Indiana Dunes) were grouped into two pairs that were themselves only about 36% similar. Cuyahoga Valley and Indiana Dunes appeared to be two parks that were unrelated floristically to the other seven, which appeared to be about 50% similar overall. Isle Royale, however, was less similar to the other six northern parks, in spite of the high individual Jaccard simi- larity with Voyageurs. Two groups of three parks each had sim- Average lichen similarity oO nN 0.1 4 Similarity = -0.0004 distance + 0.4997 R 6 300 400 500 600 700 800 900 Average distance (km) Figure 3. Relationship between average lichen flora similarity and aver- age distance from the other parks for the nine Great Lakes national parks, and fitted linear regression line. 1999] Bennett and Wetmore—Covariance of Floras 285 1600 1400 - “‘ Vascular plants = -88.418 degrees + 4841 R? = 0.4878 Lichens = 48.801 degrees - 1975 R? = 0.635 8 Number of species co So So 40 41 42 43 as 45 46 47 48 49 Latitude (degrees North) @Lichens 4 Vascular plants Figure 4. Relationships between number of vascular plant and lichen spe- cies and latitude for nine Great Lakes national parks, and fitted linear re- gression lines. ilarities of 60-70%. One of these had two parks on the southern shore of Lake Superior and one at the northern end of Lake Mi- chigan (Apostle Islands, Pictured Rocks, and Sleeping Bear Dunes), and the other consisted of three Minnesota parks (Grand Portage, Voyageurs, and St. Croix). Totals of 698 lichen species in 162 genera were found in the nine parks when the floras were aggregated. The most frequent species, i.e., those that were found in all nine parks (14 species, or 2% of all lichen species found), included Arthonia caesia, Candelariella efflorescens, Cladina rangiferina, Cladonia chlo- Table 4. Jaccard similarity (%) matrix for lichen floras of nine national parks. APIS CUVA GRPO INDU ISRO PIRO SACN SLBE PIRO bof BS 13.1 40.9 i) see SACN 43.5 20.0 33.7 19.0 364 P SLBE 42.8 20.4 31.9 pa 8 4.7 39.8 VOYA 49.8 11.0 35.6 11.1 59.1 41.7 457 30.6 286 Rhodora [Vol. 101 Table 5. Distance matrix for nine national parks. Distances are in kilo- meters between park centroids. APIS CUVA GRPO INDU ISRO PIRO SACN SLBE CUVA 950 GRPO 145 975 U 670 470 = 745 ISRO 185 930 725 PIRO 340 690 MO.2400 862 SACN 125 980: 275 620 320 435 SLBE 425 ao0-=2 7900 390 §=6410 185 480 VOYA 260 1210 260 Jae -320ic | 560 = 310 685 rophaea, C. coniocraea, C. cristatella, Flavoparmelia caperata, Lepraria finkii, Melanelia subaurifera, Parmelia sulcata, Phaeo- physica pusilloides, Physconia detersa, Punctelia rudecta, and Scoliciosporum chlorococcum. Thirty percent of lichens (210 spe- cies) occurred in only one park. Most of these (129) occurred in Isle Royale. For the vascular plants, an aggregated flora of 2102 species in 691 genera was found over all the parks, with 81 (4%) occurring in all nine parks. Twenty-eight percent of the vascular 0.70 0.60 - e, Vascular plants = -0.0002 distance + 0.4251 0.50 - . R? = 0.2612 = 0.40 | : : E 0.30 }os 0.20 “ - = 0.10 Lichens = -0.0004 distance cia 4 - 0.00 +———S=agee Cd 0 500 1000 1500 Average distance (km) 4 Vascular plants @ Lichens Figure 5. Relationships of vascular plant and lichen flora similarities to distance between parks for each of the 36 possible park pairs of the nine Great Lakes national parks, and fitted regression lines. 1999] Bennett and Wetmore—Covariance of Floras 287 Similarity (%) . 1 00 + T ! & . & oO N) 6 fe er le le Parks e 6. Cluster dendrogram for nine Great Lakes national lichen higel using Ward’s linkage and correlation for the distance meas plants (596 species) occurred in just one park each. Most of these (221) were found in Indiana Dunes. DISCUSSION At first glance it appears that for this set of nine park floras there was no obvious relationship between vascular plants and lichens. However, further examination of the parks sheds more light on the problem. Two parks, Cuyahoga Valley and Indiana Dunes, are in locations that have been subject to oxidant and sulfur oxide pollution fumigations for many years, and studies have shown that the lichen floras of these parks have decreased to about 80% of their historical floras (Wetmore 1988a, 1989). However, if these two parks were to be omitted from the analysis there would not be enough data points to draw justifiable conclu- sions about the vascular plant/lichen relationship. When the his- torical total numbers of species were used instead of current spe- cies numbers (205 for Cuyahoga Valley and 154 for Indiana Dunes), there was still no clear relationship. This appears due to the vascular plant flora numbers rather than the lichen flora num- bers. The vascular plant flora numbers were not linearly related to area (Table 3). This is probably due to confounding effects of 288 Rhodora [Vol. 101 Lichens = 1.0261 Vascular plant Lichens 100 1,000 10,000 100,000 1,000,000 Vascular plants Figure 7. Relationship between numbers of lichen and vascular plant spe- cies for 33 areas, and fitted non-linear regression line. latitude, as vascular plant diversity decreases with increasing lat- itude, the opposite of what happens with the lichen diversity (Huston 1994). Do lichen and vascular plant floras co-vary? At the scale of the Great Lakes no relationship could be seen, but what about at other scales? The flora data for the nine parks were combined with comparable data for 24 other geographic areas, including the world, and are shown in Figure 7 (Table 6). It appears that when larger geographic areas were included, a significant, positive re- lationship was found between the two floras (r = 0.9808), even when the floras of the world were omitted (r = 0.7037). When only areas less than 1000 km? were considered, the relationship broke down (r = 0.0896). Area was obviously a factor when all 33 areas were considered (Figure 8), but may not have been with areas under 1000 km? for the reasons given above. This finding differs significantly from an older study of lichens and area where no relationship was found above 1000 km? (Wetmore 1967), which was based on incomplete inventories. We conclude that any relationship between vascular plant and lichen floras is scale-dependent. For natural areas under approx- imately 1000 km?, no relationship was found for the national Table 6. Areas, numbers of vascular plant (VP) and lichen (L) species, and the ratio of the two (VP/L), for 33 locations. Vascular Reference and Location (km?) Plants Lichens VP/L Comment Homestead National Monument, NE 1 2er 19 TE.9S Bennett 1996a; Wetmore an Bennett 1997 George Washington Carver National Monument, 1 605 38 15.92 Bennett 1996a; MO Wetmore 1992a Pipestone National Monument, MN 1 439 66 6.65 Bennett 1996a; Wilson and Vinyard 1 Grand Portage National Monument, MN 3 279 182 1,53 Be 19 Wetmore 1992b Effigy Mounds National Monument, IA 6 458 73 5.80 Bennett 1996a; Wetmore and Bennett 1997 Wilson’s Creek National Battlefield, MO i) 454 88 5.16 Bennett 1996a; Wetmore and Bennett 1997 Tuxedni Wilderness, AK 23 290 214 1.36 Talbot et al. 1995; Talbot et al. 1992 Indiana Dunes National Lakeshore, IN 52 1399 62 22.56 Bennett 1996a; Wetmore 1988a Apostle Islands National Lakeshore, WI 66 509 285 1.79 Bennett 1996a; Wetmore 1990a Cuyahoga Valley National Recreation Area, OH 131 855 66 12.95 Bennett 1996a; Wetmore 1989 [6661 SPIO, JO DOURLIEAOD—sIOUTIZ AA pue WoUuUUg Table 6. Continued. Area Vascular Reference and Location (km?) Plants Lichens VP/L Comment Nantucket Island, MA 140 939 99 9.48 Dunwiddie and Sorrie 1996 (species not seen after 1960 tted) Acadia National Park, ME 190 858 397 2.16 cis 1997b; Wetmore 1984 Sleeping Bear Dunes National Lakeshore, MI 237 928 182 5.10 Bennett 1996a; Wetmore 1988b Pictured Rocks National Lakeshore, MI 255 723 245 2.95 Bennett 1996a; Wetmore 1990b St. Croix National Scenic Riverway, MN and WI 265 1165 282 4.13 Bennett 1996a; Wetmore 1991 Theodore Roosevelt National Park, ND 285 550 212 2:09 Anonymous 1997b; Wetmore 1983a Chiricahua National Monument, AZ 485 672 200 3.36 Anonymous 1997b; Wetmore and Bennett 1992 (Lichen number is doubled because flora is 50% known.) Isle Royale National Park, MI 541 698 554 1.26 Bennett 1996a; Wetmore 1985 560 603 458 1-32 Bennett 1996a Voyageurs National Park, MN Wetmore 1983b 067 elopoyy 101 ‘I°A] Table 6. Continued. Area Vascul Reference and Location (km?) Plants Lichens VP/L Comment Glacier National Park, MT 4,102 1147 425 2.70 Anonymous 1997b; Debolt and McCune 1993 Yellowstone National Park, WY 8,983 1150 236 4.87 Anonymous 1997b; Eversman 1990 Israel 21,946 2250 229 9.83 Zohary 1962; Kondratyuk et al. 1996 Netherlands 41,784 1781 633 2.81 Statistics Netherlands 97; B. Wit, pers. comm. Tasmania 67,858 1456 655 2.22 Chapman 1997; Galloway 1992 United Kingdom 229,979 2397 1,730 1.39 Palmer 1996; Galloway 1992 New Zealand 268,114 4167 1,162 3.59 Allan 1961; Moore and Edgar 1970; Healy and Edgar 1980; Webb et al. 1988; Galloway 1985 Finland 338,148 2423 1,420 Bvt Anonymous 1997a; O. Vit- Kainen, pers. comm. 1997 [6661 SvIO]J JO 90URLIeAOD—aIOUT]IO AA pue WouUEg 167 Table 6. Continued. Area Vascular Reference and Location (km?) Plants Lichens VP/L Comment California 411,049 5862 1,000 5.86 Hickman 1993; Tucker and Jordan 9 British Columbia 948,600 2850 1,600 1.78 British Columbia Ministry of Environment, Land, and Parks 199 Australia 7,682,341 14,679 2,499 5.87 Chapman 1997; Galloway 1992 China 9,596,961 30,000 1,274 23:55 Harvard University Herbaria 1997; Wei 1991 United States and Canada 21,479,211 21,757 3,600 6.04 Kartesz 1994; Esslinger and Egan 1995 World 149,702,000 243,893 17,000 14.35 Mabberley 1987; Thorne 1992; Galloway 1992 C6T eIOpouYy TOT TOA] 1999] Bennett and Wetmore—Covariance of Floras 293 1,000, 3 A 100,000 ; 3 0.2542' R? = 0.8074. 