Dodota JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL RALPH CARLETON BEAN ROBERT CRICHTON FOSTER ROLLA MILTON TRYON ? Associate Editors RADCLIFFE BARNES PIKE | LORIN IVES NEVLING, JR. ELIZABETH ANNE SHAW 2 VOLUME 72 1970 Che Nem England 3 =| totanical Club, Jur. XY Botanical Museum, Oxford St., Cambridge 38, Mass. APR 21 1970 PanLow KMawAENUER Kwenees Dodova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL 3 RALPH CARLETON BEAN ROBERT CRICHTON FOSTER ROLLA MILTON TRYON P Editors RADCLIFFE BARNES PIKE LORIN IVES NEVLING, JR. ELIZABETH ANNE SHAW Vol. 72 January-March, 1970 No. 789 CONTENTS: Western Cape Cod: Plant Notes SONNO E. DUM ansecthsithrisededestto tractabat ci eriam P cmn 1 Additions and Nomenclatural Changes in the Flora of Southern Florida — I. Robert W. Long ................................ 17 Notes from the Pringle Herbarium III P. CU BOOT srrerssecesessererssccessensnecensssestaceniabessnunntacittssssnetsantanban 47 Infraspecific Classification in the Carolina Flora Nobert- D o Wglbupu n u u... uu... S SSS SD S s s 51 New Phanerogams from the Arid Neotropics Jo, O, DIGPNGD ue sascssieaspeceosssoesassssessesashuzessossk Ras e 66 A New Species of Calamovilfa (Gramineae) from North America. Ken E. Rogers ...................................... a... 72 (Continued on Inside Cover) The New England Botanical Club, Ine. Botanical Museum, Oxford St, Cambridge, Mass. 02138 CONTENTS: — continued Saurauia Species and their Chromosomes Djaja D. Soejamrto ......... eene nennen nnn 81 Additional Chromosome Numbers in Brazilian Compositae James R. Coleman ........................... eene ener eene nana enn 94 Studies in the Compositae-Eupatorieae, XV. Jaliscoa, Mac- vaughiella, Oaxacania, and Planaltoa R. M. King and H. Robinson... eene 100 The Genus Acanthospermum (Compositae-Heliantheae-Mel- ampodinae): Taxonomic Changes and Generic Affinities Tod F. Stuessy v.cccccccccssccscessessseesscesseessessecsssesecsuessceseccesseasecseees 106 Nomenclature of the Lupinus argenteus and L. caudatus Complexes. Lloyd W. Hess and David B. Dunn ................ 110 Flora of the Wolf Islands Part III: The Marine Algae Edward J. Hehre, Joan R. Conway and Richard A. Stone 115 A Synopsis of the Genus Sessenguthia (Acanthaceae) Dieter C. Wasshausen |... eene 119 Stuart Kimball Harris Ralph C. Beam .........1........... ananassa aa 127 Bayard Long (1885-1969) John M. Fogg, Jr. cesccssssscsessecesscscsscscessesccsscsesacsccstscescssesevsccavece 130 Book Review A. R. Hodgdon ................. eee ees 137 Spiranthes tuberosa, New for Kansas Lawrence K. Magrath ....... eene 141 White Phase in Flower Development in Cypripedium acaule J. T. Baldwin, Jr. essent 142 Rhodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 72 January-March, 1970 No. 789 WESTERN CAPE COD: PLANT NOTES HENRY K. SVENSON I am not a stranger to Cape Cod. My father came to visit the Cape first about 1885, and from then on made frequent trips from Boston by train in the summers. There- fore as a small child I remember seeing, in many visits, lupine and the Deptford pink in the lichen-covered fields at East Sandwich, which were then fairly open. They are now grown up into pitch pine, and everywhere pastures and hayfields have practically disappeared. The forest has greatly increased, due also to the lessening of forest fires, and now appears mainly as an expanse of oak and pitch pine occupying the terminal moraine, the backbone of the Cape. From 1917 on, the botanical trips led by Professor Fernald to the Cape were always enjoyable and instructive occasions. Summers from 1923 to 1926, while at the Marine Biological Laboratory at Woods Hole, Dr. John M. Fogg, Jr. and I made many botanical trips into the surrounding country- side; and again in 1928, Professor Fernald and I explored some of the relatively little-known parts of western Cape Cod. Many things have changed. Some of the old roads are now blocked off, and I have been unable to locate the Chamaecyparis swamp in East Falmouth where we found Rhododendron canadense and Aster nemoralis. And I can- not find the interesting spot reported by me in Rhodora 30: 135 (1928), where Lycopodium lucidulum, L. clavatum, and 1 2 Rhodora [Vol. 72 associated plants grew. Some of these places I suspect have been covered up by the Otis Air Field. Since retiring to Cape Cod in early 1967, I have covered most of the old roads in western Cape Cod, to a large extent on foot, since many are no longer passable by automobile. The result is a fairly large number of plants scarce or pre- viously unknown on Cape Cod. Those of which I can find no previous records from literature or from the extensive collections of the New England Botanical Club, I have marked by an asterisk. Specimens have been put in the herbarium of the New England Botanical Club. In 1918 and 1919 Professor Fernald and Bayard Long found an interesting locality for plants at “Spring Hill", Sandwich, After a considerable search I found the locality, which is in East Sandwich. The beautiful maple and Chamaecyparis swamp is still intact. The sloping pastures have disappeared, and in many places have been replaced by a fearful tangle of Smilax (S. rotundifolia and S. glauca), called "catbrier" in Gray's Manual, but known more appropriately by the local inhabitants as *bullbrier" and *horsebrier". Intact also are the stands of yellow birch ( Betula lutea), the spring-heads still have Carex scabrata, and on an adjacent knoll is native Fraxinus americana, as Fernald found it, along with Carex laxiflora, Carex laxicul- mis, and C. convulata, as well as Ranunculus abortivus and R. recurvatus; and in 1967 there was a single flower of Erigeron pulchellus, which Professor Fernald’s keen eye had recognized fifty years ago on the basis of only a single leaf, The small species of Botrychium (B. matricariaejolium and P. lanceolatum) I could not find; neither was there any trace of Ophioglossum, or of the silvery spleenwort, or of Trillium in flower. The presence of this little knoll with its fragment of rich wood's vegetation is hard to explain. Scattered along the moist north slope of the terminal moraine, and at the margins of some ponds in the interior are small “forests” of beech, hornbeam (Ostrya virgin- iana), sweet birch, paper birch, flowering dogwood, hick- ories (Carya glabra, C. ovalis, and C. tomentosa), white 9 1970] Cape Cod — Svenson 3 pine, and red oak. There is much resemblance to the flora of the northern part of Long Island, New York, where these species are also present. Likewise there is a single locality for hemlock. But the relative lack of mountain laurel on Cape Cod is a noteworthy difference. Most of the ponds on Cape Cod, even the little ones, are in the process of becoming “lakes” bordered by house lots. Some of the names have changed from those on the old maps: Great or Nine-mile Pond in Barnstable (Centre- ville) is now Lake Wequaquet; Halfway Pond in Barn- stable (Hyannis) is now Mary Dunn Pond. The political divisions are confusing, too. All the Cape belongs to Barn- stable County, The village of Barnstable is on the north shore. The Town of Barnstable includes the villages of Hy- annis (now a metropolis), Centreville, Osterville, Cotuit, Marstons Mills, and Santuit. Western Cape Cod, commonly known as the “Upper Cape”, includes the separate towns of Barnstable, Sandwich, Mashpee, Falmouth (with Woods Hole and Waquoit), and Bourne (with Sagamore and Po- casset). Isoetes Tuckermani A. Br. Lovells Pond, Santuit (no. 1468). This is the common quillwort of Cape Cod ponds; it grows submerged in several feet of water. The megaspores average 520 u, with a fragile jagged-crested honeycomb sur- face. It has been collected previously in Sandwich, Fal- mouth, Harwich, Brewster, and Wellfleet. Ophioglossum vulgatum L. var. pseudopodum (Blake) Farwell. Thicket along east side of Santuit River, Cotuit (no. 811). The adder's-tongue fern has been known from Sandwich, Yarmouth, Dennis, Chatham, and Truro. Osmunda Claytoniana L. Thicket along Sandwich-Cotuit Road near Wakeby Pond, Sandwich (no, 1483) ; along rail- road near County Road, East Sandwich (no. 2046). I have seen it also along Discovery Hill Road in East Sandwich. In the New England Botanical Club Herbarium it is repre- sented from Falmouth; from near Peter’s Pond in Mashpee; and from the border of the maple swamp at “Spring Hill, 1 Rhodora [Vol. 72 Sandwich". It occurs only on “Upper” or western Cape Cod (See Fogg, Rhodora 32: 176 (1930). Botrychium virginianum (L.) Sw. A single plant (fruit- ing) on a hummock beneath Chamaecyparis, growing with marsh fern, goldenrod, and Mikania at the edge of a brack- ish marsh (no. 661), This is a most unusual habitat for the rattlesnake fern, which would be hidden in a tangle of Mikania later in the season; it is known from Cape Cod only on the basis of a leaf fragment collected by Fernald and Long at “Spring Hill" in 1918. Polypodium virginianun L. The polypody fern grows on several north-facing gravel banks on Cape Cod, just as it does on the coastal plain around Chesap:ake Bay in Mary- land. Usually I have found it on Cape Cod in groves of Ostrya where the ground slopes steeply to a pond, for ex- ample, the western side of Mashpee Pond where it is abun- dant in several large patches (no. 1798) ; Peters Pond, in South Sandwich (no. 1709); Lovell’s Pond, Santuit. It is known from Bourne, Barnstable, Sandwich (near Wakeby Pond), Brewster, and Providence. In Brewster it was noted on a stone wall by Hunnewell and Blake, and on a large boulder in Naushon by Fogg, Rhodora 32: 227 (1930). The European P. vulgare I saw growing on gravel banks facing the sea near Cuxhaven, Germany, in 1958, Polystichum acrostichoides (Michx.) Schott, The Christ- mas fern, rare on Cape Cod, is limited by Fogg (l.c) to the Upper Cape, and known only from Barnstable Village, South Sandwich, and “Spring Hill’. It also grows near Discovery Hill Road in East Sandwich (no. 668), and I have seen a single large plant in woods near Amos Pond in Mashpee. *Juniperus communis (L.) var. depressa Pursh, was re- ported from the north shore of Naushon by Fogg, where possibly introduced. It is apparently native in a sandy field on the east side of Wequaquet Lake, Centreville (no. 1325), and on a sandy roadside in West Barnstable (no. 1959), along route 6. The nearest mainland location for this plant 1970] Cape Cod — Svenson 5 seems to be near the old Darby railroad station in Plymouth, where it was collected by Sanford in 1911. “Tsuga canadensis (L.) Carr, About twenty trees on a steep bank of an island in Mystice Pond, Marstons Mills (no. 1086). The trees are considered native by all the local inhabitants; there is no reason to believe otherwise. Some are two feet or more in diameter ; several have had the tops blown off in recent hurricanes, and are branched from the lower part of the trunk. The nearest stands of hemlock ap- pear to be at the Plympton-Kingston line in Plymouth County. A single large weatherbeaten tree may be seen along rte. 3, opposite Bloody Pond in southern Plymouth, about halfway between the Cape Cod and Kingston local- ities. *Panicum amarum Ell. Margin of sand dunes, marsh side, Sandy Neck, W. Barnstable (no. 1730). Only two clumps were seen; one greenish-flowered, the other pur- plish. They grew side by side. The plant is unknown north of New Haven, Conn. It is readily distinguished from Panicum virgatum by the decumbent habit, narrow inflores- cense, and large spikelets, in this collection 6.0 mm long. The fresh anthers are lucid brown, glutinous, and 2.4-2.5 mm long in the purplish phase; 0.8-1.1 mm long in the green phase. Panicum mattamuskeetense Ashe. It is a tall plant of the southern coastal plain which reaches its northern limit on Cape Cod, where it has been known from Falmouth, Dennis, Brewster and Harwich. It was abundant on damp sandy cart roads in West Yarmouth just over the Hyannis line (no. 1508), the leaves bluish-green when fresh; spike- lets 2.3 X 1.1-1.2 mm, and the ligule 1-1.5 mm, though stated by Hitchcock to be 0.7 mm long, by Gleason 1 mm or more long, and by Fernald *less than 1 mm long." In a second locality, a sphagnous depression just north of rte 28 in Centreville (no. 1684), the spikelets were 2.0-2.1 mm long and 1.0 mm wide, and the ligule 0.5 mm long ; evidently a small-fruited phase of P. mattamuskeetense, though in some respects the habit approached P. spretum, A third 6 Rhodora [Vol. 72 collection (no. 1432), from plants abundant in the exten- sive boggy area, now being drained, west of Higgins-Cro- well Road, West Yarmouth (and a continuation of the locality from which no. 1508 was obtained), the upper as well as the lower leaves are somewhat pubescent, and the plants approach P. annulum Ashe, which is considered by Fernald as perhaps not distinct from P. mattamuskectense. The spikelets in no. 1432 are 2.5-3.0 mm long, the ligules are mainly 0.5-0.6 mm long, but may be as much as 1 mm long in the lower leaves. Typical P. annulum, with some- what velvety leaves and large spikelets is known from only a single collection on Cape Cod: cranberry-bog roads east of Slough Road, Harwich, Fernald in 1918. My fourth col- lection (no. 1464; Aug. 1, 1968) came from the gravelly bed of a brook in Marstons Mills, where it was growing with Xyris torta and X. caroliniana. It had the erect growth of P. mattamuskeetense, pubescent spikelets averaging 1.7 X 1.0 mm and ligules 0.5-0.6 mm long. The small spikelets would tend to place the collection in P. micro- carpum Muhl., but the spikelets of that species are de- scribed as “glabrous or rarely minutely pubescent" by Hitchcock & Chase; and in the 8th edition of the Manual as "glabrous (very rarely puberulent), 1.5-1.8 mm long, 0.7 mm broad". Panicum microcarpum, in which the au- tumnal form is “densely and intricately branched", is known from five collections on the Cape, ranging from Woods Hole to Orleans. I have seen it in Sandwich and Centreville, growing on moist grassy slopes. By Fogg, Rhodora 32: 211 (1930) it is considered characteristic of the “Middle Cape". The members of this Panicum-group are still puz- zling. Festuca obtusa Biehler. This grass is frequent in moist rich woods in the Boston district, but known only from Sandwich and Chilmark (Marthas Vineyard) in the south- east, In Sandwich it grows on a north-facing slope at Shawme Pond. It is now to be reported from the beech woods at the Cape Cod Museum of Natural History, Brew- ster (no. 2146). 1970] Cape Cod — Svenson T * Molinia caerulea (L.) Moench. In the report on grasses of the Flora of Massachusetts, Rhodora 49: 264 (1947), the “moor-grass” is reported only from waste land in South Boston. This handsome grass is growing well, and appar- ently spreading, on a sandy border of Forestdale Road in Mashpee (no, 1262; Nov. 1, 1967). It is common in Scot- land, but I have seen it in this country only at Scott Center, Wayne Co., in northeastern Pennsylvania, in company with Mr. W. L. Dix (Bartonia 23: 41-42. 1945). There it occu- pied nearly an acre on a dry hillside.' * Aira praecox L. Abundant on roadsides and in newly planted areas in Osterville (no. 711), forming a sort of turf in moist ground and reaching a height of 20 cm. It is also abundant along rte 28 in Mashpee and Santuit (no. 698). Previously Aira praecox has not been reported north of New Jersey, except from Nantucket in the recently pub- lished “Plants of Nantucket" by MacKeever, (1968). It commonly grows with Aira caryophyllea, which is also a native of Europe. *Sporobolus cryptandrus (Torr.) Gray. Abundant in a sandy field at the south end of Hamblins Pond, Marstons Mills (no. 1139). The spikelets are 1.9 mm long. It has previously been known from “sandy and gravelly soil; occasional in the Boston District chiefly near the seacoast, and at Scituate in Plymouth County”, Rhodora 49: 273 (1947). *Carex convoluta Mackenzie. Open woods at edge of swamp at “Spring Hill", East Sandwich, The leaves reach 2 mm in width; culms are erect; perigynia are 3.0-3.5 mm X 1.5-1.7 mm; the achenes, excluding the tips, average 19 x 1.4 mm. Mr. Mackenzie told me that he could dis- tinguish the three closely-related species (C. convoluta, C. 'Dr. Hodgdon has asked me to include here a collection of Molinia. made by David Wise and himself (no. 552) in September, 1959, on Isle au Haut, Maine. It covers a considerable area where it has per- sisted since it was first found there by Nathanial T. Kidder in 1919. I might also mention that the “moor grass" appeared, along with heather and an Erica in the artificial bog at the Brooklyn Botanical Garden in the 1930's. They came from seeds in baled European peat-moss. 8 Rhodora [Vol. 72 rosea, and C. radiata) in the field as far as he could see them, and I tend to agree with him. Measurements in the herbarium tend to overlap, and the natural appearance of the plant is commonly lost on the herbarium sheet, C. con- voluta is common in woodlands of the Boston area, and runs southward into Dighton and Fall River in Bristol County. Another and earlier collection (no. 730) from the Cape Cod locality (June 13, 1967) is not quite mature; the perigynia average 3.0 X 1.0-1.5 mm. *Carex vestita Willd. Sandy margin of a cranberry bog, southeast of Amos Pond in Mashpee (no. 699), and close to, or on, the Sandwich line. The staminate spikes of the species are especially prominent. The plant is common around Boston, and is known from Scituate and Wareham in Plymouth County. “Carex prairea Dewey. Abundant in large tufts in fresh to brackish border of Bumps R., Osterville (no, 717). It is known from a few localities in northeastern Massachu- setts, especially in the vicinity of Concord, but not from southeastern Massachusetts except on Nantucket, where it was collected by Bicknell. *Carex exilis Dewey. Dry opening in a little Chamaecy- paris swamp, near the headwaters of Scorton Creek, East Sandwich (no. 679; June 4, 1967). A single fruiting plant was seen on Aug. 17, 1967 (no, 1049) ; in the wet summer of 1968, fruiting plants were more abundant. *Carer Crawfordii Fernald. A single large clump in gravel excavations along rte 6A in Sagamore (no, 1369), very close to the Sandwich line. In southeastern Massachu- setts it has been known only from Harwich, where it was collected by Fernald and Long on June 25, 1918, and noted as “scarce, in dry sandy fields and borders of woods". C. Crawfordii is a common plant in the White Mountains, and in other parts of northern New England. Carex scabrata Schwein. Spring-heads at margin of swamp along rte 6A, West Barnstable (no. 2003) in prox- imity to Dryopteris Phegopteris and Luzula multiflora var. fusconigra, plants likewise characteristic of northern New England. C. scabrata, which frequents woodland brooks, 1970] Cape Cod — Svenson 9 extends south commonly to the Boston area. In southeast- ern Massachusetts it is known only from “Spring Hill’’, in East Sandwich. *Carex laxiflora Lam. Wooded knoll at edge of swamp, East Sandwich (no. 727; June 13, 1967, and no. 1991; May 25, 1968). This is the same location as “wooded spring- heads bordering maple swamp”, Fernald & Long 18415, July 30, 1919, determined by Fernald as C. leptonervia (likewise by Wiegand in 1922) ; and “rich wooded slope", Fernald & Long 18416, Aug. 9, 1919, determined by Fer- nald as C. laxiflora var. blanda, and in 1922 by Wiegand as C. leptonervia. Both of their collections were made late in the season, The perigynia are poorly developed and show only slight nervation, which is not produced into ribs. Vegetatively they are similar to my collections which are in a fine stage of development. C. leptonervia, therefore, can surely be deleted from the Cape Cod flora. C. laxiflora is common in the Boston region, and it extends south to North Easton and Rehoboth; it ig not known from Plymouth County. C. leptonervia extends south to the Blue Hills near Boston, and also to Attleboro in Bristol County. Both spe- cies are listed from Stoughton (Norfolk County) by S. F. Blake in The Flora of Stoughton, Mass., published by the New England Botanical Club, 1963. *Eleocharis tuberculosa Michx. In sand and peat at mar- gin of a cranberry bog, growing with Rhynchospora capitel- lata, East Sandwich (no. 1065). It is known from Plymouth County, and is of wide distribution on the coastal plain. Lemna minor L. Specimens are recorded only from Barn- stable, Brewster, Harwich, Eastham, Wellfleet, and Prov- incetown, but this little floating plant is probably of more general occurrence. Few people have seen Lemna in fruit. Fruiting collections were made by me at Santuit no at head of tide (no. 1467; July 22, 1968), and in a small dried-out Cephalanthus swamp south of Hamblins Pond, Marstons Mills (no. 1190; Oct. 12, 1967). The utricles vary from glaucous purple to green, of a color similar to the spathes of skunk cabbage, they are obovate and strongly 10 Rhodora [Vol. 72 flattened, 1 mm wide and 1 mm high. The seed lies cross- wise in the utricle and is nearly terete, 0.8 X 0.5 mm in no. 1476; 0.7 X 0.4 mm in 1190. It is greenish-white, some- what corky-inflated, strongly ribbed, obscurely cellular- reticulate, and sometimes with faint crossbars between the ribs. There is a prominent crimson-to-brown caruncle, but the operculum is inconspicuous. In Lemna perpusilla the seed is also large, commonly 0.9 X 0.5 to 1.0 X 0.4 mm; it is loosely erect within the utricle, which is elliptic. In a collection made by Torrey on Staten Island, N.Y., in 1829 (Gray Herbarium), the utricles are about 0.75 X 0.5 mm, and the style varies from erect to oblique; I did not attempt to measure the seed. Plants which I collected in the Galápagos Islands and the coast of Ecuador under the name Lemna minor Philippi have been relegated to H. perpusilla by E. H. Daubs, Univ. Illinois Mon. no. 34. 1965, but the seeds are small, only 0.85-0.4 X 0.2 mm. They are erect, terete, longitudinally ribbed with conspicuous cross partitions and completely fill the utricles. Their size and configuration is not at all that of L. per- pusilla. Clearly more taxonomic work on the utricles and seeds of Lemna is needed. Lemna valdiviana Phil. Forming small tangled submerged clumps, as is usual in the species, in a pond at Discovery Hill Road, East Sandwich (no. 1703). It has been known from Woods Hole and Hyannis. Spirodela polyrhiza (L.) Schleiden. Pond at Discovery Hill Road, East Sandwich (no. 1703A). It has been known previously from Falmouth (see Fogg, Rhodora 32: 106 (1930), and in 1938 from Marstons Mills (Seymour). * Luzula multiflora (Retz.) Lejeune var. fusconig:a Celak. Mossy hummock beneath large beech tree, at edge of a swamp along rte 6A, West Barnstable (no, 1999). The leaves are narrow, mostly less than 2 mm wide, the inflores- cence tends to be congested, and the perianth is deep brown, almost black. It is known from eastern Canada, and south- ward definitely to Old Orchard, Maine. 1970] Cape Cod — Svenson 11 Trillium cernuum L. Nearly a hundred plants in flower May 31, 1967, excavation in wooded slope, East Sandwich (no. 656). Previously it has been known from “Spring Hill” (see introduction). Goodyera pubescens (Willd.) R. Br. Depression in Cornus florida “forest”, east of Johns Pond, Mashpee (no. 2008). The only previous Cape Cod record is from Jenkins Pond, Falmouth, in 1911 and 1912. Spiranthes Grayi Ames. Sandy field, Osterville (no. 1062). This little orchid has been known from Pocasset, Falmouth, Centreville, Brewster, Harwich, Eastham, and Truro. Betula lenta L. Abundant large trees (one foot or more in diameter) in woodland, east side of Wakeby Pond, Sand- wich (no. 1179). The sweet or cherry birch, known locally as “red birch”, forms extensive stands also in the Lowell Holly Reservation in Mashpee, just to the southward. In a mimeographed list of plants compiled for the Lowell Holly Reservation in 1961 by Stephen Hamblin — whom many of us will remember at the Harvard Botanic Garden — Betula lenta is listed as a native tree. Mr. Hamblin retired to Santuit and wrote up the account for President Lowell. After Mr. Hamblin's death in 1966, the list was given to the Cape Cod Museum of Natural History at Brewster. The birch grows also on the opposite shore of Mashpee Pond at the narrows. The nearest localities for Betula lenta appear to be at the Plympton-Kingston boundary in Plymouth County. As Mr. Hamblin states, the Reservation “repre- sents the native vegetation of upper Cape Cod, undisturbed by fire or cutting for at least two or three hundred years". Betula lutea Michx. A single tree, the five trunks each about 8 inches in diameter at border of pond, Discovery Hill Road, East Sandwich (no. 674). This locality, west of the previously mentioned locality at "Spring Hill", ap- pears to be the only additional Cape Cod occurrence of the yellow birch, except at Shawme Lake, Sandwich, mentioned by me in Rhodora 31: 80 (1929). John Hay, President of the Cape Cod Museum of Natural History, and I were un- 12 Rhodora [Vol. 72 able to find the yellow birch mentioned by Mr. Hamblin in the Lowell Holly Reservation list, but we did find several trees of paper birch, omitted from the list. Betula papyrifera Marsh. There is only one Cape Cod specimen of the paper birch in the Herbarium of the New England Botanical Club, It is from the “rich wooded slope at Spring Hill, Fernald & Long, Aug. 9, 1919.” But it is more widespread. Trees have been planted along the main highway, rte 6, but native trees are scattered in many places. The finest trees are to be found on the north-facing moraine slope east of Atkins Road in East Sandwich, where they have been carefully preserved in land clearing. Trey are a little over 1 ft. in diameter, The greatest assemblage of trees appears to be at the north end of Santuit Pond in Mashpee. I have also seen a tree at the south side of Lovell’s Pond, Santuit (no. 1890), another in Osterville, and scat- tered trees in Falmouth. Most of the trecs have the appear- ance of those on Long Island, New York, where they occur on the terminal moraine at Wyandanch. Castanea dentata (Marshall Bork. Two clumps, each about 10 ft. high, vigorous though half-blighted, Old Jail Road, West Barnstable (no, 1763; Sept. 10 1968). The only specimens at the N.E.B.C. are from Yarmouthport (1907), and Bourne (1918). According to reports of older inhabitants the chestnut was only occasional on Cape Cod. Quercus rubra L. Large tree on sandspit between Middle and Mystic Ponds, Marstons Mills (no. 1961; Jan. 27, 1968). Around these ponds is probably the largest assemblage of red oaks on Cape Cod. The best time to see them is when the ponds are frozen. While ice-fishing I counted over twenty trees on the above date. There are probably many more; the trees can be most easily recognized by the large flat cups on the ground. Large tree overhanging the water, Mashpee Pond, Mashpee (no. 1180). Mr. Hay and I counted nine trees in the adjacent Lowell Holly Reservation, where the red oak was not mentioned by Mr. Hamblin. There is also a group of red oaks at Discovery Road in East Sand- wich, the largest 33 inches in diameter. The red oak is 1970] Cape Cod — Svenson 13 otherwise known on Cape Cod (specimens in the N.E.B.C.) only from Wellfleet, and from Cliff Pond and No Bottom Pond, both in Brewster. Ceratophyllum demersum L. Fruiting plants in ditch in cranberry bog, East Sandwich (no. 1068). Previously it has been collected in Chatham and Eastham. Fruiting ma- terial, rarely seen, was obtained by Fogg at Oyster Pond, Falmouth (Rhodora 32: 107 (1930) ). Thalictrum revolutum DC. Roadside, near Cotuit R., Mashpee (no. 794). It is one of the plants characteristic of the Upper Cape (Fogg, Rhodora 32: 176 (1930) ), pre- viously known from Woods Hole, Falmouth, Sandwich, and Centreville. Nasturtium officinale R. Br. There are no specimens from Cape Cod, but the plant is abundant in several streams, in- cluding Mashpee R., Mashpee (no. 705) ; Santuit R., Cotuit (no. 805) ; and Herring Brook, North Falmouth (no. 2072). The siliques of 2070 (June 8, 1968) are rather small, aver- aging 8 X 2 mm; the style 0.8 X 0.4 mm. The other col- lections were only in flower. Ribes hirtellum Michx. var. calcicola Fernald. Tidal banks of Marstons Mills R., Marstons Mills (no. 1451; July 29, 1969), in fruit. The fruits are smooth, red and glaucous, 1 cm wide when flattened. Seeds, apparently not previously described, are lucid brown, somewhat flattened, irregularly ovate, and 4-5 X 1.5 mm. They are surrounded by an ad- herent gelatinous substance. Var. calcicola was described from calcareous swamps of Gaspé, Quebec, with “young branches, petioles, and lower leaf-surfaces permanently and densely white tomentose”. Many collections from Cape Cod have been so named, but whether they exactly represent var. calcicola is open to question. Petioles in no. 1451 are pubescent, but the leaves are only hirtellous on the veins beneath. The plant is to be expected in thickets at the head of tidal streams throughout Cape Cod. *Crataegus monogyna Jacq. The European hawthorn has not been reported from Cape Cod, but small trees are abun- dant along the railroad east and west of West Barnstable 14 Rhodora [Vol. 72 station, altogether for at least a mile (no. 744, in flower June 11, 1967; in fruit, July 30, 1968). Corema Conradii Torr. Open hilltops under pitch pines north of DeGrass Road, South Mashpee (no. 1096). It is also known from North Falmouth, Osterville, Dennis, and West Brewster, but mostly it has been associated with the open sands extending from Eastham to Provincetown. In the middle and western part of the Cape it forms a springy carpet beneath pitch pines. Consequently there is no reason to question its being native in the Lowell Holly Reservation, where it was reported by Mr. Hamblin, and where it has been known for a long time. My first acquaintance with Corema was in the sands of Wellfleet and in similar situa- tions in the New Jersey Pine Barrens, where Witmer Stone (Rept. N.J. State Museum, 1910) says it is “an inhabitant of those desolate stretches of white sand which cover the most elevated portion of the Pine Barren region, stretching away for some thirty square miles." As I have seen the plant on the Maine coast in hills north of Camden, and on the promontory known as “Gertrude’s Nose", in the Shaw- angunk Mts. of southern New York, it does not grow in sand. It is listed in Gray's Manual, ed. 8, from “sandy pine- barrens, sandhills and siliceous rocks". * Rhamnus frangula L. Tree about 15 ft. high, in swamp along railroad, East Sandwich (no. 1198; Oct. 17, 1967, in fruit). Hodgdon and Krochmal (Rhodora 52: 165 (1950)), discuss the presence and spread of this species in New Hampshire. Circaea quadrisuleata (Maxim.) Franch. & Sav., var. can- «densis (L.) Hara. Moist woods along Marston Mills R., below rte 28, Marstons Mills (no. 1452). It has been found previously in Bourne, Falmouth, Sandwich, and Brewster. Rhododendron canadense (L.) Torr. Rhodora in flower, June 2, 1967, occurring over nearly an acre southeast of Johns Pond (no. 664). The area is now bisected by a power line, which may help in preservation. Kalmia latifolia L. Bushes to 6 ft. high, abundant on steep slopes at east side of Long Pond, Newtown, Barn- 1970] Cape Cod — Svenson 15 stable (no. 655). The mountain laurel has not been re- ported previously from Cape Cod, but Miss Eda M. Roos of East Sandwich informs me that it has been known as a native plant for more than a half century near the north end of Wakeby Pond, which is close by. It is surprising that this bush, so abundant on Long Island, should be of such limited occurrence on Cape Cod. Gentiana crinita Froel. The fringed gentian, as in most other places, tends to be sporadic in occurrence. It has been known for a long time at East Sandwich, sometimes in profusion, at other times scarce. I collected it on Nov. 9, 1966, For the past two years it has not made an appear- ance, but will probably do so when conditions for growth are favorable. It is known from several localities in Ply- mouth County. Bartonia paniculata (Michx.) Muhl. Chamaecyparis swamp, South Main St., Centreville (no. 1053; Aug, 20, 1967). The material is slender, and greatly resembles the illustration of B. tenella in Rhodora 2: pl. 15, f. 6 (1900). The leaves are subulate, purplish on the lower part of the stem, green on the middle and upper part. Anthers very from purplish to dark yellow. Bicknell (Bull. Torr. Cl. 42: 33, 1915, and 46: 423, 1919) found similar intermediate plants on Marthas Vineyard and Long Island. On Cape Cod, Bartonia paniculata has been previously known from Barnstable and Dennis. *Myosotis scorpioides L. Densely covering extensive areas in Santuit R., Cotuit (no. 752), The European for- get-me-not has not been recorded from the Cape. *Orobanche uniflora L. Springy hummock below Chamae- cyparis, at edge of brackish marsh, Bumps R., Osterville (no. 678). Known southward to the Taunton and New Bed- ford area in Bristol County. Jasione montana L. Sandy fields, Osterville (no. 817). For many years this handsome blue-flowered plant was known only in the vicinity of the cemetery at East Sand- wich, It now appears to be spreading rapidly, and is abun- dant at the south end of Hambling Pond in Marstons Mills. 16 Rhodora [Vol. 72 Erigeron pulchellus Michx. On knoll at edge of swamp at “Spring Hill", East Sandwich (no. 720). In the dry sum- mer of 1927, the plants occupied an area about 2 feet square, and I found a single flower. In 1968, a wet summer, every- thing was overgrown and I could find no trace of the plants. The nearest locality is New Bedford. OSTERVILLE, MASS. 02655 ADDITIONS AND NOMENCLATURAL CHANGES IN THE FLORA OF SOUTHERN FLORIDA — I. ROBERT W. LONG! The purpose of this paper is to validate certain nomen- clatural changes, and to record additions and certain taxo- nomic reinterpretations in the vascular flora of southern Florida. These changes and additions have resulted from preliminary studies that have been part of the preparation of a manual of the flora of tropical Florida. The flora of southern Florida is a rich one for so small an area (Long, Lakela, and Broome, 1969), and it was of special interest to J. K. Small. Numerous nomenclatural and taxonomic re- visions have been necessary, but it will not be possible to report here all the changes from names used by Small in his manuals dealing with this flora (1903, 1913a, 1913b, and 1933). Other additions or changes requiring validation may be necessary as work progresses; the present paper includes references only for certain dicotyledonous families. RANUNCULACEAE Clematis The populations of Clematis baldwinii are highly variable in peninsular Florida, where the species is endemic, but two recognizable morphological extremes occur. These are sep- arable on the basis of sepal length and leaf shape, but they are connected by morphologically intermediate plants. Gray (1895) made reference to the broad leaf form of C. bald- winii but did not name it. Erickson (1943) placed the spe- cies in his monotypic subsect. Baldwinianae, sect. Viorna, separated from its apparent nearest relative the western ‘Contribution No. 47 from the Botanical Laboratories, University of South Florida. Research was supported in part by grant No. GB-2742 from the National Science Foundation. Much of this investigation was carried out at the Arnold Arboretum and Gray Herbarium, Harvard University, while the author held a Mercer Re- search Fellowship in the Arnold Arboretum, 1968. 17 18 Rhodora [Vol. 72 C. fremontii S. Wats. which has all leaves simple and ovate to orbicular in shape. Additional field work and an analysis of population vari- ability would be very desirable in C. baldwinii. A possible third variety characterized by very small flowers, sepals less than 2 cm long and with linear leaves, occurring on the white sand scrub of central Florida and extending into southern Florida, may also deserve varietal recognition. It is possible that var. latiuscula may represent hybrids with broad leaf forms of C. crispa L. or C. reticulata Walt., or may even represent remnants of wholly different population systems. Sepals 3-4 em long; leaves mostly linear to narrowly lanceolate. ...... ssssnsseessesesscassosncaresccoensescsvonssseeseeavebasscusossvsssebeess C. baldwinii var. baldwinii Sepals 4-5.5 em long; leaves mostly elliptic-lanceolate to narrowly ovate. .............. r nennen nnne een een nennen nns C. baldwinii var. latiuscula CLEMATIS BALDWINII var. BALDWINII T. & G., Fl. N. Am. 1: 8. 1838. Viorna baldwinii Small, Fl. SE. U. S. 439, 1331, 1903. DADE CO.: east boundary Everglades National Park, Dec. 23, 1955, F. C. Craighead and L. E. Arnold, s.n. (GH); HILLSBOROUGH CO.: northwest of Tampa near junction of Waters street and Hulsey road, May 19, 1962, O. Lakela 25969 (GH, USF); LEE CO.: occasional erect in pine-palmetto flatwoods 10 mi wsw Salvista, May 13, 1958, R. Kral 6542 (GH); POLK CO.: partly cleared pineland close to RR tracks, just off Lakeland city limits, infrequent, March 12, 1961, O. Lakela 23836 (GH, USF). CLEMATIS BALDWINII var. latiuscula R. W. Long, var. nov. A typo differt foliis integris; lamina ovato-attenuata, 7.5-8.0 em longa, 2.0-2.4 lata, breviter petiolata; supera folia sessilia; lamina manifeste palmatim trinervis, reticu- lata; sepala longa 5.0-5.4 cm, lata 6-8 mm; pedunculi 18- 22 cm longi, nudiusculi. TYPE, LEVY CO. FLA. In low, open pineland, S 34, T 168, R 16E, April 25, 1959, G. R. Cooley 6549 and R. J. Eaton. In Gray Herbarium, isotype in USF. CITRUS CO.: 2 mi w Sumter Co. line on hwy 48 between Bushnell and Floral City on shoulder of road and edge of wood, April 29, 1958, G. Cooley 6069, C. Wood and K. Wilson (GH, USF) ; HENDRY CO.: near 1970] Southern Florida — Long 19 Goodno, hammock, April 8, 1940, D. E. Eyles 6819 (GH); HILLS- BOROUGH CO.: sand barren beyond Ballast Point, W. Tampa, May 30, 1923, J. R. Churchill s. n. (GH); SARASOTA CO.: Sarasota, Dec. 25, 1942, A. E. Perkins, s.n. (GH); VOLUSIA CO.: Port Orange, May 13, 1895, F. C. Straub 145 (GH). POLYGALACEAE Polygala The genus is well-represented in southern Florida by 17 Species and a number of infraspecific taxa. Several are endemic to the limestone soils that support pineland and hammock vegetation. Polygala boykinii, for example, in- cludes three well-marked varieties in southern Florida, two of which are endemic. They may be separated as follows: 1. Lower-most leaves elliptic-obovate to suborbicular, often mucro- nate. 2. Leaf-blades elliptic-obovate, upper blades linear. ........................ Oe Su tum 1s Eu scorsi P. boykinii var. boykinii 2. Leaf-blades suborbicular, upper blades usually narrowly lanceo- |e? arent Pee nn ag n NC S P. boykinii var. suborbicularis 1. Lower-most leaves subulate to linear or linear-lanceolate. ............ P. boykinii var. sparsifolia POLYGALA BOYKINII var. BOYKINII Nutt. Jour. Acad. Phil. 7: 86. 1834. Pinelands, Fla. to La, Ga. HILLSBOROUGH CO.: along country road from 30th st. and Skipper Ave., low seasonally wet, weedy roadside, June 17, 1962, O. Lakela 25131 (USF); MONROE CO.: low area near Fla 94, Pinecrest, ca 6 mi w of Dade-Monroe county line T54S, R 34 E, Dec. 29, 1959, D. B. and S. S. Ward 1176 (USF). POLYGALA BOYKINII var. suborbicularis R. W. Long, var. nov. A typo differt foliis infernis suborbicularibus 0.8-1.4 cm longis, 0.6-0.8 em latis, superis foliis anguste lanceolatis raro linearibus, 0.9-1.2 longis, 0.3-0.5 em latis. TYPE, LEE CO. FLA. Pinewoods, 10 mile camp, near Ever- glades, March 23-26, 1905, A. A. Eaton 1384. In Gray Herbarium. HERNANDO CO.: west of Weeki Wachee springs, frequent, March 19, 1958, G. R. Cooley 5650 and J. Monachino (GH, USF); LEE CO.: vicinity of Fort Myers, in hammock, April 3, 1916, Jeanette P. Standley 129 (GH) (A smaller leaf-form but similar to the type); MONROE CO.: Big Pine Key in limerock outcropping under Pinus elliottii and Sabal palmetto, May 3, 1958, G. Cooley 6195, et al. (GH, UsF). 20 Rhodora [Vol. 72 POLYGALA BOYKINII var. SPARSIFOLIA Wheelock, Mem. Torr. Club 2: 121. 1891. TYPE, MONROE CO., FLA. Coral soil, Cudjoe Key, A. H. Curtiss, s.n. (Isotype, GH!). P. praetervisa Chodat, Mem. Soc. Phys. Geneve 31(2): 140, 1891. P. flagellaris Small, Bull. N. Y. Bot. Gard. 3: 427. 1905 (p.p. incl. type). P. sparsifolia Small, Fl. SE. U. S. 686, 1535. 1905. Endemic to extreme southern peninsular Florida and the Florida Keys, this is perhaps the most distinctive variety in the species. DADE CO.: Everglades National Park near Homestead, Miami oolite and solution holes, hammock and pineland association, Oct. 16, 1962, G. Cooley 9410, et. al. (GH, USF) (Intermediate form to typical variety); MONROE C0.: Big Pine Key, pine-palm woods, May 24, 1952, W. B. Robertson 274 (GH); Big Pine Key, pineland, April 19, 1942, C. E. and M. S. Eyles 8201 (GH); Big Pine Key, pinewoods, Feb, 21, 1936, E. P. Killip (GH). A second species that includes two endemic varieties in Florida is Polygala grandiflora, Small described a number of endemic species from this complex that may be referred to one or the other of the varieties described below. 1. Wings greenish or purplish-tinged. 2. Leaf blades elliptic or elliptic-lanceolate. .................ee NEN P. grandiflora var. grandiflora 2. Leaf blades linear or linear-lanceolate. ............. eee NENNEN P. grandiflora var. angustifolia 1. Wings dark purple. ............ een P. grandiflora var. leiodes POLYGALA GRANDIFLORA var. GRANDIFLORA Walt., Fl. Car. 179. 1788. P. cumulicola Small, Bull. Torr. Bot. Club 51: 9381. 1924. P. miamiensis Small, in Blake, N. Am. Fl. 25(5) : 340. 1924. Small stated that P. cumulicola differs from P. grandiflora in having deep-rose purple wings and a short raceme, which is true (isotype GH!). Additional study may warrant recog- nition of this race as a variety of P. grandiflora, however at present these characters do not seem to mark a definite 1970] Southern Florida — Long 21 taxon within the species sufficiently to necessitate taxonomic recognition. POLYGALA GRANDIFLORA var. ANGUSTIFOLIA T. & G., Fl. N. Am. 1: 671. 1840. P. grandiflora var. leptophylla Chodat, Mem. Soc. Phys. et Hist. Nat. Genèv. 31: 57. 1898. P. corallicola Small, Bull. N. Y. Bot. Gard. 3: 425. 1905. POLYGALA GRANDIFLORA var. LEIODES Blake, N. Am. Fl. 25: 339. 1924. P. grandiflora var. grandiflora is found throughout a num- ber of the Southeastern states, but P. grandiflora var. an- gustifolia and P. grandiflora var. leiodes are generally con- fined to extreme southern Florida and the West Indies. EUPHORBIACEAE Croton Six species occur in southern Florida, and the most vari- able of these is Croton glandulosus, a wide-spread tropical weed species that includes two varieties endemic to the dry pinelands. The typical variety appears to be more common in the maritime sands suggesting ecotypic separation of the varieties. However, C. glandulosus var. glandulosus may be found in various sites, as pinelands, old fields, sandy soil, and disturbed sites. The varieties may be separated as fol- lows: 1. Calyx and fruit pubescent. 2. Leaf blades stellate-pilose on both surfaces, margins serrate to dentate Misi. Qu u a su, C. glandulosus var. glandulosus 2. Leaf blades densely stellate-pubescent on both surfaces, mar- eins crenate-serrate. .................... C. glandulosus var. simpsonii 1. Calyx and fruit smooth or nearly so. .......... ee C. glandulosus var. floridanus eeesssesosssosssssstsossosessosesosooosemeeeosessoestoee nn CROTON GLANDULOSUS L. var. GLANDULOSUS COLLIER CO.: beach area of Marco Island, off US 92; construction site with destruction of native vegetation, July 1, 1967, O. Lakela 30839 (USF); vicinity of Naples, recently cleared weedy sandlot with Centrosema, Euphorbia, Cassia, weed in vacant lot, Oct. 9, 1962, G. R. Cooley 8990, R. Eaton, and O. Lakela (USF); DADE CO.: sandy beach, Key Biscayne, Nov. 2, 1965, F. C. Craighead (USF); HIGHLANDS 22 Rhodora [Vol. 72 co.: frequent in grove by SAL RR, 2.1 m n. of Sebring on US alt 27, T 34S, R 29E, June 8, 1964, D. Burch 384 (USF); PALM BEACH co.: w. side of Dreher Park, West Palm Beach, disturbed ground, sand, full sun, 7 dm tall, Aug. 29, 1967, Paul Cassen 180 (USF). CROTON GLANDULOSUS var. SIMPSONII Ferg., Rep. Mo. Bot. Gard. 12: 51. 1901. BREVARD CO.: Melbourne Beach, n of park area, with escaped cultigens, June 7, 1964, O. Lakela 27206A (USF); DADE CO.: pineland associated with Acalypha, Neptunia, Schrankia, pubescence coarse, Old Cutler Road, near Deering Hammock, July 17, 1964, O. Lakela 27275 (USF). CROTON GLANDULOSUS var. floridanus (Ferg.) R. W. Long, stat. nov. C. floridanus Ferguson, Rep. Mo. Bot. Gard. 12: 50, pl. 15. 1901. This variety is readily separable from the other ones by means of the glabrous or nearly glabrous sepals of the pistillate flowers, and glabrous fruits. All populations of C. glandulosus in southern Florida appear to be distinct from the northern and central Florida C. glandulosus var. septentrionalis Muell. Arg. COLLIER CO.: under trees, Vanderbilt Beach, n Naples, leeward side of dunes, Sept. 8, 1967, O. Lakela 31069, (USF); Marco Island, beach area; plants frequent in sandy openings, Sept. 27, 1964, O. Lakela 27509 (USF); MANATEE CO.: Anna Maria Island, coastal dunes and white sand beaches, Jan. 15, 1967, R. W. Long 2367, R. Broome, C. Croley (USF); MONROE CO.: Murray Key, Florida Bay off Flamingo, open drier site with salt grasses and succulents, May 8, 1965, O. Lakela 28599, R. W. Long, and F. Craighead; PINELLAS CO.: Fort DeSoto Park, s end, capsules glabrous, Sept. 10, 1963, R. Thorne, R. Long, O. Lakela 26282 (usr). MYRTACEAE Myrcianthes The two species of Myrcianthes reported as endemic for Florida (Wilson, 1961) are very probably merely local variants of M. fragrans, a wide-ranging species in the Amer- ican tropics. They differ from the typical form chiefly in having slightly longer petioles and by producing cymes usually 7-flowered instead of 3-flowered. However, there 1970] Southern Florida — Long 23 is some justification for maintaining M. simpsonii because the cymes are several-flowered, usually 10-14; this would then be our only endemic variety of M. fragrans. There is less justification for maintaining M. dicrana, and it is therefore placed in synonomy under the typical variety. MYRCIANTHES FRAGRANS (Sw.) McVaugh — Shrubs or trees up to 20 m tall, usually less, with reddish-brown or light- brown bark. Leaves narrowly obovate to elliptic-cuneate 1-8 em long, acute or obtuse, pale green above, slightly paler beneath. Flowers in cymes with 3-14 flowers; sepals about 1-2 mm long, corolla about 1 cm wide. Berry globose or ellipsoidal. Hammocks, s Fla, tropical Amer. Corolla less than 1 cm wide, cymes usually 3-7 flowered. .................... MES EIER E e adieesttasveinbsfeosnr irte TN s S EE T OS var. fragrans Corolla over 1 cm wide, cymes several flowered, usually 10-14. ........ var. simpsonii Oe eee eee eee eee ere er errr rere rere ricer errr rrr errr iri) MYRCIANTHES FRAGRANS (Sw.) McVaugh var. FRAGRANS, Fieldiana: Bot. 29: 485, 1963. Anamomis dicrana Britton, Britton and Shafer, N.A. Trees, 728, fig. 668. 1908. (Eugenia dicrana Berg.) DADE CO.: Everglades, Paradise Key, May 5, 1908, E. A. Bessey (A); Everglades National Park near Long Pine Key, 5% miles w, 2 m n of Paradise Key, altitude 6 ft. in hammock in pinewoods, limerock with pockets of soil in cracks, considerable organic litter, assoc. with Ficus brevifolia, Metopium toxiferum, Rapanea, Dipholis, tree up to 7 ft. irregularly branched, April 22, 1958, A. Traverse 586 (GH); INDIAN RIVER CO.: near Vero Beach, abundant in dense scrub; scrub or tree up to 6 m tall, bark flaky, brownish; galled fruits orange, July 2, 1949, L. J. Brass 20582 (GH). MYRCIANTHES FRAGRANS var. simpsonii (Small) R. W. Long, comb. nov. Anamomis simpsonii Small, Torreya 17: 222. 1917. Myrcianthes simpsonii (Small) K. A. Wilson, Jour. Ar- nold Arb. 41: 276. 1961. DADE CO.: Arch Creek, May 1918, C. J. Simpson (A) ; hammock along Arch creek, May 12, 1917, J. K. Small 8287 (GH); MONROE CO.: North Key Largo ca 3 m from left turn of US I, coral limestone, Sweitenia, Gouania, Dalbergia, 4 m high, Oct. 13, 1962, G. R. Cooley 9293, J. R. Eaton, O. Lakela (GH, UsF). 24 Rhodora [Vol. 72 PLUMBAGINACEAE Limonium Two Limoniums have been described from Florida ma- terial: L. carolinianum (Walt.) Britt. var. angustata A. Gray, and L. nashii Small. In his monograph Blake stated that L. carolinianum has a glabrous calyx and deltoid-obtuse lobes. Gray based his variety (type, GH!) on the narrow- ness of the leaves. Small described L. nashii as having pubescent calyx lobes, at least at the base; L. trichogonum Blake supposedly has still more strongly pubescent calyx. Finally, to complicate further the taxonomy is L. obtusi- lobum (type, GH!) supposedly with a shorter calyx that is pilose-pubescent. Blake stated that the chief characters for specific discrimination in Limonium were pubescence, size and form of calyx and shape of the calyx lobes, shane of flowering bractlets, and aggregation of flowers. Ahles incorrectly reduced L. obtusilobum to a variety of L. carolinianum. The former is characterized by a line of pubescence along the sepal and is hirsute on the blunt- shaped sepals; the latter species is entirely glabrous and is so characterized. Ahles then made L. carolinianum var. angustata, which was originally characterized by narrow- ness of the leaves, a variety of L. nashii, a species that was originally based on a sparingly pubescent calyx tube. The aggregation of flowers is a poor character for sep- arating taxa because there is continual variation from one extreme to another. Pubescence of the calyx is also unstable. After examination of type collections in GH and of collec- tions from southern Florida, it is apparent that only two varieties of L. carolinianum are worthy of recognition: L. carolinianum var. angustata (Gray) Blake representing a narrow leaf, dwarfish race endemic to the Florida Keys and southern peninsular Florida, and L. carolinianum var. carolinianum the larger more robust race. Leaves lanceolate-elliptic to obovate-spatulate; floral internodes less than 5 mm long. .............. Innen var. carolinianum Leaves linear-lanceolate to narrowly elliptic; floral internodes about 5 mm long. ................ a eene nennen var. angustatum 1970] Southern Florida — Long 25 LIMONIUM CAROLINIANUM (Walt.) Britt. var. CAROLINIA- NUM L. nashii Small, Bull. Torr. Bot. Club 24: 491. 1897. L. trichoganum Blake, Rhodora 18: 61. 1916. L. nashii var. trichoganum Blake, Rhodora 25: 58. 1923. L. carolinianum var. nashii (Small) Boiv. Nat. Canad. 93: 643. 1966. LIMONIUM CAROLINIANUM var. ANGUSTATUM (A. Gray) Blake, Rhodora 25: 56. 1923. Statice brasiliensis var. angustatum A. Gray, Synopt. Fl. N. Amer. 2(1) : 54. 1878. L. angustatum (A. Gray) Small, Bull. Torr. Bot. Club 24: 488. 1897. L. obtusilobum Blake, Rhodora 18: 63. 1916. L. carolinianum var. obtusilobum (Blake) Ahles, Journ. Elisha Mitch. Sci. Soc. 80: 173, 1964. L. nashii var. angustatum (A. Gray) Ahles, Journ. Elisha Mitch. Sci. Soc. 80: 173. 1964. TYPE: Pine Key. Blodgett. In Gray Herbarium! LEE CO.:: on wet mucky soil of mangrove swamp, Bokeelia Island, Dec. 31, 1956, R. Kral 3901 (GH); MONROE CO.: Big Pine Key, saline flats south of Inn, Dec. 29, 1953, E. P. Killip 43400 (GH); PINELLAS CO.: salt flats, Snug Harbor, Nov. 14, 1959, R. K. Godfrey 59192 (GH). SAPOTACEAE Bumelia The United States species of Bumelia were revised by Clark (1942) and later by Cronquist (1945). Bumelia an- gustifolia was described by Nuttall from a tree that was common in Key West according to Blodgett. It has leaves unusually small and narrow linear-oblong and obtuse, with the blade 2.5-3.0 em long and up to 1 cm wide that come out in clusters from the center of the preceding bud. Nuttall was the first to recognize the distinctiveness of this race. All specimens seen of this taxon occur in southern Florida, and apparently it is endemic to the lower Florida Keys and the Cape Sable area. Bumelia celastrina var. celastrina also occurs in this area but not nearly so commonly, al- 26 Rhodora [Vol. 72 though it is abundant farther north in peninsular Florida. The two races intergrade although the extremes are easily distinguished. B. celastrina var. angustifolia, with its much reduced, narrower leaves, appears to be a distinct ecotype adapted to coastal hammocks and pinelands. The type for B. celastrina (photo, GH!) is from Mexico and is of a large-leaved plant with blades 3-4 em long. Possibly, Cronquist drew broad, inclusive circumscriptions that incorporate many otherwise distinctive races, and probably population structure is considerably more com- plex than his treatment would infer. Certainly the Florida Keys race is morphologically separable from the typical variety. BUMELIA CELASTRINA H.B.K. Trees or shrubs up to 8 m tall with glabrous stems. Leaves evergreen, fascicled, blades oblanceolate to oblanceolate-spatulate, 1-4 em long, glabrous. Flowers few-several in axillary cymes or umbels; calyx lobes 2 mm long, ovate, corolla 3-4 mm wide, lateral lobes irregularly toothed; staminodia 2-3 mm long, lacerate. Styles 2.5-4 mm long. Berry cylindrical 7-13 mm long. Leaf blades oblanceolate to spatulate 2-4 em long, slightly pubescent when DAMM var. celastrina Leaf blades linear-oblong, obtuse, mostly 1-2 cm long, glabrous when MO T MM var. angustifolia BUMELIA CELASTRINA var. angustifolia (Nutt.) R. W. Long, stat. nov. Bumelia angustifolia Nutt., N. Am. Sylva 3: 38, pl. 93. 1849. CHARLOTTE CO.: Boca Grande, golf links, small tree, Jan. 20, 1920, J. S. Ames (A); DADE CO.: East Cape, Nov. 1912, C. T. Simpson (A) ; 2 mi e of Cape Sable, Nov. 1912, C. T. Simpson; MONROE CO.: Sum- merland Key, July 2, 1956, C. K. Brizicky 355, & W. L. Stern (A); Little Torch Key, hammock, tree 15-18 ft high, 7 in. dbh, June 29, 1956, G. K. Brizicky 344 & W. L. Stern, (A); Big Pine Key, small tree 10 ft high, 3 in diam, dry pinelands near Myrtle and Wesley's place, June 22, 1956, G. K. Brizicky and W. L. Stern. LOGANIACEAE Cynoctonum Three species, including one new one, occur in southern 1970] Southern Florida — Long 27 Florida, and they may be distinguished as follows: Leaf blades narrowly lanceolate to ovate-orbicular, sessile. ................ ROI MERC UU EO ee TURN e cR Le 1. C. sessilifolium Leaf blades narrowly lanceolate to ovate-lanceolate or ovate, short- petioled. Lower leaves ovate-lanceolate or ovate, fleshy, larger blades tending to cluster at the base in a rosette; inflorescence mostly Z4 cm long. eoo BO arisen iSi ss 2. C. succulentum Lower leaves lanceolate to elliptic or ovate, not clustered at the base; inflorescence mostly 4-6 cm long. ........... 3. C. mitreola 1. CYNOCTONUM SESSILIFOLIUM (Walt.) J. F. Gmel. Three varieties are present in southern Florida: 1. Leaf blades elliptic to ovate or orbicular. 2. Leaf blades mostly 2-4 cm long, apex obtuse. ............................ me HUI HUI HUNE var. sessilifolium 2. Leaf blades mostly 1-1.5 cm long, apex acute. uns... RU SOS sS MEN Cd, ol ca cane Lace te Ro DA qata var. microphyllum 1. Leaf blades narrowly lanceolate, .................... var. angustifolium CYNOCTONUM SESSILIFOLIUM var. SESSILIFOLIUM. Ditches, low ground, wet places, a coastal plain plant, Fla to La, Tex, and Va. Widely distributed in Fla. CYNOCTONUM SESSILIFOLIUM var. microphyllum, R. W. Long, var. nov. A typo differt caule simplici, foliis sessilibus, parvulis, 1- 1.5 em longis, pro parte maxima minus quam 1 em longis, apicis acutis; laminis ellipticis vel late lanceolatis ; inflores- centiis aliquot ramosis, ramis minus quam 2 cm longis per anthesin. TYPE: BREVARD CO. FLA.: 4 miles west of Melbourne, oc- casional on moist sandy peat of pine flatwoods; flowers white. August 22, 1958, R. Kral 7961. In Gray Herbarium, isotype in USF. Stems simple, leaves sessile, small, the blades 1-1.5 cm long, mostly less than 1 cm long with acute to broadly acute apex ; blades elliptic to broadly lanceolate, inflorescence few- branched, the branches less than 2 cm long at flowering. FLORIDA: CALHOUN CO.: grassy swamps, Sept. 1, 1889, Chapman. Herb. 2824" (GH); CITRUS CO.: 5 mi s Homosassa; frequent on moist sandy peat of clearing near cypress dome, August 10, 1958, R. Kral 7764 (GH, USF); DADE CO.: Miami, July 1877, A. P. Garber (GH); 28 Rhodora [Vol. 72 DUVAL CO.: Baldwin, August 6, 1895, G. V. Nash (GH); INDIAN RIVER CO.: 7 mi. west of Vero Beach, frequent on moist sandy peat of grass-sedge bog, August 29, 1957, R. Kral 5587 (GH, USF); LEE CO.: around flatwood ponds, Ft. Myers, July-Aug. 1900, A. S. Hitch- cock 214 (GH); LEVY CO.: along Rt. 24, 3 mi southwest of Otter Creek, sandy soil along highway right-of-way, Sept. 13, 1954, R. E. Perdue Jr. 1732 (GH); MONROE CO.:: Key West, Blodgett (GH); OSCEOLA CO.: pond on prairie, Aug. 4, 1903, A. Fredholm 5947 (GH). GEORGIA: DOUGLAS CO.: acid bog-meadow 2 mi e of Villa Rica, flowers white, July 27, 1948, A. Cronquist 5563 (GH). LOUISIANA: BEAUREGARD PARISH: wet sandy soil along shallow ditch about 10 mi south of De Ridder, July 21, 1938, D. S. and H. B. Correll 9673 (GH); VERNON PARISH: 2 mi. w of Leander, grassy field with wet areas from springs; woody vegetation removed except for a few pines, frequent, flowers white, July 7, 1950, G. L. Webster and R. L. Wilbur 3235 (an). MISSISSIPPI: JACKSON CO.: Ocean Springs, July 27, 1896, C. L. Pollard (GH). NORTH CAROLINA: CARTARET CO.: savannah, at Newport, Aug. 6, 1938, R. K. Godfrey 5786 (GH). TEXAS: HOUSTON CO.: Grapeland, sandy open bogs, Sept. 22, 1917, E. J. Palmer 12849 (au). SOUTH CAROLINA: HORRY co.: Conway, savannah, Sept. 1, 1940, P. O. Schallert (Gn). This variety appears as rigidly erect, simple plants with much reduced leaves. In distribution it appears to be sim- ilar to that of the typical variety although it apparently is ecotypically adapted to wet, sandy or more often peaty soil, rather than the chiefly pineland habitat of the typical race, Both morphologically and ecologically it is readily separable from the other two varieties. CYNOCTONUM SESSILIFOLIUM var. ANGUSTIFOLIUM T. & G. Mitreola sessilifolia B angustifolia T. & G., Fl. N. Amer. 2: 45. 1841. Cynoctonum angustifolium Small, Bull. Torr. Bot. Club 23: 129. 1896. Wet soil, Fla to Ga on the coastal plain. FLORIDA: DADE CO.: southwest of Homestead, Everglades National Park, oolite depressions, trailside, Oct. 16, 1962, O. Lakela and F. C. Craighead (USF); Everglades National Park, pinelands, plant red- dish before drying, Oct. 12, 1962, G. R. Cooley 9223, R. Eaton, J. Ray, R. Long, and O. Lakela (USF); HERNANDO CO.: one mile n of Weeki 1970] Southern Florida — Long 29 Wachee Springs, occasional on moist sandy pineland pond border, June 9, 1958, R. and Mabel Kral 6778 (USF). 2. Cynoctonum succulentum, R. W. Long, sp. nov. Planta annua 2-4 dm alta, caulis simplex, raro ramosus super. Folia elliptica ad ovata, 3-5 cm longa, 1.5-2.0 lata, glabra, succulenta, infima rosulata, caulina folia vix demi- nuta, omnia petiolis brevissimis. Inflorescentia cyma ramosa Spicis brevibus pro parte maxima 2-4 cm longis, dense secunda ; sepala 0.5-0.75 mm longa, corolla 1-1.5 mm longa, bractea 1-1.5 mm longa paulatim decrescens, plerumque suc- culenta. Capsula plerumque mitriformis. TYPE: In pine barrens, near Manatee, Florida, June 11, 1890, J. H. Simpson. In Gray Herbarium. Annual herbs with smooth stems 2-4 dm tall, simple, seldom branched above. Leaves elliptic to ovate, 3-5 cm long, 1.5-2.0 cm wide, acuminate, smooth, succulent ; lower leaves forming a basal rosette, petioled, upper stem leaves not much reduced in size, short-petioled or subsessile; stip- ules much reduced. Inflorescence a terminal, peduncled cyme, flowers spiked along one side of the branches, the branches usually 2-4 cm long, flowers densely arranged along the axis; sepals 0.5-0.75 mm long, corolla white, 1.0- 1.5 mm long, somewhat funnel-form to urceolate; stamens 5, included; ovary 2-locular, styles 2, very short and fused at the tip to form a common stigma; bracts 1.0-1.5 mm long, gradually tapering, usually succulent, Capsule 3-4 mm long, exserted, somewhat mitriform, many- seeded. In drier sites, pinelands, less commonly in moist soil, glades, endemic to southern peninsular Florida. COLLIER CO.: abundant on wet muddy edges of marsh; corolla white, East Henson Marsh, Feb. 14, 1946, L. J. Brass 15972 (GH); DADE co.: Royal Palm Hammock, June 29, 1915, J. K. Small, C. Mosier, G. Small (Gu) ; Everglades, 10 mi nw of Hialeah on route 27, common, associated with Asclepias incarnata; flowers white, August 24, 1951, Grady Webster 4237 (GH); glades south of Long Pine Key, not common, April 24, 1952, W. B. Robertson, Jr. 211 (GH); LEE CO.: vicinity of Fort Myers, in pineland, May 22, 1916, Jeanette Standley 200 (GH); PINELLAS CO.: sandy soil near Maximo point, St. Peters- burg, Sept. 10, 1954, R. F. Thorne 15407 (GH). 30 Rhodora [Vol. 72 Cynoctonum succulentum appears strikingly different from the related, widespread C. mitreola because of its thicker, more succulent leaves, the larger ones often clus- tered towards the base forming a rosette, and because the leaves have very short petioles, or are subsessile, the base almost truncate, and because of its more restricted inflores- cence. Although C. mitreola (photo of type in GH!) is ad- mittedly a highly variable species, its leaves are more often lanceolate, acute, and definitely petioled, and never clus- tered at the base; in addition, it is often branched near the top, and the inflorescence is usually widely branched. The leaves are membranous, and cymes often arise from the lower axils. The two species appear to be ecologically dis- tinct: C. succulentum is found in pine barrens and drier soil more frequently than not, and C. mitreola is typically found in drainage ditches, swamps, and moist sites. 29. CYNOCTONUM MITREOLA (L.) Britt. Mem. Torr. Bot. Club 5: 258. 1894. Low ground, wet places, Fla to Va, Ark, Mex, WI. This is the most common species of Cynoctonum in southern Florida, and is the most variable, particularly in leaf shape and size. Apparent hybrids with C. sessilifolium are found in Florida, also. In practice, these putative hybrids are usually referred to C. mitreola. APOCYNACEAE Urechites The tropical genus Urechites is represented in the south- ern Florida flora by the extremely variable U. lutea (L.) Britt., a vine or scrambling shrub with bright green leaves and tubular yellow corollas, Two readily separable varieties can be identified, however, representing apparently ecotyp- ically differentiated races: Leaf undersurfaces, sepals, and follicles glabrous or barely pubescent. tteeeseeeeeeeeeeeeesssseeeeessesesaneeessssesensaesessessenssessesssastsaestesetsrsssensttensscs var. lutea IRR var. sericea URECHITES LUTEA (L.) Britt. var. LUTEA Bull. N. Y. Bot. Gard. 5: 316. 1907. 1970] Southern Florida — Long 81 Twining vines, generally in hammocks, mangroves asso- ciations, southern Florida, Florida Keys, West Indies. URECHITES LUTEA var. sericea R. W. Long, var. nov. A typo differt foliis tomentosis vel quidem pilosissimis ; calyce et pedunculis hirsutis, corollae tubo pilis mollibus. TYPE: vicinity of La Valle, Tortue Island, Haiti. Twining on shrubs to height of 15 ft., flowers yellow, thicket east of harbor. Dec. 28, 1928, Jan. 9, 1929, E. C. and G. M. Leonard. 11642. In Gray Herbarium. Urechites jaegeri Muell.-Arg. Linnaea 30: 444, 1860, pro parte. Urechites pinetorum Small, Addisonia 4: 21, pl. 131. 1919. Differing from the typical variety by having leaves tomen- tose underneath, or at least thickly pubescent; calyx and peduncle hirsute, corolla tube with soft hairs; fruits cov- ered with soft, silky pubescence. The new variety is abundantly distinct, at least in south- ern Florida, and apparently in the West Indies ; var. sericea frequently occurs in rocky often limestone soil, while var. lutea is found more frequently in moist soil. Morphological intermediates are found but they apparently are not com- mon. FLORIDA: DADE CO.: pinelands west of Coconut Grove, May 15, 1918, J. K. Small (GH); Coconut Grove, 1910, Miss C. Rotham (^); Homestead, pineland area between Homstead and Florida City near S. R. 27, June 4, 1966, R. W. Long 1674 (USF). BAHAMAS: ANDROS ISLAND, near Niccols Town, in pine yards and along roadsides, June 14-16, 1965, R. W. Long 1367 (USF). JAMAICA: sT. CATHERINE PARISH: Mt. Diablo, site of Blue mountain, August 21, 1965, H. A. Hespenheide 1379 (GH); Manchester, Red Mould Forest Reserve, s of Troy in open place, climber to 12 ft; corolla yellow, April 23, 1956, W. T. Stern 917 (GH); ST. ANN PARISH: Moneague, Jan. 23, 1952, F. W. Hunnewell 19786 (GH); limestone turn-around area alt. 1500, Minea area near Lydford p. o. Dec. 20, 1953, R. A. Howard and G. R. Proctor 13489 (GH); near Casteltou, June 26, 1915, W. Harris 12085 (GH). CUBA: ISLE OF PINES, swamp about 2 km due n of Nueva Gerona, Mar. 18, 1953, E. P. Killip, 43140 (GH); Buenas Aires, in brushland, July 10, 1929, F. M. Salvoza (GH); Soledad, Cienfuegos, Aug. 4, 1927, J. G. Jack 5230 (GH); rocky pasture, Soledad Cienfuegos, Sept. 2, 1927, J. G. Jack 5413 (GH); Havana, camino del Morro, 32 Rhodora [Vol. 72 Santiago de Cuba, June 1944, Hno. Clemente, 3689 (GH); Santa Clara, brushy field, July 13, 1936, L. B. Smith, A. R. Hodgdon, (GH); Oriente Province, Cerro de Miraflores, Cananova, July, 1942, Hno. León 21106 (GH). HISPANIOLA: SANTO DOMINGO: in thickets, May 13, 1929, E. L. Ekman, 12453 (4); MONET CRISTI: semiscandent plant, elevated coral reefs near Rio Yaqee del Norte, Oct. 23-24, 1946, R. and E. S. Howard 9584 (GH). CONVOLVULACEAE Bonamia Although Myint has pointed out that Stylisma and Bon- amia may be separated by means of differences in the shape of cotyledons, otherwise the two genera are closely similar and there appears to be no sound reason for maintaining them as separate taxa. Intermediate species serve to form a continuous pattern of variation between the two groups. BONAMIA abdita (T. Myint) R. W. Long, comb. nov. Stylisma abdita T. Myint, Brittonia 18: 107. 1966. Stictocardia A specimen of S. tiliifolia Hall. f. from Key West, Fla. in the Gray Herbarium suggests that this species may be naturalized in the Florida Keys or southern peninsular Florida. No other collections have been seen, however. BORAGINACEAE Heliotropium Three heliotropes are found in southern Florida, H. an- giospermum Murr., H. curassavicum L., and H. polyphyllum Lehm. The latter species is represented by two readily dis- tinguished varieties: Stems erect, often strict. .......1.222.......... n... var. polyphyllum Stems spreading-decumbent or prostrate. .................... var. horizontale HELIOTROPIUM POLYPHYLLUM Lehm, var. POLYPHYLLUM in Neue Schr. Naturf. Ges. Halle iii, II, 9. 1817. including H. polyphyllum var. leavenworthii (Torr.) A. Gray, Proc. Amer. Acad. Arts & Sci. 10: 49. 1874. This is a tropical species that occurs in coastal areas, low 1970] Southern Florida — Long 33 hammocks, and pineland in moist soil; var. leavenworthu is merely the form with yellow rather than white corollas, said to be "strict", but not appearing to be worthy of tax- onomic distinction from the typical variety. HELIOTROPIUM POLYPHYLLUM var. horizontale (Small) R. W. Long, stat. nov. Heliotropium horizontale Small, Bull. N. Y. Bot. Gard. 3: 435. 1905. TYPE: pinelands between Cutler and Camp Longview, Nov. 1903, Small & Carter 742 (NY, photo GH!) DADE CO.: pinelands about Cox hammock, June 24, 1915, J. K. Small 6579, C. Mosier, and G. Small (GH); LEE CO.: Seacoast, Sanibel, July-Aug 1900, A. S. Hitchcock 221 (GH); MANATEE CO.: north end of Longboat Key near Long Beach, 15 miles nw of Sarasota, sprawling perennial herb in sand, corolla white with yellow center, Aug. 1, 1950, R. L. Wilbur and G. L. Webster 2510 (GH); PALM BEACH CO.: north of Palm Beach, suffruticose, 1-2 ft. corolla yellow, April 29, 1920, Alfred Rehder 853 (GH); PINELLAS CO.: Fort De Soto Park, Mullet Key, south of Gulfport, beaches north end of key, stems depressed, trailing, cccasional, corolla white, Oct. 5, 1963, O. Lakela 26527 (GH, USF); SARASOTA CO.: Siesta Key, Anne E. Perkins, Jan. 28, 1943 (GH). This variety appears to be a maritime ecotype distin- guished by its prostrate spreading habit and apparent pref- erence for habitats very near the coast in peninsular Florida where it is apparently endemic. In addition to the three species cited, two other helio- tropes may occur in southern Florida: H. fruticosum L. (H. phyllostachyum Torr., H. myosotoides Chap.) was collected by Blodgett at Key West, and later by Garber and by Palmer. No recent collections have been seen, however. Heliotropium indicum L. (Tiaridium indicum (L.) Lehm.), a hispid annual native to the Old World may occur in dis- turbed sites in southern Florida, but apparently it is un- common. VERBENACEAE Lantana Lantana ovatifolia Britt, is distinguished from other spe- cies in southern Florida by its unarmed stems and heads 34 Rhodora [Vol. 72 with bracts not formed into an involucre. Two well-marked varieties occur although the typical one appears to be less common. Stems erect; leaves up to 9 CM long. .............. a... var. ovatifolia Stems decumbent or reclining; leaves up to 4 cm long. .. var. reclinata LANTANA OVATIFOLIA var. OVATIFOLIA, Britton in Bull. N. Y. Bot. Gard. 4: 123. 