4A A — 2 ns @ oe nm ee Le z 000 ont ae noma “a a Se crys Out 2 a, my \ ei a4 cr eee ae ts oe = ei Lichens = 50.827 area?*P ‘eee ee R? = 0.8453 . 1 1 e 10 : u 16-01 16400 1.6401 1.6402 1.6403 16+04 1.6405 1.6406 1.6407 1.6408 1.E+09 Area (km’) Figure 8. Relationships between numbers of lichen and vascular plant species and area for 33 areas, and fitted non-linear regression lines. parks and areas in this study. This is partly due to the confound- ing effect of latitude, which affects vascular plant and lichen rich- ness in opposite ways. A second reason is that we deliberately chose a set of natural areas in a relatively homogeneous ecolog- ical region so as not to introduce significant habitat diversity. The third reason is that vascular plants and lichens occupy microhab- itats that are unrelated because of the scale differences between the two groups of plants. Lichens occupy habitats at the centi- meter scale, while vascular plants could be said to occupy habitats at the meter scale. In addition, lichens can grow in some micro- habitats where vascular plants are not found, e.g., house roofs or rocks. However, when the range of natural areas was expanded to include those larger than 1000 km’, a relationship between vascular plant and lichen floras was found. This is most likely due to the effect area has on increasing ecosystem diversity, which produces more overall habitat diversity and hence species diversity. The similarity clusters of the parks based on the two types of floras were not completely congruent. Both separated Indiana Dunes and Cuyahoga Valley from the other seven parks, although St. Croix was grouped with them for the vascular plants. This separation of Cuyahoga Valley and Indiana Dunes is probably 294 Rhodora [Vol. 101 related to geographic distance, but may also be confounded by the profound effects of air pollutants on the lichen floras. In the remaining seven parks, Isle Royale appeared rather unique for lichens in this region, but not for vascular plants. In fact, for vascular plants, very little splitting of the cluster of seven was apparent (Bennett 1996b). Lichen floras in this area appeared to be more unique geographically than vascular plant floras, prob- ably due to the diversity of species in the Great Lakes, not the distribution of species per se. The ratio of six vascular plant species to one lichen species for North America appears to hold on average for the 33 areas as well (Table 6): the overall average ratio for these areas was 6.20 (SD = 5.89). The ratio found for the world floras, however, was an exception: it was over double the average ratio for North America. No logical explanation can be proposed at this time for this anomaly In conclusion, although average similarities of these nine parks, based on vascular plants and lichens, were comparable (just over 30%), this generalization hides the fact that the two types of floras do not appear related to one another at this scale. There was a closer relationship among the vascular plant floras than among the lichen floras, suggesting that geographic affinities between em are weaker for lichens. Thus, management strategies to con- serve the vascular plant floras may not conserve the lichen floras on an equal basis. Lichen biodiversity may have to be managed differently than vascular plant biodiversity. ACKNOWLEDGMENTS. Funding for the lichen field work de- scribed in this study came from the Midwestern Regional Office, National Park Service, Omaha, Nebraska, and the Biological Re- sources Division, U. S. Geological Survey, Madison, Wisconsin, and is gratefully acknowledged. LITERATURE CITED ALLAN, H. H. 1961. cio of New Zealand, Vol. 1. Government Printer, Wel- lington, New Zeal ANONYMOUS. 1997a. kes of the Vascular Plants of Finland. Internet URL: www.helsinki.fi/kmus/chklst.html. 1997b. Flora and fauna of the national parks. Internet URL: endeav- edu/NPS. our. lee ucdavis. 1999] Bennett and Wetmore—Covariance of Floras 295 BENNETT, J. P. 1996a. Floristic summary of 22 national parks in the mid- western United States. Nat. Areas J. 16: 295-302. . 1996b. Similarities and life cycle distributions of floras of 22 national parks in the midwestern United States. Nat. Areas J. 16: 303-309. . 1997. Nested taxa-area curves for eastern United States floras. Rho- dei 99: 241-251. BRITISH COLUMBIA MINISTRY OF ENVIRONMENT, meine AND ParKS. 1997. In- ternet URL: www.env.gov.be.ca/wld/biodiver. pres Az 1997. Seonety Statistics on the: flora of repre Internet URL: html DEBOLT, ‘. AND B. MCCUNE. 1993. Lichens of Glacier National I Park, Mon- tana. Bryologist 96: 192-204. DUNWIDDIE, P. W. AND B. A. Sorrie. 1996. A flora of the vascular and non- vascular plants of Nantucket, Tuckernuck, and Muskeget Islands. Rho- dora 98: 94-98. Ecan, R. S. 1987. A fifth checklist of the lichen-forming, lichenicolous and allied fungi of the continental United States and Canada. Bryologist 90: 77-173. EsSLINGER, T. L. AND R. S. EGAN. 1995. A sixth checklist of the liche orming, lichenicolous, and allied fungi of the continental United sae and Canada. Bryologist 98: 467-549 EVERSMAN, S. 1990. Lichens of Yellowstone National Park. Bryologist 93: 197-205 Faitu, D. BP. AND P. A. WALKER. 1996. How do indicator groups provide information about the relative biodiversity of different sets of areas?: On hotspots, complementarity and pattern-based approaches. Biodiversity Letters: 3: 18—25. GaLLoway, D. J. 1985. Flora of New Zealand, Lichens. P. D. Hasselberg, Government Printer, Wellington, New and. . 1992. Biodiversity: A lichenological perspective. Biodiversity and Conservation 1: 312-323. Gaston, K. J. 1996. Spatial covariance in the species richness of higher taxa, Cha. 12. In: M. E. Hochberg, J. Clobert, and R. Barbault, eds., Aspects of the Genesis and Maintenance of Biological Diversity. Oxford Univ. Press, Oxford, England. Gicpert, O. L. 1977. Lichen Conservation in Britain, Cha. 11. Jn: M. R. D. Seaward, ed., Lichen Ecology. Academic Press, New York. HarVARD UNIVERSITY HERBARIA. 1997. Internet URL: herbaria. harvard. e , Hawxksworth, D. L. 1991. The fungal dimension of biodiversity: Magnitude, significance, ona conservation. Mycol. Res. 95: 641—65 HEALY, A. J. AND E. Epcar. 1980. — of New Zealand, Vol. 3. Government Printer, Wellington, New Zealan ares J. E., ed. 1993. The aaa cere Higher Plants of California. Univ. of California Press, Berkeley, C Huston, 'M. A. 1994. Biological cnc ag "Cambridge Univ. Press, Cam- bridge, England. Kartesz, J. T. 1994. A Synonymized Checklist of the Vascular Flora of the 296 Rhodora [Vol. 101 United States, Canada, and Greenland, 2™4 ed., Vol. 1—Checklist. Timber Press, Portland, ap Ps ahienly S. Y., I. L. NAvrotskayA, S. D. ZELENKO, S. P. WASSER, AND E. NEvo. 1996. The First Checklist of Lichen-Forming and Lichenicol- ous Fungi of Israel. Kholodny Inst. Of Botany, Univ. of Haifa. Peledfus Publ. House, Kyiv, Israel. bigger og D. J. 1987. The Plant-Book, A Portable Dictionary of the Higher Plants. Cambridge Univ. Press, Cambridge, Englan eee B. AND M. J. MEFFORD. 1997. PC-ORD. Multivariate Analysis of Moore, L. B. AND E. Epaar. 1970. Flora of New Zealand, Vol. 2. Govern- ment Printer, Wellington, New Zealand. PALMER, M. A. 1996. A strategic approach to the conservation of plants in the United Kingdom. J. Appl. Ecol. 33: 1231-1240. PHARO, E. J. AND A. J. BEATTIE. 1997. sy ce and lichen diversity: A co STATISTICS NETHERLANDS. 1997. Internet ae Be Ses TALBOT, S. S., S. L. TALBOT, AND J. W. THOMSON. 1992. Lichens of the Tuxedni Bivens Area, Alaska. Bryologist 95: 20-30. . L. WELSH. 1995. Botanical Reconnaissance of the Tuxedni Wilderness Area, Alaska. Biological Science Report 6, National Biological Service, U. S. Department of the Interior, Washington, DC. THORNE, R. T. 1992. Classification and geography of the flowering plants. Bot. Rev. oa 225-348. Tucker, S. C. AND W. P. JorpaNn. 1979. A catalog of California lichens. Ee: ‘WEBB, ret W. R. SYKES, AND P. J. GARNOCK-JONES. 1988. Flora of New Ze d, Vol 4. cys Division, D. S. I. R., Christchurch, New Zealand. WEI, anaes 1991. An Enumeration of Lichens in China. International ademic peng Beijin WETMoRE, C. M. 1967. Lichens of the Black Hills of South Dakota and Wyoming. Publication of the Museum, Michigan State Univ., Biological Series, Vol. 3, No. 4, East Lansing, MI. . 1983a. bien and Air Quality in Theodore Roosevelt National Park. Final Report. Botany Dept., Univ. of Minnesota, St. Paul, MN . 1983b. Lichens and Air Quality in Voyageurs National Park. Final Report to the National Park Service. Botany Dept., Univ. of Minnesota, St. Paul, MN. . 