1905, TYPE: Eight mile Rocks, Grand Bahama, Britton & Mill- spaugh 2450 (NY). DADE CO.: sand dunes near head of Indian creek, opposite Miami, Nov. 22, 1912, J. K. Small 3872 (GH); pinelands, Miami, May 8, 1930, F. Duckett (GH); LEE CO.: frequently flowers yellow, shrub 160 cm tall, May 11, 1954, G. R. Cooley 2674 (GH); PINELLAS CO.: five miles south of Port Richey, white sandy scrub bordering inlet from Gulf, slash pine, saw palmetto, scrub oaks, Aug. 1960, J. D. Ray 9988 (GH, USF). LANTANA OVATIFOLIA var. reclinata R. W. Long, var. nov. A typo differt ramis prostratis, foliis parvis, plerumque 1-4 cm longis fructu 2.5-3.5 mm longis. TYPE: FLA, DADE CO.: Fuch's hammock, Homestead, corolla yellow, plant decumbent. George R. Cooley 9324, R. J. Eaton, F. C. Craighead, O. Lakela, Oct. 14, 1962. In Gray Herbarium, isotype in USF. L. depressa Small, in Bull. N. Y. Bot. Gard. 3: 436. 1905. Shrubs with reclining or decumbent stems, leaves mostly 1-4 em long, elliptic to ovate, serrate, scabrous, pubescent beneath, fruit 2.5-3.5 mm long, pinelands, endemic to south- ern Florida. DADE CO.: corolla yellow, leaves dark green above, open field, Home- stead, Sept. 17, 1952, J. Hardin, W. Humphrey, W. Duncan 14049 (GH); pinelands about Sykes hammock, March 4, 1915, J. Small 5667, C. Mosier, E. Small (GH); rocky pinelands near Goulds, April 13, 1942, D. E. and M. S. Eyles 8082 (GH); low calcareous ground, 4 miles sw of Royal Palm hammock, May 22, 1925, E. Palmer, (GĦ); Brickell hammock, Miami, April 27, 1920, A. Rehder 811 (A); pine- lands, Long Pine Key, common, April 14, 1952, W. B. Robertson 156 (GH). In recognizing this Lantana Small noted that to him it differed from L. camara L. in producing prostrate unarmed 1970] Southern Florida — Long 35 branches and smaller flowers. The plant appears to be more closely related to the West Indian species L. ovatifolia from which it differs chiefly in producing reclining or prostrate branches, smaller leaves and somewhat smaller fruits. It is apparently endemic to a very small portion of extreme southern Florida judging from collections. RUBIACEAE Ernodea In Florida Small distinguished two species, E. littoralis Sw. and E. angusta Small, the latter separated by its sup- posedly narrower leaves, smaller flowers, and corolla pink or reddish rather than pinkish-white or white. Experience in the field suggests that the two taxa are connected by intermediates although they apparently are different eco- types. The typical variety generally occurs in the coastal areas, in sandy or rocky soil, while var. angusta is more commonly found inland in pinelands and in drier sites. The two may be keyed as follows: Leaf-blades linear oblong to elliptic, calyx mostly 7-8 mm long. ........ MAI S S s var. littoralis Leaf-blades linear; calyx mostly 5-6 mm long. .............. var. angusta. ERNODEA LITTORALIS var. LITTORALIS Swartz, Prod. Veg. Ind. Oct. 29, T788; Corolla usually pinkish-white or white, coastal areas, southern Florida, Florida Keys, and West Indies. ERNODEA LITTORALIS var. angusta (Small) R. W. Long, stat. nov. E. angusta, Small, in Bull. N. Y. Bot. Gard. 3: 438. 1905. Corolla usually pink or reddish, usually less than 1 cm long, in pinelands, southern Florida Keys, and West Indies. TYPE: FLORIDA, DADE CO.: in pinelands between Cutler & Camp Longview, Nov. 1903, Small 870 and Carter (NY). DADE CO.: open sandy stretch of disturbed hammock and pineland, with remnants cf original vegetation north of Post Office, Homestead; fruits shiny, orange, July 19, 1964, O. Lakela 27299 (GH, USF); small trailing shrub, flowers red, Coconut Grove, Jan. 26, 1932, J. G. Jack (A); open pinelands, limestone outcrop near Rockdale, May 22, 1925, E. J. Palmer 27511 (A); Coconut Grove, Feb. 3, 1933, 36 Rhodora [Vol. 72 H. O'Neill 8158 (A); MONROE CO.: vicinity of Watson’s hammock, edge of pinelands along roadside, scrambling shrub, fruits yellow, March 25, 1958, W. L. Stern 304 and K. L. Chambers (GH); Big Pine Key, Feb. 4, 1940, R. J. Seibert 1289 (A). The relationship of Ernodea, a small genus of the West Indies principally in the Bahamas and Florida, with the Hispaniolan genus Isidorea A. Rich ex A. P. DeCandolle is apparent through intermediate species, such as I. leon- ardii Urban and E. littoralis. It is probable the two plants are congeneric, but until further field studies are made it is advisable to maintain the genera separately. Generally, the morphological variation of Isidorea appears to be broader than Ernodea, and could be further broadened to include the latter genus. Spermacoce Two species occur in southern Florida, S. tenuior var. floridana and S. tetraquetra. From examination of speci- mens, it appears doubtful that S. tenuior var. tenuior is present although it does occur in northern Florida. Our specimens are mostly procumbent or decumbent with small leaves, often 1-2 em long, annual, and with smooth hypan- thia. Typical S. tenuior plants have hirsute hypanthia and generally iarger, more ovate or elliptic leaves (photo, type GH!), It appears, however, that the two varieties are con- nected by morphological intermediates. Spermacoce tetra- quetra, our other species, is a distinctive form because of its generally larger habit, usually erect, and hirsute-white pubescent leaves and stems. Stems and leaves glabrous or nearly so. .... S. tenuior var. floridana Stems and leaves pubescent with whitish hairs. .......... S. tetraquetra SPERMACOCE TENUIOR L. var. floridana (Urban) R. W. Long, stat, nov. Spermacoce floridana Urban, Symb. Ant. 7: 550. 1913. Spermacoce keyensis Small, Fl. Fla. Keys 141, 155. 1915. Spermacoce floridana Gandoger, Bull. Soc. Bot. Fr. 65: 35. 1918. Spermacoce portoricensis A. Gray, Syn Fl. N. A. 1(2): 34. 1884, non Balbis = Hemidiodia ocimifolia. 1970] Southern Florida — Long 37 TYPE: in locis apertis arenosis Ins. Key West, Florida austr. Rugel 298, Feb. 1846 (isotype, GH!) DADE CO.: waste ground, Dec. 15, 1902, A. Fredholm 5642 (GH); Miami, door yards, A. H. Curtiss 1109 (GH); MONROE CO.: hammocks, Key West, March 20, 1915, J. K. Small 5990 and C. A. Mosier (aH); Key West, Jan. 7, 1892, J. H. Simpson 477 (GH); Key West, Aug. 1877, A. P. Garber (GH); Key West in umbrosis, Ins. Key West, Florida austr, Rugel, Feb. 1846 (cited in original description), (GH). SPERMACOCE TETRAQUETRA A. Rich. DADE CO.: hammocks between Miami and Coconut Grove, Nov. 26- Dec. 20, 1913, J. K. Small 4667 and G. K. Small (GH); Everglades National Park, near Homestead in Miami oolite and solution holes, hammock and pineland association, hirsute hairs whitish, coarse, Oct. 16, 1962, G. R. Cooley 9404, J. Eaton, F. Craighead and O. Lakela (GH); in hammocks between Coconut Grove and Cutler, Nov. 13-23, 1908, J. K. Small 1207, J. Carter (GH). CUCURBITACEAE Melothria The variation of Melothria pendula in Florida is partic- ularly difficult to understand taxonomically. Small described two new species from collections by G. V. Nash, and both of them are obviously closely related to M. pendula. Cog- niaux described several variants of M. pendula in his two monographs. In all instances leaf pubescence and texture, leaf blade and size and shape together with the morphology of the fruiting peduncles and fruit were all considered tax- onomically important. Variation of M. pendula in southern Florida is partic- ularly complex. Our common plants are small, rough-leaved forms that can be referred to M. pendula var. aspera; forms with thick leaves and stout peduncles and large fruits are referrable to var. crassifolia; much less commonly found is the typical variety with large, thin, pubescent leaves, At this time it is not possible to identify any ecological differ- ences between the varieties, and all are apparently con- nected by intermediates. The relationships may be seen in the following key: 1. Leaf blades thin, often 5-8 em wide, hirtulous. ........ var. pendula 1. Leaf blades thicker, often 2-4 cm wide, scabrous or pubescent with thick stout hairs. 38 Rhodora [Vol. 72 2. Leaf blades mostly 2-3 em wide, very scabrous on both sides, deeply lobed; berries globose. ................... eee var. aspera 2. Leaf blades mostly 3-4 cm wide, rough pubescent, shallowly lobed; berries oblong. ................ een var. crassifolia MELOTHRIA PENDULA L. var, PENDULA JACKSON CO.: vine, branches spreading on the ground and running up trunks of rough-barked trees; flood plain forest along Apalachicola River, w of Chattahoochee, Oct. 17, 1958, R. K. Godfrey 57835 (GH) ; LEE C0.: Ft. Myers, marshes, July-Aug. 1900, A. S. Hitchcock 107 (GH); PALM BEACH CO.: Jupiter Island, July 28, 1956, G. Cooley 4874, E. West and T. Daggy (GH). MELOTHRIA PENDULA var. ASPERA Cogn. in DC. Monog. Phan. 3: 587. 1881. incl. M. pendula var. microcarpa Cogn. DC. Monog. Phan. 3: 587. 1881. M. nashii Small, Bull. Torr. Bot, Club 25: 483. 1898. M. microcarpa Shuttlew. ex Small, Man. S. E. Fl. U. S. 1284. 1933. CITRUS CO.: sandy soil dominated by herbs, fruit oblong, July 12, 1962, O. Lakela 25162 (GH, USF); DADE CO.: glades farm lands south of Long Pine Key, common, April 23, 1952, W. B. Robertson 198, (GH); LEE CO.: Sanibel Island, climbing perennial of damp waste places, Feb. 23, 1954, G. R. Cooley 2519 (GH); PALM BEACH CO.: Kelsey City, sandy bank alone ditch, Dec. 4, 1920, Fannie Randolph 116 (GH). MELOTHRIA PENDULA var. CRASSIFOLIA (Small) Cogn. Pflan- zenreich 4, Fam. 275, I: 88, 1916. Melothria crassifolia Small, Bull. Torr. Bot. Club. 25: 483. 1898. TYPE: LAKE CO.: collected in vicinity of Eustis, stems creeping on ground, moist soil, June 1-15, 1894, G. V. Nash (isotype, GH!). COLLIER CO.: north of Deep Lake, route 29, Canal bank, and margin of hammock w side of the road, Oct. 1962 G. Cooley 9382 (GH, USF); DADE CO.: Cape Sable, April 13, 1923, A. Harris C23352 (GH); MANATEE CO.: ad ostia fl. Manatee, Florida, austr. occ. June 1845 legit Rugel 260. ASTERACEAE Kuhnia Examination of herbarium and field collections leads me to conclude that K. mosieri is another geographic variety 1970] Southern Florida — Long 39 of the widespread K. eupatorioides, and is closely related to K. eupatorioides var. pyramidalis Raf. The southern Florida population is narrowly endemic to Dade County; Shinner’s (1946) citation of a Chapman specimen from Aspalaga, Fla. is apparently an error, and the plant in question is referrable to var. pyramidalis. KUHNIA EUPATORIOIDES L. var. floridana R. W. Long, var. nov. A typo differt foliis linearibus, minus quam 3 mm latis, plerumque minus quam 3 em longis. TYPE: DADE CO.: Miami, Florida, May 13, 1904, S. M. Tracy 9046. In Gray Herbarium. Stems simple, up to 7 dm tall, leaves linear to less com- monly oblong-entire, scabrous-pubescent above, densely resin-dotted beneath, larger blades 2-4 em long, 1-3 mm wide, the upper leaves gradually becoming smaller. In- florescence corymbose or corymbose-paniculate; involucres 8.5-10.5 mm high, florets 9-13, corollas 6-6.5 mm long, achenes 3.5-4.5 mm long. Kuhnia mosieri Small, Man. S. E. Fl. 1329 and 1508, 1933. Two names, K. eupatorioides var. gracilis T. & G. and K. paniculata Cass. have been applied incorrectly to slender forms of K. eupatorioides var. pyramidalis and to K. eupa- torioides var. floridana. DADE CO.: rocky pinelands between Bay Biscayne and the Everglades, A. H. Curtiss 1187 (GH); open grassy area, near intersection of Redlands and 308th St., Homestead, frequent in unbroken ground, Aug. 10, 1963, O. Lakela 26137 (USF); Sunset Dr. and SW 71 Ct., Miami, vacant lot, pineland, sandy soil, S26, T54S, R40E, Sept. 5, 1968, G. Avery 459 (USF). Aster The genus is represented in southern Florida by nine species, Small described a number of new species from this area, but examination of collections has required some nomenclatural changes. ASTER CONCOLOR L. var. simulatus (Small) R. W. Long, stat. nov. 40 Rhodora [Vol. 72 Aster simulatus Small, Bull. Torr. Bot. Club 51: 388. 1924. TYPE: DADE CO., FLA., Silver Palm Schoolhouse, Nov. 26, 1913, J. K. Small in NY Herbarium, Small stated that this plant is related to A. concolor but differed by having upper leaf-blades scale-like, smaller, and more copiously puberulent bracts and involucres; A. con- color var. concolor has leaves ovate-lanceolate to lanceolate and stems silky pubescent. BROWARD CO.: pinelands w of Haledale, Dec. 16, 1923, J. Small 11090, J. B. DeWinkeler, and C. Mosier (GH); DUVAL CO.: dry pine barrens near Jacksonville, Dec. 1, 1898, A. H. Curtiss 6484; HILLSBOROUGH co.: ca. 1 m ne of Fletcher Ave., and Morris Bridge Rd, in fencerow, Dee. 2, 1962, O. Lakela 25598 (GH, USF); ORANGE CO.: dry woods, Winter Park, May 19, 1927, F. W. Hunnewell 10,533 (GH). ASTER PATENS Ait. var. floridanus R. W. Long, var. nov. A typo differt caulibus et foliis scabris; capitulo circa 2 cm latis. TYPE: COLLIER CO., FLA., abundant on pine and cabbage palm islands in Big Cypress, Brown Lake, 15 mi south of Immokalee; 60-80 cm high; leaves scabrous, heads about 2 em diameter, showy, rays pale violet, Dec. 12, 1945, L. J. Brass, 15780. In Gray Herbarium, Aster fontinalis Alexander ex Small, Man. S. E. Flora 1382. 1933. This variety is definitely related to Aster patens, but it represents a race that is endemic to peninsular Florida. It is separable from the typical form on the basis of scabrous stems and leaves, and larger heads. FLAGLER CO.: west of Bunnell, tall branching, Nov. 30, 1943, Oakes Ames 410 (GH); near Bunnell, rays deep lilac, Nov. 30, 1943, Oakes Ames 406 (GH) ; LEON CO.: open old field pine woods, 2 mi e of Talla- hassee, Nov. 4, 1955, R. K. Godfrey 54305 (GH); MADISON CO.: dis- turbed roadside sands bordering long leaf pine turkey oak woods, 4 mi w of Madison, April, 1957, R. Kral 4590 (GH). ASTER TENUIFOLIUS L. var. aphyllus R. W. Long, var. nov. A typo differt foliis linearibus ad subulilinearibus, demin- utiis vel destitutis super; caulibus tenuibus. TYPE: HILLSBOROUGH CO, Florida, northwest of Tampa, 1970] Southern Florida — Long 41 south of State Route 580 and west of Rocky Creek, Dec. 24, 1962, O. Lakela 25610. In Gray Herbarium, isotype in USF. A. bracei Britton ex Small, Fl. Miami 190, 1913, type, New Providence, Britton and Brace 394, in NY. This variety resembles A. tenuifolius var. tenuifolius (photo, GH!) in producing stolons, in general habit, and in ecological preferences. It differs in producing smaller, much reduced linear or linear-subulate leaves that are apparently absent in upper stems; it also has smaller heads. Both varieties, however, are connected by intermediate forms. Apparently A. tenuifolius var. aphyllus represents a West Indian-Florida population segregate of the more northern A. tenuifolius. BAHAMAS: Eleuthera, palmetto-land near airstrip ca 2.5 miles n of Rock Sound, 76 degrees 10 min W, 25 degrees, 52 min N common; rays white or pink, July 27, 1960, G. Webster and T. Williams 10719 (GH); Grand Bahama Island, July 1904, G. Allen 17 (GH); New Providence, pine barrens, in bogs, Feb. 28, 1888, Eggers 4311 (GH); New Providence, saline rocky flats, near Nassau, Feb. 18, 1903, A. H. Curtiss 84. FLORIDA: COLLIER CO.: plentiful on pine and cabbage palm islands in Big Cypress, erect 30-70 cm, leaves somewhat fleshy, heads little more than 1 cm diam, Brown Lake 15 mi s of Immokalee, Dec. 12, 1945, L. J. Brass 15775; HILLSBOROUGH CO.: about 6 mi se of Tampa, weedy waste flat formerly in longleaf pine-cabbage palmetto, Oct. 28, 1960, J. D. Ray 10573 (intermediate to typical var.); MONROE CO.: Big Pine Key, Feb. 26, 1936, E. P. Killip 31621; DADE CO.: sand dunes near head of Indian Creek, opposite Miami, Nov. 22, 1912, J. K. Small 3879 (GH); PINELLAS CO.: sandflats near Pasadena Golf Course, Gulfport, Dec. 21-28, 1949, R. F. Thorne 9401 (GH); WAKULLA CO.: brackish marsh, St. Marks Wildlife Refuge, Oct. 6, 1955, R. Kral and R. Godfrey 54084. ASTER DUMOSUS var. SUBULAEFOLIUS T. & G., Fl. N. A. 2: 128. 1841, incl. A. simmondsii, Small Fl. Miami 190, 200. TOTIS. A. sulznerae, Small, Fl. Miami 190. 1913. A. dumosus var. gracilipes Wiegand, Rhodora 30: 166. 1928. Small's two species represent mixtures of smallish, stiff, linear-leaved forms, and larger more lax-leafy forms, judg- ing from his annotations. The former may be referred to 42 Rhodora [Vol. 72 A. dumosus var. subulaefolius for which specimens are cited below. The latter, occurring less commonly in southern Florida, may be referred to A. dumosus var. coridifolius (Michx.) T. & G. DADE CO.: Everglades w of Peters, Nov. 20, 1916, J. K. Small 7922, (GH); New River canal beyond head of New River, Nov. 11-25, 1913, J. Small 4431 and G. Small (GH); Royal Palm hammock, Feb. 20, 1915, J. K. Small 5428 and E. Small (GH); Deering Snapper Creek hammock, Dec. 10, 1919, J. Small et. al. (GH); VOLUSIA CO.: 1 mi w of Ormond, in open field, Nov. 23, 1943, Oakes Ames 11 (GH). Flaveria The evidence for considering F. latifolia as representing stabilized diploid hybrids in southern Florida has been pre- sented elsewhere (Long & Rhamstine, 1968). The correct citation for the hybrid was not given, however, and is given here for validation. FLAVERIA X LATIFOLIA (J. R. Johnston) Rydb. (pro sp.) (F. floridana X F. linearis) F. linearis var. latifolia J. R. Johnston, Proc. Amer. Acad. Arts Sci. 39: 289. 1903, (pro parte, including the type). F. latifolia (J. R. Johnston) Rydb. North Amer. Flora 34: 145. 1915. (pro parte). Heterotheca The transfer of Chrysopsis to Heterotheca (cf. Shinners, 1951; Wagenknecht, 1960) requires a number of new com- binations and nomenclatural revisions in Florida species. HETEROTHECA hyssopifolia (Nutt.) R. W. Long, comb. nov. Chrysopsis hyssopifolia Nutt., Jour. Acad. Natural Sci- ences, Phil. 7: 67. 1834. The species was described by Nuttall from a specimen collected in “West Florida" as smooth with narrow or linear leaves. The specimens from southern Florida are not so leafy as the northern and west Florida plants, but otherwise they are very similar. BAY CO.: coarse sand, scrub oak barren, Tyndall Field, Military Reservation, Oct. 15, 1955, R. K. Godfrey 54182 and R. Kral (GĦ); GULF CO.: Apalachicola, Chapman Herb. 1959, dry pine barrens, 1970] Southern Florida — Long 43 Sept. 18, 1895; HARDEE CO.: common on dry prairie; apex of bracts often purplish; rays and disk yellow, about 3 mi e of Zolfo Springs, July 23, 1949, L. J. Brass 20613 (a good example of the south Florida race) (GH); HERNANDO CO.: scrub-oak and sand ridge, 4.5 mi n of Brooksville, Oct. 16, 1950, R. K. Godfrey 50858 (GH); INDIAN RIVER CO.: dry pine barrens, A. 'H. Curtiss 1364 (GH); dry pine barrens, Melbourne, Aug. 3, 1896, A. H. Curtiss 5736. HETEROTHECA HYSSOPIFOLIA (Nutt.) R. W. Long var. sub- ulata (Small) R. W. Long, stat. and comb. nov. Chrysopsis subulata Small, Man. Fl. S.E. U.S. 1338. 1933. TYPE: between Avon Park and Sebring, Florida, J. K. Small 11495, in NY Herb. The variety is similar to the typical one except the bracts of the involucre are prolonged and have conspicuous, curved, or bent, caudate tips. The leaf blades are somewhat broader than the typical plant, but they are very similar otherwise. Apparently H. hyssopifolia var. subulata is another of the endemic races of central Florida, although specimens may be found farther north and south in the state. FRANKLIN CO.: pine savannah !4 mi south of Ochlockonee River by U. S. 319, T 6S, 3W, Sec. 2, Oct. 3, 1954, J. B. Morrill and R. K. Godfrey (GH); HIGHLANDS CO.: scrub between Avon Park and Sebring, July 17, 1924, J. K. Small, J. W. Small, and J. DeWinkeler, (topotype) (GH); INDIAN RIVER CO.: dry pine barrens near Eau Gallie, A. H. Curtiss 1364; ORANGE CO.: dry pine barrens, July 7, 1902, A. Fredholm 5356; POLK CO.: pine palmetto flatwoods, near the southern end of Crooked Lake, Aug. 3, 1955, R. K. Godfrey 53835 and R. Kral (GH). HETEROTHECA GRAMINIFOLIA (Michx.) Shinners, var. traceyi (Small) R. W. Long, stat. and comb. nov. Chrysopsis traceyi Small, Fl. S. E. U, S. 1182, 1339. 1903. TYPE: Tracy, Pls. Gulf St. 7713. In herb. Ny. This variety is distinguished by its longer ray flowers that are about 1 cm long, and by lower leaf blades that are much elongated, Otherwise, it is similar to the typical variety; H. graminifolia var. traceyi is endemic to penin- sular Florida, especially southern Florida. COLLIER CO.: Big Cypress, 6 mi w of Miles City, frequent on low pinelands, leaves silvery, flower heads up to 3.5 em in diam, florets a rich yellow, Jan. 14, 1946, L. J. Brass 15883 (GH); DADE CO.: Krome hammock near Homestead, April 11, 1924, J. K. Small 11,162, 44 Rhodora [Vol. 72 J. DeWinkeler, C. Mosier, (GH); FLAGLER CO.: south of Bunnell on Route 4, on banks of railroad ditch in full sun, heads up to 3.5 cm across, all plant prevalently dwarf, Dec. 6, 1943, Oakes Ames 233; MONROE CO.: Big Pine Key, rays bright yellow, 6-10 mm long, pine- palm woods northeast of Inn, Nov. 25, 1951, E. P. Killip 41459 (GH). HETEROTHECA floridana (Small) R. W. Long, comb. nov. Chrysopsis floridana Small, Fl. S. E. U. S. 1185, 1339. 1903. Chrysopsis mariana var. floridana (Small) Fern. Rho- dora 39: 455. 1957. TYPE, Pl. Gulf St. Tracy 7344, Bradenton, Fla. In NY herb. Isotype GH! This is a distinctive species with heavy pubescence, tomentose leaves that are spatulate near the base and ovate to ovate-lanceolate near the top of the stem. It is endemic to Florida, apparently to the lower Gulf coast and presum- ably in southwestern Florida. The species appears to be related to H. scabrella T. & G. but since so few collections have been made, it is not possible to place the taxon def- initely in relation to other species of the genus. HILLSBOROUGH CO.: near Ruskin, in openings of white sand in scrub oaks and Ceratiola, low flat terrace south of Little Manatee River Bridge, rays golden yellow, glandular on phyllaries and peduncles. Plants suffrutescent, perennial, Nov. 11, 1961, O. Lakela 24826 (GH, USF). Cirsium Cirsium vittatum was described by Small as differing from C. pinetorum (= C. horridulum Michx.) in having narrow, elongated, basal leaf blades which are not pin- natifid, and anthers twice as long as the filaments. The plants are often smaller than the typical variety, and are endemic to southern Florida. Although corollas are usually cream-colored or yellowish, they may be purplish. The two races apparently hybridize and intergrade freely, and may occur together in the same area. A third taxon, C. nuttallii (DC.) A. Gray, morphologically distinct with heads not involucrate, also occurs in southern Florida. 1970] Southern Florida — Long 45 CIRSIUM HORRIDULUM Michx. var. HORRIDULUM Cirsium smallii Britton, Britt. & Millsp. Bahama Fl. 458, 1930. Carduus smallii (Britt.) Ahles, Jour. Elisha Mitchell. Sci. Soc. 90: 173. 1964. DADE CO.: southwest of Homestead, Everglades National Park, com- mon throughout, Mar. 30, 1963, O. Lakela 25736 (leaves heavily tomentose) (USF); HILLSBOROUGH CO.: seasonally wet pineland & palmetto 2.4 m nw of Trout Creek, April 17, 1964, O. Lakela 27156 and A. Burdette (USF); MONROE CO.: Big Pine Key, common in flat pinelands, growing with C. vittatum, Nov. 13, 1964, O. Lakela 27897 (USF); PINELLAS CO.: cleared roadside largely of white sand, Jct. US 19, and Fla 52, May 23, 1963, O. Lakela 26015 (USF). CIRSIUM HORRIDULUM var. vittatum (Small) R. W. Long, stat. and comb. nov. Carduus vittatum Small, Bull. N. Y. Bot. Gard, 3: 439. 1905. Cirsium vittatum Small, Man FI. S. E. U. S. 1483. 1933. COLLIER CO.: Big Cypress, Fahkahatchee, south of Miles City, glade and cypress margin, Mar. 6, 1965. O. Lakela 28197 and R. W. Long (intermediate between C. horridulwm and C. vittatum) (USF); MON- ROE COo.: Big Pine Key, Watson's hammock and surrounding pine- land, Aug. 6. 1966, R. W. Long 2099 (USF); Everglades National Park, near Homestead, G. R. Cooley 9419, et. al. Oct. 16, 1962 (GH); Everglades National Park, southwest of Homestead, outcrops of oolite, common, Mar. 30, 1963, O. Lakela 25720 (USF). DEPARTMENT OF BOTANY AND BACTERIOLOGY UNIVERSITY OF SOUTH FLORIDA TAMPA, FLORIDA 33620 LITERATURE CITED CLARK, R. B. 1942. A revision cf the genus Bumelia in the United States. Ann. Mo. Bot. Gard. 29: 155-182. CRONQUIST, A. 1945, Studies in the Sapotaceae, III. Dipholis and Bumelia, Jour. Arnold Arb. 26: 432-471. ErIcKson, R. O. 1943. Taxonomy of Clematis, section Viorna. Ann. Mo. Bot. Gard. 30: 1-62. GRAY, A. 1895. Synop. Fl. N. Amer. I, pt. 1, 1-208. New York. Lonc, R. W., O. LAKELA, and R. BROOME. 1969. Some preliminary statistics of the Flora of southern Florida. Rhodora (in press). and E. RHAMSTINE. 1968. Evidence for the hybrid origin of Flaveria latifolia (Compositae). Brittonia 20: 238-250. 46 Rhodora [Vol. 72 SHINNERS, L. H. 1951. The North Texas species of Heterotheca, including Chrysopsis. Field & Lab 18: 66-71. SMALL, J. K. 1903. Flora of the Southeastern U. S. New York. 1913a. Flora of the Florida Keys. New York. 1913b. Flora of Miami. New York. 1933. Manual of the Southeastern Flora. Univ. of N.C. Press, Chapel Hill. WAGENKNECHT, B. L. 1960. Revision of Heterotheca, Section Heterotheca (Compositae). Rhodora 62: 61-76; 97-107. WILSON, K. A. 1961. The genera of Myrtaceae in the Southeastern United States. Jour. Arnold Arb. 41: 270-278. NOTES FROM THE PRINGLE HERBARIUM III F. C. SEYMOUR In the second installment of Notes from the Pringle Her- barium (Rhodora 69: 377-380. 1967), reference was made to “other names left for subsequent mention." Some of these are recorded here. 1. Potamogeton Berchtoldii Fieber var. lacunatus (Hag- stróm) Fern. Growing in stagnant water, 25 June 1966, Alpine Heights, Warren, Vermont, Frank C. Seymour 23,883 (vT). This variety, rather infrequent at best, appears to be new to Vermont. The name, Alpine Heights, though rather pretentious, seems well justified. Near by, at the water's edge was Carex canescens L. var. canescens, (Seymour 23,744) (Seymour Herbarium) a species which M. L. Fer- nald (Gray's Man. ed. 8, p. 313) describes as “circum- boreal.” Not far away (1/4 mile) is a station where Listera auriculata Wieg. has been collected. 2. Anthoxanthum odoratum L. f. giganteum P. Junge was growing in a sugarbush in Jericho, Vermont, 15 Aug. 1968, George B. Keiser & Frank C. Seymour 27,058 (vT). This form, growing side by side with f. odoratum (Seymour 21,056, (VT)), presented such a striking contrast that they seemed like different species. 3. Bromus racemosus L. This species, so far as I can ascer- tain, was first collected in Vermont by Cyrus G. Pringle in 1880 in Charlotte. Prof. William D. Countryman has re- cently found it in Vernon, Vt. It was a surprise to find it, not by a roadside, but far from any road near Lake Bomo- seen in Castleton on 14 June 1968, on a field trip of the Vermont Botanical and Bird Club (Seymour 20,901 (vT)). It was among fragments of slaty rock with Potentilla ar- guta Pursh near a trail where many hikers have passed by. It seems rare enough to mention. 4. Eragrostis multicaulis Steudel, growing along a road- side, 15 Aug. 1968, in Jericho, Vermont, Frank C. Seymour & George B. Keiser 27,054 (vT). This grass has become a 47 48 Rhodora [Vol. 72 very common roadside weed in some parts of New England but has not been reported previously from Vermont. 5. Panicum dichotomiflorum Michx. Spreading from some locality of its earliest establishment in this country, another weed has reached Vermont. Two varieties, var. dichotomi- florum and var. geniculatum (Wood) Fern., were growing near together at the border of a parking lot and along a walk on the campus of the University of Vermont. There is nothing surprising about this collection except that neither variety was known before in Vermont. Var. dichot- omiflorum: Burlington, Vt., 30 Aug. 1968, Seymour 27,085 (VT) and 18 Sept. 1968, Seymour 27,269 (VT). Var. genicu- latum: same date and place as the last, Seymour 27,270 (VT). 6. Eleocharis tenuis (Willd.) Schultes. That this has not been reported previously from Vermont seems strange. lt was found growing on the shore of Hardwick Pond in Hard- wick, Vt., 13 Aug. 1966, Frank C. Seymour (24,513) and Susan C. Gates (Seymour Herbarium). Nearby in the water was found an abundance of another unusual species, Pota- mogeton pectinatus L., Sago. Although there are a number of other localities in Vermont where the latter has been found, this writer considers it quite uncommon. 7. Eleocharis nitida Fern. On a collecting trip when Mr. Charles E. Nichols was the delightful host, he drew special attention to an Eleocharis which looked unusual because of its capillary stems. It proves to be Eleocharis nitida Fern- ald from Enosburg, Vermont, Seymour & Nichols 26,503 (Seymour Herbarium). Dr. Henry K. Svenson has very kindly verified the identification, commenting “I believe it is new for Vermont." 8. Orchis spectabilis L. f. Willeyi forma nova, labio roseo. TYPE: St. Johnsbury, Vt., 22 May 1968, Herman D. Willey, s. n. (VT). Named in recognition of its collector, Herman D. Willey. M, L. Fernald in Gray's Man. ed. 8, p. 468, men- tions that the lip may be *rarely pink", but he does not give it à name. Mr. Willey sent to the Pringle Herbarium a beautiful transparency of the form with pink lip taken on 1970] Pringle Herbarium — Seymour 49 June 11, 1967. The next year he very kindly sent the speci- men, which is the type. Mr. Willey writes that he has been watching this form for several years and that the several plants continue constantly to have pink lips. 9. Salix cordata Michx. is a name not new to Vermont botany. E. J. Dole in the 3rd. ed. of the Flora of Ver- mont, published in 1937, lists S. cordata Michx. but omits Salix rigida Muhl. Not only formerly were these two con- sidered to be one species but still there is much doubt as to whether they are distinct. Since S. cordata differs more from typical Salix rigida than does Salix rigida var. an- gustata (Pursh) Fern., there is good reason to treat them as distinct. To distinguish these two species, see the clear description by M. L. Fernald in Rhodora 48: 33. 1946. Doubtless because S. rigida Muhl. was considered to be a synonym of Salix cordata Michx. (not S. cordata Muhl.), specimens of Salix cordata Michx. occurring in Vermont have not been reported. The following specimens from Ver- mont in the Pringle Herbarium belong in S. cordata Michx. Burlington, 24 July 1940, L. A. Charette 316; Leicester, 19 Sept. 1909, D. Lewis Dutton, s. n.; specimen without definite locality but probably collected in Middletown Springs, Vt., 4 May and 21 July 1902, Dana S. Carpenter. Var. abrasa Fern., also has been found in Roxbury, 2 Oct. 1968, Mar- garet F. Clark (VT). 10. Salix ridida Muhl. var. angustata (Pursh) Fern. Col- lected in Vt, Waterville, 1 June 1966, Seymour 23,801 (Seymour Herbarium). This also has not been reported previously from this state. 11. Salix X *subsericea (Andersson) Schneider has been found previously in Brandon, Vermont. A specimen which this writer believes is this hybrid was collected by him in Eden, Vt, at Eden Pond, 24 Sept. 1966, Seymour 25,250 (Seymour Herbarium). 12. Polygonum Persicaria L. var. ruderale (Salisb.) Meis- ner. How could a plant appeal more eloquently for recog- nition than did this plant, growing right in the crevices of the cement walk in front of the Pringle Herbarium? There 50 Rhodora [Vol. 72 were a number of specimens which persisted up to the time of frost in 1968. The first collection was made on 15 Oct. 1967 (Seymour Herbarium). A few more specimens (Sey- mour 27,083, (VT)) were taken on 29 Aug. 1968. This au- thor finds no earlier specimens or records of its presence in Vermont. 19. Lupinus polyphyllus Lindley. Roberta G. Poland (Mrs. Burdette) discovered a colony on the lower levels near the trail of Monadnock Mt., in Lemington, Vermont, on 24 June 1967. She collected a few specimens, one of which is in the Pringle Herbarium. On visiting the spot a few days later, the writer found many plants growing tall and rank, This plant, indigenous in northwest America, is well established in Lemington. 14. Pycnanthemum tenuifolium Schrader var. Laurencei varietas nova, foliis similibus Pycnanthemo tenuifolio Schrader sed caule copiose pubescente. TYPE: Massachu- setts, Acton, 28 July 1966, Laurence E. Richardson (VT). A specimen very kindly given me by Mr. Laurence E. Rich- ardson bears a strong resemblance to Pycnanthemum pilosum Nutt. in its densely hairy stem. However, the leaves are very narrow, the widest only 3 mm wide as in P. tenuifolium, In typical Pycnanthemum tenuifolium the stems are glabrous. That it is a hybrid of P. pilosum is unlikely in as much as P. pilosum is not known east of southern Michigan. Therefore I designate it as a new variety in honor of the collector Laurence E. Richardson. 