1984. Lichens and Air Quality in Acadia National Park. Final Re- port to the National Park Service. Botany Dept., Univ. of Minnesota, St. Paul, . 1985. Lichens and Air Quality in Isle Royale National Park. Final Report to the National Park Service. Botany Dept., Univ. of Minnesota, St. Paul, MN. - 1988a. Lichens and air quality in Indiana Dunes National Lakeshore. Mycotizon 33: 25-39. . 1988b. sprog of Sleeping Bear Dunes National Lakeshore. Michi- gan Bot. 27: 111-118. 1999] Bennett and Wetmore—Covariance of Floras 297 . 1989. Lichens and air quality in Cuyahoga Valley National Recrea- tion Area, Ohio. Bryologist 92: 273-281. . 1990a. ae of Apostle Islands National Lakeshore. Michigan Bot. 29: 65-7 1990b. iaisaa of Pictured Rocks National Lakeshore. Michigan Bot. 29: 19-26. . 1991. Lichens and Air Quality in St. Croix National Scenic River- way. Final Report to the National Park Service. Botany Dept., Univ. of Minnesota, St. Paul, MN. . 1992a. Lichens and Air Quality in George Washington Carver Na- diaiat Monument. Final Report to the National Park Service. Plant Bi- ology Dept., Univ. of Minnesota, St. Paul ——. 1992b. Lichens and Air Quality in Grand Portage National Monu- ment. Final Report to the National Park Service. Botany Dept., Univ. of Minnesota, St. Paul, MN. AND J. P. BENNETT. 1992. NPLICHEN: A National Park Service Li- chen Data Base. Final Report. Plant Biology Dept., Univ. of Minnesota, St. Paul, MN. AN . 1996. 1995 Lichen Studies in Apostle Islands National Biological Service. Botany Dept., Univ. of Minnesota, St. Paul, MN. AND 997. Lichens and Air Quality in Three Prairie Parks Final Report. Biological Resources Division, U. S. Geological Survey, Madison, Witson, G. D. AND T. W. VINYARD. 1986. Changes in the lichen flora of ‘ipestone National Monument. Prairie Nat. 18: 9-14 ZOHARY, M. 1962. Plant Life of Palestine. Ronald Press, New York. RHODORA, Vol. 101, No. 907, pp. 298-299, 1999 NEW ENGLAND NOTE NOTES ON THE HABITS AND LIFE-HISTORY OF BIDENS DISCOIDEA: AN EPIPHYTE IN MASSACHUSETTS FLOODPLAIN PONDS MATTHEW G. HICKLER University of Massachusetts, Department of Biology, Amherst, MA 01003 Bidens discoidea (T. & G.) Britton is uncommon or under- collected in Massachusetts and in need of attention from field botanists to clarify its status in the state (Sorrie 1990; P. Somers, pers. comm.). It resembles the common B. frondosa L. for which it is easily mistaken. However, the two species are readily iden- tified in the field by the number of outer involucral bracts: three to five in B. discoidea, and five to ten in B. frondosa. Bidens discoidea is an annual whose range covers much of the eastern half of the United States and adjacent Canada (Gleason and Cronquist 1991). Wiegand (1899) considered the species to be “quite rare in New England” and more common in the western part of its range where it grows on logs and stumps in lakes and bogs. In the southeastern Coastal Plain it is found “usually on fallen logs and stumps in swamps” (Fernald 1936) and is “‘con- spicuous” in freshwater tidal swamps (Beaven and Oosting 1939). Sorrie (1990) commented that he was aware of four extant Massachusetts stations for Bidens discoidea and noted habitats as ponds, oxbows, and buttonbush (Cephalanthus occidentalis 5) swamps. He also noted an occurrence of it growing as an epi- phyte, but provided no further details. Robert Bertin (pers. comm.) recently found previously undocumented populations of B. discoidea on the shores of a cluster of beaver ponds in Worces- ter County, Massachusetts, most commonly growing on partially submerged stumps and logs, but occasionally on shoreline soils. Bidens discoidea is common on floodplain ponds along the Na- shua River on land now or formerly part of the Fort Devens Mil- itary Reservation in Worcester and Middlesex Counties, Massa- chusetts. Here, the species is found almost exclusively growing as an epiphyte on Cephalanthus occidentalis and was observed on 13 out of 15 floodplain ponds inventoried in 1995 and 1996. In these ponds, water levels typically decline a meter or more between spring and late summer, but levels fluctuate during the growing 298 1999] New England Note 299 season in response to mse patterns (M. Hickler, unpubl. data). Shoreline vegetation in these ponds often includes dense thickets of C. occi eon At least three species of moss [Cli- maceum americanum Brid., Drepanocladus aduncus (Hedw.) Warnst. var. kneiffi (Schimp. in B.S.G.) M6nk., and Dichelyma capillaceum (With.) Myr.] grow on Cephalanthus stems, where ey form dense tufts, concentrated around the normal high water Bidens discoidea has a strategy for avoiding the stresses as- sociated with unpredictable hydrology and light competition from Cephalanthus in this habitat. The awned achenes, which are well adapted for dispersal by hitchhiking on passing animals, are also perfectly pre-adapted to lodge in the thick tufts of moss. Achenes germinate in their mossy seedbeds after water levels begin to recede in the spring but long before ground-level soil has drained. The roots then follow the receding water toward the ground (M. Hickler, pers. obs.). Perched high in the Cephalanthus stems, the aerial portions of the plants are above the influence of fl ng and in position to penetrate through the dense Cephalanthus can- opy to where there is ample sunlight. By late summer, B. discoi- dea plants can be seen throughout the Cephalanthus swamps with their tops above the shrub canopy in full sunlight and roots pro- jecting down, sometimes a meter or more, to the soil. ACKNOWLEDGMENTS. Thanks to Roberta Lombardi and Sarah Cooper-Ellis for moss identifications. LITERATURE CITED BEAVEN, G. FE AND H. J. Oostinc. 1939. Pocomoke Swamp: A study of a cypress swamp on the Eastern Shore of Maryland. Bull. Torrey Bot. Club 66: 367-389. FERNALD, M. L. 1936. Plants from the outer coastal plain of Virginia. Rhodora 38: 376—4. GLEason, H. A. AND A. CRONQUIST. 1991. Manual of Vascular Plants of North- eastern United States and Adjacent Canada, 2nd ed. The New York Bo- tanical pape Bronx, Sorrig, B. A. 1990. “Watch List”: Uncommon or Rare Massachusetts Plants. naeteree No. 16, 260-10-300-4-C.R., Natural Heritage and Endangered Species Program, Massachusetts Division of Fisheries and Wildlife, Bos- ton, MA. WIEGAND, K. M. 1899. Some species of Bidens found in the United States and Canada. Rhodora 8: 399-422 RHODORA, Vol. 101, No. 907, pp. 300-301, 1999 NEW ENGLAND NOTE A NEW BARNSTABLE COUNTY, MA, RECORD FOR ASCLEPIAS PURPURASCENS DONALD G. SCHALL ENSR, 95 State Road, Buzzards Bay, MA 02532 Mario J. DiGREGORIO AND PAMELA POLLONI Sabatia Inc., 107 Goeletta Drive, Hatchville, MA 02536 While inspecting the status of a small population of Rhodo- dendron canadense (L.) Torr. present in a moist woodland habitat on the edge of a shallow wetland depression in Falmouth, Mas- sachusetts, we discovered three individuals of Asclepias purpur- ascens L. (Asclepiadaceae) in the understory ground cover. The species is found from New Hampshire to Virginia, west to Iowa, Kansas, and Oklahoma, and north to Wisconsin (Gleason and Cronquist 1991). Purple milkweed is identified as belonging to a “Division 2; Regionally Rare Taxa” category in “‘Flora Conser- vanda: New England” by Brumback and Mehrhoff et al. (1996) with fewer than 20 current occurrences in New England. Purple milkweed is a State Threatened taxon in Massachusetts with two extant occurrences. In New Hampshire, Connecticut, and Rhode Island, purple milkweed is listed as a State Historic taxon with no extant populations. The taxon is not reported from Maine or Vermont. Previous records for A. purpurascens on Cape Cod and the Islands are rare with only a few occurrences documented dur- ing the past decade (P. Somers, Massachusetts Natural Heritage and Endangered Species Program, pers. comm.). A single speci- men was collected in 1986 from a dry heathland community on the Miacomet Plains on Nantucket (Sorrie and Dunwiddle 1996). Two early records of A. purpurascens from dry oak forest habitat in West Tisbury and Chilmark are referenced in The Flora of Martha’s Vineyard published by the Martha’s Vineyard Sandplain Restoration Project (1998). Despite repeated surveys performed in conjunction with the New England Plant Conservation Pro- gram (NEPCoP) sponsored by the New England Wild Flower Society in Framingham, Massachusetts, the authors have been unable to relocate a Barnstable County population discovered by 300 1999] New England Note 301 Richard LeBlond in 1989 under a transmission line corridor in Brewster, Massachusetts. This species is not mentioned in ‘‘Notes on the rare flora of Massachusetts’’ (Sorrie 1987). The Barnstable County plants were first observed on July 8, 1998. A single terminal umbel comprised of 24 flowers atop a pubescent peduncle was observed on the taller (SO cm height) specimen. The smaller specimens did not produce inflorescences. Distinguishing features included the bright magenta hoods (7 mm), which turned darker purple-red with age. The erect hoods clearly surpassed the gynostegium and the short, incurved horns. The opposite, elliptical leaves (15 cm) had distinct petioles, trans- verse veins and an acuminate leaf tip. The stem was slightly hairy while the undersurface of the leaf was covered with short, downy hairs. The upper surface of the leaf was glabrous. Associated species in the ground cover included Pyrola rotun- difolia L. var. americana (Sweet) Fern., Gaultheria procumbens L., Vaccinium angustifolium Aiton, and Lycopodium obscurum under an open canopy of Pinus rigida Miller, Quercus alba L., and Q. velutina Lam. A locally rare fern, Osmunda claytoniana L., occurred on the upland slope of the wetland depression inter- mixed with O. cinnamomea L LITERATURE CITED BRUMBACK, W. E. AND L. J. MEHRHOFF, in collaboration with R. W. ENSER, S. C. GAWLER, R. G. Popp, P. SOMERS, AND D. D. SPERDUTO, with assis- tance from W. D. COUNTRYMAN AND C. B. HELLQUIST. 1996. Flora Con- servanda: New England. The New England Plant Conservation Program (NEPCoP) list of plants in need of conservation. Rhodora 98: 233-361. 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. MARTHA’S VINEYARD SANDPLAIN RESTORATION PROJECT. 1998. The Flora of Martha’s Vineyard. The Mary R. Wakeman Conservation Center, Vine- yard Haven, MA. Sorrig, B. A. 1987. Notes on the rare flora of Massachusetts. Rhodora 89: 113-196. Sorrig, B. A. AND P. W. Dunwippie. 1996. The Vascular and Non-Vascular Audubon Society, Massachusetts Natural Heritage and Endangered Spe- cies Program, Nantucket Maria Mitchell Association, and the Nature Conservancy, Nantucket, RHODORA, Vol. 101, No. 907, pp. 302, 1999 NEW ENGLAND NOTE CYPERUS MICROIRIA: A NEW ADDITION TO THE FLORA OF CONNECTICUT TAD M. ZEBRYK 63 Hillside Drive, East Longmeadow, MA 01028 Cyperus microiria Steudel (Cyperaceae). Hartford County: Windsor, Huckleberry Road, waste area at town landfill, moist acidic loamy fine sand, rare, 18 Sep 1998, Zebryk 5760 (NEBC, EIU). Determined by Gordon C. Tucker, Eastern Illinois Univer- sity. While conducting a floristic inventory and rare plant survey at Northwest Park and vicinity for the Town of Windsor, Connec- ticut, an unusual Cyperus was noted among other ruderal species at the adjacent Windsor Town Landfill. Bearing a distinct resem- blance to C. iria L., a somewhat invasive Eurasian species com- mon throughout the southeastern U.S., this unknown specimen was determined to be C. microiria, an annual native to China, Korea, and Japan. Apparently a rare adventive in eastern North America, Cyperus microiria has previously been reported in New York City, Phi- ladelphia, and recently in Kentucky. The collection from Windsor is only the second known from New England (G. Tucker, pers. comm.), the first being from Wayland, Massachusetts (P. Ada- konis s.n., 13 Sep 1979, Wayland, Middlesex County; MASs). The landfill at Windsor serves as a haven and probable point of dispersal for a large number of invasive introduced and native species, including Cyperus esculentus L., Froelichia gracilis (Hook.) Mog., Setaria glauca (L.) P. Beauv., S. geniculata (Lam.) P. Beauv., S. viridis (L.) P. Beauv., Microstegium vimineum (Trin.) A. Camus, and Cirsium arvense (L.) Scop. At the present time, little is known about the potential for Cyperus microiria to be- come a troublesome invasive weed, similar to C. rotundus L. in the South, or the widespread C. esculentus. Certainly the small population at Windsor, if it persists, is worth monitoring to eval- uate the potential impact of this species in our region. 302 RHODORA, Vol. 101, No. 907, pp. 303-305, 1999 BOOK REVIEW Proceedings of a Symposium on the Recovery and Future of the Northeastern Forest, Connecticut College, April 12, 1997 ed- ited by Robert A. Askins and Glenn D. Dryer. 1998. Pub- lished in the journal Northeastern Naturalist, Volume 5, Number 2, pp. 95-172. ISSN 1092-6194 $10.00 (paperback). Published by collaborative effort based at the Humboldt Field Research Institute, Steuben, ME (available from http:/ /maine.maine.edu/~eaglhill; mention this review and receive a 20% discount). On April 12, 1997, the Center for Conservation Biology and Environmental Studies at Connecticut College, New London, sponsored a symposium for land managers, conservationists, and educators in environmental science, designed to assist in the de- termination of whether biological diversity and ecological func- tioning is restorable and sustainable. The seven presenters, a com- mendable ensemble of some of the more notable ecologists in the field today, have produced a symposium of papers which is both thoughtful and informative. The overall viewpoint is one of in- tegrative ecology (the belief that complex systems such as eco- systems must be looked at as a whole, over long periods, to be understood), however the reductionist perspective is also repre- sented, as well as suggestions from environmental economists. The take-home message of this symposium is clearly that northeastern forests need to be evaluated and a balance struck between their ecological and subjective social value, and then proactively managed to foster and protect such states. The point that nature is dynamic and therefore there is no historical state which should serve as our restorative target, is often repeated. Lesser themes of the conference include the need for long-term research, the effects of air pollution on forest ecosystems, and the need to improve the processes by which ecological research is disseminated to managers and policy-makers. The conference ad- mirably wrestles with the coexisting evidence of forests in met- abolic decline, systemically speaking, while many forest species are on the increase. William A. Niering provides a good start with his overview of the natural and anthropogenic agents affecting regional forests in his paper ‘‘Forces that shaped the forests of the northeastern Unit- 303 304 Rhodora [Vol. 101 ed States.”” The impacts of human settlement, by far the predom- inant influence, is recounted from the Indian to the post-colonial period, and current threats such as pollution, introduction of ex- otic species, and fragmentation of forests are cited. David R. Foster and Glenn Motzkin follow by asserting the importance of a historical perspective in the interpretation of for- est landscapes for ecological conservation, management, and de- velopment in “‘Ecology and conservation in the cree landscape of New England: Lessons from nature’s history.” The authors claim that while climate and elevation were the primary forces shaping regional forests in precolonial days, past land-use and edaphic factors exert the reigning influences on vegetative com- position and structure today. They introduce the point that nature has no static ideal; vegetative communities have no record of long-term consistency, nor have regional floras shown any ten- dency to revert to floras of the past. Mankind thus has strongly influenced both past and future forest states. ene E. Likens and Kathleen E Lambert discuss the critical need for sustained research in their paper ‘““The importance of long-term data in addressing regional environmental issues.” They strongly present the danger of using short-term data to eval- uate long-term patterns, the need for safeguarding long-term data and research sites in perpetuity, and recommendations that ex- perimental designs be adaptable to future concerns and not “‘ques- tion-driven.’’ Also advocated is the use of long-term watershed ecosystem research to guide sustainable forestry policy. John M. Skelly takes the road less traveled with his clearly reductionist perspective in “‘A brief assessment of forest health in northeastern United States and southeastern Canada.” He ar- gues that while air pollution may be a forest stressor, there has been no hard evidence that regional forests are in decline. In support of this view are cited two short-term and one ten-year forest agencies study, which found no clear cause-effect relation- ship between the tree mortality and pollution deposition. “The epidemiology of forest decline in eastern deciduous forests,” Orie L. Loucks elegantly and effectively refutes the re- ductionist viewpoint. What begins as a discussion of the appro- priate research framework for the study of complex systems ends in a well-presented scientific argument concluding that air pol- lution has inde en the cause of large-scale tree mortality in the region. Along the way, the integrative versus reductionist ap- 1999] Book Review 305 proaches are discussed, and the systematic symptoms of “forest decline”’ are defined. Dina Franceschi and James R. Kahn present the economist’s perspective in ‘“‘The potential contribution of economics to the recovery of northeastern forests.”’ Their point is that the social benefits of healthy forests need to be determined, and then poli- cies providing economic incentives to landowners and managers need to reflect these goals. As long as private benefits exceed the public good, forests will continue to be developed, fragmented, and polluted. Lastly, John Kricher delivers a summary of sorts with *‘Noth- ing endures but change: Ecology’s newly emerging paradigm,” in which he states that human stewardship of natural systems is essential. The concept of “‘nature’s balance”’ is a western philo- sophical construct which must be dismissed so that proactive goals for ecosystem management can be determined and achieved. Clearly, the northeastern forests of tomorrow are being shaped by the perspectives of land managers, conservationists, and educators of today. This symposium does a commendable job of representing these evolving perspectives. —LESLIE TEELING, Department of Plant Biology, University of New Hampshire, Durham, NH 03824. RHODORA, Vol. 101, No. 907, pp. 306—308, 1999 BOOK REVIEW Discovering the Unknown Landscape: A History of America’s Wetlands by Ann Vileisis. 1997. xii+433 pp. illustrations, photos, appendix, map, index. ISBN 1-55963-314-X $27.50 (cloth). Island Press, Washington, DC. In the preface (p. xi—xii) of Discovering the Unknown Land- scape: A History of America’s Wetlands, author Ann Vileisis states that, ““The matter of wetlands and their conservation is not a matter of science alone—but one of culture as well.” Building upon this thesis, Vileisis examines the interactions between Americans and their neighboring wetlands from colonial to modern times. Throughout the narrative, readers learn how the 221 million acres of wetlands that were once distributed across America were sys- tematically reduced to approximately 102.3 million acres by the 1990s. Efforts to develop and exploit wetland habitats as well as the more recent desire to conserve wetlands are examined within the prevailing political, economic, and environmental trends of erican society. From the harvesting of salt marshes by New England colonists to the levee systems of the United States Army Corps of Engineers, readers learn how various attitudes and poli- cies permanently altered the complexion of the continent. Concepts of wetland science are frequently discussed in the text. However, the objective of the narrative is not to explain details of wetland biology and hydrology, but instead to chart the actions of society as the ecological values of wetlands became better understood. Throughout Discovering the Unknown Land- scape, the complex interactions of government agencies, politi- cians, and private citizens are illustrated. The eighteen chapters have titles such as, ““A Landscape on the Periphery,” “‘Wetlands Portrayed and Envisioned,” “Federal Bulldozers and Draglines,” “The Reagan Agenda Challenges Wetland Gains,” and “The Promise of Restoration.” Each chapter is divided into several subtopics relating different aspects of the chapter theme. Regions that are revisited throughout the book include the Florida Ever- glades, the bottomland swamps of Louisiana, the riparian wet- lands of the Mississippi drainage, the prairie potholes of the upper Great Plains, and the Central Valley of California. Some of the topics discussed include: 306 1999] Book Review 307 * the early perceptions of wetlands, including those of writers and artists * the consequences of the Swamp Land Acts of the mid-1800s contradictory federal policies, some of which promoted wet- land reclamation for agriculture and economic development, while other legislation encouraged wetland conservation the continued degradation of the nation’s wetlands by a swell- ing post-World War II population, and the increasing efforts of conservationists at local, state, and federal levels the Federal Water Pollution Control Act of 1972 (the Clean Water Act) and the debate over the regulation of wetland development through the interpretation of Section 404 * the environmental impacts of the Peagan administration, the “no net loss’’ wetland directive of the Bush administration, and the policies of the Clinton administration as well as the 104th Congress Vileisis has provided a thorough summation of an extensive and complex topic within American environmental history. The narrative is organized well and reads smoothly despite the abun- dance of agency acronyms and references to legislation that po- tentially could make prose cumbersome. As one decade sets the tone for the next, the reader gains an awareness of the historical precedents that have contributed to contemporary wetland issues. Discovering the Unknown Landscape has been extensively re- searched. The 64 pages of notes organized by chapter are a useful resource. The notes contain a wealth of information cited from diverse sources ranging from technical government documents and academic histories to the popular press. A 12-page index, numerous archival photographs and illustrations, and the occa- sional map and graph also enhance the text. The one page appendix entitled “Some Common and Scientific Names of Wetland Plants” consists of a wide spectrum of species with both cosmopolitan (e.g., Phragmites australis and Typha latifolia) and regional (e.g., Avicennia germinans and Cladium jamaicense) distribution patterns. Among this list of plant species there are some editorial problems, including a few misspellings such as Lythrum salarica [sic] and some taxonomic confusion. The scientific name Scirpus acutus includes the author citation when all other species have the author omitted. Spartina pectinata is placed in the list three times under various names; it is cited 308 Rhodora [Vol. 101 > 66 with the common names “‘black grass,” “‘sloughgrass”’ (under the synonym S. michauxiana), and “‘prairie cordgrass.”” The common name prairie cordgrass was paired with the apparently non-exis- tent scientific name S. pectiana (see Kartesz 1994). Regarding other common names, Nymphaea odorata is cited oddly as ‘‘wa- ter lily tuber,’ and “sheep laurel” (Kalmia angustifolia), is er- roneously paired with the scientific name for mountain laurel (K. latifolia), an upland laurel species. Discovering the Unknown Landscape has provided an important service by placing our scientific knowledge of wetland values with- in the framework of our nation’s history. Vileisis (p. 350) con- cludes by stating that, “Informed by history, we can remember the trade-offs already made and turn away from the mistakes and mis- understandings of a time when we knew no better.”” This enhanced perspective is a valuable complement to the wetlands dialogue that too frequently finds human economics and development at odds with environmental concerns. While reading the story of our American wetlands it becomes painfully apparent that as a nation we have not really “‘discovered the unknown landscape.”’ Instead, by using our technological and engineering prowess, we have at- tempted to conquer a vital feature of our natural topography that too often has been viewed as a blight rather than a blessing. Discovering the Unknown Landscape is an informative addi- tion to the literature of the history, politics, and public perceptions of American wetlands. At the conclusion of the first chapter, Vi- leisis (p. 10) observes that, ‘‘Perhaps a newfound awareness of wetlands can inspire and nourish a vision of stewardship for these long-abused and misunderstood landscapes.” Discovering the Unknown Landscape contributes significantly to the ongoing ef- fort to elevate the importance of wetlands in our nation’s con- science. Readers will gain a clear appreciation for the issues that have defined the relationships between American culture and the wetlands that are finally beginning to be appreciated and better understood. For educators, conservationists, consultants, politi- cians, or any reader with a general interest in the history of Amer- ican wetlands, Discovering the Unknown Landscape is a thought- provoking synthesis and a highly recommended resource. LITERATURE CITED Kartesz, J. T. 1994. A Synonymised Checklist of the Vascular Flora of the United States, Canada, and Greenland. Timber Press, Portland, OR. —C. Eric HELLQuIsT, 391 West Road, Adams, MA 01220. RHODORA, Vol. 101, No. 907, pp. 309-310, 1999 BOOK REVIEW Flora of Maine: A Manual for Identification of Native and Nat- uralized Vascular Plants of Maine by Arthur Haines and Thomas FE Vining. 1998. 