15. Sonchus uliginosus Bieb. On the same trip with George P. Keiser mentioned above, a Sonchus growing by the road- side in Essex, Vermont, 15 Aug. 1968, attracted my atten- tion. Whether Sonchus uliginosus Bieb. had previously been found in Vermont I did not know until I returned to the Herbarium. These specimens, Seymour & Keiser 27,025 (VT), appear to be the only ones from Vermont. Specimens of these collections, in cases in which there is a duplicate, are being given to the New England Botanical Club. PRINGLE HERBARIUM BURLINGTON, VT. 05401 INFRASPECIFIC CLASSIFICATION IN THE CAROLINA FLORA ROBERT L. WILBUR* The taxonomic treatment of infraspecific variation is one of the more troublesome problems confronting the syste- matic botanist. Judging from numerous recent articles concerned with this subject, the topic is no less controversial among zoologists nor is it apparently any closer to satis- factory resolution. For example, Wilson and Brown (Syst. Zoo. 2: 100., 1953) “are convinced that the subspecies con- cept is the most critical and disorderly area of modern sys- tematic theory — more so than taxonomists have realized or theorists have admitted.” A comparison of the infraspecific taxa recognized in two recently completed floras of temperate North America will emphasize something of the diversity in current taxonomic practice. Steyermark (1963) recognized 517 subspecies or varieties within Missouri and an additional 297 forms. Rad- ford et al. (1968) included 174 subspecies or varieties with- in North and South Carolina but no forms. The Carolinas possess a total of 3360 recognized species compared to 2438 known from Missouri. Although the Carolinas together have an area approximately one fifth larger than Missouri, their greater number of species is perhaps better attributed to their greater diversity in topography, geology, climate and ecology than exists in Missouri. There is a ratio in the Carolina Flora of 18.7 species to every subspecies or variety recognized while the Missouri Flora has only 4.7 times as many species as it has subspecies or varieties. If one included all infraspecific taxa recognized in Steyer- mark’s Missouri flora, there is an accepted subspecies, variety or form for every three species included. It is safe to conclude that the greater richness of the Missouri flora in *Grateful acknowledgement is made to the National Science Foun- dation for a grant of research funds to Duke University (NSF-Grant 18799) which was held during the preparation of this paper. 51 52 Rhodora [Vol. 72 recognized infraspecific taxa in comparison to that of the Carolinas is due more to the difference in taxonomic out- look of the respective authors than it is to differences in the frequency of infraspecific variation within the two areas. It would not appear that either of the two taxonomic pub- lications is exceptional in their treatment of infraspecific variation but merely illustrate something of the diversity existing in current taxonomic practice. As has been sug- gested elsewhere (Wilbur, 1968), greater effort should be made towards achieving a greater degree of uniformity by taxonomists before such major works as the projected Flora North America and Flora Neotropica are prepared. Three infraspecific categories are most frequently en- countered in botanical taxonomic literature: subspecies, variety and form, It is no more possible to define the proper use of each of these taxonomic ranks than it would be to summarize completely the great diversity in current tax- onomic usage. The following statements perhaps sketchily outline the principal trends in botanical practice. 1) infraspecific taxa are incipient species and the tax- onomist can and should utilize the hierarchy of subspecies, varietas, and forma to present an estimate of how far the population has diverged towards the status of species. 2) it is futile to attempt categorizing the great array of variation encountered within most wide-ranging species, hence, one should restrict oneself to naming formally only recognizable populations occupying either different geo- graphic areas or ecological sites. a) Since the category variety has been applied to so many different kinds of variation from slight ge- netical variant, or even habitat modifications to the morphologically distinctive representative of the species in a major geographic and/or ecologic area, its continued scientific use is rendered undesirable since it is too imprecise. Consequently one should designate these geographically and/or ecologically recognizable units as subspecies — a practice 1970] Carolina Flora — Wilbur 53 which also has the added virtue of being in closer accord with the prevailing zoological procedure. b) The category variety ought to be employed for these geographically and/or ecologically recogniz- able units since it has been historically far more frequently employed botanically, Consequently the necessary bibliographic aud nomenclatural paper- work would be reduced to a minimum in contrast to the numerous changes required if one were to adopt subspecies — a category whose botanical usage has been just as varied and imprecise as variety. 3) Both subspecies and varieties are useful infraspecific categories for geographically and/or ecologically distinctive populations with the category subspecies employed for par- ticularly distinctive taxa while variety is to be applied to those somewhat less strongly delimited or, perhaps less sub- jectively, the subspecies might be employed as a collective taxon for a group of varieties apparently more closely re- lated to each other than to another variety or a group of varieties within the same species. It would appear that the taxonomic category forma has undergone a sharp decline in botanical esteem and as a re- sult there are far fewer minor variants such as color forms of flowers named in recent years than was the custom three or four decades ago. Taxonomists doubtless have wisely concluded that their task is overwhelming enough if they limit themselves to categorizing populations and dis- regard the sporadic, although often conspicuous, individual variations. Consequently one should not be disappointed that the various authors of the Carolina Manual did not emulate the author of the Flora of Missouri in naming formae. Certainly most taxonomists would agree that if a hier- archy of infraspecific taxa is utilized then the ranks em- ployed should indicate degrees of divergence in the named populations. It is illogical to call apparently equivalent populations subspecies in one genus and varietas in another. 54 Rhodora [Vol. 72 Yet this easy going practice is openly acknowledged by Munz (1959) in the preface of his “A California Flora.” The Carolina Manual apparently follows the same practice since subspecies are recognized only in those few genera (e.g. Nuphar, Hydrangea, Acer, Asclepias, and Phlox) in which the monographer, whose work was adopted, employed that category rather than variety (or rarely used both). The category subspecies as the sole infraspecific unit is far less frequently encountered in the literature of eastern North American botany than it is in the West but, unfor- tunately, that nomenclatural blight is spreading eastward with increasing momentum, Although doubtless not a precise standard of measure- ment compared to those employed by the Newtonian physi- cists, the “biological species concept” has introduced a use- ful level of objectivity in helping taxonomists to delimit species in sexually reproducing organisms, One might also hope that eventually the infraspecific categories might con- vey a more uniform indication of similar biological signif- icance. A small sample of the varietas recognized within the Carolina Flora will, I believe, disillusion anyone as to the biological meaningfulness of the category as employed. There, varieties are an extremely heterogenous assemblage. I would like to discuss some of them under the following headings. 1) In some cases the recognized varieties should not be recognized formally as they are either trivial genetical variants, modifications perhaps due to habitat or portions or a morphological continuum. 2) Other varieties are accepted or proposed which give evidence of being better considered as biological species. 2) And there are species possessing morphologically rec- ognizable populations with distinct geographical ranges which are treated as taxonomically indivisable. 1. EXAMPLES OF VARIETIES LACKING TAXONOMIC MERIT. a) Monotropsis odorata var. lehmaniae (Burnham) Ahles, Jour. Elisha Mitchell Sci. Soc, 80: 172. 1964. (Man- ual, p. 797) 1970] Carolina Flora — Wilbur 55 Baldwin's own observations (Rhodora 59: 259-262. 1957) coupled with the detailed study of Wolf (Amer. Midl. Nat. 8: 104-187. 1922) has, as summarized by Wood (Jour. Arnold Arb. 42: 65. 1961), “effectively disposed of M. leh- maniae Burnham as the immature fall phase of M. odorata". To my knowledge nothing has been published to refute the conclusions reached in the above papers. If new and con- tradictory knowledge is available, its publication would be of far greater biological interest than the reduction from specific to varietal status of what apparently has been shown to be the fall stage of growth of the spring flowering Mono- tropsis odorata. The striking fragrance of the flowers is detected only at anthesis in the spring and all other dif- ferences suggested as distinguishing the supposed two taxa are apparently equally transitional or ephemeral including the differences in the proportions of corolla tube and lobes. Ahles states in the Manual (p. 797) that M. odorata var. odorata flowers in February through April in the Carolinas and fruits in May and June, The var. lehmaniae flowers in the fall from September into November and is said to fruit in October and November. The supposedly diagnostic fea- tures employed by Ahles in the Manual are set forth as follows: Leaves yellowish to tawny, chaffy ; flowers fragrant ...... var. odorata. Leaves purplish or lavender, fleshy; flowers not fra- Oia PU E Iu audae MARI RE. var. lehmaniae. In the Carolina Manual (p. 795) only the typical element is mapped but maps of both are presented in the Atlas (Rad- ford et al. p. 88. 1965). b) Calycanthus floridus var. laevigatus (Willd.) T. & G. (Manual, p. 476). Nicely concluded (Castanea 30: 67. 1965) that within C. floridus there exist “two populations, or varieties, which are more or less distinguishable by the glabrous or pubes- cent undersurface of the leaf" with ‘‘each having a more or less distinct geographical range." His map (p. 70) shows a considerable overlapping range of plants with or without 56 Rhodora [Vol. 72 pubescence. Personal observations and Niceley’s own ad- mission that both the twigs and leaf undersurface of var. laevigatus could be slightly pubescent would suggest that the two are at least not “ideal” varieties. Populations ad- mittedly exist in which “both extremes of pubescence as well as various intermediates are present." It would appear that the differences were of a slight genetical basis and the taxonomic utility of recognizing such as taxa on the basis of existing knowledge seems highly questionable. It seems unlikely that we are concerned here with biologically dis- tinctive populations. Instead the pubescence types are ap- parently no more deserving of taxonomic recognition than the striking glaucous form of Fothergilla major L. (some- times segregated as F. monticola Ashe) or the non-glaucous form of Zenobia pulverulenta (Bartr. ex Willd.) Pollard (formerly often designated Z. cassinefolia (Vent.) Pol- lard), which now are both rightfully not allotted taxonomic recognition. c) Clethra alnifolia var. tomentosa (Lam.) Michx. (Man- ual, p. 793). A recent study (Jour. Elisha Mitchell Sci. Soc, 83: 82-88. 1967) demonstrated, at least to my satisfaction, that “there seems no basis whatsoever to recognize taxa of any rank" within C. alnifolia. The variation in amount of pubescence on the abaxial leaf surface ranges from glabrate or spar- ingly strigillose along the principal veins to so copiously stellate-tomentose that the lower epidermis is masked from view. That study also indicated that the reputed differences in pubescence of style, capsule size and orientation and shape of sepals which in the past had been thought to be correlated with the striking difference in foliar pubescence were either not diagnostic or based upon faulty observa- tions. The extremes of variation are so striking that it is not surprising to find taxonomists disinclined to submerge the tomentose individuals with the more abundant sparingly strigillose types. There seems, however, to be no biolog- ically meaningful criteria by which the pubescence types can be recognized as separate taxa. There is an earlier 1970] Carolina Flora — Wilbur 57 varietal name, C. alnifolia var. pubescens Ait., which should be employed by those who wish to persist in naming seg- ments of a continuum, 2. EXAMPLES OF VARIETIES BETTER TREATED AS SPECIES. a) Sabatia dodecandra (L.) B.S.P. (Manual, pp. 838-839). Ahles (Jour. Elisha Mitchell Sci. Soc. 80: 173. 1964) re- duced to varietal status without discussion the very distinc- tive Sabatia kennedyana Fern. (= S. dodecandra var. kennedyana (Fern.) Ahles) and S. bartramii Wilbur (= S. dodecandra var. coriacea (Ell) Ahles). Ahles also placed S. harperi Small, a synonym of S. foliosa Fern., in the syn- onymy of S. dodecandra var. dodecandra. Four taxa of which three were considered species in the most recent monograph of the genus (Rhodora 57: 1-23, 43-71, 78-104. 1955) were consequently reduced by Ahles to three varieties of a single species. It now appears probable that I erred in reducing S. foliosa to varietal status since it differs from S. dodecandra morphologically and possesses a distinctive range and a very different habitat, Perry's studies (1967) suggest that the chromosome number of the two taxa (S. dodecandra s.s. and S. foliosa) differs as well. Consequently I now believe all current evidence would favor the recogni- tion of four species in this complex and not three as I rec- ognized in 1955 and certainly not one species and three varieties as Ahles suggests, Sabatia bartramii and S. foliosa are largely sympatric but natural hybrids are unknown. They differ greatly both morphologically and apparently ecologically. Perry's extremely thorough work has shown that their chromosome numbers differ and that artificial crosses between the two result in vigorous but sterile progeny. These facts are certainly more suggestive of spe- cific than varietal status. Sabatia dodecandra s.s. is a plant of brackish marshes from Connecticut as far south at least as North Carolina while S. kennedyana is a plant of open stream banks and margins of fresh water ponds with a three-parted, disjunct range: 1) Nova Scotia, 2) Massa- chusetts and Rhode Island, and 3) North Carolina and northeastern South Carolina. Morphologically the two seem 58 Rhodora [Vol. 72 readily distinguishable. Natural hybrids are unknown. Again Perry has shown that the two taxa differ in chromo- some number and that their artificially produced progeny are sterile. To treat as does Ahles four taxa such as these as three varieties of a single species obscures their biological or evolutionary divergence, b) Utricularia inflata var. minor (L.) Chapm. (Manual, p. 969). The parenthetical inclusion of Linnaeus in the above tri- nominal is an error, for Linnaeus’ taxon is a very different species; Chapman’s varietal name was new and not a trans- fer. Reinert and Godfrey (Amer. Jour. Bot. 49: 213-220. 1962) have, I believe, admirably demonstrated that U. in- flata and U. radiata Small (= U. inflata var. minor Chapm.) are specifically distinct. According to Reinert and Godfrey’s thorough and beautifully illustrated study, the two taxa are sympatric throughout most of their collective range and often grow intermixed in the same pond. They found the two taxa to be “easily distinguishable” “on the basis of 5 hitherto-neglected, morphological characters .. . and on the basis of 3 reinvestigated quantitative criteria." The two taxa are not known to intergrade and the differ- ences were maintained in culture. Eight distinguishing characters between two taxa is in itself impressive to any- one not a numerical taxonomist but more significant in de- termining their status in the taxonomic hierarchy is the obvious and highly effective reproductive isolation existing between the two. Infraspecific classification is not intended to be applied automatically to taxa which are difficult for the biologist to distinguish but for populations which lack the hallmark of a species. Utricularia radiata clearly seems to be, as Reinert and Godfrey have demonstrated, specif- ically distinct; there seems to be ample indirect evidence in the lack of intermediates that the two are most effectively reproductively isolated. 3. SPECIES TREATED AS INDIVISIBLE IN WHICH INFRASPECIFIC TAXA OUGHT TO BE RECOGNIZED. 1970] Carolina Flora — Wilbur 59 Silene caroliniana Walt. appears in the Carolina Manual without designated infraspecific taxa. In fact S. caroliniana var. pensylvanica (Michx.) Fernald is placed in synonymy and the inclusion of Alabama in the southeastern distribu- tion of the species suggests that even S. caroliniana ssp. wherryi (Small) R. T. Clausen is synonymized since it is the only element of the complex known from that state. ame Map 1. Distribution of Silene caroliniana. S.c. var. caroliniana, large solid dots; S.c. var. pensylvanica, small solid dots; S.c. ssp. wherryi, open circles with small central dot. 60 Rhodora [Vol. 72 The infraspecific variation of S. caroliniana was the sub- ject of a very thorough study by Professor Clausen (Rho- dora 41: 575-584. 1939) whose findings have not been fun- damentally enlarged upon in the past three decades. In the following year Fernald transferred Clausen’s subspecies to varietal status without additional botanical comment. That paper (Rhodora 42: 239-276, 281-302. 1940) was prefaced, however, by a lengthy discussion of the proper and improper use of subspecies, variety and form, The tenor of Fernald’s arguments can be gleaned from the following quotation: “The modern fad of certain botanists, to substitute the here- tofore clear term subspecies, erroneously used and often misunderstood by them, for the long established varietas, as used correctly for more than two centuries, is. . . a practice which cheapens the status of true subspecies and makes for inaccuracy and misunderstanding." Hitchcock and Maguire (Univ. Washington Publ. Biol. 13: 1-73 1947) accepted Clausen's concept of S. caroliniana and also treated the three major variants as subspecies, They stated, however, that “in the absence of obvious inter- mediate material and confluence of characters, it is not im- probable that this (ssp. wherryi) might ultimately have to be recognized as a distinct species...” In passing it might be noted that their almost impossibly small maps with many different symbols are most confusing — so much so that at least two errors are apparent for S. caroliniana (Map 1) : Silene caroliniana ssp. wherryi is mapped for the inner coastal plain of North Carolina where it is not known and ssp. caroliniana is shown as occurring in the mountains of Tennessee where only ssp. pensylvanica has been found. Kruckeberg (Brittonia 16: 95-105. 1963) found, except- ing S. caroliniana and S. virginica, that “the species east of the Mississippi River have undergone sufficient genetic divergence to have developed salient morphological gaps reinforced by strong internal barriers to crossing." Artifi- cial crosses result in vigorous but sterile hybrids, Although S. caroliniana and S. virginica may be artificially crossed (and a few natural crosses are known) Kruckeberg felt it 1970] Carolina Flora — Wilbur 61 best to “defer to common regional floristic practice and keep them as separate species.” Kruckeberg stated that “crosses among the three sub- species of S. caroliniana are likewise highly fertile” but his table of crosses shows that only two subspecies were avail- able to him: ssp. wherryi and ssp. caroliniana. The identity of the last seemed unlikely as S. caroliniana s.s. is not known from central North Carolina, (In fact no specimen of S. caroliniana s.l. is yet known to me from Orange County, the alleged source of the collection). Another collection re- ported upon by Kruckeberg in an earlier paper (Madroiio 15: 205-215. 1960) as ssp. caroliniana from Franklin Co., N. Carolina is ssp. pensylvanica, unless it differs from all other collections known from that area. Kruckeberg felt that the subspecies of S. caroliniana “are still valid as they coincided with geographic and morphologically distinct variants within the species.” Therefore in spite of the hopes expressed by Clausen, biosystematic studies have not as yet provided a new dimension to our understanding of the com- plex. Such studies have apparently confirmed the fact that the three taxa are more closely related to one another than they are to the other eastern species and that the morpholog- ical differences are maintained primarily through geo- graphic isolation. Unsurprisingly enough my examination of more than eight hundred herbarium specimens* has confirmed the ex- istence of three morphologically distinguishable taxa within the species and reaffirmed the allopatry of each of these taxa. Unlike Clausen, who found that “intermediates do occur, indicating continuity in the series”, I found the three taxa to be remarkably clearly delimited. I was not as im- pressed with the differences in shape of the apex of the basal leaves as have been Clausen and others and found that the specimens could be better correlated geographically if *I am indebted to the curators of the herbaria listed below who loaned the specimens which made my study possible. The herbaria are indicated by their Index Herbariorum symbols: DUKE, GA, GH, KY, MO, NCU, NSC, NY, PAC, TENN, US, VBD, VPI and WVA. 62 Rhodora [Vol. 72 the importance of the shape of the apex was not weighed as heavily as the pubescence on the surface of the basal leaves. The difference in pubescence of the calyces seems to me to be more fundamental. I therefore feel that the dif- ference between the wherryi is greater than that between the eastern taxa, caroliniana and pensylvanica. Consequently neither rigid adherence to the category sub- species advocated by Clausen nor to that of varietas prac- ticed by Fernald would allow one to indicate the apparent greater evolutionary divergence between wherryi and the two eastern taxa in contrast to the lesser differentiation between caroliniana and pensylvanica. Silene caroliniana appears then to be an excellent example in which two levels of the taxonomic infraspecific hierarchy are required to express our present knowledge of the biological relationship. Key to the variants of Silene caroliniana. 1. Calyces densely glandular-pubescent and narrowly tubu- lar; primarily east of the Appalachian Mts. .................. 1. ssp. caroliniana. 2. Basal leaves rather densely spreading short-pubes- cent on both surfaces, mostly narrowly to broadly spatulate and obtuse „sss la. var. caroliniana. 2. Basal leaves glabrous on both surfaces, mostly nar- rowly oblanceolate and acute .. 1b. var. pensylvanica. 1. Calyces densely pubescent with white eglandular tri- chomes and broadly tubular; occurring west of the Ap- palachian Mts. 2. ssp. wherryi. 1. Silene caroliniana ssp. caroliniana Calyces, pedicels and upper portion of stems densely glandular-pubescent. Calyx narrowly tubular, Surface of basal leaves glabrous or spreading short-pubescent, Range: primarily confined to the Appalachian Mountains or to the area to the east of the Mountains i. e. the Atlantic drainage slope. There are two morphological variants of this subspecies and they ean best be recognized by the spreading short- pubescence on the surfaces of the basal leaves of the more 1970] Carolina Flora — Wilbur 63 southern element in contrast to the glabrous surfaces of the basal leaves of the northern populations. These seem clearly differentiated and, since I do not feel that variation in the apices of the basal leaves is evidence of hybridization or introgression but part of the expected range of variation, the morphologically delimited populations are clearly al- lopatric. la. S. caroliniana ssp. caroliniana var. caroliniana. Basal leaves usually broadly oblanceolate to spatulate and typically with a broadly rounded apex, often (0.5)1.5-9 cm wide, rather densely spreading short-pubescent on both sur- faces. Calyces 1.5-1.8(2.0) cm long. Blades of petals re- portedly 0.8-1.3 em long, claws slightly longer than the calyx. There has been considerable discrepancy in the reports concerning the morphological features of the supposed type in Walter’s collection in the British Museum. Fortunately the description is exceptionally full in Walter’s Flora for even the obtuse and pubescent basal leaves are mentioned. Hence there can be no doubt as to the taxon to which the name should be applied. Small (Torreya 26: 67. 1926) stated that the specimen in Walter’s herbarium “has the calyx distinctly though sparsely glandular.” Clausen (Rho- dora 41: 579. 1939) reported that Weatherby’s notes on the specimen in Walter’s Herbarium indicated “that the basal leaves of this specimen are oblanceolate and acutish and that the pubescence is most nearly matched by a collec- tion from .. . Missouri" (i. e. a specimen of ssp. wherryi). The reference to “pubescence” is not clear as to whether that of the basal leaves (as Clausen assumed) or that of the calyx is being indicated. In any event Walter's descrip- tion is unmistakable and would take precedence over any specimen in Waiter's collection that is not in accord with the written account just as Shinners (Castanea 27: 71. 1962) has shown for Bonamia aquatica (Walt.) Gray. lb. S. caroliniana ssp. caroliniana var. pensylvanica (Michx) Fernald, Rhodora 42: 260. 1940, 64 Rhodora [Vol. 72 S. pensylvanica Michx., Fl. Bor. Am. 1: 272. 1803. S. caroliniana ssp. pensylvanica (Michx.) Clau- sen, Rhodora 41: 580. 1939. Basal leaves usually narrowly oblanceolate and typically with an acute apex, usually 0.5-1.5 (2.0) cm wide, glabrous on both surfaces. Calyces (1.0)1.3-1.8 em long. Blades of petals reportedly 0.8-1.5 cm long, claws usually slightly longer than the calyx. 2. S. caroliniana ssp. wherryi (Small) Clausen, Rhodora 41: 582. 1939. S. wherryi Small, Torreya 26: 66. 1926. S. caroliniana var. wherryi (Small) Fernald, Rhodora 42: 260. 1940. Calyces, pedicels and upper portion of stems densely pubescent with tapering hyaline, eglandular, wide-spread- ing, pilose trichomes. Calyx more broadly tubular. Surface of basal leaves glabrous. Range west of the Appalachian Mountains in south central Ohio and north central Ken- tucky, southern Missouri and the northern half of Alabama. In contrast to the above contention that both subspecies and varieties can be used to advantage in expressing the degree of variation within a species, Raven (Contr. U.S. Nat. Herb. 37: 167-168. 1969) approvingly paraphrases H. L. Mason's belief that although “two of any three taxa will always be more closely related to one another than they are to the third" and questions whether it is “practical to recognize all such shades of relationship in formal tax- onomy." Obviously, although it would be impossible to ex- press hierarchially every shade of variation, I believe we can better express degrees of significant infraspecific varia- tion if we don't assume arbitrarily that only one infra- specific category is to be recognized. Although I have not personally worked with plants in which I have found a need for more than two, I would not arbitrarily suggest that two is the absolute upper limit. Raven also claims that the Nomenclatural Code “states explicitly that subspecies are the primary unit into which 1970] Carolina Flora — Wilbur 65 species are divided, and it is therefore incorrect to use any other infraspecific taxon, such as variety, for the primary division of a species." Such an interpretation is, in my opinion, a complete misrepresentation of the Code. Article 4 lists twenty two categories that usually suffice in classify- ing plants. It is true that subspecies is listed first in the descending series of categories after species just as sub- family is listed first after family and subgenus after genus. Surely one does not conclude from such a hierarchical listing that families can not be divided into tribes unless they are first divisible into subfamilies or that sections are admis- sible only in genera which have recognizable subgenera. Consequently there is nothing implicit or explicit in the Code to suggest that is is mandatory to use the category subspecies if only one infraspecific taxonomic level is em- ployed, DEPARTMENT OF BOTANY DUKE UNIVERSITY DURHAM, NORTH CAROLINA 27706 REFERENCES MUuNZ, PHiLIP A. 1959. A California Flora. Univ. Calif. Press. PERRY, JAMES D. 1967. Biosystematic Studies in the North Ameri- can Genus Sabatia (Gentianaceae). Unpublished dissertation, Duke University. RADFORD, A. E., H. E. AHLES and C. R. BELL. 1968. Manual of the Vascular Flora of the Carolinas. University of North Carolina Press, Chapel Hill. 1965. Atlas of the Vascular Flora of the Carolinas. N. C. Agri. Expt. Sta. Tech. Bull. 165. STEYERMARK, JULIAN A. 1963. Flora of Missouri. Iowa State Uni- versity Press, Ames. WILBUR, ROBERT L. 1968. The status of Hedyotis procumbens var. hirsuta (Rubiaceae). Rhodora 70: 306-311. Table I Comparison of Infraspecific Usage in Two Recent North American Floras Ssp. Species/ Species/ Species & var. Formae ssp. & var. ssp., var., formae The Carolinas: 3360 174 O0 18.7 18.7 Missouri: 2438 517 297 4.7 2.99 NEW PHANEROGAMS FROM THE ARID NEOTROPICS R. C. BARNEBY In course of field work in Mexico in connection with a projected revision of Dalea, and in routine identification at New York Botanical Garden, several undescribed phanero- gams have come to light. Descriptions and diagnoses of these are presented: Psoralea holosericea Barneby, sp. nov., P. mexicanae (L. f.) Vail affinis, ob pubem densam sericeo-tomentosam omnium fere partium solum cum ejus var. Trianae (Vail) Macbr. comparanda, sed ab hac (in Ecuadoria australi sympatrica) pube longiori (pilis ad 1.2-1.6 nec 0.5-1.1 mm usque longis) et praesertim calycis albo- nec fusco-hirsuti tertia parte longioris dente dorsali subduplo longiori petala subsuperanti absimilis. Verosimiliter frutex 1-2-metralis habitu P. mexicanae simillima, praeter foliorum paginam superiorem tantum secus nervos subap- presse pilosulam pilis patulis undique molliter cano-hirsuta; foliorum petiolus 1-1.5 em longus, foliola ovato-acuminata (3) 3.5-5.5 cm longa; racemi spiciformes paniculatim dispositi ineunti anthesi densiflori 3-3.5 em longi, + 1.5 cm diametro, breviter pedunculati et subsessiles; calycis barbato-sericei eglandulosi pube ablata membranacei viridis + 7 mm longi tubus ad anthesin 2.5-3 mm longus, dens dorsalis aliis paullo longior fere 4 mm longus petala aequans vel breviter superans; petala azureo-marginata, vexillum alaeque 5.7 mm, carina 5 mm longae; calyx fructifer verosimiliter accrescens cum legumine ignotus, ECUADOR. Azuay: locally common, km 83 out of Cuenca on road to Loja, alt. 3000 m, September 25, 1959, Bassett Maguire & Celia K. Maguire 44,307. — Holotypus, NY; iso- tvpi, K, US, USM. In general habit, in shape and size of leaves, stipules and bracts, and in form of calyx and petals, P. holosericea closely resembles P. mexicana, (L. f.) Vail as interpreted by Mac- bride (in Field Mus. Bot, 133: 358-360. 1943). Because of the dense, soft, hirsute (and on the stems also subtomen- tose) pubescence of all parts except the upper leaf surface which is pubescent only in lines along the principal nerves, the new species suggests P. mexicana var. Trianae (Vail) 66 1970] Phanerogams — Barneby 67 Macbride; but the vesture of var. Trianae, although similar in quality, is much shorter, and in the inflorescence differs in color. The bracts and calyces of P. holosericea are shaggy with long, soft, lustrous, white hairs, whereas those of var. Trianae are hirsutulous with shorter and stiffer, mostly ascending and black or fuscous (mixed with a few longer pale) hairs among which arise many stalked or sessile glands. However the presence of glands is a variable feature in the Andean psoraleas and cannot be relied on as a dif- ferential character. More important differences are found in the relative proportions of the calyx and petals. In P. mexicana (sens, lat.) the petals are +5 mm long, but at early anthesis project well beyond the small, relatively short- toothed calyx; in P. holosericea they are of nearly the same size but are included in the much longer calyx. In conse- quence the young flowering spike of P. holosericea appears substantially thicker than that of any form of P. mexicana. The differences may be summarized: Stem (at least upward), lower leaf-surface and inflores- cence densely silky-hirsute with spreading hairs up to 1.2- 1.6 mm long; calyx at anthesis + 7 mm long, pale green beneath the white vesture, eglandular or nearly so, the dorsal tooth +4 mm long, equaling or slightly surpassing Wie petals te |i os e ey ee P. holosericea Stem, lower leaf-surface and inflorescence hirsutulous with antrorsely ascending and spreading hairs up to 0.5- 0.9(1.1) mm long; calyx at anthesis +4.5 mm long, livid beneath the largely or partly black vesture, usually glandu- lar, the dorsal tooth 2.2-2.5 mm long, shorter than the petals. P. mexicana var. Trianae *...... Polygala (§ Adenophora) evolvulacea Barneby, sp. nov., P. macra- deniae Gray affinis sed caulibus gracillimis flexuosis et praesertim floribus subdimidio brevioribus, alis 2-2.7 (nec 4-5.5) mm longis carinae brevioribus nec aequilongis vel longioribus, ulterius ab ejus var. glanduloso-pilosa (Chod.) Blake cum qua pube parciuscula brevi congruit foliis anguste obovatis duplo latioribus absimilis. Caules gracillimi flexuosi debiles prostrati ac incurvi 1-2.5 dm longi ut folia pilis incurvis + 0.15 mm longis puberuli virides nunc pur- pureo-tincti; folia 2.5-4.5 mm longa secus ramulos sparsa nec conferta 68 Rhodora [Vol. 72 nec imbricata anguste obovata vel oblanceolata acutiuscula 1-1.9 mm lata glandulis pellucidis utrinque obsessa; racemi brevissime 1-4-flori folio suffuleranti superati; sepala ovata cymbiformia 0.6-0.9 mm longa 0.4-0.5 mm lata extus puberula; alae obovato-cuneatae obtusae 2.2.1 mm longae, 0.9-1.5 mm latae extus pilosulae, totae vel supra medium violascentes; carinae exsertae 2.6-3.6 mm longae luteae lamina oblongo-quadrata obtusa integra 1.3-1.7 mm longa; capsula semenque nisi arillus paullo brevior iis P. macradeniae similes. MEXICO. Tamaulipas: vertical bank of soft crumbling rotted limestone, +300 m (1000 ft.), brushy hillside on the first bench of Sierra Madre Oriental +6.5 km (4 mi) W of Antiguo Morelos, November 22, 1966, Ripley & Barneby 14,755. — Holotypus, NY; isotypi, CAS, GH, MICH, US. The center of diversity of Polygala subsect. Adenophora Blake (Cont. Gray Herb., New Ser. 42: 54. 