837 pp. with 10 blank, numbered pages for notes. ISBN 0-9664874-0-0 $45.00 plus shipping (paperback). V. EF Thomas Co., Bar Harbor, ME. Botanists in Maine can claim a number of luminaries over the past two centuries. We begin with John Josselyn, who compiled the first list of plants observed in Scarborough in 1672. Merritt L. Fernald and his colleagues named our state botanical society in honor of Josselyn a few years before joining with others in Cambridge, Massachusetts, to create the New England Botanical Club. The authors of the recently published Flora of Maine, Ar- thur Haines and Thomas F Vining, acknowledge a nearly unbro- ken thread of field-oriented catalogs and listings that have pro- vided information for their work. This latest manual is the first (nearly) complete flora. Two years and two months seem hardly enough time to pro- duce a state flora, yet Haines and Vining have called on years of experience in the field, in university classrooms and laboratories, and amongst a large number of colleagues and advisors to pro- duce a credible and accessible volume. It is an essential volume for the libraries of New England botanists and institutional li- braries everywhere. The most important element of a flora, for this reader, is its keys. Every manual treats a different segment of the plant world defined by a combination of factors, including political geogra- phy, topography, taxonomic group, or the particular interests and needs of readers. Haines and Vining have prepared keys to the vascular plants native or naturalized to Maine, and they warn that such a scope allows them to use characters for identification that might not be appropriate elsewhere. Nevertheless, the keys to families (following Judd et al. 1999) are sound and usable: well written, segmented into logical groups, and based on easily rec- ognizable field characters. Keys to genera and species comprise essential characters wher- ever possible. Complete descriptions of genera and species are not found here as they are in Gray’s Manual (Fernald 1950) or the Manual of Vascular Plants of Northeastern United States and 309 310 Rhodora [Vol. 101 Adjacent Canada (Gleason and Cronquist 1991). Yet the reader can easily build such complete descriptions from the information used to create the keys. Special notes appear for taxa that have a designated rank of rarity following the criteria of the Maine Nat- ural Areas Program. The Flora of Maine should certainly be found on the desk or in the satchel of Maine’s resident and visiting botanists. It may be a model for botanists in neighboring New England states, and perhaps it will inspire the long-overdue revision of our most im- portant regional flora, Gray’s Manual (Fernald 1950). The Uni- versity of Maine should be proud of the work of authors Arthur Haines and Tom Vining. LITERATURE CITED FERNALD, M. L. 1950. Gray’s Manual of Botany, 8" edition (corrected print- ing 1993). Dioscorides Press, Portland, OR. GLEASON, H. A. AND A. CRONQUIST. 1991. Manual of Vascular Plants of featieiern United States and Adjacent Canada. The New York Botan- k Jupp, W. S., C. S. CAMPBELL, E. A. KELLOGG, AND P. S. STEVENS. 1999. Plant Systematics: A Phylogenetic Approach. Sinauer Associates, Inc., Sun- derland, MA. —W. DONALD Hupson, Jr., Chewonki Foundation, Wiscasset, ME 04578-4822. RHODORA, Vol. 101, No. 907, pp. 311-317, 1999 NEBC MEETING NEWS March 1999, Thomas J. Rawinski, Director of Ecological Man- agement at Massachusetts Audubon’s Center for Biological Con- servation, addressed the Club on the topic ‘Travels through Vir- ginia: Botanical Wonders and Conservation Victories.”’ It was a partial accounting of his seven years away from New England, or in his words, ‘‘a report back to the hive,”’ regarding his em- ployment as an ecologist with the Virginia Division of Natural Heritage, part of the state’s Department of Conservation and Rec- reation. Using slide images and recounting many botanical dis- coveries and new natural areas protected as a result of the Divi- sion’s efforts, he gave us an overview of the state’s natural areas and botanically diverse ecosystems. He reminded us of how much L. Fernald had loved Virginia, noting that Fernald added more new species to his 8th edition of Gray’s Manual of Botany from Virginia than from Newfoundland, another area he had explored extensively. Contrasting Virginia with New England, Rawinski said Virgi- nia has only two natural lakes, lacks some northern taxa such as Chamaedaphne, and is nearly devoid of paper and gray birch. As evidence of a richer flora than New England, he noted that there were 25 oaks, 8 pines, 6 magnolias, 25 Rhynchospora spp., 13 Vaccinium spp., and 10 Trillium spp., plus representation by fam- ilies such as the Bromeliaceae and Loganiaceae. Virginia endem- ics include three species of Clematis: C. addisonii, which occurs on dolomite; C. coactilis, a species found on both dolomite and shale; and C. viticaulis, a narrow endemic found only on shale barrens. Other endemics include Helenium virginicum, a species of acid ponds in the Shenandoah Valley, and /liamna corei, a species for which they found fire worked well as a management tool. The clearly defined physiographic provinces of the state helped Rawinski orient us to the state’s major geological and climatic regions and to the locations of unique habitats for plants within them. Elevations range from the 5700 ft. high Mt. Rogers in the Blue Ridge Mountains in southwestern Virginia to sea level where one can find extensive tidal marshes. He compared the Piedmont of Virginia to Worcester County, Massachusetts, point- ing out that neither is diverse in habitat types, but that each pos- sesses some interesting ones, such as those with diabase bedrock. 311 312 Rhodora [Vol. 101 Botanical hot spots are clustered in habitats such as shale barrens, sea-level fens, and dolomite outcrops: types found primarily in the eastern or western parts of the state. Using habitat character- istics as primary indicators, he and colleagues located 30 new state records, such as Carex arctata and Sporobolus heterolepis, between 1990 and 1997. Recent funding of 11.5 million dollars through a bond bill has allowed the Department to add over 20 new natural areas to its preserve system. Rawinski highlighted certain ecosystems. More than 500 po- tential shale barrens were identified by the Heritage Division; they typically occur on steep hillsides undercut by streams. An abundance of limestone in the Shenandoah Valley and elsewhere has resulted in some notable dolomitic cliffs and, in the south- western corner of the state, some calcareous glades with taxa such as the newly described clover species, Trifolium calcaricum. Do- lomite glades supported the globally rare Echinacea laevigata, which can be found growing in loamy, dolomitic soil with prairie taxa such as Castilleja coccinea and Senecio plattensis. A priority for protection in Rawinski’s eyes was an ultramafic barren (i.e., a serpentine-like area with high magnesium levels) in the Pied- mont supporting several disjunct and rare species. Buffalo Moun- tain, a monadnock in the Blue Ridge, supported a diversity of vegetation associations and herbaceous species, including nine rare species for the state. Rawinski postulated that the thin soil mantle and open glade habitat is maintained by a natural defi- ciency of clay, which facilitates lateral movement of water and washing of any deposited soil. Ultramafic fens are another botan- ical hot spot in Virginia, providing habitat for 20 state-listed rare species, including a state record discovery: Tofieldia glutinosa. A truly significant area in the Piedmont is Fort Pickett Military Reservation, where frequent fires have maintained a population of the very rare sumac, Rhus michauxii, it occupies hundreds of acres, making it by far the largest known population. The Coastal Plain, as in New England, has many ponds, but in Virginia they are sinkhole depressions over 100,000 years old. Here one can find large overcup oaks, Quercus lyrata, and rare herbs such as Carex joorii, Hottonia inflata, Sabatia campanu- lata, and Chelone cuthbertii. In an especially dry year, ten new Fimbristylis perpusilla records showed up at pond sites. Other coastal plain communities of special interest are sand hills with longleaf pine, pocosins or shrub bogs, sea level fens, and cypress- 1999] NEBC Meeting News 313 tupelo swamps, where — big-eared bats can be found in cavities of the ancient tree For more information on Mirginta natural areas and biota, Tom Rawinski advised seeking out the Virginia Department of Con- servation and Recreation’s web site. April 1999, Michael Donoghue introduced the ‘“‘NEBC 1999 Distinguished Speaker,”’ Dr. Peter Raven, Director of the Missouri Botanical Garden. Accomplishments and qualities mentioned in- cluded (1) his ability to organize and galvanize people around ideas and visions, (2) his authorship of The Biology of Plants, (3) his collaborative research on the Onagraceae as a model for systematic botany, (4) his leadership in coordinating the Flora of China project, and (5) his leadership in conservation of biological diversity, including serving on the President’s Commission on Science wa Technology and co-authoring a paper entitled ““Teamin Life,’ a statement on the need and mechanisms for preserving biodiversity. Dr. Raven addressed the topic, “Plant Conservation Globally and Locally.”” He explained that his approach would be to paint the broadest picture possible of the current crisis in biological conservation around the world and why the crisis exists. His main objective was to stimulate our thinking about these issues and invite a dialogue about strategies for combating the problem. Ra- ven’s broadest picture included a review of 3.8 billion years of biological evolution on the earth and the five major extinction events that have influenced its pathways to the present. The first three extinction events occurred when life was restricted to the marine world. He pointed out that terrestrial life began 430 mil- lion YBP, at a time equivalent to 90% of the way through the time-line of earth’s existence. He emphasized the importance of cyanobacteria in making colonization of terrestrial habitats pos- sible by changing the earth’s atmosphere to an oxidizing