1916, subse- quently treated as sect. Adenophora in N. Am. Fl. 25: 334-5, 1924) extends through central and southeastern San Luis Potosí into Nuevo León, on the limestones of the Sierra Madre Oriental and the contiguous plateau. All three spe- cies known to Blake are either endemic to this region or occur in it. Of the three, P. glandulosa H. B. K. and P. phoenicistes Blake have broadly obovate leaves and flowers much larger than those of P. evolvulacea, which is therefore likely to be confused only with P. macradenia Gray and in particular, because of its green, sparsely and shortly pube- rulent foliage, with the relatively southern and more nearly sympatrie P. macradenia var. glanduloso-pilosa already mentioned in the diagnosis. The new species differs from P. macradenia as a whole in its smaller, differently propor- tioned flower in which the wings are both absolutely and relatively shorter. Furthermore the pliantly flexuous, dis- tally incurved or trailing, very slender and sparsely leafy stems are markedly different from the crowded, stiff and usually erect, densely leafy tufts of P. macradenia. From var. glanduloso-pilosa, still known only from two Purpus collections from west of the Sierra Madre crest at over 1000 m in San Luis Potosí, it differs in its leaves of lance- obovate rather than almost linear outline, and in its habitat 1970] Phanerogams — Barneby 69 in a quite alien vegetational belt at only 300 m in the Sierra’s eastern foothills. Polygala ($ Eurhinotropis) erythrorrhiza Barneby, sp. nov. P. Tweedyi Britt. proxime affinis sed caulibus debilibus flexuosis ex ipso radicis carnosae rubro-aurantiacae collo (nec e caudice suffru- tieuloso) ortis, foliisque latioribus iis racemo oppositis ellipticis membranaceis 2-4 (nec linearibus vel lineari-ellipticis 1-1.5 mm latis absimilis. Caules graciles omnino herbacei (1) 1.5-3.5 dm longi, ut folia pilis mollibus incurvis minutim densiuscule puberuli virides; folia nisi ima et summa breviora subhomomorpha lanceolato-elliptica vel ellip- tica acuta mucronata margine revoluta pinnatim paucinervia, saltem media 1-2.2 cm longa 2-5 mm lata; racemi oppositifolii subsessiles geniculati 5-12-flori; flos, capsula semenque fere ut in P. Tweedyi; alae pallide roseae glabrae; petala interiora basin versus ciliata, carinae rostro obtusissimo vix 0.5 mm longo. MEXICO. Guerrero: hanging down out of crevices of soft gypseous-conglomerate rock face, low cliff + 18 km (11 mi) S of Iguala, 870 m (2600 ft), November 9, 1964, Ripley & Barneby 13,737. Holotypus, NY; isotypi, CAS, K, MICH, US. The small-flowered herbaceous polygalas of subsect. Eurhinotropis Blake (Cont. Gray Herb., New Ser. 42: 71. 1916, raised to rank of sectio in N. Am. Fl. 25: 310 (key) + spp. 99-103 incl. 1924) are a critical group, all of the more widely dispersed members being variable in pubescence of the foliage and flower-parts as well as in size of the individ- ual flower. Specific lines are hard to draw, for the available differential criteria are seldom absolute. The center of spe- ciation lies on the arid altiplano of northern Mexico and ad- joining United States; no species has been reported hitherto from south of Hidalgo. The discovery of P. erythrorrhiza in the middle Balsas valley south of the Transverse Volcanic Belt thus marks a substantial range extension for the group. The closest relative of P. erythrorrhiza is probably the dis- tantly allopatric P. Tweedyi. Both have puberulent stems and leaves, and the flower of P. Tweedyi, although com- monly a little larger, varies in size down to a small extreme hardly different from that of P. erythrorrhiza. The best diagnostic features are found in the foliage and habit of 70 Rhodora [Vol. 72 growth. Mature plants of P. Tweedyi develop a woody root and suffruticulose caudex, and either all leaves or at least those leaves borne above the middle of the stem and opposed to the racemes are linear to linear-elliptic in outline and of firm, subcoriaceous texture. The stems of P. erythrorrhiza spring directly from the truncate apex of a fleshy, coralloid, orange-red taproot, and the homomorphic leaves are all elliptic and of relatively thin texture. In general facies P. erythrorrhiza suggests the primarily Texan P. Lindheimeri Gray, ordinarily characterized by a vesture of longer spread- ing hairs, and the too closely related P. nitida T. S. Bdg., which seems to be little more than a puberulent phase of it. However this may be, P. Lindheimeri and P. nitida differ collectively from P. erythrorrhiza in the strong tertiary nervature of the leaves, which become prominently reticu- late when mature, not merely penninerved. The other de- scribed members of subsect. Eurhinotropis are more easily distinguished by their small leaves. Anisacanthus stramineus Barneby, sp. nov. ob bracteas amplas foliaceas calyci amplexo longiores cum A. abdito T. S. Bdg. necnon A. ochoterenae Miranda comparanda, ab illo foliis floralibus sub- sessilibus et corolla straminea, ab hoe corolla bracteisque dimidio minoribus facile distincta. Frutex submetralis, cortice vetustiori albido, caulibus novellis pilis patulis villosissimis eglandulosis, foliis pilis multicellularibus sparsis cum brevioribus capitato-glandulosis crebris commixtis viscido-pubes- centibus; folia difformia, caulina (ad anthesin caduca) majuscula petiolata (petiolo ad 1 cm usque longo), lamina ovata vel ovato- lanceolata basi late cuneata ad 2 cm usque longa dorso elevatim costata et pinnatim nervosa, ea ramulorum axillarium flores sufful- crantia oblongo-lanceolata subsessilia vix 1 cm attingentia; bracteae foliaceae lineari-oblongae obtusae calvcem amplectentes 6-9 mm longae fere 1 mm latae; calyx 4.5 mm longus, laciniis lineari-attenuatis 3.5 mm longis glanduloso-puberulis; corollae + 2.5 cm longae gla- berrimae stramineae tubus cylindricus 0.95-1.1 cm longus 1-1.2 mm diametro ad fauces paullum ampliatus, labia subaequilonga posterius integrum dorso apicem versus pallide violaceo-pictum anterius fere ad basin 3-partitum lobis 4.5 mm latis; filamenta 10-11 mm longa basi incrassata glaberrima; antherarum thecae subexacte collaterales subaequilongae + 1.6 mm longae muticae; capsula glabra; semen pallide roseum 1.7 mm longum. 1970] Phanerogams — Barneby TA MEXICO. Puebla: arid hills, 1110 m (3700 ft.), near Cox- cotlan, November 18, 1966, H. D. Ripley 14, 731. Holotypus, NY; isotypus, US. The only close relative of A. stramineus described in the last revision of Anisacanthus (Hagen in Ann. Mo. Bot. Gard. 28: 385-404. 1941) is A. abditus T. S. Bdg. of the Pacific slope in Sinaloa, The two species are alike in the large bracts which surpass and conceal the calyx; but A. abditus differs greatly in having foliage glabrous below the inflorescence, petioled leaves between the flowers, and a longer, red corolla. An apparently nearer relative has been described since: A. Ochoterenae Miranda (in Ann. Inst. Biol. Mex. 12: 606. 1941), from Morelos. Through the kind- ness of Dr. A. Gómez-Pompa I have had access to the holo- type (Miranda 1327, MEXU) and several subsequent collec- tions of A. Ochoterenae which resembles A. stramineus in having yellow flowers but is easily distinguished by its nar- rower corolla, This is a little over 5 em long, and the strap- shaped free segments of the lower lip are nearly 3 cm long but only 3 mm wide. The corolla of A. stramineus is only 2.5 cm long, with oblong free segments about 1.5 em long and 4.5 mm across, that is 3-4 not 10 times longer than wide. THE NEW YORK BOTANICAL GARDEN BRONX, NEW YORK 10458 A NEW SPECIES OF CALAMOVILFA (GRAMINEAE) FROM NORTH AMERICA: KEN E. ROGERS Calamovilfa arcuata K. E. Rogers, sp. nov. — Perennis, usque ad 15 m alta vel altior; rhizomata gracilia, brevia; nodi barbati sub annulum eum confertis mollibus adscendo-appressis pilis usque ad 8.0 mm longis; vaginae firmae, confertae, parce pubescentes in dorso cum mollibus longis in tubercula adfixis pilis vel glabrescentes; margines, praecipue apice, dense pilosi; collum circulus densus e longis albis pilis compositus; panicula aperta, purpuracea, 15-45 cm longa, 8-40 cm lata; rami primarii patentes denum adscendentes, superne apice pilosi; pulvini pilosi vel pubescentes; spiculae anguste lanceolatae, purpureo colore tinctae, 6.0-7.4 mm longae; gluma prima ovato- lanceolata, acuta, acuminata vel aristo-aculeata, plerumque arcuata, 2.7-4.1 mm olnga; gluma secunda ovato-lanceolata, acuminata vel etiam aristo-aculeata, arcuata, 4.2-5.4 mm longa; lemma lanceolatum, attenuatum, arcuatum, 5.5-7.0 mm longum, usque ad 2 mm paleam superans; palea lanceolata, attenuata, 5.4-6.2 mm longa, binervis, inter nervos sulcata, pubescens vel pilosa in atque inter nervos, apice breviter bifid scaberuloque. Perennial, up to 1.5 m or more tall; rhizomes slender, short, brown- ish, as much as 5.0 mm thick; culms densely tufted, erect, unbranched, round or slightly subcompressed, smooth or slightly scaberulous below the nodes; nodes bearded below the annulus with soft ascending- appressed hairs up to 8.0 mm long; sheaths firm, close, sparsely pubescent on the surface with long, soft tubercle-based hairs or glabrescent, the hairs persistent and dense along the margins and at the apex; sheaths overlapping and persistent at the base, becoming shorter than the internodes above, mostly 6-15 cm long (or the upper- most as much as 22 cm long); collar a dense ring of long, white hairs; ligule erose-ciliate, whitish, convex, 0.2-0.7 mm long, the cilia less than 0.5 mm long; blades flat, linear, attenuate to a slender, thread-like involute point, articulated with the sheaths by a narrow line, 30-85 em long, 1.5-6.5 mm wide, firm, the nerves approximate, the upper surface bearing soft hairs up to 5.0 mm long, these dense behind the ligule, the surface becoming glabrate toward the apex, the lower surface sparsely pilose toward the base or glabrous; margins of the blades narrowly white cartilaginous, antrorsely scabrid except at the ciliate base. Panicle terminal, open, purplish, 15-45 cm long, 8-40 cm wide; peduncle 15-38 em long; primary branches solitary and alternate or ‘Contribution from the Botanical Laboratory, The University of Tennessee, N.S. 308. 1970] Calamovilfa — Rogers 73 some of them paired at the nodes, slender, spreading or ascending, pilose or pubescent in the axils, more or less scabrid on the angles, the lowermost as much as 22 cm long; secondary branches ascending, 2-11 cm long; tertiary branches ascending, 2-8 cm long; rachis angled, the angles scaberulous or scabrid; pulvini pilose or pubescent; spike- lets paired and solitary, appressed-ascending, narrowly lanceolate, acuminate, tinged with purple, 6.0-7.4 mm long; pedicels scabrid to nearly smooth, the longer of a pair 2.5-8.0 mm long, the shorter 1.0- 2.0 mm long; glumes, lemma and palea firm; first glume ovate- lanceolate, acute, acuminate or awn-pointed, usually arcuate, 2.7-4.1 mm long, 1-nerved, the nerve scaberulous on the upper portion; second glume ovate-lanceolate, acuminate or awn-pointed, arcuate, 4.2-5.4 mm long, 1-nerved, the nerve scaberulous on the upper portion; lemma lanceolate, attenuate, arcuate, 5.5-7.0 mm long, 1-nerved, pubescent or pilose on and along the midnerve for % to 2/3 its length; callus bearded with white hairs, the hairs 2.0-2.5 mm long, 0.32 to 0.42 the length of the lemma; palea to 2 mm shorter than the lemma, lanceolate, attenuate, 5.4-6.2 mm long, 2-nerved, furrowed between the nerves and pubescent or pilose on and between the nerves, the apex shallowly bifid and scaberulous; stamens 3.0 mm long. TYPE: TENNESSEE. CUMBERLAND CO.: about 8 miles north- northeast of Crab Orchard, off the old gravel road from Crab Orchard to Rockwood, downstream about 600-800 feet from Antioch Bridge over Daddy’s Creek, in a shrub-dom- inated community, October 3, 1968, Rogers, Sharp, Del- gadillo, and Meijer 42409 (Us 281796). Isotypes at TENN, US, TAES, ISC, LAF. Other Specimens Examined: OKLAHOMA. PUSHMATAHA CO.: Pushmataha Wildlife Ref- uge, SE Oklahoma, Porter s.n., Sept. 1968 (Us). The genus Calamovilfa Hack. has previously been treated as comprising four species in North America. One species, C. curtissii (Vasey) Scribn., is endemic to northern Florida. Two other species, C. longifolia (Hook.) Scribn. and C. gigantea (Nutt.) Scribn. and Merr., occur from southern Canada, Michigan, Indiana and Missouri west to Colorado and Idaho, and south to Texas and Arizona. A fourth spe- cies, C. brevipilis (Torr.) Scribn., is a coastal plain species reported to occur from New Jersey to South Carolina (Hitchcock, 1951; Gould, 1968). 74 Rhodora [Vol. 72 ty | Z > id a ` Ms ; 4 y , da^ | S a) t wu £d k IN Galsmoavilfa brevipilie (Torri) Serin, è i Daddy's Greek from asan VO tent pueg Dagsa inis cut PR sets i edax a mcs neige, j * COEM e FLORA TENNESSEGNSIS e Üwberiand County Bad LEGO? LI Figure 1. Photograph of Calamovilfa arcuata (Rogers et al 42409). 1970] Calamovilfa — Rogers 75 In the fall of 1968, on a plant collecting trip with other botanists to the Cumberland Plateau in Tennessee, I col- lected a grass which was initially thought to be either a Panicum or some unfamiliar grass (Figure 1). Perhaps intuitively, enough material was collected for several her- barium specimens. Later examination of the material led to a tentative identification of it as Calamovilfa brevipilis. The discovery of this species on the plateau, although a little surprising, was not inconceivable since a number of Coastal Plain elements occur there. I was also aware that this material was atypical of C. brevipilis, if not distinct from it. A number of specimens were sent to agrostologists for observation. Most agreed that it was probably an atypical form of C. brevipilis. Dr. Thomas R. Soderstrom, Curator of Grasses at the United States National Museum, after examining my material and a single specimen essentially identical to it from southeast Oklahoma, suggested that I describe the grass as a new species. A careful comparison was made of C. arcuata with her- barium material of other species in Calamovilfa. I found that C. arcuata was apparently most closely allied to C. brevipilis and C. curtissii, and between these two it was most closely related to the former. Thieret (1966) in his Synopsis of the genus Calamovilfa (Gramineae) regarded the genus as comprising two sec- tions, Section Calamovilfa and Section Interior. Features of plants in the former section are short rhizomes, ligular hairs less than 0.5 mm long, a line of articulation between sheath and blade, and a distribution in the Coastal Plain of eastern United States. Extensively creeping rhizomes, ligular hairs 0.75 mm or more long, line of articulation not present between sheath and blade, and a distribution in the interior United States and Canada characterize plants of the latter section. After examining the material of Cala- movilfa from the Cumberland Plateau in Tennessee, I con- cluded that it belonged to the sect. Calamovilfa. Utilizing measurements taken both from herbarium speci- mens and those appearing in the literature, a comparison 76 Rhodora [Vol. 72 was drawn between the species in sect. Calamovilfa (Table 1). nodes sheaths indumentum collar ligule, length blades, indumentum panicle pulvini spikelet, length glumes, shape lemma length shape palea, length lemma/palea callus hairs, length TABLE 1. C. brevipilis glabrous glabrous or slightly pubescent at the apex glabreus 0.2-0.4 mm upper surface sparsely pilose at base open; branches spreading to ascending glabrcus 4.0-5.8 mm acute, not arcuate 4.0-5.4 mm acuminate, not arcuate 3.8-5.8 mm subequal or nearly so, or palea longer than lemma 1.5 mm C. arcuata pilose margins pilose; surface pilose or glabrescent pilose 0.3-0.7 mm upper surface densely pilose at base, the rest of the surface pilose to glabrescent open; branches spreading to ascending pubescent or pilose 6.0-7.4 mm acute to acuminate, arcuate 5.8-7.2 mm attenuate, arcuate 5.8-6.2 mm lemma longer than palea 2.0-2.5 mm C. curtissii glabrous sparsely pilose or glabrescent glabrous or sparsely pilose 0.3-0.4 mm upper surface pilose at base, the rest of the surface pilose to glabrescent contracted; branches appressed glabrous 3.7-5.6 mm aeute, not arcuate 3.5-5.2 mm acute, not arcuate 3.5-5.0 mm subequal 1.0-1.5 mm 1970] Calamovilfa — Rogers 77 The above comparison of species illustrates that C. ar- cuata is clearly separated from C. curtissii by the open panicle with spreading to ascending branches and the larger spikelets which are 6.0 mm. or more long. C. curtissii has a contracted panicle with appressed branches and spikelets not more than 5.6 mm long. C. arcuata is distinguished from C. brevipilis, and in most cases from C. curtissii, by the pilose nodes, the pilose sheaths (at least marginally), the denser indumentum behind the ligule at the base of the blade, the pubescent or pilose pulvini, the arcuate glumes and lemma, the longer lemma and palea, the lemma exceed- ing the palea rather conspicuously, the longer callus hairs, and the larger spikelets. The specific epithet arcuata refers to the unique out- wardly curved glumes and lemma (Figure 2). In the single caryopsis of C. arcuata available for study it was clear that the pericarp and testa were not adherent upon soaking in water. This observation corroborates that of Reeder and Ellington (1960) with regard to caryopses in the genus. Knowledge of the habitat of C. arcuata is limited by the absence of precise information about the ecology of the Oklahoma material. However, I have had an opportunity to observe closely the habitat of the Tennessee population, both at the time it was first collected, and later when I revisited the area in December, 1968. Thus far, only a single popula-: tion of C. arcuata has been found in Tennessee. It appears to be a well established population, numbering some 25 to 30 or more individuals growing separately and gregarious- ly. The plants are growing on rather thin, recent siliceous alluvium overlying sandstone rock. This area projects into Daddy’s Creek about 30-60 feet and has a length of 100 feet or so, and is dominated by shrubs under which there are numerous herbaceous plants. There is only a scattering of small trees. The surrounding area has a maturely dissected topography on which there are rich deciduous woods inter- spersed with pine, A list of the principal plants associated with C. arcuata, arranged approximately in their decreasing 78 Rhodora [Vol. 72 Imm Figure 2. Photograph of drawings of Calamovilfa arcuata: a, spikelet, X25; b, c, floret, X25. 1970] Calamovilfa — Rogers 79 order of importance in the plant community, is given in Table 2. TABLE 2. List of principal flowering plants associated with C. ar- cuata, Daddy’s Creek, Cumberland County, Tennessee Shrubs Trees Cornus amomum Miller Liquidambar styraciflua L. Alnus serrulata Acer rubrum L. (Aiton) Willd. Pinus virginiana Miller Hypericum prolificum L. Nyssa sylvatica Marshall Itea virginica L. Diospyros virginiana L. Sambucus canadensis L. Fraxinus sp. Ilex verticillata (L.) Gray Rosa palustris Marsh Rhododendron sp. (azalea) Herbs Andropogon gerardii Carex sp. Vitman Bromus purgans L. Andropogon scoparius Elymus villosus Muhl. Michx. Lycopus americanus Muhl. Panicum microcarpon Muhl. Oxypolis rigidior (L.) Raf. Panicum clandestinum L. Scutellaria sp. Solidago rugosa Miller Muhlenbergia sylvatica Eupatorium fistulosum (Torr.) Torr. Barratt Xyris torta Smith Lobelia cardinalis L. Aster dumosus L. ACKNOWLEDGMENTS I wish to express my sincere thanks to Dr, Thomas R. Soderstrom, Curator of Grasses, United States National Museum, Washington, D.C., for examination of type speci- mens and for helpful observations; Dr. Edward E. Terrell, Botanist, New Crops Research Branch, U. S. Department of Agriculture, Beltsville, Maryland, for the loan of the 80 Rhodora [Vol. 72 Oklahoma specimen; and Mrs. Marie Hicks, Graduate Stu- dent, Department of Botany, The University of Tennessee, Knoxville, for preparing the illustrations. THE UNIVERSITY OF TENNESSEE DEPARTMENT OF BOTANY KNOXVILLE 37066 LITERATURE CITED GOULD, FRANK W. 1968. Grass Systematics. McGraw-Hill, New York. 382 p. HITCHCOCK, A. S. 1935. Manual of the Grasses of the United States. U. S. Dept. Agr. Misc. Publ. 200, 1040 pp., illus. (rev. 1951 by Agnes Chase, 1071 pp., illus.). REEDER, J. R. and M. A. ELLINGTON. 1960. Calamovilfa, a Mis- placed Genus of Gramineae. Brittonia 12(1): 71-77. THIERET, JOHN W. 1966. Synopsis of the Genus Calamovilfa (Gramineae). Castanea 31: 145-152. SAURAUIA SPECIES AND THEIR CHROMOSOMES! DJAJA D. SOEJARTO Saurauia is a widespread genus of flowering plants, about 250 species of which have been described ( Melchior, 1964). The members are represented in both Old and New World tropics and subtropics. The American range extends from Central Mexico in the north to Central Bolivia in the south, through Andean South America, It is not known from the West Indies, and no records indicate its presence in the Guianas and Brazil. According to a recent study by Hunter (1966), 22 species occur in Mexico and Central America, and my present study indicates that 49 species are represented in South America. The cytology of the genus has never been studied. In his treatment of the Peruvian species of Saurauia, Macbride (1955) remarks that ". .. specific characters to this day are not understood and a satisfactory treatment will prob- ably be possible only with cytological as well as morpholog- ical data." Hunter (1966), without the benefit of cytological data, was able to show that the species (Mexican and Central American) could be classified comprehensibly on a morphological basis alone. As a non-cytologist, I was stimulated by Macbride’s remark, and I set to work to collect cytological “pickles” in the field during the summer of 1963 in the Narifio and Putumayo regions of Colombia. Later, upon examination in the laboratory, the materials which were thought to be in the right developmental stages for the study of meiosis of the pollen mother cells turned out to be very old. Provoked by this failure I collected more cytological materials repeatedly from various parts of South America (Colombia, Ecuador, Peru) during the summers of 1964 and of 1965 and in May and June of 1966 in Colombia. It is from these more adequate materials that the observations here reported were made. "This paper is a minor part of a thesis (*Studies of South American Saurauia") submitted to the Department of Biology of Harvard Uni- versity in partial fulfillment of the requirement for the degree of Doctor of Philosophy. 81 "Wl 000E “BO [8 3sə4oj urejunour prumy 'euejog enbsog 'ojseq eaoqe ‘ounen "(Sui "q9noj Sp Aog əy} orga 'si9Aog Əpym YIM Gnays) ^3[nuog "| `M vprydozyorwo DINDINDG ` a4eld ` : ` > T `. » ` > " 21 -i e Lai a w — © A e c ma ^ — 1970] Saurauia Chromosomes — Soejarto 83 š ew Plate 2. Saurauia bullosa Wawra. Nariño, above Pasto, Bosque Botana, second growth vegetation, alt. ca. 2800 m. Note the wheel-like pattern of insertion of the primary branches. 84 Rhodora LLLLELITLTLTTTUTTEETTITPTTTT © Plate 3. Meiotic configurations in Saurauia. A, S. bullosa, early anaphase I, equatorial view, n=30. B, S. omichlophila (voucher: Soejarto 914), diakinesis, n—30. C, S. isoxanthotricha (voucher: Soe- 1970] Saurauia Chromosomes — Soejarto 85 MATERIALS AND METHODS Flower buds at different stages of development were col- lected in the field at various times of the day and were killed and fixed immediately in Carnoy’s solution (3 parts absolute alcohol, 1 part glacial acetic acid by volume) for from 24 to 72 hours. Since the sepals in most species form a thick cover over the bud, these were first removed or, when the buds are small, were cracked in several places (without removing the sepals), before dipping them into the fixative so that the liquid could penetrate the tissue rapidly. Small vials were used as containers. At the end of the period in- dicated above, the fixative was substituted by 70% ethyl alcohol. Due to the nature of travel, no serious attempt was made to keep these vials in refrigeration. Only after arrival in the United States, were these vials containing the fixed materials kept in the refrigerator until ready for examination. For each cytological specimen, a voucher was collected. A complete set of the vouchers is deposited at the Gray Herbarium of Harvard University. Permanent microscope slides are deposited in the Harvard Botanical Museum. RESULTS Nineteen individuals representing fifteen species have been observed and the chromosomes counted. The chromo- some behavior at meiosis in all but one of the individuals studied appears normal, and the chromosome morphology and number are remarkably stable. Representative meiotic configurations are shown on Plate 3 and Plate 4. The gametic chromosome number of the fifteen species observed is n = 30, and the chromosome size during the first meta- phase (meta-anaphase) varies but little, as shown by the cut-ups on Plate 3. The chromosome length at this stage ranges from 3.5-6.54, the mean length being 5p. jarto 1563), diakinesis, n—45 (42II + 61). D, cut-ups of A arranged linearly to show size range and variation of the early anaphase (meta-anaphase) chromosomes. All the same scale, approx. 1600X. A and B photomicrographs, C camera lucida drawing. 86 Rhodora [Vol. 72 ww" 9 Plate 4. Meiotie configurations in Saurauia. A, S. biserrata (Soejarto 1426). B, S. Stapfiana (Soejarto 2046). C, S. pastasana (Soejarto 1344). D, S. tomentosa (Soejarto 1433). E, S. brachybotrys (Soejarto 2052). F, S. Humboldtiana (Soejarto 913). G, S. aequa- toriensis (Soejarto 1342). H, S. isoxanthotricha (Soejarto 1531). I, S. portachuelensis (Soejarto 1158). All about 1600X. A, B, C, E, H camera lucida drawings; D, F, G photomicrographs; I a tracing of a photomicrograph. 1970] Saurauia Chromosomes — Soejarto 87 In the course of the study, it was noted that the size of the pollen mother cells is relatively stable in all the nineteen individuals the diameter being 35-504. A slight increase in diameter is observed in Soejarto 1563, the diameter being 45-604. For technical reasons, no detailed study of the meiotic process was attempted, yet some remarks regarding the successive stages observed during the chromosome counts are given below. The size of the flower buds in which meiosis was observed is not constant in all the species studied; this is due, pri- marily, to the different sepal thickness and the stamen number in the various species. To cite examples: the flower buds in which meiosis was observed for S. bullosa and S. tomentosa (with 100-200 stamens and relatively thick sepals) have a diameter of about 10 mm, whereas for S. chiliantha and S. portachuelensis (with 15-30 stamens and relatively thin sepals) the diameter is about 3 mm. How- ever, anther size, in which meiosis was observed, is essen- tially the same, being 0.5-1.0 mm in length and about 0.25- 0.5 mm in width. Examination of various anthers from the same flower bud from which meiosis was observed shows that the developmental stages within one bud are not uni- form. The outer whorls of the anthers (especially those species with high stamen number) were observed to show earlier stages of meiosis than those of the inner whorls. Although I made no quantitative study, I believe that this phenomenon agrees with Brown’s study (1935) of the cen- trifugal maturation of the stamens in Saurauia subspinosa. The pollen mother cells are globose to subglobose and at the resting stage each possesses a rather large nucleus, con- taining one or two nucleoli (when only one, it is either centric or excentric). During the early phases of meiotic prophase, there is an increase in cell size (from about 30, to 40-504), and the chromosomes appear as densely inter- twined threads which are very unsuitable for analysis. Chiasma formation begins to be seen in diplotene rather clearly, but the materials are not favorable for the exact determination of its frequency, both in this stage as well 88 Rhodora [Vol. 72 as in later stages; furthermore, the chromosomes in this stage appear slightly too diffuse for an exact delimitation of the chiasmata. At diakinesis, the chromosomes attain the greatest contraction, and staining reaction with aceto- carmine is good. They appear quite dark and are distributed rather evenly in the cell, which facilitates counting. Both ring and rod bivalents were observed, but they tend to ap- pear as dark blobs, making exact determination difficult. The cytoplasm in most species examined also takes stain, though lightly, which occasionally renders correct counting unfavorable. In some species, such as S. peduncularis, the nucleolus is quite late in disappearing, but in the majority, it becomes invisible during diakinesis. At the first meta- phase, the chromosomes become arranged on the equatorial plane which is usually impossible to analyze from a lateral view, and the spindle fibers are only barely discernible. As a result of squashing, the chromosomes in the early first anaphase (meta-anaphase) frequently become well sep- arated (especially in S. tomentosa and S. bullosa) and good for counting, above all when the pollen mother cells are well separated as a result of thorough maceration. Countings reported here were made either from diakinesis and/or early first anaphase configurations, at an average of 15 cells per preparation. During the early first anaphase, the bivalents are held together by one or two terminal chiasmata or one interstitial chiasma. In this stage, the position of the centromeres could also be observed, either median to submedian or subterminal, the majority being subterminal. No lagging chromosomes were observed during the first anaphase in the individuals with n — 20, though, very rare- ly, bridge formations were noted. In Soejarto 1563, how- ever, the univalents usually move to the poles earlier than the bivalents, though occasionally they lag behind; bridge formation was also observed. It should be noted that at the metaphase plate, in the case of Soejarto 1562, the univalents are distributed randomly. At the dyad stage, presumably at the beginning of homotypie division, the chromosomes are sometimes distributed rather evenly to facilitate counting. 1970] Saurauia Chromosomes — Soejarto 89 At the second metaphase and anaphase, the two sets of the spindle fibers lie either in the same plane or at right angles to each other. Finally, cytokinesis is of simultaneous type and the tetrad arrangement of the pollen grains is tet- rahedral. The developmental stage within a single anther is not uniform (synchronous) in all the species examined; al- though usually one or two meiotic stages are represented, very frequently more were observed, e.g., diakinesis is ob- served among the numerous pollen mother cells, either in the first metaphase and/or anaphase, even in telophase to tetrad stage. The only explanation which I can offer is that this phenomenon may have been the result of the different collecting time of the specimens. In all the individuals studied, I always observed pieces or complete packets of rhaphides among the pollen mother cells. Presumably, these are present within the anther tissue and became broken and separated during maceration and squashing. The results of chromosome counts are summarized in the following Table, where the order of the species is listed geographically, in a north-to-south and west-to-east order, corresponding alphabetically to the order of the species in the accompanying map. DISCUSSIONS AND CONCLUSIONS This study has made clear that chromosome number and morphology in Saurauia appear to be of little use in specific classification and grouping. At least 14 species have highly similar chromosome morphology, with gametic chromosome complement of n = 30. However, any statement at the present stage of our knowledge should be considered tenta- tive, and a final conclusion must await further study, pref- erably including comparative data of representatives from Central America, Mexico, and Asia. The existence of an individual with a gametic chromo- some number of » — 45 (39-42 bivalents plus 12-16 uni- valents) can be explained as a result of interspecific hybridization. Mitotic figures obtained from root tip smear [Vol. 72 Rhodora 92 ‘s}UNOD ourosOoulOIq2 JO SYNSƏY I ATAVL ovyoulyg | "Sue4dg IIOS 00€z 9Aoqe 'oonueng 'o3d([ nəq 9caFI Oox19[ooq Cd 3 WD) 9imaas2siqcS CO EIN [PYS 'ezejseq | IOE 00&I -OLEN 'A044 *iopeno"] FRET ojiu(ooq SNQ vuDiD4q cS ^N souug | ondeaddg IIOS 0081 ‘enyeinsuny, Aoig “lopenay| SPELT oxiefoog sisuaiLozDNbav `Ç "NN o[ognuo?j.l0q “01D | "IMS T'Y II0& 008 ‘oABUININ, “WON 'erquio[o)) ELET ojte(oog | Siuofipwn]nd "gy I o[ənuov140d “01g | "[nu»s Hp IIO 004€ *o&ewnjnq Wop "erquio[o))| gcrp ojiefoog | sisuajomu2nj4od `Ç H ouvoud Ta | “Sueids (HHH) II0S 0082 ‘OUMEN "O3d([ “BIQUIO[OD gEpPT oqiefoog | DSOJUIWO} 'S "[ vugjog ənbsog | IIOS 0067 ‘OUNEN 'Ojd([ 'erquio[0) | gjpp ojiefeoog | CIMEM Dso]pmq "S I (19- IZT) o[anyor}.10g "019 | + GISF-II6€) 008 'ofeumjnq “WON *eiquio[o?)| e9c[ OZIB fƏOS | 'osng DYNLJOYJUNLOSI `ç o|[ənuot1404 '047) | TI0& 008Z ‘oAeungymndq 'ur) ‘erquoop| TEE, oelaog | sng D04214)0:]4D0rO081'S ^H vuvjog onbsog S6FI (YIS "ar II0S 0008 ‘OUEN 'O3d(p 'erquio[0) *pp6 ojiwíeog | »judojioruo "S. `D eseng ory 9ebT | IIOS OOST ‘OUEN 'O3d(q 'erquio[0))| ‘ggg oyefeog | dP p 520]nounpod S ^4 Bulepuanbay, ap OLS IIOS OOLZ volvweuipuny 'o3d([ 'erquio[o?) &I6 O}Ie laog sug DuDjpjoqunH `Ç `" uosuog mus “AA IOS 008€ "unbonguy 'o3dq 'erquio[o))| LEZ ojiwfoog | 0109 ^g. `A uosuos 9F06 IIOS 0086 ""mibonuy ‘oid *erquio[o)| epog oiefoog sng nwpnydvjS 'S “OD seu[eq sey IIOS 000€ "emiboguy 'o3d([ ‘erquiojog| opoz ol4e[əos Id PLL rusun’ cg Bus[eH EIS 'ur[[opo]q IIOS 0006 ‘etnboyuy 'o3d([ ‘erquiojog) geog oxiefeog "zo4ng s/i4309ffonaq `Ç ^V ONIUHIVd (ur) NAWIOddS OLLOIMIN AC OLILTV NISIUO YUAHONOA SWIOMWdS 1970] Saurauia Chromosomes — Soejarto 93 Agricola of Universidad de Nariño, and Dr. Alvaro Fer- nández P., director of the Botany Department, Instituto Ciencias Naturales, Universidad Nacional, Bogotá. Finally, I wish to thank my wife, Mariela, for her con- stant encouragement, and for typing the manuscript. Any errors and/or misinterpretations found in this paper, how- ever, are my sole responsibility. DEPARTMENTO DE BIOLOGÍA UNIVERSIDAD DE ANTIOQUIA MEDELLIN, COLOMBIA LITERATURE CITED BRowN, E. G. S. 1935. The floral mechanism of Saurauia sub- spinosa Anth., Transact. Bot. Soc. Edinburgh, 31: 485-497. Hunter, G. E. 1966. Revision of Mexican and Central American Saurauia (Dilleniaceae). Ann. Missouri Bot. Gard., 53: 47-89. MACBRIDE, J. F. 1955. Flora of Peru, pt. 3A, vols. 1 & 2: 679. Field Mus. Natur. Hist. Publ, Bot., vol. 13. MELCHIOR, H. 1964. In ENGLER, A., Syllabus der Pflanzenfamilien, ed. 12 (Band II): 161. SoEJaRTO, D. D. 1968. Studies of South American Saurauia (Acti- nidiaceae). Unpublished doctoral thesis submitted to the Depart- ment of Biology, Harvard University, 458 pp. + 61 plates. ADDITIONAL CHROMOSOME NUMBERS IN BRAZILIAN COMPOSITAE JAMES R. COLEMAN While serving as a research taxonomist at the Instituto de Botanica of São Paulo, Brazil, I had the opportunity to collect material of Brazilian Compositae for cytological study. A previous report on this study has been published (Coleman 1968). In the present paper 40 counts are reported for 35 species of 18 genera (Table 1). Initial reports are given for 29 Species and 2 genera. The initial generic reports are for Elvira, n = 12, and Ichthyothere, n = ca 33. The methods used are identical to those described by Cole- man (1968). The material of Eupatorium ballotaefolium, E. organense and E. tremulum was determined by Dr. Ly- man B. Smith and Simsia dombeyana by Dr. J. Cuatrecasas. My appreciation is sincerely expressed, All other identi- fications are my own. A complete set of voucher specimens is deposited in the University of Georgia Herbarium and 2 nearly complete set in the Herbarium of the Instituto de Botanica of São Paulo, Brazil. VERNONIEAE The report of n = 16 for Centratherum punctatum agrees with an earlier report for that species based on Panamanian material (Turner and King 1964). No other counts have been reported for the genus. EUPATORIEAE The count of n = 10 for Ageratum campuloclinoides is the initial report for the species. Robinson (1913) consid- ered A. campuloclinoides a doubtful Ageratum and sug- gested its affinities to be with Trichogonia. Since the counts reported to date for Trichogonia agree with those of A gera- tum in being based on x = 10, chromosome number is no aid in better defining the relationship of Ageratum campulocli- noides. 94 1970] Brazilian Compositae — Coleman 95 The report of n = 10 for Alomia fastigiata agrees with a previous report for this species (Coleman 1968). The Central American Alomia microcarpa (Benth.) Rob. has also been reported as n — 10 (Turner and King 1964). Counts of n = 9 are reported for two populations of Alomia longifolia (Gardn.) Rob. This species was originally de- scribed in the genus Ageratum, but was subsequently trans- ferred to Carelia and later Alomia. No reports are available for Carelia, but previous reports for both Ageratum and Alomia are based on x — 10. North American species of Stevia have been reported as n — 11, 12 and 17 (Powell and Turner 1963). With the report of n = 12 for Stevia organensis, counts of n — 11 and 12 have now been reported for Brazilian species. The count of n — 10 for Eupatorium laetevirens agrees with an earlier report for the species. The report of n = 10 for E. ballotaefolium concurs with a count presented for Colombian material of that species (Powell and King, 1969). I have previously reported n = 30 for E. ballotae- folium from southern Brazil (Coleman, 1968). ASTEREAE Counts of n = 9 for species of Baccharidastrum and Bac- charis are in agreement with previous reports for these genera. HELIANTHEAE Blainvillea rhomboidea, n = 17, is the second species of the genus to be reported. Blainvillea latifolia DC. has been reported as » — 39 (Mehra et al 1965). An approximate count of n = 24 + 2 is reported for Spilanthes acmella. Previous counts of n = 7 (Malik and Ahmad in Cave 1964) and n — 12 (Mehra et al 1965) have been reported. Turner et al (1962) have shown a diploid, tetraploid and hexaploid series to exist in S. americana Hieron. A similar situation evidently occurs in S. acmella. The counts of n = 12 for Elvira biflora and n = ca 33 for Ichthyothere rufa are the initial reports for these gen- era. 96 Rhodora [Vol. 72 ANTHEMIDEAE The report of n — 9 for Anthemis nobilis is consistent with previous reports for the genus. SENECIONEAE Counts reported for species of Senecio are consistent with the majority of previous reports for that genus in being based on x — 10. THE UNIVERSITY OF GEORGIA DEPARTMENT OF BOTANY ATHENS 30601 LITERATURE CITED CAVE, M. S. (ed.). 1964. Index to plant chromosome numbers. Uni- versity of North Carolina Press, Chapel Hill. COLEMAN, JAMES R. 1968. Chromosome numbers in some Brazilian Compositae. Rhodora 70: 228-240. MEnRA, P. N., B. S. GILL, J. K. MEHTA and S. S. SIDHU. 1965. Cyto- logical investigations on the Indian Compositae. I. North-Indian taxa. Caryologia 18: 35-68. POWELL, M. A. and R. M. KING. 1969. Chromosome numbers in the Compositae: Columbian species. Amer. Jour. Bot. 56: 116-121. POWELL, M. A. and B. L. TURNER. 1963. Chromosome numbers in the Compositae. VII. Additional species from the southwestern United States and Mexico. Madrono 17: 128-140. ROBINSON, B. L. 1913. Revision of Alomia, Ageratum and Oxylobus. Proc. Amer. Acad. Arts and Sci. 49(8): 438-491. TURNER, B. L., M. POWELL and R. M. KING. 1962. Chromosome numbers in the Compositae. VI. Additional Mexican and Guate- malan species. Rhodora 64: 251-271. TURNER, B. L. and R. M. KiNG. 1964. Chromosome numbers in the Compositae. VIII. Mexican and Central American species. Southwest. Nat. 9: 27-39. 97 Brazilian Compositae — Coleman 1970] chp ‘lvondes op ojuog ovg :o[neq ovg jo o3e3g TLV "envy op [uuoroe N onbivg 'equioive]y :odrəugp ap ory Jo oye3g SLS 'so£31() sop VIIIS jeuoney onbaeg :oirouef ap on] Jo 9)%4S PSS '"eunueop Jo oidrorunjy *oquurzoepa1ioA :veIQeg jo 2484S 8LS '"Ienej[9p [euor€eN enbreq :oireuef ap ory Jo 9484S 66€ 'OBpIof op sodure;) jo N wy G ed :o[neq ous Jo o1e3g 609 'o€piof op sodureO jo ASH Wy OT-g :o[neq ous Jo 9484S LLP ‘9I JO ordrorunjg *exro[ouuer) eeg Jo 34S 99h ‘“eyuedurey jo M Wy GT. 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AVaHLNVITAH ‘ds steye g ‘Od SeployyuBUOYyIe} stieuooeg y ‘Od LINIUOSIU SIIEYIM Y y ‘Od LUNJSNII srieuooeg y 'pnojg seprourtgwee[o SLIeqooeg, ‘Od *uogjrnounog4p “jo SEYDEL EY y layeg MPNA slaeyoorg y ‘OC EMOJI SEYI səroədç 99 Brazilian Compositae — Coleman 1970] pop '"o?piof op sodure;) Jo TSA wy OT-g :ojneq ogs jo 93038 ESP EWEA 9p [£uoro€ N anbaeg ‘Leder op ofaag :SIEIOr) SVUIJT JO 91%1S 89 “Blvonpueluey Jo t p WY Gc Bd :sIeJor) seurpy JO 93e3g T9p “Blvonpuvuley) jo Y Wy GZ Bd :SIEIIN SEUL JO 3484S Lop 'OgpIof op sodtuv;) Jo ASH Wy OT-g :o[neq ogg go 9784S G ‘SOBSIQ, SOP VIIIS [euoro€wN anbieg :odpouef ep Oy Jo nen A|snotAedd poj31odaz JOU €12uor), y {[snoraedd pojrodox you SIIAS y ‘Od snsso[Soot orəəuəç,, ‘Od snsso[8oot oroeuog., owe !gpropueurumu OUIS y ewe ng.opueurumWu oroeuosg.. ‘TPA St[Iqexoprsop oroəuəs,, 'upier) sni[ogreuno Oroeuag, AVANOIOUNGAS STUDIES IN THE COMPOSITAE-EUPATORIEAE, XV. JALISCOA, MACVAUGHIELLA, OAXACANIA, AND PLANALTOA. R. M. KING AND H. ROBINSON Continuing studies of the genera of the Eupatorieae have shown the need for the following revisions and clari- fications. In the case of Oaxacania, it seems fitting to make comparison with the closely related Carterothamnus. Mac- vaughiella and Jaliscoa, each with two described species, have proved to have too narrow a species concept. Planaltoa has shown a number of previously undescribed useful char- acteristics when observed microscopically. Jaliscoa S. Watson Proc. Amer. Acad, Arts Sci. 25: 153. 1890. (Type species, Jaliscoa pringlei S. Wats.) Suffrutescent wcody shrubs to 3 meters tall, ternately branched above. Stems erect, appressed-puberulous, very faintly striate. Leaves opposite or ternate, thin, very obscurely puberulous; blades ovate to broadly ovate, the margins crenulate-serrulate to sharply serrate, up to 16 em long and 11 em wide. Inflorescence a cymose-panicle; heads discoid, campanulate, up to 5 mm broad and ca 9 mm high (including style branches), ca 15 flowered, receptacle paleaceous, slightly convex or flat, phyllaries ca 12, imbricated, lanceolate, ca 3.5-4.0 mm long, apex obtuse or acute; corollas white, campanulate, ca 5 mm long (including lobes), 5-lobed, glabrous; stamens 5; anthers appendaged, the latter ca 225 u long; style branches 2, greatly exserted at ma- turity, papillose, style slightly glandular; pappus an obscure, entire, callus border or a lacerate-fimbriate crown; achenes ca 2.5 mm long, dark brown, 4 angled, sparsely setose, setae biseriate; base of achene forming a distinct carpopodium; pollen spherical, tricolpate, dis- tinctly spinose, ca 20-23 u in diameter, chromosome number not determined. Jaliscoa pringlei S. Watson, Proc. Amer. Acad. 25: 153. 1890. [Mexico: JALISCO: talus of cool ledges, bluffs of the Rio Grande de Santiago, Oct. 12, 1889, C. G. Pringle 2491. (Holotype GH!, Iso- types G!; MEXU!; MO!; NY!; PH!; UC!; US!)] Jaliscoa pappifera S. F. Blake, Contr. U.S. Nat. Herb. 22: 587. 1924. [MEXICO: MORELOS: barranca near Cuernavaca, 4500 ft, C. G. Pringle 9931. (Holotype us!, Isotypes F!; GH!; MO!; NY!)] 100 1970] Compositae-Eupatorieae — King and Robinson 101 Additional specimens examined: MEXICO: JALISCO: Puente San Pedro, 5 mi. SW of Tecalitlan, 1200 m., McVaugh & Koelz 1299 (MICH); 11-12 mi. SW of Autlan, ca. 1000 m., McVaugh & Koelz 873 (MICH) ; 2.5-4.0 mi. above (north of) La Cuesta, road to Talpa de Allende, 800-1000 m., McVaugh 21201 (MICH); barranca of Guadalajara, 4500 ft., Pringle 11544 (F, MICH, MO, US). Michoacan: Püerto Cruces, Coalcoman, Hinton 12374 (MICH). Morelos: barranca near Cuernavaca, 5000 ft., Pringle 6158 (F, ENCB, MEXU, MO, NY, PH, UC, US). State either Guerrero or Michoacan, Sierra Madre, Langlassé 575 (G, MICH, US). After investigating a number of specimens, we find that only one species can be recognized in this complex. The stems of this species seem to be particularly sus- ceptible to insect damage, and the resulting numerous per- forations are almost a characteristic. Macvaughiella R. M. King and H. Robinson Sida 3 (4): 282. 1968. Schaetzellia Sch.-Bip. Flora 33: 419. 1850. Not Schaet- zellia Klotzsch, Allgemeine Gartenzeitung 1849: 82. 1849. (Type species Schaetzellia mexicana Sch-Bip.) Perennial or suffrutescent herb to one meter tall, sparingly branched. Stems mostly single, erect, pubescent, striate. Leaves thin, pubescent, deltoid to rhomboid, the margins serrate to dentate; blades up to 3 em broad, up to 3.8 em long. Inflorescence corymbose; heads discoid, campanulate to turbinate, 2-4 mm broad, ca 7 mm tall (including style branches), 16-25 flowered, receptacle conical, naked, phyllaries ca 10, imbricated, ovate-lanceolate, ca 5 mm long, ca 1.25 mm wide, apex acuminate; corollas white, funnelform, ca 3 mm long (including lobes), 5 lobed (lobes slightly longer than wide) outer surface of lobes bearing both glands and nonglandular uniseriate hairs; stamens 5; anthers appendaged, the latter ca 100 u long and ca 150 » wide around base; style branches 2, exserted at maturity, style bearing occasional glands ventrally; pappus consisting usually of 2 more rarely 1 or 3-4 setae, usually equaling the length of the corolla; achenes when dry very flat, dark brown, ca 2 mm long, base of achene forming a distinct carpopodium; pollen spherical, tri- colpate, distinctly spinose, ca 18 u in diameter; chromosome number not determined. Key to varieties of Macvaughiella mexicana. Phyllaries essentially glabrous ................ M. mexicana var. mexicana Phyllaries: pubescent -aana y... u A. M. mexicana var. standleyi Macvaughiella mexicana (Sch.-Bip.) R. M. King and H. Robinson var. ‘mexicana 102 Rhodora [Vol. 72 Schaetzellia mexicana Sch.Bip., Flora 33: 419. 1850. [MExICO: VERACRUZ: Bord de la rivière à Acasonica, 1200 ft, Jan 1839, J. Linden 1168. (Holotype B, destroyed, Isotype P! Photo us!) ] Macvaughiella mexicana (Sch.-Bip.) R. M. King and H. Robinson, Sida 3(4): 282. 1968. Additional specimens examined: MEXICO: VERACRUZ: Mirador, Liebman 63 (C); Puente Nacional, Purpus 14284 (A, UC); Zacuapan, Fortin, Purpus 2855 (US); near Rancho Viejo, Purpus 16220 (A); Barranca de Tenampa, Purpus 2189 (F, GH, MO, P, UC, US); without precise locality, Galeotti 2302 (P, w). Macvaughiella mexicana (Sch.-Bip.) R. M. King and H. Robinson var. standleyi (Steyermark) R. M. King & H. Robinson stat. nov. Schaetzellia standleyi Steyermark, Publ. Field Mus. Nat. Hist., Bot. 23: 107. 1944. [GUATEMALA: CHIQUMULA: Montana Castilla, vicinity of Mon- taña Cebollas, along Rio Lucía Saso, 3 mi SE of Quezaltepeque, alt. 1200-1500 m, Nov 6, 1939, J. A. Steyermark 31844 (Holo- type F! Photo us!) ] Macvaughiella standleyi (Steyermark) R. M. King and H. Robinson, Sida 3(4): 282. 1968. Additional specimens examined: MEXICO: VERACRUZ: Zacuapan, Sulphur Spring, Purpus 2189a (F, NY). GUATEMALA: JALAPA: vicinity of Jalapa, Standley 76699 (F, US), 77406 (r); Jutiapa: quebrada above Ovejero, on road between Monjas (Dept. Jalapa) and El Progreso, alt. ca 1400 m, Standley 77645 (F); Lago Retana, between Ovejero and Progreso, Steyermark 32041 (F). EL SALVADOR: LA LIBERTAD: road to La Libertad, Molina, Burger & Wallenla 16687 (F); between Guayabal and Río Guaza, Calerón 1936 (US); SAN VICENTE: vicinity of San Vincente, Standley & Padilla 3639 (F, GH). HONDURAS: CHOLUTECA: vicinity of San Marcos de Colón, Standley 15903 (F, GH, US). EL PARAISO: along or near Rio California, between Rio Choluteca and Jacaleapa, Standley 17176 (F); Rio Choluteca at Ojo de Agua, L. O. Willliams 17310 (F, GH, US), Standley 4683 (F, GH). MORAZAN: El Quebracho, above El Zamorano, Standley 357 (F); quebrada El Horno, entre El Frijolar y Tabla Grande, Molina 833 (F); Matorrales secos de Puente Colorado, al norte de Teguci- galpa, Molina 3885 (F); near Río El Quebracho above El Jicarito, Standley 27758 (F, GH, MO, US); drainage of the Río Yeguare about Longitude 87° W. and Latitude 14° N., L. O. Williams 17039 (F, GH, 1970] Compositae-Eupatorieae — King and Robinson 103 MO, US), Molina 1758 (F, GH, MO, US), L. O. Williams & Molina s.n. (F, US); slopes of Cerro de Uyuca, along trail between Hoya Grande and Valle Encantado, Standley 15246 (F, GH, NY, US); region of El Jicarito, above El Zamorano, Standley 25134 (GH). It should be noted that the ranges of the two varieties do overlap in Veracruz as indicated by collections cited as Purpus 2189. Oaxacania B. L. Robinson & J. Greenm. Amer. Journ, Sci. III. 50: 151, 152. 1895. (Type species, Oazacania malvaefolia B. L. Robinson & J. Greenm.) Sprawling herbs or sub-shrubs up to one meter long, somewhat woody at base, many branched. Stems striate, covered with glandular hairs. Leaves alternate, blades thin, orbicular or reniform, 3-7 cleft two thirds or more toward base, lobes oblong, glandular pubescent on both surfaces, up to 3.5 em wide and long, petioles 2-3 cm long. In- florescences numerous, short peduncled, monocephalic; heads discoid, campanulate, 1-1.2 cm high (including style branches), ca 100 flowered, receptacle paleaceous, convex, phyllaries ca 30-40, in 5 series, lanceolate, 3 nerved, corollas white, campanulate, ca 6.5 mm long (including lobes), 5 lobed, outer surface covered with numerous long stalked glands; stamens 5, anthers appendaged, the latter ca 60 u long, filaments with distinct collars developed just below the anther; style branches 2, greatly exserted at maturity, papillose, style gla- brous; pappus obsolete or a short lacerate crown; achenes linear- oblong, strongly compressed, 4 angled, setose, setae biseriate; base of achene forming a distinct carpopodium; pollen spherical, tri- colpate, very distinctly spinose, ca 25 u in diameter, spines up to 2 u long, chromosome number not determined. Oaxacania malvaefolia B. L. Robinson & J. Greenm. Amer. Journ. Sci. III. 50: 151. 1895. [MEXICO: OAXACA: dry cliffs, Tomellin Canyon, 3,000 ft Dec. 22, 1894, C. G. Pringle 6117 (Holotype GH!, Isotypes G!; ENCB!; MEXU!; MICH!; MO! NY!; UC!; us!)] Additional specimens examined: MEXICO: OAXACA: Chiltapin; Oaxaca side of Río Santa Lucia, Purpus 3084 (F, MO, NY, UC, US); De Almoloyas a Santa Catarena, Conzatti 1654 (F, MEXU, Us). Oaxacania bears considerable resemblance to Cartero- thamnus R. M. King, but is clearly distinguished by char- acters indicated in the following table: 104 Oaxacania achenes flattened long stalked glands on co- rolla obsolete pappus thin hardly expanded tips of the style small firm cells forming col- lar at the base of anther Pianaltoa P. Taubert Engl. Rhodora [Vol. 72 Carterothamnus symmetrical achenes corolla glabrous pappus of numerous scales and one long seta thick knoblike tips of the style very lax cells forming collar at base of anther Bot. Jahrb. 20: 454. 1896. (Type species, Planaltoa salviifolia Taubert) Woody shrubs or sub-shrubs, Leaves simple, alternate, sessile. Peduncles not strongly differentiated. Inflorescence polycephalic. Heads turbinate or cylindrical, 4-5 flowered. Phyllaries subequal, in 2-3 series, lanceolate. Corollas slender, tubular, regular, 5 lobed, outer surface with both glands and hairs. Hairs and stalks of glands on corolla often biseriate. Receptacle flat, naked. Anther appendages rather large with slightly reflexed margins. Pollen spherical, tri- colpate, distinetly spinose. Style branches elongate, exserted at maturity, narrowly obtuse; basal node of style covered with non- glandular hairs. Pappus lacking. Achenes 5-6 angled, glabrous. The species of Planaltoa may be distinguished by the following key: 1. Leaf blades oblong-elliptical, most hairs of plant with glandular tips, inner surface of corolla glabrous. ......................- P. salviifolia 1. Leaf blades lanceolate, slightly auriculate, hairs of plant without glandular tips, lobes with shorter uniseriate hairs both on the inner and outer surfaces. .................................... P. lychnophoroides Planaltoa salviifolia Taubert, Engl. Bot. Jahrb. 21: 454. 1896. [BRaziL: GOIÁS: Serra dos Pyrenos, 1892-1893, Ule 2971 (B, de- stroyed) ] Plants ea. one meter tall? Stems few-branched, terete. Stems, leaves, phyllaries, and outer surface of corolla covered with dense long pubescence, hairs bearing minute secretory cells apically. Leaves rigid, sessile. Blades oblong-elliptical, 6-10 mm wide, apex obtusely pointed; margins with minute recurved teeth in the upper half. Heads ca 15 mm high (including style branches), ca 4-5 flowered. Phyllaries 5-7, imbricated, in 2-3 series, ca 12 mm long. Florets ca 8 mm long. Lower outer surface of corolla with a few blunt, uni- seriate, pitted, nonglandular projections; outer surface with long biseriate hairs bearing minute secretory cells apically, upper ?4 of outer surface with scattered subsessile glands, inner surface of corolla 1970] Compositae-Eupatorieae — King and Robinson 105 glabrous. Stamens 5. Anthers (including appendages) ca 3 mm long and 250 & wide; appendages 250 u wide and 200 u long. Style branches papillose, slightly broader apically, exserted at maturity. Achenes prismatie, dark brown at maturity, ca 3 mm long, ca .75 mm wide. Pollen 25 u in diameter. Chromosome number not deter- mined. Additional specimens examined: BRAZIL: GOIÁS: Pirineus, Corumba, Macedo 3715 (MO, NY, US); without precise locality, Glaziou 21618 (G, GH, Us). Without precise locality, Voyage d' Auguste de Saint-Hilaire de 1816 à 1821 (F). Planaltoa lychnophoroides G. M. Barroso, Sellowia 17: 79. 1965. [BRAZIL: GorÁS: Cachoeira de Vargem Grande, 1894-95, Glaziou 21594. (Holotype G!; Isotype F!)] Plants ca. one meter tall. Stems branched, terete. Stems, leaves, phyllaries, and outer surface of corolla with dense long nonglandular pubescence. Leaves rigid, sessile, lanceolate, slightly auriculate, 2.5-3.0 em long, 3.0-5.0 mm wide; lower surface with occasional sub- sessile glands; margins reflexed, serrate. Heads ca 10 mm high (including style branches), 4-5 flowered. Phyllaries 5-7, imbricated, in two series, ca 10 mm long; outer surface with glands. Florets ca 7-8 mm long. Outer surface of corolla with long uniseriate, non- glandular hairs; lower outer surface of corolla with many biseriate non-glandular setae intermixed with hairs; lobes with shorter uni- seriate hairs both on the inner and outer surfaces; subsessile glands sparse on the upper outer surface of corolla. Stamens 5. Anthers (including appendages) ca 2.5 mm long and 450 u wide; appendages 250 u wide and 300 z long. Style branches long-papillose, exserted at maturity. Achenes prismatic, dark brown at maturity, ca 3 mm long, ca .75 mm wide. Pollen 25 » in diameter. Chromosome number not determined. Additional specimens examined: BRAZIL: GOIÁS: 7 km W of Veadeiros, Irwin, Greer, Souza, Santos 12909 (MO, TEX, US). DEPARTMENT OF BOTANY SMITHSONIAN INSTITUTION WASHINGTON, D.C. 20560 THE GENUS ACANTHOSPERMUM (COMPOSITAE- HELIANTHEAE-MELAMPODINAE) : TAXONOMIC CHANGES AND GENERIC AFFINITIES Top F. STUESSY' TAXONOMIC CHANGES Blake (1921) listed eight good species in his revision of the genus Acanthospermum. One year later he himself added two more species to the genus, A. brachyceratum and A. leptolobum, both from the Galápagos Islands. Another earlier described species, A. lecocarpoides (Robinson and Greenman, 1895), was also from these same islands. The genus has stood unchanged with 10 species until just re- cently Cronquist and Stuessy (in Cronquist, 1969) removed the Galápagos species into the related genus Lecocarpus based upon total morphology (see Cronquist, 1969, for ad- ditional information regarding Lecocarpus and these trans- fers). In the present study based on herbarium material of Acanthospermum,? A. donii is treated as being conspecific with A. microcarpum which further reduces the number of species to six. Of the three sections of the genus recognized by Blake (1921), section Lecocarpopsis is now defunct due to removal of the Galápagos species. The following list includes the presently recognized taxa and correct sectional placements : Section Ceratochlaena DC. A. hispidum DC A. humile (Sw.) DC. (photograph of B isotype, TEX!) A. microcarpum Rob. (holotype, GH!) "Thanks are due Dr. A. Cronquist for helpful suggestions regarding the final manuscript. *Appreciation is expressed to the curators of the following herbaria from which loans of specimens were made (abbreviations after Lan- jouw and Stafleu, 1964): ARIZ, F, GH, MICH, NY, SMU, TEX, UC, UPS, US. 106 1970] Acanthospermum — Stuessy 107 A. simile Blake (isotypes, F! NY[2]!; frag. of BM holo- type, GH!; photograph of holotype, GH!; photograph of holotype and frag. of K isotype, US!; photograph of F isotype, F!) Section Acanthospermum (Section Xanthoides DC.) A. australe (Loefl. Kuntze A. consobrinum Blake (frag. of BM holotype, GH ! ; photo- graph of holotype, US!). Blake's sectional characters, key to species, descriptions, and synonymy for these remaining six species of the genus are adequate and are not duplicated here. Both Acanthospermum donii Blake (type probably from Ecuador; frag. of BM holotype, GH!; photograph of holo- type, US!) and A. microcarpum (type from Galápagos Is.) are known from only a very few collections. After examin- ing available material, I consider the fruit? differences (mainly size), emphasized by Blake as specific criteria, to be intergrading and of trivial taxonomic significance. This inclusion removes A. microcarpum from the numbers of endemic Galápagos Is. species. GENERIC AFFINITIES There are three genera in the Compositae (all subtribe Melampodiinae) that possess vascularized inner involucral bracts each tightly enclosing and fused with a single ray achene: Acanthospermum, Lecocarpus, and Melampodium. Bentham (1873) felt that both “The east tropical Acantho- spermum and the Galapagian Lecocarpus might be included in the widely spread Melampodium" (p. 434). As already mentioned, there has been some misunderstanding of the generic limits of Acanthospermum and Lecocarpus, but Acanthospermum and Melampodium rarely have been con- fused. Despite similarities the 3 genera can be distinguished as follows (in part after Bentham and Hooker, 1873, and Hoffman, 1890) : “Fruit” is defined as the ray achene and the enclosing inner in- volucral bract (Robinson, 1901). 108 Melampodium (1) Fruit apex hooded or not hooded, sides smooth to tuberculate. (2) Leaves entire to variously lobed. (3) Organs only rarely glandular. (4) Plants annual herbs or suffruticose perennials. (5) Distribution main- ly in Mexico and Cen- tral America. Rhodora Acanthospermum (1) Fruit apex not hooded, sides with straight or hooked prickles. (2) Leaves entire to variously lobed. (3) Organs agandu- lar. (4) herbs. Plants annual (5) Distribution mainly in North and South America [Vol. 72 Lecocarpus (1) Fruit apex rim- med with a broad wing or with one to several long prickles, sides smooth. (2) Leaves markedly pinnatifidly cleft or divided. (3) Organs markedly glandular. (4) Plants shrubby perennials (annuals?). (5) Distribution re- stricted to Galapagos Is. (in- troduced to Old World), one non- endemic species on Galápagos Is. (A. microcarpum). The classical distinction between Melampodium and Acanthospermum has been based mainly on fruits without prickles versus the prickled condition, respectively. How- ever, there is one Melampodium species, M. longifolium, that possesses “horns” (enlarged apical extensions of the enclosing bract) similar to the apical prickles of A. simile and A. microcarpum. Nevertheless, this character serves with other characters to successfully delimit the two genera. Melampodium has been surveyed extensively for chromo- some numbers (for a review see Stuessy, 1969) and n = 10 is found in the largest number of species in the genus. The few reported counts of Acanthospermum are n = 10: and n = 11°; these support the presumed close phyletic relation- *A. australe, Carlquist (1954). 5A. hispidum, Miège (1960, reported as 2n = 22); A. australe, Turner and Irwin (1960) and Coleman (1968). 1970] Acanthospermum — Stuessy 109 ship of the two genera. Cytological information regarding Lecocarpus is lacking. ACADEMIC FACULTY OF ORGANISMIC AND DEVELOPMENTAL BIOLOGY AND THE HERBARIUM THE OHIO STATE UNIVERSITY COLUMBUS 43210 LITERATURE CITED BENTHAM, G. 1873. Notes on the classification, history, and geo- graphical distribution of Compositae. Jour. Linn. Soc. Lond. Bot. 13: 335-577. , and J. D. HOOKER. 1873. Acanthospermum, Leco- carpus and Melampodium. Genera Plantarum 2: 348-349. BLAKE, S. F. 1921. Revision of the genus Acanthospermum. Con- trib. U. S. Nat. Herb. 20: 383-392. CARLQUIST, S. 1954. In Documented chromosome numbers of plants. Madrono 12: 210. COLEMAN, J. R. 1968. Chromosome numbers in some Brazilian Compositae. Rhodora 70: 228-240. CRONQUIST, A. 1970. Compositae. I» I. L. Wiggins and D. M. Porter, Flora of the Galápagos Islands. Stanford University Press, Stanford (in press). HOFFMANN, O. 1890. Melampodinae. In A. Engler and K. Prantl, Die natürlichen Pflanzenfamilien 4(5): 214-219. LANJOUW, J., and F. A. STAFLEU. 1964. The herbaria of the world. Index Herbariorum. Part 1. Ed. 5. Regnum Vegetabile 31: 205-228. MiEGE, J. 1960. Troisième liste de nombres chromosomique d'espéces d'Afrique occidentale. Ann. Fac. Sci. Univ. Dakar 5: 75-85. ROBINSON, B. L. 1901. Synopsis of the genus Melampodium. Proc. Amer. Acad. Arts & Sci. 36: 455-466. , and J. M. GREENMAN. 1895. On the flora of the Galápagos Islands, as shown by the collection of Dr. G. Baur. Amer. J. Sci., Ser. 3, 50: 135-149. STUESSY, T. F. 1970. Chromosome studies in Melampodium. (Com- positae-Heliantheae). Madroño (in press). TURNER, B. L., and H. S. Irwin. 1960. Chromosome numbers in the Compositae. II. Meiotic counts for fourteen species of Brazilian Compositae. Rhodora 62: 122-126. NOMENCLATURE OF THE LUPINUS ARGENTEUS AND L. CAUDATUS COMPLEXES LLovp W. HESS! AND DAVID B. DUNN The genus Lupinus has long been recognized as one of the very complex groups of plants and the many names pub- lished add to the confusion. Among the groups of lupines, the L. argenteus complex, which occupies the Great Basin and the Rocky Mountain regions, has been one of the most difficult. Hybridizing studies on this complex were con- ducted at the Rocky Mountain Biological Laboratory over a five year period, These studies demonstrated that hy- bridization between the L. argenteus and the L. caudatus complexes can explain many of the variations which were named and that the two form polymorphic interacting taxa covering much of the same geographic area. The long lists of synonymy add little to the basie understanding of the problem and will be included in the monographic paper in the University Museum Contribution Series, Only the basic names of the group are presented here along with the rea- sons for the suggested changes in rank. The remaining names of related taxa, in which no change is suggested, are omitted. Since the key to all of the taxa of the group will be in the monograph it is not included in the present paper. The two complexes presented here are very closely re- lated through introgressive hybridization with L. caudatus at the xerophytic extreme and some of the subspecies of L. argenteus at the mesophytic extreme. The flower shape of both is very similar with the banner generally reflexed well above the midpoint. There is a patch of pubescence in the central area of the banner on the dorsal side which may be covered by the upper lip of the calyx. In a few taxa this patch of pubescence is absent. There is a well developed spur at the base of the upper lip of the calyx in L. caudatus Research supported by NSF-GB-5572, at the University of Mis- souri-Columbia, Columbia, Mo. Present address of Hess, SREL, Aiken, South Carolina 29801. 110 1970] Lupinus argenteus — Hess and Dunn 111 and the trait extends into the L. argenteus complex. The flowers of the taxa treated here are usually from 8-12 mm long and this separates them from the smaller related L. parviflorus complex which will be treated separately. There are 80 synonyms involved in the complete nomenclature so only the basonyms will be cited below. l. Lupinus argenteus Pursh (subsp. argenteus var. ar- genteus) Fl. Am. Sept. 2: 468, 1814. Type: Banks of the Kooskoosky, — now considered to be the Clearwater River, M. Lewis 4. (Holotype: K, former Lambert Herb.). The closest matching material presently is in Montana, on the eastern base of the Rocky Mountains, The habitat is arid plains and sagebrush from Canada to Arizona. la. Lupinus argenteus subsp. argenteus var. tenellus (Dougl. ex. G. Don) Dunn, Leafl. W. Bot. 7: 254, 1955. Type: Vicinity of Grand Rapids of Columbia River, Douglas 277. (Holotype: CGE). An outcropping genome, completely sympatric with argenteus, the flowers narrow viewed lat- erally and the leaflets often very narrow. The taxon repre- sents introgression from more xerophytic taxa from an- other complex. It is more abundant than argenteus in the more arid areas but it has not demonstrated dominance of an ecological area. lb. Lupinus argenteus subsp. rubricaulis (Greene) Hess & Dunn, comb. nov. Basonym: L. rubricaulis Greene, Pl. Baker. 3: 35, 1901. Type: Crested Butte, Colorado, Baker 5342. (Holotype: ND; Isotypes MIN, MO, RM). The taxon is an altitudinal subspecies associated with montane forests, commonly in the open park areas in the spruce-fir zone but intergrading in numerous places through the Rocky Moun- tains with argenteus or tenellus at its lower limits and with spathulatus at its upper limits. le. Lupinus argenteus subsp. spathulatus (Rydb.) Hess & Dunn, comb. nov, Basonym: L. spathulatus Rydb., Bull. Torr. Bot. Club 29: 204, 1902. Type: Wasatch Mountains in 1869, S. Watson 225, (Holotype: NY). The taxon is an- 112 Rhodora [Vol. 72 other altitudinal subspecies at the subalpine zone, commonly growing under and among spruce-fir forests up to timber- line. It grades into rubricaulis completely and either of the subspecies may simulate the appearance of the other by ecological modification. The broad flat leaflets may be found at lower elevations in shaded areas in aspen groves, while the narrow leaflets may be produced at high elevations in exposed locations. However, there is as much as a month’s difference in the flowering times at the different elevations, requiring genetic alterations to accommodate the ecological differences. The material from the type locality appears quite distinct but can be matched by numerous specimens from other areas so that the same process appears to be functional throughout the range of subspecies spathulatus. 2. Lupinus X alpestris A. Nels. Hybridity suggested (L. caudatus X L. argenteus). Bull. Torr. Bot. Club 26: 127, 1899. Type: Medicine Bow Mts., Wyoming, prob. Univ. summer camp, E. Nelson 5070. (Holotype: RM; Isotype: MO). The original material is closest to subsp. rubricaulis but the upper surface of the leaflets is finely pubescent, indicating introgression from L. caudatus. The multiple locations where hybridization has taken place have pro- duced a whole range of intermediate forms throughout the geographie region in which caudatus and the argenteus complex occur. Some of these have become relatively stable intermediates at various levels between the two taxa with numerous names applied by various investigators. In most areas the argenteus traits dominate and the taxon is gen- erally sympatric with argenteus, However in the Great Basin, where there is greater aridity, the vegetative char- actistics of caudatus prevail, while the floral characteristics of argenteus tend to be dominant. It seems preferable to retain the hybrid binomial designation to indicate what is involved, even if the entity is highly variable, rather than reduce it to one of the meaningless varietal names which have been published. 