one. The resulting increase in oxygen produced by their photosynthetic ac- tivities over 3 billion years made possible the production of a stratospheric ozone layer that allowed the ancestors of the four groups dominant on land at the present time (arthropods, fungi, terrestrial vertebrates, and plants) to colonize terrestrial habitats. The fourth great extinction event occurred at the end of the Perm- ian, about 280 YBP, impacting the earth’s first forests and early dinosaurs. In the following Mesozoic Era, dinosaurs and cycads 314 Rhodora [Vol. 101 flourished and angiosperms evolved, making life much more di- verse than previously. About million YBP, at the end of the Cretaceous Period, the fifth great extinction occurred, presumably as the result of a large meteorite crashing into the earth off what is now the Yu- catan Peninsula. The collision created an opaque cloud around the globe that impeded photosynthesis and, according to estimates by David Raup et al., eliminated two-thirds of terrestrial species in a short period of time. At that point, Raven estimated loosely that the number of eucaryotic organisms remaining may have numbered between 500,000—700,000. It took approximately ten million years for life to recover, and the resulting evolutionary pathways led to the evolution of most current groups of organ- isms. Today, according to a 1997 paper by Sir Robert May pre- sented at the National Forum on Biodiversity at the National Academy of Sciences, the number of eucaryotic species can be estimated conservatively at about seven million. Of these, only about one in four has a valid name. In the tropics, the ratio is much less, around one in twenty. Even for the described species of organisms our knowledge is extremely limited; many are known only from a single specimen at the bottom of a museum vial. No one can give a plausible estimate of the number of pro- caryotic organisms. One gram of soil in a Norwegian beech forest is estimated to have 5,500 species of bacteria, more than the total number of species recognized formally from the entire world, and how these figures relate to other ecosystems around the world, or to the total number of bacterial species, is unknown. Even more poorly known, Raven says, is the multitude of relationships that mediate the flow of energy through the globe’s ecosystems and other aspects of their functioning. Bringing humans into the picture, Raven drew attention to our Homo erectus ancestors who migrated out of Africa and discov- ered fire-making, causing some low-scale disturbances approxi- mately 1.5—2 million YBP. Homo sapiens appeared on earth about 200,000 YBP and arrived in the New World 14,000—16,000 YBP. The cultivation of crops began at about 10,000 YBP at a number of widely-scattered centers, when there may have been only sev- eral million people globally, a population equivalent to that of the Greater Boston area today. Agriculture allowed for a reliable food source, and by the time of Christ the earth’s population had grown to around 120—150 million. By the Renaissance, it had 1999] NEBC Meeting News ats grown to 0.5 billion, and by the time of Thomas Malthus, who speculated at the start of the Industrial Revolution that human populations would outstrip agriculture’s ability to produce food, about 1 billion people were walking the earth. By 1950, our num- bers had increased to 2.5 billion, and now 50 years later, the earth’s population has reached nearly 6 billion, bringing even greater importance to questions of sustainability. *‘Sustainability and biodiversity are two sides of a coin,’’ Ra- ven said. Can the human world sustain itself while maintaining global biological diversity? The results of the last 50 years are not encouraging. One-fourth of the world’s topsoil has been lost and one-third of the world’s forests have been destroyed since World War II. There has been a drastic increase in extinction rates. It is estimated that extinction rates have increased 100 X since the Renaissance and are 1000 X background rates, based on fossil records. The rates are accelerating, according to extinc- tion models developed by Stuart Pimm et al. using island bio- geography theory, and may reach 10,000 X in the next century. Over the next 25 years, they estimate that one-third of tropical organisms will become extinct or be on the way to extinction, and that by the end of the next century, three-fourths of all trop- ical organisms and two-thirds of the worldwide total will be gone or on the way to extinction. Homo sapiens, Raven says, is driving an extinction event comparable to the scope of the fifth major extinction event 65 million years ago. Nothing could be as dam- aging to the future of our species, primarily because of our direct dependency on other living organisms. The 350,000 species of photosynthetic organisms—plants, algae, and a few bacteria—are responsible for all productivity on earth, and human beings obtain all their food from plants directly and indirectly. Another anthro- pocentric reason for preserving biodiversity is that three-fourths of humanity depends directly on biodiversity, mostly plants, for its pharmacopoeia while the remaining quarter go to drug stores and derive such benefits indirectly. Why say the 21st century will be the “Age of Biology,”’ Raven asks, if all of the organisms are going down the drain? Where will we get all of the organisms to create new products or to use in new sustainable systems, or for that matter to use in bio-engineering new biological products or life forms? He cautions us to heed Aldo Leopold’s advice and follow the “‘first rule of intelligent tinkering” by ‘saving all of the cogs and wheels.” 316 Rhodora [Vol. 101 Raven asks, ‘“‘What things are in the way of fixing the prob- lem?’ One is the sovereignty and pride of nations that inhibit our ability to pay attention to one another internationally and establish global solutions. He cites the United States’ failure to sign the ‘International Convention on Biological Diversity’ as a prime example. A second appears to be greed and waste by the devel- oped world, because Raven described the industrialized nations as possessing 20% of the world’s population and 85% of the world’s wealth, while creating 80—90% of its pollution. In the world today, 360 billionaires have wealth equal to what two bil- lion of the poorest people earn in a year. The people at the low end of the economic spectrum are totally disenfranchised, which denies the rest of the world the potential benefit of their wisdom and creativity. We simplify when we say “‘over-population”’ is the problem globally. For instance, Brazil and Mexico have had population control policies for 20 years and have slowed popu- lation growth and improved education for women, whereas the U.S. has no population policy, despite a doubling of its population in the past 50 years and a standard of living 30—40 times that of Brazil or Indonesia. Because of our consumptive standard of liv- ing, the impact on the environment of doubling the U.S. popu- lation (adding 135 million people in 50 years) is equivalent to adding 4 billion people to populations in Brazil or Indonesia. We do not need to waste at the levels we do. The United States wastes twice as much per capita as in Europe. We are living as if there is no tomorrow and in denial that our economy is related to the rest of the world. What can we do to help create a more sustainable world and slow the loss of biological diversity? Ra- ven’s answers include: (1) be leaders in showing the way rather than demanding that the third world lead the way, (2) pay atten- tion to internationalism by convincing others that the 80% of the world’s people living in non-industrialized countries are of deep and profound importance to us, (3) vote and encourage others to do so, (4) support conservation groups you are congenial with Se aie and then support sustainability, and (5) think t what you are doing personally and make wise choices. Pa says, “Think about what is just, what is stable, what is sustainable. Think about a world within which biological diver- sity, cultural diversity, beauty, music, poetry, philosophy, litera- ture, and all the things we value and cherish can coexist. What do we do to create a world that will maintain these things?” We 1999] NEBC Meeting News a7 can be optimistic or pessimistic amidst the ‘gloom and doom” that we face. “Yes, the world will become more homogeneous and less diverse,” Raven acknowledges. But the future depends on how we live now, he says. “If you want to be optimistic, do so because of your own determination to do something about it,” says our 1999 saci pala Speaker, as he leads the Club for- ward into its second century —PAuL Somers, Recording Secretary ANNOUNCEMENT NEW ENGLAND BOTANICAL CLUB GRADUATE STUDENT RESEARCH AWARD The New England Botanical Club will offer $2,000 in support of botanical research to be conducted by graduate students in 2000. This award is made annually to stimulate and encourage botanical research on the New England flora, and to make pos- sible visits to the New England region by those who would not otherwise be able to do