3. Lupinus caudatus Hell. (subsp. caudatus) Proc. Calif. Acad. 2: 197, f. 61, 1862. Type: Carson Valley, Kellogg. 1970] Lupinus argenteus — Hess and Dunn 113 (Holotype: CAS 62286). Associated with sage brush, ex- tending into the arid pine zones along the east side of the Sierra Nevada Mts. The plants have a densely sericeous hair covering throughout and the upper lip of the calyx has a well developed spur at the base. The spur almost disappears in a clinal gradient eastward to the Laramie area, The genetic markers of ciliation near the claws of the wings and keel, on the margins and laterally, are gen- erally present but become less frequent eastward. 3a. Lupinus caudatus subsp. montigenus (Heller) Hess & Dunn, comb. nov. Basonym: L. montigenus Heller, Muh- lenbergia 6: 109, f. 16, 1910. Type: Mt. Rose, Nevada, Heller 9880. (Holotype: RENO; Isotypes: MO, RM, UC). This taxon is viewed as an altitudinal subspecies, restricted to the higher mountains of the east side of the Sierra Nevada Mts. The flowers are the largest of the caudatus complex and at the higher peaks have very little develop- ment of the calyx spur, with the banner reflexing at the midpoint but the gradation into caudatus at the lower zones appears complete, with the spur becoming very pro- nounced. 3b. Lupinus caudatus subsp. cutleri ( Eastw.) Hess & Dunn, comb. nov. Basonym: L. cutleri Eastw., Leafl. W, Bot. 4: 192, 1945, Type: 18 mi NW Fort Defiance, Arizona, Cutler 2141. (Holotype: CAS. Isotypes: DS, MO). This taxon appears restricted to the mountains of the basin area of southeastern Utah, northern Arizona, and north western New Mexico. It has predominantly caudatus characteristics and appears to intergrade with subspecies argophyllus but also appears to have characteristics derived from introgres- sion from an undetermined source. 3c. Lupinus caudatus subsp. argophyllus (Gray) Phillips, Res. Stud. Wash. St. Coll. 23: 200, 1955. Basonym: L. decumbens Nutt. var. argophyllus Gray, Mem. Am. Acad. 4: 37, 1849. Type: Santa Fe, New Mexico, Fendler. (Holo- type: GH). This taxon is the product of early introgression between caudatus and argenteus. Floristically the traits 114 Rhodora [Vol. 72 are predominantly those of caudatus, with all the char- acteristic caudatus markers, including the ciliation near the claws of the wings and keel, and a well developed spur on the calyx. Vegetatively the characteristics are predom- inantly those of argenteus, including the short petioles, and the leaflets are frequently nearly glabrous above. Subspe- cies argophyllus is quite distinct from any of the material treated as X alpestris, and appears to be a stabilized taxon. It does appear to intergrade into caudatus in southwestern Colorado, 4. Lupinus X inyoensis Heller (pro species). Hybridity suggested (L. caudatus X L. palmeri). Muhlenbergia 2: 211, 1906. Type: Foothills W of Bishop, California, Heller 8312, (Isotypes: CAS, ISC, MO, UC). The only suggested change is the insertion of the X to indicate hybrid origin. Three of the four isotypes had spreading hairs, the main trait derived from palmeri, while the fourth had the typical appressed hairs of caudatus suggesting that the original col- lection represented a mixed population. In several field population samples made in 1968 in Nevada there were mixed plants of L. palmeri and L. caudatus with occasional hybrid plants. The population of > inyoensis appears re- stricted to Inyo and Mono Cos., California. The authors wish to express their appreciation to the curators of the several herbaria indicated in the text by the codes designated in Index Herbariorum. Citations of their specimens will be in the monograph. Appreciation is also expressed to Rocky Mountain Biological Laboratory for the use of their facilities in summer. BOTANY DEPARTMENT UNIVERSITY OF MISSOURI-COLUMBIA COLUMBIA, MISSOURI 65201 FLORA OF THE WOLF ISLANDS PART III: THE MARINE ALGAE EDWARD J. HEHRE,' JOAN R. CONWAY’, AND RICHARD À. STONE? The vascular plants of the Wolf Islands have been investi- gated thoroughly by A. R. Hodgdon and R. B. Pike (Rho- dora, 65:82-96, 364-365, 1963; 66:61-62, 140-155, 413-416, 1964; 71:297-302, 1969). At their suggestion we have un- dertaken a floristic study of the marine algae of these islands. We have made three visits (4 June and 13 August, 1968 and 22 February, 1969) to South Wolf Island, as it affords the most accessible intertidal regions. Numerous small coves offer a variety of protected habitats. However, most of the coast is exposed. There is a great expanse of intertidal region — the average tidal amplitude is about 17 feet. Ninety-one taxa are recorded from South Wolf Island, including 27 Chlorophyceae, 26 Phaeophyceae and 38 Rho- dophyceae. This compares with 114 taxa recorded from Campobello Island during a three year study (Stone, et al., in press). Future trips will include the other Wolf islands and may yield additional species. The algae were collected and preserved in a brine solution for laboratory identification and processing. Voucher speci- mens are deposited in the Herbarium of the University of New Hampshire and Southampton College, Long Island, New York. The nomenclature of Parke and Dixon (1963) has been applied in most cases, except for the Acrochae- tiaceae (Papenfuss, 1947). ‘present address: Division of Natural Sciences, Southampton Col- lege, Southampton, L.I., New York. "present address: Department of Botany, University of Massachu- setts, Amherst, Mass. ‘present address: Department of Biology, University of Massachu- setts, Boston, Mass. 115 116 Rhodora [Vol. 72 CHECKLIST CHLOROPHYCEAE: Chaetomorpha melagonium (Weber et Mohr) Kiitzing Codiolum gregarium A. Braun C. petrocelidis Kuckuck C. pusillum (Lyngbye) Kjellman Epicladia flustrae Reinke Enteromorpha groenlandica (J. Agardh) Setchell et Gardner . intestinalis (L.) Link . linza (L.) J. Agardh . marginata J. Agardh . micrococea Kützing . minima Nägeli Monostroma fuscum (Postels et Ruprecht) Wittrock forma blyttii (Areschoug) Collins M. grevillei (Thuret) Whittrock M. leptodermum Kjellman M. pulchrum Farlow Prasiola stipitata Suhr in Jessen Pseudendoclonium marinum (Reinke) Aleem et Schulz Rhizoclonium riparium (Roth) Harvey var. implerum (Dillwyn) Rosenvinge R. tortuosum Kützing Spongomorpha arcta (Dillwyn) Kützing S. lanosa (Roth) Kützing S. spinescens Kützing Ulva lactuca L. Urospora collabens (C. Agardh) Holmes et Batters U. penicilliformis (Roth) Areschoug U. speciosa (Carmichael) Leblond et Hamel U. wormskjoldii (Mertens in Hornem) Rosenvinge PHAEOPHYCEAE: Agarum cribrosum (Mertens) Bory Alaria esculenta (L.) Greville Ascophyllum nodosum (L.) Le Jolis Asperococcus echinatus (Mertens) Greville Chorda tomentosa Lyngbye Chordaria flagelliformis (O. F. Müller) C. Agardh Desmarestia aculeata (L.) Lamouroux Dictyosiphon foeniculaceus (Hudson) Greville Ectocarpus confervoides (Roth) Le Jolis Elachista fucicola (Velley) Areschoug Fucus distichus L. ssp. distichus Powell F. distichus L. ssp. edentatus (de La Pylaie) Powell q q q 1 q 1970] Marine Algae — Hehre, Conway and Stone 117 F. distichus L. ssp. evanescens (C. Agardh) Powell F. spiralis L. F. vesiculosus L. Isthmoplea sphaerophora (Carmichael) Kjellman Laminaria digitata (Hudson) Lamouroux L. longicruris De la Pylaie L. saccharina (L.) Lamouroux sensu Wilce Leathesia difformis (L.) Areschoug Myrionema strangulans Greville Petalonia fascia (O. E. Miller) Kruntze Pilayella littoralis (L.) Kjellman Ralfsia fungiformis (Gunnerus) Setchell et Gardner Scytosiphon lomentaria (Lyngbye) Link Sphacelaria cirrosa (Roth) C. Agardh RHODOPHYCEAE: Ahnfeltia plicata (Hudson) Fries Antithamnion cruciatum. (C. Agardh) Nägeli A. floccosum (Müller) Kleen Audouinella membranacea (Magnus) Papenfuss Bangia fuscopurpurea (Dillwyn) Lyngbye Ceramium deslongchampsii Chauvin in Duby var. hooperi (Harvey) Taylor C. rubrum (Hudson) J. Agardh Chondrus crispus Stackhouse Choreocolax polysiphoniae Reinsch Clathromorphum circumscriptum (Strømfelt) Foslie Corallina officinalis L. Cystoclonium purpureum (Hudson) Batters Euthora cristata (C. Agardh) J. Agardh Gigartina stellata (Stackhouse) Batters Halosaccion ramentaceum (L.) J. Agardh Hildenbrandia prototypus Nardo Kylinia alariae (Jónsson) Kylin K. secundata (Lyngbye) Papenfuss Lithophyllum corallinae (Crouan frat.) Heydrich Lithothamnium glaciale Kjellman Membranoptera alata (Hudson) Stackhouse Petrocelis middendorfii (Ruprecht) Kjellman Peyssonelia rosenvingii Schmitz in Rosenvinge Phycodrys rubens (L.) Batters Phymatolithon lenormandi (Areschoug) Adey Plumaria elegans (Bonnemaison) Schmitz Polyides rotundus (Hudson) Greville Polysiphonia lanosa (L.) Tandy P. urceolata (Lightfoot ex Dillwyn) Greville 118 Rhodora [Vol. 72 Porphyra leucosticta Thuret in Le Jolis P. linearis Greville P. miniata (C. Agardh) C. Agardh P. umbilicalis (L.) J. Agardh P. umbilicalis (L.) J. Agardh f. epiphytica Collins Ptilota serrata Kützing Rhodochorton purpureum (Lightfoot) Rosenvinge Rhodomela confervoides (Hudson) Silva Rhodymenia palmata (L.) Greville ACKNOWLEDGEMENTS The authors wish to thank Dr. A. C. Mathieson for his assistance in the identification of several plants; Radcliffe and Sumner Pike for their hospitality and generosity which made this study possible. We are grateful to Drs. A. R. Hodgdon and R. B. Pike for financial support for travel from the “Wolf Island Fund” throughout this study. DEPARTMENT OF BOTANY UNIVERSITY OF NEW HAMPSHIRE DURHAM 03824 REFERENCES PAPENFUSS, G. A. 1947. Further Contributions toward an Under- standing of the Acrochaetium-Rhodochorton Complex of the Red Algae. Univ. of the Calif. Publ. Bot. 18:433-447. PARKE, M. and P. S. Drxon. 1968. Checklist of British Marine Algae — Second Revision. J. mar. biol. Ass. U.K. 48:783-832. STONE, R. A., E. J. HEHRE, J. R. Conway, and A. C. MATHIESON, A Preliminary Checklist of the Marine Algae of Campobello Island, New Brunswick, Canada. Rhodora (in press) TAYLOR, W. R. 1957. The Marine Algae of the Northeast Coast of North America. VIII & 509 pp. Univ. Michigan Press, Ann Arbor. A SYNOPSIS OF THE GENUS SUESSENGUTHIA (ACANTHACEAE) DIETER C. WASSHAUSEN During the course of routine identifications of Acantha- ceae from South America, I have found two undescribed species of the genus Suessenguthia Merxmiiller. In an at- tempt to make proper placement of the species, the mono- graph of Sanchezia and related genera (Leonard & Smith, 1964) was used. In this paper Sanchezia leucerythra was described as a new species. After examining the type, its four fertile appendaged stamens indicate that it should be transferred to the genus Suessenguthia, The two new species are described and the new combination is made. A key is also provided. The loan of material from the New York Botanical Garden is gratefully acknowledged. SUESSENGUTHIA H. Merxmiiller, Mitteilungen der Botanischen Staatssamm- lung Miinchen [1], Heft 6: 178, pl. 1953. (Type species, Suessenguthia trochilophila Merxm.) Shrubs; stems erect, quadrangular, sulcate ; leaves oppo- site, short-petioled, cystoliths conspicuous; flowers solitary or fascicled, usually large and conspicuous, borne in heads, spikes or cymes, the flower cluster subtended by small or often large and partly connate bracts, these sometimes colored ; calyx 5-parted, the segments subequal ; corolla lilac or red, the tube cylindric, 5-lobed, the lobes equal; stamens 4, the longer two exserted, the shorter two included or barely reaching the throat of the corolla; anthers 2-celled, longitudinally ciliate, mucronate at base; ovary disc minute, sessile; style glabrous; ovules 4 per locule. The species of Suessenguthia can be distinguished by the following key: a. Inflorescence capitate or cymose. b. Inflorescence capitate; bracts ovate, acuminate, corolla minutely pubescent t S nas L ennt stint nenne tnn 1. S. leucerythra b. Inflorescence cymose; bracts obovate, caudate; corolla densely Bae Pi SEE o t D n IL NE au NOS. ers 2. S. vargasii 120 Rhodora [Vol. 72 10, 20: 4 ) O- S. leucerythra | N A M- S. vargasii yor eT A- S. cuscoensis F ` S A- S. trochilophila YY ` D J); BN M 30 — — ` =>" Z 80 70 Fig. 1. Distribution of Suessenguthia. a. Inflorescence spicate. c. Fascicles all sessile in elongated spikes; bracts subtending the floral cluster elliptic ..........ssse ees 3. S. cuscoensis c. Basal fascicles of flowers pedunculate; bracts subtending the floral cluster triangular-cuspidate ................ 4. S. trichilophila 1, Suessenguthia leucerythra (Leonard & Smith) Wass- hausen, comb. nov. Sanchezia leucerythra Leonard & Smith, Rhodora 66: 318. 1904. 1970] Suessenguthia — Wasshausen 121 PERU: Junin: dense forest near Perene Bridge, Rio Paucartambo Valley, 700 m, June 19, 1929, E. P. Killip & A. C. Smith 25271 (Holotype NYv!). Shrub 2-5 m tall, stem quadrangular, glabrous, sulcate ; leaf blades lanceolate, up to 18 cm long and 5 cm wide, acuminate, the tip itself obtuse, the upper surface glabrous, minutely papillose, cystoliths numerous, 0.25 mm long, on the lower surface the costa and lateral veins (ca 8 pairs) conspicuous, pilose, the hairs appressed, the margins short repand-crenate; petiole up to 2 cm long, appressed-pilose ; inflorescence capitate, 3.5 em long, 3 cm wide; lowermost bracts lanceolate, up to 4.5 cm long, 1.5 cm wide, acuminate, pilose, the hairs appressed, 0.5 mm long, bracts subtending the flower cluster ovate, 3 cm long, 1.3 cm wide, acuminate, sericeous, the hairs appressed and 1 mm long, distally deep pink-purple, basally white; bractlets 3 cm long, 1 cm wide, acuminate, sericeous, the hairs appressed, 0.5 mm long, brown; calyx segments 27-30 mm long, 2-6 mm wide, subequal, linear-lanceolate, slender-acuminate, sericeous ; corolla pale red, basally white, 4-4.5 cm long, basally 3-5 mm wide, 10 mm wide at throat, minutely pubescent with recurved hairs, the lobes ca 10-15 mm long, broadly rounded and short-emarginate, erect, densely retrorsely pubescent ; stamens 4, cream white, the two longer exserted 7-10 mm beyond the throat of the corolla, the two shorter included and barely reaching the throat of the corolla; ovary pilose ; fruit not seen. OTHER SPECIMENS EXAMINED: PERU: Junin: open place in forest, Rio Negro, 800 m, August 14, 1960, F. Woytkowski 5794 (Us) ; Cusco: Prov. de La Convencion, *Subiendo a Ichigurato; Alto Urubamba", April 16, 1966, C. Vargas C. 17235 (CUZ, US). 2. Suessenguthia vargasii Wasshausen, sp. nov. PERU: Cusco: Prov. de Quispicanchis, forest, between Quince Mil and San Lorenzo, 700 m, July 25, 1957, C. Vargas C. 11732 ( Holo- type US! Isotype CUZ). Frutex, caulibus quadrangularibus ; folii lamina oblongo- elliptica acuminata, apice curvata, basi angustata, glabra, integra vel undulata, costa et venis prominentibus; in- florescentia pauciramosa, cymosa, floribus in fasciculis; 122 Rhodora [Vol. 72 bracteae obovatae, caudatae, sericeae; bracteolae anguste oblongae, acuminatae; calycis segmenta lanceolata vel oblanceolata, subaequalia, sericea; corolla pallide rubra, dense sericea, lobis late rotundatis et leviter emarginatis; stamina 4, longiora exserta, breviora sub-exserta; ovarium dense sericeum. Shrub 2-5 m tall, stem quadrangular, glabrous, sulcate; leaf blades oblong-elliptic, 10-16 cm long, 3.5-5 em wide (the lowermost probably larger), gradually narrowed to a slender curved apex (the tip blunt), narrowed at base, glabrous, entire or undulate, the costa and lateral veins (6 to 8 pairs) prominent beneath; petioles 5-12 mm long, connate at base by a small ridge or narrow band of leaf tissue; flowers borne in fascicles in a few-branched cymose inflorescence, 4 cm long, 2 cm wide; lowermost bracts obovate, up to 2 em long, 1.3 em wide, caudate, sericeous, the hairs appressed, yellowish ; bracts subtending the flower cluster obovate, up to 3 cm long, 1.3 em wide, caudate, densely sericeous, the hairs appressed, yellow; bractlets narrowly oblong, 3 cm long, 7 mm wide, acuminate, densely sericeous, the hairs appressed, distally deep pink purple; calyx segments up to 30 mm long, subequal, 4-5 mm wide, lanceolate to oblanceolate, slender-acuminate, sericeous ; corolla drying pale red, 4-6 cm long, basally 4 mm wide, 8 mm wide at throat, densely sericeous with appressed hairs, these 2.5 mm long, the lobes 10-15 mm long, broadly rounded and short-emarginate, erect or spreading, serice- ous, the hairs spreading; stamens 4, the longer two exserted, the shorter two barely reaching the throat of the corolla; style 5 cm long, glabrous: ovary densely sericeous; fruit not seen. OTHER SPECIMENS EXAMINED: PERU: Cusco: Prov. de Paucartambo, forest, Kosnipata, between Pileapata — Atalaya, 450-550 m, August 5, 1956, C. Vargas C. 11289 (cUz, US); Madre de Dios: Prov. de Manu, "Synduya?", 400 m, October 6, 1966, C. Vargas C. 17831 (CUZ, US). 3. Suessenguthia cuscoensis Wasshausen, sp. nov. PERU: Cusco: Prov. de Paucartambo, forest, Kosnipata, between Pileapata — Atalaya, 450-550 m, August 5, 1956, C. Vargas C. 11288 (Holotype us! Isotype cuz). Frutex, caulibus quadrangularibus pilosis suleatis; folii 1970] Suessenguthia — Wasshausen 123 Fig. 2. Suessenguthia vargasii (a-f): a-—inflorescence X 1⁄2; b—leaf blade X %; c— lowermost bract X 1; d—bract X 1; e — bractlets X 1; f— calyx X 1; g— corolla X %; Suessenguthia cuscoensis (h-l): h— inflorescence X %; i —-bract X 1; j — bract- let X 1; k— calyx X 1; 1— corolla X %. 124 Rhodora [Vol. 72 lamina elliptica acuminata, apice ipso obtuso, ad basim in petiolum decurrens, supra glabra, subtus parce pilosa costa et venis lateralibus obscuris; spicae terminales et axillares, floribus pluribus in fasciculis sessilibus congestis ; bracteae ellipticae, acuminatae, parce sericeae; bracteolae lanceolatae, acuminatae, parce sericeae; calycis segmenta lanceolata, subaequalia, parce sericea; corolla laete rubra sericea, lobis late rotundatis et leviter emarginatis; sta- mina 4, vix exserta; ovarium dense sericeum. Shrub 2-5 m tall, the stem quadrangular, sulcate, pilose, the hairs appressed; leaf blades elliptic, up to 16 cm long and 7 em wide (the lowermost probably larger), acuminate, the tip itself obtuse, attenuate at base and decurrent on the petiole, firm, the upper surface glabrous, minutely papillose, cystoliths numerous, on the lower surface the costa and lateral veins rather inconspicuous, sparingly pilose, the hairs appressed, the margins short repand-crenate; petioles up to 1.5 em long, connate at base by a small ridge or narrow band of leaf tissue; flowers borne in sessile fascicles in axillary and terminal spikes, these lax, internodes 2-3 cm long, sericeous, the hairs appressed; lowermost bracts ovate, 3-9 em long, 1.5-2.5 cm wide, acuminate, thinly sericeous, the hairs appressed; bracts subtending the flower cluster elliptic, 2.8-3.5 cm long, 0.7-1.3 cm wide, acuminate, thinly sericeous, the hairs white, appressed, 1 mm long; bractlets lanceolate, 2.3 cm long, 4 mm wide, acuminate, thinly sericeous, the hairs appressed; calyx segments lanceolate, 1.8 cm long, subequal, 3-4 mm wide, acuminate, thinly sericeous, the hairs white, appressed; corolla drying pale red, 4 cm long, basally 3 mm wide, 7 mm wide at throat, sericeous with appressed hairs 1 mm long, the lobes 7-9 mm long, broadly rounded and short-emarginate, erect, sericeous, the hairs appressed; stamens 4, barely exceeding the throat of the corolla; style glabrous; ovary densely sericeous; fruit not seen. 4. Suessenguthia trochilophila Merxm. in Mitteil. Bot. Staatss. Miinchen [1], Heft 6: 178, pl. 1953; Rhodora 66: 343. 1964. 1970] Suessenguthia — Wasshausen 125 BOLIVIA: at the foot of the Andes near Rurrenabaque, on the right bank of the Rio Beni, September 23, 1951, G. Niethammer 208 (Holotype M). Shrub 1.5-2 m tall, stem quadrangular, suleate, glabrous ; leaf blades broadly lanceolate, up to 25 cm long and 10 cm wide, subacute, basally narrowed and decurrent on the petiole, firm, the upper surface glabrous, minutely papillose, cystoliths numerous, on the lower surface the costa and lateral veins (ca 8-10 pairs) prominent, the margins repand-crenate; petioles up to 1.5 em long, connate at base by a small ridge or narrow band of leaf tissue; flowers borne in sessile fascicles in terminal 2-3-branched spikes, these lax, the internodes 1.5-2.5 cm long, sparingly sericeous, tne hairs spreading; lowermost bracts broadly ovate, 2-2.5 cm long, 1.5-2 em wide, acuminate, glabrous or very spar- ingly sericeous; bracts subtending the floral cluster tri- angular-cuspidate, 2.6-2.8 cm long, 1-1.3 em wide, sparingly pilose, the hairs appressed, 1 mm long; bractlets narrowly lanceolate or subulate, 1.8 cm long, 4 mm wide, long acuminate, thinly sericeous, the hairs appressed; calyx segments lanceolate, subequal, 2.2.9.4 em long and 0.3-0.5 cm wide, acuminate, thinly sericeous, the hairs white, appressed; corolla rose-lilac, 4.5 em long, basally 3 mm wide, 1 cm wide at throat, glabrous, the lobes 12 mm long, apically rounded, spreading not revolute, glabrous; stamens 4, the two longer sub-exserted, the two shorter included in the corolla tube; style 3.7 cm long, long exserted, glabrous; ovary ovate, apically pilose; fruit not seen. OTHER SPECIMENS EXAMINED: BOLIVIA: Beni: Rurrenabaque, 300 m, November 1921, H. H. Rusby 850 (Ny, Us); La Paz: w~ sand near river, in shade, near Coroico, Yungas, August 2, 1894, M. Bang 2367 (GH, NY, US); Guanai, 600 m, May 1886, H. H. Rusby 1119 (vs); woods, San Carlos, Mapiri, 700 m, August 1907, O. Buchtien 1403 (Ny, US) ; San Bartolome (near Calisaya), basin of Río Bopi, Prov. S. Yungas, 750-900 m, July 1929, B. A. Krukoff 10186 (US); Asunta (near Evenay), Prov. S. Yungas, 690-750 m, July 1939, B. A. Krukoff 10585 (vs). SMITHSONIAN INSTITUTION UNITED STATES NATIONAL MUSEUM, WASHINGTON, D.C. 20560 126 Rhodora [Vol. 72 Stuart Kimball Harris outside of Lake of Clouds Hut of Appala- chian Mountain Club on Spring Flower Walks, June 1967. Photo- graph taken by Wesley N. Tiffney. The peruvian type headgear was made by Dr. Harris. STUART KIMBALL HARRIS Again the New England Botanical Club has lost one of its most active members. Stuart Kimball Harris died July 30, 1969, after a sickness of several months. The Club misses a comrade and friend who was always willing to help, and a fellow-worker in its many different activities. He is survived by his wife, Calista Crane Harris, a daugh- ter, Sally, a son, Kimball, and two grandchildren. Dr. Harris was born in Haverhill, Massachusetts, the son of Hayden B. and Vina Harris. After High School he went to Boston University from which he received three degrees, a Bachelor of Business Administration in 1927, Bachelor of Science in 1930, and Master of Arts in 1932. Evidently his life-interest began to assert itself and he earned at Harvard University an M.A. and under Prof. Fernald a Ph.D. in 1936 in the field of Botany. He wrote more than forty botanical papers, earlier on lichens, and later on the flowering plants. Several of these professional papers he edited and they were later published by the Appalachian Mountain Club as the principal text of the book, “Mountain Flowers of New England" in 1964. Through his careful proof-reading and editing, the revision of Prof. Pease’s “Flora of Northern New Hampshire” was published in 1964. Since its publication Dr. Harris had collected extensively in northern New Hampshire to add new localities to those which Prof. Pease had, as well as species not before listed in the area. For many years he had been collecting data to ad to a revision of Robinson's “Flora of Essex County, Massachu- setts" which material was practically ready for publication. Another of his botanical projects was a check-list of the plants of Rockingham County, New Hampshire, giving the species, where and when it was found and by whom. Dr. Harris had been a member of the New England Botanieal Club since 1930 and spent many hours in the study and identification of plants not only from New Eng- land but from all over the United States and even some 127 128 Rhodora [Vol. 72 from Honduras and Mexico. He travelled extensively and always brought back a good lot of plants to be worked over. He was repeatedly in California and other western states. One fruitful trip was to the Big Bend area in Texas. In 1944 he joined a scientific expedition to Alaska by the Alcan Highway where he served as cook. He was a Member of the Plant Distribution Committee of the New England Botanical Club whose reports appear in Rhodora from time to time. He was for thirty three years an Associate Editor of Rhodora, the publication of the New England Botanical Club. He personally contributed plants both to the herbarium of the New England Botanical Club and to the Gray Herbarium, and supervised their iden- tification. He was also Recording Secretary of the Club from 1935 to 1964 and was President at the time of his death. Dr. Harris was for 36 years a member of the Faculty of the College of Liberal Arts of Boston University where he taught courses in Biology but was happiest and at his best in the botany courses, especially with the upper class men and the graduate students where his stimulating and en- thusiastie teaching was greatly appreciated. He was always ready and willing to help and advise students outside the classroom. As already indicated, Dr. Harris was far from what some scientists disparagingly called “a closet botanist.” His in- terest and energy were evident in his years of association with the Appalachian Mountain Club. From the beginning as hutman at Lake of the Clouds, he went on to become the Club Naturalist and for a number of years he led the annual Spring Flower Tours in the Presidential Range. In all his walks and tours he remained enthusiastic and tireless. Another phase of his interest in Natural History until his last illness was the study and identification of birds. He revised the most recent edition of the “Field List of the Birds of Essex County, Massachusetts”. During his active membership in the Essex County Ornithological Club he held various offices including the presidency. For over 1970] Stuart Kimball Harris — Bean 129 thirty years he joined in the annual Club canoe trip on the Ipswich River. He belonged to the Nuttall Club of Cam- bridge, the oldest ornithological club in the United States, and the Massachusetts Audubon Society, For a number of years he made a Christmas bird count in the vicinity of his camp in northern New Hampshire, which was published by the National Audubon Society. This count was made what- ever the weather conditions. Also on most days when he was at home he took an extensive bird walk before he drove to Boston. All this led him to take an active interest in the Ipswich River Sanctuary of the Massachusetts Audubon Society where he served on the Development Committee. He was a member and past chairman of the Conservation Commission in the town of Boxford where he made his home. There he worked to preserve the beauty of the woods, fields, ponds and streams. As if these activities were not enough to make use of his leisure time, he had other interests to which he gave real thought and careful planning, He was an enthusiastic sil- versmith and also did worthwhile work in weaving. Even in the last weeks before he had to give up much activity, he came to the Gray Herbarium and worked on the identifi- cation of his last summer’s collections. He will be sorely missed in many places and by many friends. His work for the New England Botanical Club must command the greatest respect. There are many who enjoy one avocation but Dr. Harris was interested and active in many phases of Natural History beside Botany. RALPH C. BEAN 48 EMERSON STREET WAKEFIELD, MASS. 01880 BAYARD LONG (1885-1969) Bayard Henry Long was born on September 22, 1885, and died on June 9, 1969. His father was John Luther Long, a Philadelphia lawyer who was born and raised in Hanover, York County, Pennsylvania. The elder Long’s sister had gone as a missionary to Japan in the early 1890’s and had married the Rev. J. H. Correll. A few years later the Corrells returned to Phila- delphia where Bayard’s father had established a law prac- tice. Over dinner one evening the Corrells related to Long the story of an American naval officer who had married a geisha girl in Nagasaki. Such marriages were then con- sidered temporary arrangements, which could easily be an- nulled. The bride in this case who had given birth to a child, took a different view of the matter and, when her “husband” returned to Japan with his American wife, killed herself. John Luther Long, who had turned to writing as a side- line, added a few embellishments to this story and it was published in the Century Magazine in 1898. Later David Belasco adapted it as a play which was a success on Broad- way in 1900 and at Belasco’s suggestion Giacomo Puccini used the story as the basis for his opera “Madame kutter- fly" which is recorded as a failure in its premiere in Milan in 1904, but later, after revision, became a great success and is still a standard offering in the repertoire of the worlds great opera companies. The elder Long had established himself as a novelist and playwright and one of his most successful achievements was “The Darling of the Gods" produced by Belasco and starring Blanche Bates. Long died October 31, 1927. Very little is known of the early years and young man- hood of John Luther Long’s son, Bayard. We know that he attended Cheltenham High School, not far removed from his father’s house in Ashbourne (now Elkins Park), a sub- urb of Philadelphia, and that upon his graduation in 1904 he was the Orator of his class. He then attended the Uni- 130 ‘OP6T j5nzny '&osiof MƏN JLT uapdwey “uo paekeg 131 Bayard Long — Fogg 1970] 132 Rhodora [Vol. 72 versity of Pennsylvania, where he majored in Botany and in June, 1908, was awarded the degree of Bachelor of Science in Biology. He later registered in Graduate School of Arts and Sciences at the University, but apparently did not receive a graduate degree. Just when Bayard Long became affiliated with the Phila- delphia Academy of Natural Sciences the Academy’s rec- ords do not reveal, but it was probably while he was still taking courses in Botany at the University. We know that he became a member of the Philadelphia Botanical Club in 1906 and that in 1913 he was elected Curator of the Club’s Local Herbarium, a position to which he was annually re-elected until the year of his death. Thus, his custodian- ship of one of the country’s finest local herbaria spanned an interval of 56 years. At no time during his career did Bayard accept any salary from the Academy. The Philadelphia Botanical Club had been founded in December, 1891, by a group of amateur botanists who were enthusiastically devoted to studying the flora of the Phila- delphia local area. By their definition this area comprised all of southern New Jersey and southeastern Pennsylvania, as well as Harford County, Maryland and New Castle County, Delaware. For many years members of the Club explored and col- lected in the numerous plant habitats included in this area, e.g; the Pine Barrens of New Jersey, the glaciated portions of Northampton County, the serpentine outcrops of Chester and Lancaster Counties, etc. Long was a frequent member of these expeditions and early demonstrated his keenness as an observer and his superb ability as a collector. The early issues of Bartonia, the official publication of the Club, were filled with accounts of the exciting finds made on these trips and the cases in the Local Herbarium (housed at the Academy) were soon bulging with specimens from all corners of the local area. It is doubtful that anyone ever possessed a higher stand- ard for the quality of an herbarium specimen than Bayard Long. Every leaf had to be laid out flat, every inflorescence 1970] Bayard Long — Fogg 133 properly displayed, every flower part clearly shown. Extra flowers and loose fruits and seeds were placed in pockets affixed to the sheet. Root systems (collected in their en- tirety whenever possible) were scrupulously clean, habitats were accurately described and localities were identified to the nearest tenth of a mile and closest compass point. All of this seems the more remarkable when it is realized that Long collected close to 80,000 numbers, not including col- lections made as a member of Fernald’s expeditions. It was inevitable that a man of Long’s keen intelligence and critical judgement should come to the attention of Professor M. L. Fernald, In 1911 Fernald wrote, “since Mr. Long has on other occasions shown keen discrimination in his study of the Cyperaceae, it is a pleasure permanently to associate his name with the new Scirpus Longii n.sp." (Rhodora 13. 6. 1911) During the years that followed Fernald named nine other species in Long's honor. Between July 2 and September 9, 1920, Long joined the Gray expedition under Fernald to Nova Scotia (see Rhodora 23: 94. 