so. It is anticipated that two awards will be given, although the actual number and amount of awards will depend on the proposals received. The award will be given to the graduate student submitting the best research proposal dealing with systematic botany, biosyste- matics, plant ecology, or plant conservation biology. Papers based on the research funded must acknowledge the NEBC’s support. Submission of manuscripts to the Club’s journal, Rhodora, is strongly encouraged. Applicants must submit three copies of each of the following: a proposal of no more than three double-spaced pages, a budget, and a curriculum vitae. Two letters in support of the proposed research, one from the student’s thesis advisor, should be sent directly to the Awards Committee by sponsors. All materials should be sent to: Awards Committee, The New England Botan- ical Club, 22 Divinity Avenue, Cambridge, MA 02138-2020. Pro- posals and supporting letters must be received no later than March 1, 2000. The recipient(s) will be notified by April 30, 2000. Two Graduate Student Research Awards were given in 1999. Joel Gerwin of the University of Massachusetts at Boston re- ceived support for his proposal entitled “‘Long-term effects of forest fragmentation on genetic diversity of red oak (Quercus rubra L.): A comparison of old-growth and secondary forests.” Also chosen for an award was Julie Ellis of Brown University, whose proposal was entitled ‘“The role of nesting seabirds in structuring New England coastal plant communities.” 318 ANNOUNCEMENT HUMBOLDT FIELD RESEARCH INSTITUTE PRIZE The New England Botanical Club is offering a prize for the best essay of 500 words or fewer on the theme ‘‘How I would benefit from an Eagle Hill Seminar.’”’ This prize is offered in conjunction with the Humboldt Field Research Institute, and con- sists of tuition for a week-long Eagle Hill Field Seminar or Work- shop in Steuben, Maine. In return for a donation from the Club of a set of Rhodora back-issues, the Institute has reciprocated with the donation to the Club of this prize. The contest is open to any member of the NEBC. Please submit three copies of your essay before December 31, 1999. Essays should be mailed to: The New England Botanical Club, ATTN: Eagle Hill Essay, 22 Divinity Avenue, Cambridge, MA 02138. The winner will be notified in advance and announced at the March, 2000, Annual Meeting. 219 INFORMATION FOR CONTRIBUTORS TO RHODORA Submission of a manuscript implies it is not being considered for publication simultaneously elsewhere, either in whole or in part. GENERAL: Manuscripts should be submitted in triplicate. The text must be double-spaced throughout, including tables, figure legends, and literature citations. Use a non-proportional font throughout and do not justify the right margin. Do not indicate the style of type through the use of capitals, underscoring, or bold, except for names of genera and species which should be in italics or underscored throughout. Do not underline punctuation. All pages should be num- bered in the upper right-hand corner. For guidance in matters not addressed here, consult the editorial office by phone at (603) 862- 3205, FAX (603) 862-4757, or e-mail: janets@christa.unh.edu. Brev- ity is urged for all submissions. Submit manuscripts to the Editor-in- Chief. TITLE, AUTHOR(S), AND ADDRESS(ES): Center title, in capital letters. Omit authors of scientific names. Below title, include au- thor(s) name(s), affiliation(s), and address(es). If “current address” is different, it should follow immediately below, not as a footnote. ABSTRACT: An abstract and a list of key words should be included with each paper, except for shorter papers submitted as Notes. An abstract must be one paragraph, and should not include literature citations or taxonomic authorities. Please be concise, while including information about the paper’s intent, materials and methods, results, and significance of findings. TEXT: Main headings are all capital letters and centered on one line. Examples are: MATERIALS AND METHODS, RESULTS, and DIS- CUSSION. Do not title the Introduction. Do not combine sections of the paper (such as Results and Discussion), or use Conclusions or Summary. Second level headings should be indented, bold, upper and lower case, and end with a period. Taxonomic authorities should be cited for all species names at their first usage in the text, or in a referenced table. Cite each figure and table in the text in numerical der. Each reference cited in the text must be in the Literature Cited. Cross-check spelling of author(s) name(s) and dates of publication. Literature citations in the text should be as follows: Hill (1982) or (Hill 1982). For two or more authors, cite as follows: Angelo and Boufford (1996) or (Angelo and Boufford 1996). Cite several refer- ences alphabetically by first author, rather than chronologically. With- in parentheses, use a semicolon to separate different types of citations (Hill 1982; Angelo and Boufford 1996) or (Figure 4; Table 2). FLORAS AND TAXONOMIC TREATMENTS: Specimen citation should be selected critically, especially for common species of broad 320 INFORMATION FOR CONTRIBUTORS 321 distribution. Keys and synonymy for systematic revisions should be taxon should carry a Latin diagnosis (rather than a full Latin descrip- tion), which sets forth succinctly how the new taxon differs from its congeners. LITERATURE CITED: All bibliographic entries must be cited in the paper, unless a special exception has been made by the Editor (such papers will be allowed a REFERENCES section). Verify all entries against original sources, paying special attention to spelling and de- tails of publication. Cite references in strict alphabetical order by first author’s surname. Do not write authors’ names in all capital letters. References by a single author precede multi-authored works of same senior author, regardless of date. Use a long dash when the author(s) is the same as in the entry immediately preceding (see recent issues). Refer to Botanico-Periodicum-Huntianum (B-P-H 1968) and B-P-H/ Supplement (1991) for standardized abbreviations for journals. TABLES: Tables must be double-spaced. Tables may be continued on an extra page, if necessary. As much as possible, the title should be self-explanatory. Do not use footnotes; instead, add notes after the end of the table title. Broadside tables should be avoided, if pos- sible. Each table should be cited in the text in numerical order. FIGURES: Illustrations must be either black and white half-tones (photographs), drawings, or graphs. Illustrations must be camera- ready; flaws cannot be corrected by the Editor or the printer. Add symbols or shading with press-on sheets. The printed plate will be 4 X 6 inches; be sure that illustrations are proportioned to reduce correctly. Allow space for a caption, if possible. Magnification/re- duction values should be calculated to reflect the actual printed size. Maps must indicate scale and compass direction. The double-spaced list of legends for figures should be provided on a separate page. Each figure should be cited in the text in numerical order. THE NEW ENGLAND BOTANICAL CLUB 22 Divinity Avenue Cambridge, MA 02138 The New England Botanical Club is a nonprofit organization that promotes the study of plants of North America, especially the flora of New England and adjacent areas. The Club holds regular meetings, and has a large herbarium of New England plants and a library. It publishes a quarterly journal, RHO- DORA, which is now in its 101st year and contains about 400 pages per volume. Visit our web site at http://www.herbaria. harvard.edu/nebc Membership is open to all persons interested in systematics and field botany. Annual dues are $35.00, including a subscrip- tion to RHODORA. Members living within about 200 miles of Boston receive notices of the Club meetings. To join, please fill out this membership application and send with enclosed dues to the above address. Regular Member 35.00 Family Rate $45.00 Student Member $25.00 For this calendar year For the next calendar year Name Address City & State Zip Phone FAX email Special interests (optional): ia Elected Officers and Council Members for 1999—2000: President: David S. Conant, Department of Natural Sciences, Lyndon State College, Lyndonville, VT 05851 Vice-President (and Program Chair): Lisa A. Standley, Vanasse Hangen Brustlin, Inc., 101 Walnut St., P.O. Box 9151, Wa- tertown, MA 02272 Corresponding Secretary: Nancy M. Eyster-Smith, Department of Natural Sciences, Bentley College, Waltham, MA 02154- 4705 Treasurer: Harold G. Brotzman, Box 9092, Department of Bi- ology, Massachusetts College of Liberal Arts, North Adams, MA 01247-4100 Recording Secretary: Paul Som Curator of Vascular Plants: a Angelo Assistant Curator of Vascular Plants: Pamela B. Weatherbee Curator of Nonvascular Plants: Anna M Reid Librarian: Leslie J. Mehrhoff Councillors: W. Donald Hudson, Jr. (Past President) Arthur V. Gilman 2000 Karen B. Searcy 2001 David Lovejoy 2002 Jennifer Forman (Graduate Student Member) 2000 Appointed — avid E. Boufford, Associate Curator ae R. Sullivan, Editor-in-Chief, Rhodora