1921) and in 1924 and 1925 he was a member of the party which explored Newfoundland. It was in his "Journal of the Summer of 1925" that Fernald stated, “If there is a keener collector or discoverer of native plants than Bayard Long, I have yet to meet him”. (Rhodora 28: 94, 1926.) I first met Bayard Long in the autumn of 1921. At that time I had had no formal training in botany, but was be- ginning to be interested in plants and was attempting to identify the species that I encountered in the suburbs of Philadelphia. I had purchased a copy of the 7th edition of Gray’s Manual and found it rather hard going. In despera- tion I took some of my puzzles in to the Academy of Natural Sciences and inquired of a guard where I would find a botanist. I was directed to the Herbarium of the Philadel- phia Botanical Club. There I met Mr. Long who quickly solved my problems, encouraged me to continue collecting and advised me to join the Philadelphia Botanical Club, which I promptly did. Thus began an association which 134 Rhodora [Vol. 72 lasted for more than 40 years —a friendship cemented by countless field trips in the local and more distant areas. As an undergraduate student in the early 20's I had become fascinated by the writings of M. L. Fernald and had read every word he had published in the pages of Rhodora. In particular was I excited by the accounts of his expeditions to New Brunswick, the Gaspé Peninsula, Nova Scotia and Newfoundland. I begged Long, who had already been on several of these trips, to ascertain whether there would ever be an opportunity for me to become a member. It was entirely through his intercession that Professor Fer- nald invited me to join Long and him on a brief excursion to Newfoundland in 1926. This was the summer of the Fourth International Con- gress at Ithaca, so we were unable to leave for Newfound- land until late in August. Three years later, in the summer of 1929, I was again privileged to join Fernald (with whom I had now taken my Doctor's degree) and Long on a much longer tour. The events of these two trips were summarized by Fer- nald in an article entitled, Recent Discoveries in the New- foundland Flora", which occupied all twelve numbers of Rhodora for 1933 (Vol. 35). In the 1930's I accompanied Fernald and Long on two collecting trips to the Coastal Plain of southeastern Vir- ginia, The results of these trips were also published by Fernald in Rhodora. Bayard Long's intellectual qualifications were such that he could doubtless have succeeded in any field of endeavor which appealed to him. He was an ardent philatelist, his specialty being British Colonies, a subject area which calls for keen powers of observation. His tastes in literature and music were exalted and he had a deep appreciation of natural beauty. With it all Long was an extremely modest and almost painfully self-effacing individual. He appeared to shrink from human contacts, but once convinced of a person's sincere desire to learn, no effort was too great, no demands 1970] Bayard Long — Fogg 135 on his time too exacting for him to render assistance and share his prodigious knowledge, Certainly no one has ever possessed his intimate acquaintance with the local flora, but he was equally at home in the field in Newfoundland and Virginia. Long's first serious illness occurred in the early 1960's, although he continued to work at the Academy until the fall of 1962. Afterwards he was confined to his home in Elkins Park, where almost until the very end he continued by letter and telephone to answer the questions of those who called upon him for help. PUBLICATIONS OF BAYARD LONG 1. Range extension of Scirpus Smithii var. setosus. Rhodora 12: 155-156. 1910. 2. Pinus serotina Michx. in southern New Jersey and other local notes. Bartonia. 2:17-21. 1910. 3. Certain species becoming well established at Ashbourne and else- where near Philadelphia. Bartonia. 3:22-25. 1911. 4. Galium labradoricum in Pennsylvania. Rhodora. 14:199, 1-200. 1912. Some results of recent field work in the Cape May peninsula. Bartonia. 4:14-19. 1912. 6. Range extension in Antennaria. Rhodora. 15:117-122. 1913. T. Ludwigiantha brevipes Long n.sp. Britton & Brown Illus. Flora Ed. 2. II:586. 1913. 8. (With Fernald) The American variations of Potentilla palustris. Rhodora. 16:5-11, pl. 106. 1914. 9. On the occurrence of Keeled Garlic in America. Bartonia. 7:6-16. 1915; : 10. Discovery of Prunus cuneata in southern New Jersey. Rhodora. 18:66-70. 1916. 11. A belated correction. Rhodora. 18:142-143. 1916. 12. Delphinium consolida in America, with a consideration of the status of Delphinium Ajacis. Rhodora. 18:169-177. 1916. 13. Range of Carex novae-angliae extended into Pennsylvania. Rhodora. 19:96-100. 1917. 14. History of the American record of Scirpus muconatus. ,Rhodora. 20:41-48. 1918. 15. Eragrostis peregrina a frequent plant about Philadephia. Rhodora. 20:173-180. 1918. 16. Jasione montana a conspicuous weed near Lakewood, New Jersey. Rhodora. 21:105-108. 1919. j ct 136 Rhodora [Vol. 72 17. The specific characters of Eragrostis peregrina and its two allies. Rhodora. 21:133-141. 1919. 18. Notes on the American occurrence of Crepis biennis. Rhodora. 21:209-214. 1919. 19. Regarding Gentiana Andrewsii in the coastal plain of New Jersey. 22:104-110. 1920. 20. A further note on Crepis biennis. Rhodora. 22:192, 193. 1920. 21. A station for Croton glandulosus in New Jersey. Rhodora. 23: 221-223. 1921. 22, Muscari comosum a new introduction found in Philadelphia. Rhodora. 24:16-20. 1922. 23. Naturalized occurrence of Prunus Padus in America. Rhodora. 25:169-177. 1923. 24. Some changes in the aspect of the list of the Philadelphia flora. Bartonia. 8:12-32. 1924. 25. Some noteworthy indigenous species new to the Philadelphia area. Bartonia. 10:30-52. 1929. PLANTS NAMED FOR BAYARD LONG Scirpus Longii Fernald, Rhodora 13:6-8. 1911. Cardamine Longii Fernald, Rhodora 19:91. 1917. Carex Longii Mackenzie, Bull. Torrey Bot. Club 29:373. 1923. Braya Longii Fernald, Rhodora 28:202. 1926. Antennaria Longii Fernald, Rhodora 28:237. 1926 (1927). Bryum Longii E. B. Bartram, Rhodora 30:6, 7. 1928. Antennaria Bayardi Fernald, Rhodora 38:402. 1936. Lycopus americanus Muhl., var. Longii Benner, Bartonia 16:46. 1934 (1935). Malaxis Bayardi Fernald, Rhodora 38:402. 1936. Juncus Longi! Fernald, Rhodora 39:397. 1937. Rubus Longii Fernald, Rhodora 40:434. 1938. Carex Bayardi Fernald, Rhodora 44:71. 1942. Xyris Bayardi Fernald, Rhodora 48:56, 1946. In the preparation of this account I have been greatly as- sisted by Dr. E. T. Wherry and Mrs, Nellie Erisman. JOHN M. FOGG, JR. ARBORETUM OF THE BARNES FOUNDATION MERION, PA. 19066 BOOK REVIEW IMPORTANT WORK ON THE FLORA OF NEW ENGLAND! A. R. HopGpon The experience of having used Seymour’s “Flora of New England” rather constantly for over a year has provided this reviewer with an unusual awareness of both the strengths and weaknesses of the book. At present, when dealing with New England plants, I am inclined to rely nearly as much on Seymour’s “Flora” as on Fernald’s 8th edition of Gray’s Manual. This is not to say that it replaces the “Manual”, but it does supplement it nicely in certain particulars and in nomenclature tends to bring the older book up to date for New England. “The Flora of New England”, as indicated in the subtitle, is more than a flora in that it has keys to genera and spe- cies and brief diagnoses of families and genera, thus making it available as a laboratory text for college classes dealing with New England plants. In this respect, then, it is a "Manual". Blake and Atwood (Geographical Guide to Floras of the World, Part I, U. S. Dep. Agr., Misc. Publ. 401, 1942, pp 8- 9) have made some suggestions concerning the kinds of in- formation to be included in a “flora”, many but not all of which have been satisfactorily followed by Seymour. In addition it would have been desirable, for an area so long worked over as New England, to include a listing, perhaps only a selected one, of the local floras of the region, par- ticularly since many of these provide additional and much more detailed information about the occurrences of plants locally. For example, it seems neglectful not to have listed all Vascular Plants including Ferns and Fern Allies and Flowering Plants growing without Cultivation in New England, by Frank Conkling Seymour. Charles E. Tuttle Co. Rutland, Vermont, 1969. xvi + 596 pp. illus. $12.50. 138 Rhodora [Vol. 72 Unfortunately, the “profit motive" has to be a prime con- sideration in the preparation of a “flora”. In order to “sell” the book has to be kept within certain limits of size, and has to be useful to a variety of persons such as college students in botany courses. Recent state floras and now this regional flora reflect a trend to incorporate in them keys, brief de- scriptions and numerous illustrations as well. It certainly is often in order for a flora to provide in one package a body of information relating to the places where taxa are to be found along with the means to identify and recognize these taxa. But to do both of these tasks well or even passably well for an area the size of New England is a very large order. It is obvious that the author had to make many compromises to produce a book that could at once be both “flora” and “manual”. To me, it still seems a matter of wonder that the result of these concessions by Seymour is, nonetheless, a book of considerable merit. However, at this point I would urge that the authors of floras now in the making should adhere more strictly to the ideal concept of a flora, and that sponsors strive a bit harder to provide financial support. If the current trend is continued we may expect for the “Gray’s Manual Range" that there will be an almost endless series of local, state and sectional “man- uals” with greatly overlapping and repetitive information to the detriment of the peculiar information about ranges, locations, collectors and records that we have good reason to expect from a carefully documented “flora”. There is little point in describing the book's contents. By now it has been widely distributed and must indeed be famil- iar to many of the readers of Rhodora and should, of course, be in the hands of all those seriously interested in the flora of New England. Moreover, although I can't agree with all of Mr. Seymour's choices of scientific names or all his taxonomic decisions, these are often matters of opinion for which no competent author should be severely criticized. It is, then, at certain of the floristic aspects of “The Flora of New England" that I will direct my attention, a few 1970] Book Review — Hodgdon 139 points seeming to invite discussion and possible disagree- ment. On page XIII in the Introduction we encounter the following statement: “Im determining what species consti- tute the flora of New England, all which give evidence of having grown in this region without cultivation within twenty-five years are included." My own feeling about this is quite at variance with that of Mr. Seymour; if there is good evidence that a species grew naturally in historic time in any part of New England it is still part of the flora, and if it is believed not to grow there now, it still should be listed and its present status indicated. For phytogeograph- ical considerations certainly a flora would include such spe- cies. I don’t know whether the occasional discrepancies between the ranges of certain species, in the listing of States for example, as given by Fernald in Gray’s Manual and by Seymour, are attributable to the latter’s interpretation of what constitutes a flora, but in working up ranges of New England species for phytogeographical projects I always resort to Gray’s Manual as well as Seymour. Mr. Seymour is certainly to be commended for the care that he has taken in personally examining specimens for his records, and for his avoidance of unsubstantiated reports. In many instances he has visited localities of botanical in- terest and thereby verified interesting records. One’s con- fidence is heightened by the knowledge that he has made a number of interesting discoveries in the New England flora. A great many of the published records of New England species appearing in Rhodora through the years and in other journals to a lesser extent, as well as in local floras and in special treatments of ferns, orchids, woody plants, etc., certainly have much to contribute to the “flora” of a region, even though in some instances the specimens cited or men- tioned in these works may be lost or buried in some small herbarium. It is my recollection that Professor Fernald paid attention to all serious reports that came to his attention, though it must be admitted that he accepted these only when he had confidence in the author’s critical capacity. In any n 4 140 Rhodora [Vol. 72 revision of “The New England Flora”, I would hope that the author will thoroughly examine the voluminous body of literature dealing with the vascular flora of New Eng- land, sifting out of it pertinent data to make the record complete. I know that Frank Seymour visited the herbarium of the University of New Hampshire sometime during the years while he was preparing his book, but our herbarium is grow- ing fast and, had he checked it a year or two before the manuscript was published, he would have had a very large number of additional townships in New Hampshire and some new State records. In the matter of citing localities where specimens have been seen, I have a few comments to make. Somewhat arbitrarily Seymour decided to list the township whenever a species has been collected only once in a particular county, but otherwise to list the county when two or more town- ships in that particular county are represented. However, there is no map showing the townships in each state, nor is there any listing of towns in their respective counties. Thus one needs a good atlas to plot the ranges of species. In order to provide information about ranges of all taxa effectively (which I would think might be a major considera- tion in a flora), it would be necessary either to supply range maps at least for critical taxa, or to provide many more township records and a ready means of locating townships. Thus the book would require considerable amplification of content to provide anything approaching exact data on dis- tribution for a great many taxa. Moreover, many counties such as Coós County in New Hampshire and particularly several of the counties in Maine, are so large that any in- dication of presence by county is not enough. The importance of a scientific work is determined by the role that it plays in the progressive development of the sci- ence, For my own work on the study of the plants of North- eastern America, this book will stand as a sound foundation. Its weaknesses will serve to show us a better way, and its 0 1970] Spiranthes tuberosa — Magrath 141 many strengths will provide the base on which those of us who are working on facets of the flora of New England can build. DEPARTMENT OF BOTANY UNIVERSITY OF NEW HAMPSHIRE DURHAM 03824 SPIRANTHES TUBEROSA, NEW FOR KANSAS — During a study of the family Orchidaceae in Kansas, it was discovered that several specimens of Spiranthes tuberosa Raf. had been collected in southeastern Kansas (Bourbon, Cherokee and Woodson Counties) and mis- takenly identified as the more common S. gracilis (Bigel.) Beck. This orchid has previously been reported from Massachusetts, through the Atlantic States, south to south- central Florida, west to eastern Texas, Arkansas, Kentucky, Missouri and also Michigan. These specimens represent an extension northwestward of the range of the species. REPRESENTATIVE SPECIMENS EXAMINED: Bourbon Co.: open wooded hillside, rocky clay soil, not common, 2 mi s of Uniontown, 2 October 1955, MeGregor 11110 (KANU). Cherokee Co.: sandy rocky soil under Quercus alba and Q. stellata, not common, 5% mi e of Baxter Springs, 3 October 1955, MeGregor 11055 (KANU). Woodson Co.: open oak wooded bank, sandy soil, rare, Lake Fegan, Woodson Co. State Park, 25 September 1955, Lathrop 1884 (KANU). LAWRENCE K. MAGRATH DEPARTMENT OF BOTANY UNIVERSITY OF KANSAS, LAWRENCE 66044 WHITE PHASE IN FLOWER DEVELOPMENT IN CYPRIPEDIUM ACAULE — Flowering plants of Cypri- pedium acaule were unusually abundant in the Williams- burg, Virginia, area in April and May of 1969: populations of hundreds of specimens were frequent, On April 28th a friend showed me a young plant with a white lip. I had not noted this condition before, but subse- quent search over the next few days revealed the lins of all young flowers in the area to be white and to become pink with maturity (Fig. 1). Low temperatures for the period involved ranged from the mid-30's to the mid-40’s. Could these unseasonably low temperatures account for the white phase in the develop- ment of these flowers? We shall attempt to get an answer another year by bringing young plants into temperature- controlled chambers. Is there a likelihood that f. a/biflorum of this species is temperature induced? It is at least provocative that in certain areas near the White Mountains the white form replaces the pink (Anderson, 1936) and that f. albiflorum has been reported in the mountains as far south as North Carolina (Correll, 1950). We found pollen from ten plants in the white phase and from ten in the pink to be normal. LITERATURE CITED ANDERSON, EDGAR. 1936. Color variation in eastern North Ameri- can flowers as exemplified by Hepatica acutiloba. Rhodora 38: 301-804. CORRELL, DONOVAN S. 1950. Native Orchids of North America. Chronica Botanica Co., Waltham, Mass. J. T. BALDWIN, JR. COLLEGE OF WILLIAM AND MARY WILLIAMSBURG, VIRGINIA 23185 1970] Cypripedium acaule — Baldwin 14: Fig. 1. White and pink phases of Cypripedium acaule, April 29, 1969, Williamsburg, Virginia. Photo by Col. D. W. Noake, printed two thirds natural size and reduced in publication. Volume 71, No. 788, including pages 495-606, was issued December 31, 1969. Volume 72, No. 789, including pages 1-143, was issued March 31, 1970. FüRLOW REFERENCE LIBRARY JUL 21 1970 Hovova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL ^ RALPH CARLETON BEAN ROBERT CRICHTON FOSTER ROLLA MILTON TRYON Associate Editors RADCLIFFE BARNES PIKE LORIN IVES NEVLING, JR. ELIZABETH ANNE SHAW J Vol. 72 April-June, 1970 No. 790 CONTENTS: The Taxonomy of Vernonia acaulis, V. glauca and V. nove- boracensis (Compositae). Samuel B. Jomes oons 145 Distributional History of Epilobium hirsutum (Great Hairy Willow-Herb) in North America. Do zii T: Stube ks esna E IC A 164 A Re-evaluation of Polygonum meisnerianum in North America. Richard S. Mitchell... 182 Eragrostis plana in South Carolina F. W. Gould and R. I. Lonard .......... eerte 188 An Undescribed Species of Astragalus (Leguminosae) from Utah. Stanley L. Welsh eee 189 (Continued on Inside Cover) The New England Botanical Club, Ine. Botanical Museum, Oxford St., Cambridge, Mass. 02138 CONTENTS: — continued Investigations of New England Marine Algae III: Com- position, Seasonal Occurrence and Reproductive Peri- odicity of the Marine Rhodophyceae in New Hampshire. Edward J. Hehre and Arthur C. Mathieson .................. 194 Hybrids in Rubus Subgenus Eubatus in New England. Frederic L. Steele and A. R. Hodg dom ..............3.. 240 Chrysanthemum lacustre Brotero New to Gray's Manual Range. Charles E. Stevens .......22. eee 250 Soil Preferences and Variation in Flavonoid Pigments in Species of Asters. Warren G. Abrahamson and Otto T. Solbrig ............. 251 Biosytematics of Setcreasea brevifolia. S. A. Faruqi and K. L. Mehra .... eese 264 New and Overlooked Species of the Virginia Flora. A. M. Harvill, Jv. ................... eene nennen nnne nete 272 Listera ovata (L.) R. Br. in the Bruce Peninsula, Ontario. H. Vincent Elliott and Frank S. Cook ................................ 274 Zonation of Bryophytes in the Headwaters of a New Hamp- Shire Stream. Janice M. Glime ........... eene 276 Vernon Black Gum Swamp. F. R. Fosberg and Terry Blunt ............5. eee 280 Recent Plant Records for Nebraska. Robert J. Lemaire ceccccccccccccccsssscsssscsssecsssssvcsscsseecccssseeseneetseesenens 283 An Assemblage of Halophytes in Northern Ohio. Allison W. Cusick euenit 285 A Range Extension for Odontites serotina (Lam.) Dum. (Scrophulariaceae). Neil A. Harriman esere 286 A New Argentine Sisyrinchium. Robert C. Foster eese eene eene ette 287 Rhodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 72 April-June, 1970 No. 790 THE TAXONOMY OF VERNONIA ACAULIS, V. GLAUCA AND V. NOVEBORACENSIS (COMPOSITAE) SAMUEL B. JONES Delimitation and identification of all taxonomic units are extremely difficult in the genus Vernonia since there are numerous, intermediate, perplexing forms (Ekman, 1914). One postulation is that the genus underwent rapid evolution in North America following the Cretaceous Period and that it is probably still evolving (Jones, 1964). This has resulted in the formation of population systems that appear some- what morphologically disinct yet taxonomically difficult. Because the reproductive barriers among many of the spe- cies are incomplete (Jones, 1966, 1967), and because hy- bridization occurs commonly where species show biotic or adjacent sympatry, the genus is of interest from both evo- lutionary and taxonomic viewpoints. The three species Vernonia acaulis (Walt.) Gleason, V. glauca (L.) Willd. and V. noveboracensis (L.) Michx. are the most eastern in the United States. They occur from Massachusetts to eastern Kentucky and Tennessee, and into Florida and Alabama along the Coastal Plain, Pied- mont and mountains. The number of synonyms for the latter two species is an indication of the uncertainty which has characterized their treatment during the past 200 years. It is also obvious that the ranges of the three species have been poorly understood. The reason for this is that their 145 146 Rhodora [Vol. 72 ranges were often copied from earlier floras which were erroneous. These taxa were chosen for systematic study because of the conspicuous variability within the taxa, es- pecially in V. noveboracensis, The usual field and herbarium studies were supplemented by biosystematic investigations. In the course of this investigation, assistance has been re- ceived from many persons. It is not possible to thank them all specifically, but my sincere thanks are none the less real. This work was supported by National Science Foundation grant GB 7208 and the University of Georgia. Specimens for examination were obtained from the following herbaria: ALU; AUA; FSU; FLAS; GA; GH; NCU; NY; OS; PH; SMU; US; TENN; VDB; WIS; WVA. Annotated specimens of the three taxa may be found in all of these herbaria. HISTORICAL ACCOUNT Walter (1788) applied the name Chrysocoma acaulis, and he was clearly referring to a plant with a basal cluster of leaves. Michaux (1803) described the same taxon and gave it the name Vernonia oligophylla. In 1891 Kuntze applied the generic name of Cacalia. In 1906 Gleason recognized the name of Walter and made the combination Vernonia acaulis (Walt.) Gleason. Vernonia georgiana Bartlett has been shown to be a hybrid between V. acaulis and V. an- gustifolia Michx. (Jones, 1967). It is obvious from an examination of the microfiche of the Linnaean Herbarium that Linnaeus’ (1753) Serratula nove- boracensis refers to Vernonia noveboracensis. Apparently his Serratula praealta differed from the type only in the shorter appendages of the involucral bracts (Gleason, 1906). Walter’s (1788) Chrysocoma tomentosa shows no essential difference from V. noveboracensis. Michaux (1803) trans- ferred the species to the genus Vernonia making the combi- nation V. noveboracensis (L.) Michx. He also recognized V. praealta, Hill (1768) applied the generic name Behen, and Miller in 1768 used the name Serratula caroliniana Mill. Walter's (1788) Chrysocoma tomentosa was recognized by Elliott (1824) ; however, Elliott was not certain that his 1970] Vernonia — Jones 147 plant was the same as that described by Walter. More re- cently, Steele (1901) discussed the matter, and regarded the name V. tomentosa Ell. to be a plant of wet places from the south Atlantic Coast. Gleason (1906) studied Steele’s specimens and felt that they did not warrant its separation as a distinct species. According to Gray (1852), V. ruge- liana Shuttlew. is the ordinary form of V. noveboracensis. The next major addition to the synonomy was that of V. harperi Gleason (1906). It is based on Harper 1424 from Coffee County, Georgia. Gleason distinguished it from V. noveboracensis because of its larger heads and more nu- merous flowers. Serratula glauca L. was based on a plant collected by Clayton, studied by Linnaeus and fragments were given to Asa Gray in 1839 (Fernald, 1941). Willdenow (1804) made the combination Vernonia glauca (L.) Willd. Gray (1884) included V. glauca as part of V. noveboracensis var. lati- folia, Britton’s (1898) description of V. noveboracensis tomentosa (Walt.) Britton refers to V. glauca. In 1941 Fernald described V. glauca (L.) Willd. forma longiaristata Fernald which has longer bract tips. SYSTEMATIC TREATMENT Key to Species 1. Principal leaves basal, the stem leaves usually much reduced ........ Eum nl d a ann S d d ER EI 1. V. acaulis l. Principal leaves cauline, the stem leaves usually not reduced ex- Cept near IN MOTERCOTICE «arriere erae ren hoops ce er n s ss aa Ud Danos 2 2. Pappus straw-colored to white, lower surface of leaves paler than upper; leaves usually rather abruptly narrowed at base; plants of well drained soil; normally flowering in July ........... pde mM dE Ee ctc 2. V. glauca 2. Pappus purple or brownish-purple, lower surface of leaves not much paler than upper; leaves gradually narrowed at base; plants of moist soil; normally flowering in August and Sep- tempek nin As stet P 3. V. noveboracensis 1. Verononia acaulis (Walt.) Gleason, Bull. N.Y. Bot, Gard. 4: 222. 1906. Chrysocoma acaulis Walt., Fl. Car. 196. 1788. Type: Not examined. Photograph of type in PH. 148 Rhodora [Vol. 72 Vernonia oligophylla Michx., Fl. Bor.-Am. 2: 94. 1803. Cacalia acaulis Kuntze, Rev. Gen. 968. 1891. Stems erect, glabrous to thinly puberulent, simple to the inflorescence, 3-10 dm high. Leaves mostly basal, basal leaves 20 (12-30) cm long, 7 (4-10) em wide, oblong to obovate, sparsely pubescent above, sparsely pubescent to nearly glabrous below, tip acute, base attenuate, margins coarsely and irrgularly serrate, cauline leaves bractlike, In- florescence loose and open branched. Heads 42 (31-59) flowered. Involucre broadly campanulate, 6.6 (5.0-8.5) mm high, 7.0 (5.7-8.5) mm wide. Bracts lanceolate to linear elliptie. Bract tips acuminate to long acuminate. Bracts greenish-purple, thinly puberulent on back. Achenes sparse- ly pubescent, strongly ribbed, resinous, 3 (2.7-3.2) mm long. Pappus straw colored to white, bristles 5.5-9.0 mm long, scales narrow, 0.5-1.0 mm long. Flowering in July. Chromo- some number n — 17. Habitat: uplands, sandy woods, flat- Figure 1. Distribution map of Vernonia acaulis based on county distributions. 1970] Vernonia — Jones 149 woods and sandhill ecotones. Distribution: in part of Coastal Plain and Piedmont of North Carolina, South Caro- lina, Georgia (see Fig. 1). Vernonia acaulis is quite distinct and is easily identified. Occasionally, it does hybridize with V. noveboracensis and V. angustifolia Michx. The F, hybrids are easy to recognize as they are intermediate between their parents. This hy- bridization does not seem to have blurred the spacies lines. Local population samples were made in Elbert Co., Ga. (15736) and Montgomery Co., N. C. (15783). Transplants were grown in the garden and the greenhouse from these two locations. Table 1 presents the sample means, standard deviations, coefficients of variation and ranges of certain morphological characters from the North Carolina popula- tion. There is probably no more variation between local populations than one would normally expect in outbreeding population systems. Experimental hybrids were made among V. acaulis, V. noveboracensis, V. glauca and V. angustifolia. The F, hybrids were intermediate. They had good pollen stainability with aniline blue in lactophenol and good pairing of the chromosomes at meiosis. TABLE 1 Vernonia acaulis: population sample means (X), standard deviations (s), coefficients of variation (C) and ranges of seven morphological characters; n — 25; population sample 15783, Montgomery Co., N. C. Character X 8 C Range lower leaf width cm 6.5 1.4 0.2 3.7 - 10.0 lower leaf length cm 20.2 4.0 0.2 12 -30 middle leaf width cm 0.7 0.4 0.6 0.2- 2.0 middle leaf length cm 8.7 3. 0.4 1.7 - 15.6 involucre width mm 7.0 0.7 0.1 5.7- 8.5 involucre length mm 6.6 0.9 0.1 5.0- 8.5 flowers per head 41.8 7.0 02T 59 150 Rhodora [Vol. 72 Normally, Vernonia acaulis is ecologically isolated from V. angustifolia. The latter usually grows in drier places than the former. It is also ecologically as well as seasonally isolated from V. noveboracensis. Vernonia acaulis flowers in July, while V. noveboracensis flowers in August. The habitat in which V. noveboracensis grows is often quite wet, while V. acaulis grows in well-drained soil. Isolation, however, is not always completely effective as the flowering periods may overlap or the ecological niches may be within a few meters of each other. 2. Vernonia glauca (L.) Willd., Sp. Pl. 3: 1633. 1804, Serratula glauca L., Sp. Pl. 818. 1753. Type: According to Fernald (1941), it is based on a specimen collected by Clayton, studied by Linnaeus and fragments given to Gray in 1839. Not examined. Suprago glauca Gaertn., Fruct. 2: 402. 1791. Vernonia noveboracensis latifolia A. Gray, Syn. Fl. N. Am. 1: 89, in part. 1884. Vernonia noveboracensis tomentosa Britton, Britt, & Brown. Ill. Fl. 3: 302, in part. 1884. Vernonia glauca (L.) Willd. forma longiaristata Fern., Rhodora 43: 652. 1941. Stems erect, glabrous, simple, 10-16 dm high. Leaves numerous, 15 (12.5-19) cm long, 5 (3.4-7.4) em wide, ovate- lanceolate to almost ovate, glabrous or scabrellate above, pale and thinly puberulent below, tips short-acuminate, abruptly narrowed at the base, margins irregularly dentate to nearly entire. Inflorescence compact. Heads 40 (32-48) flowered. Involucre broadly campanulate 7.3 (5.0-8.8) mm high, 7.4 (6.1-8.5) mm wide. Bracts, inner lance-ovate, outer lanceolate, bract tips acute or acuminate to long acuminate. Bracts greenish-purple, glabrous or puberulent. Achenes nearly glabrous, ribbed, 3 (2.7-3.3) mm long. Pappus straw-colored to white, bristles 6.7 mm long, scales narrow 0.4-0.8 mm long. Flowering in July. Chromosome number, n = 17. Habitat: edge of well-drained upland oak- hickory woods. Distribution: mountains and piedmont from Pennsylvania to Alabama (see Fig. 2). 1970] Vernonia — Jones 151 Figure 2. Distribution map of Vernonia glauca based on county distributions. Local population samples were collected in Clarke Co., Ga. (15146) and in Orange Co., N. C. (15788). Transplants were grown in the greenhouse and garden from the above locations and also from Gwinnett Co., Ga. (15349). Ex- perimental hybrids were produced with a number of other species of Vernonia, including V. noveboracensis and V. acaulis. The F, hybrids had good chromosome pairing at meiosis, and the pollen stained well with aniline blue in lactophenol, Although misinterpretation of Vernonia glauca versus V. 152 Rhodora [Vol. 72 noveboracensis is common, V. glauca is morphologically and ecologically distinct from the latter. There is some variation within and between the populations ; however, this variation is probably no more than is normally expected in heterozy- gous individuals and populations. This normal variation includes Fernald's (1941) forma longiaristata, and it is certainly not worthy of a name. Table 2 presents some morphological measurements from one sample. TABLE 2 Vernonia glauca: population sample means (x), standard deviations (s), coefficients of variation (C) and ranges of six morphological characters; n — 25; population sample 15146 Clarke Co., Ga. Character x 8 C Range middle leaf width cm 4.9 1.0 0.2 3.4- 7.4 middle leaf length cm 15.7 1.7 0.1 12.5- 19.0 involucre width mm 7.4 0.6 0.1 6.1- 8.5 involucre length mm 7.3 0.9 0.1 5.0- 8.8 flowers per head 40.5 4.4 0.1 32 -48 ratio middle leaf length/width 3.3 0.6 0.2 2.4- 5.0 Natural hybrids between Vernonia glauca and V. flaccidi- folia Small were collected in Gwinnett Co., Ga. (15349). The parents and the hybrids were at the edge of oak-hickory woods and in flower at the same time. During the course of this study no specimens of natural hybrids between V. glauca and V. noveboracensis were unequivocally desig- nated. They should, however, be looked for. These two species are normally seasonally and ecologically isolated. Vernonia glauca flowers in July, while V. noveboracensis blooms in August. In addition, the former grows in well- drained upland woods and is rarely found in the lowlands, while the latter is found in wet meadows, low roadsides, and stream banks and found often in standing water. Ex- perimental F, hybrids are intermediate between their par- ents in the key characters of pappus color and leaf shape. 1970] Vernonia — Jones 153 Figure 3. Distribution map of Vernonia noveboracensis based on county distributions. 3. Vernonia noveboracensis (L.) Michx. Fl. Bor.-Am. 2: 95. 1808. Serratula noveboracensis L., Sp. Pl. 818. 1753. Type: Not examined, photograph of type in microfiche of Linnaean Herbarium. Photograph of type in GH. Serratula praealta L., Sp. Pl. 818. 1753. 154 Rhodora [Vol. 72 Behen noveboracensis Hill, Hort. Kew. 68. 1768. Behen praealtum Hill, Hort. Kew. 68. 1768. Serratula caroliniana Mill., Gard. Dict. ed. 8. Serratula no. 7, 1768. Chrysocoma tomentosa Walt., Fl. Car. 196. 1788. Vernonia praealta Michx., Fl. Bor.-Am. 2: 95. 1803. Vernonia tomentosa Ell., Bot. S. C. & Ga. 2: 288. 1821. Vernonia noveboracensis (L.) Michx. var. praealta Wood, Classbook 183. 1845. Vernonia rugeliana Shuttlew.; A. Gray, Pl. Wright. 1: 82. 1852, Vernonia noveboracensis (L.) Michx. var. latifolia A. Gray, Sym. Fl. N. Am. 2: 89. 1884. Cacalia noveboracensis Kuntze, Rev. Gen. 324. 1891. Vernonia noveboracensis tomentosa Britton, Porter & Britton, Mem. Torrey Bot. Club 5: 311. 1894. Vernonia harperi Gleason, Bull. N. Y. Bot. Gard. 4: 221. 1906. Stems erect, glabrous to thinly pubescent, simple to the inflorescence 10-25 dm high. Leaves numerous, 18 (12-28) em long, 3.4 (1.3-5.5) cm wide, lanceolate, glabrous or scabrellate above, thinly tomentose below, tip acuminate, base attenuate, margins nearly entire. Inflorescence loose and spreading. Heads 44 (30-65) flowered. Involucre cam- panulate, 12 (7-17) mm high, 7.7 (6-10) mm wide. Bracts triangular-ovate. Bract tips acuminate to long acuminate with a filiform appendage. Bracts greenish-purple, pubes- cent and sometimes ciliate. Achenes sparsely pubescent, ribbed, 4.1 (4-4.5) mm long. Pappus brownish-purple, bristles 6.5 (6-7) mm long, scales linear, 0.4-0.8 mm long. Corollas whitish and pappus pale in forma albiflora Brit- ton, 1890, Bull. Torrey Bot. Club 17: 124. Corollas lilac in forma lilacina Oswald, 1957, Phytologia 5: 465. Flowering in August and September, Chromosome number, » — 17. Habitat: low, wet roadsides, wet meadows, creek banks. Distribution: piedmont, coastal plain and mountains from Massachusetts to eastern Kentucky and eastern Tennessee into Alabama and Florida (see Fig. 3). 1970] Vernonia — Jones 155 Of the three taxa, Vernonia noveboracensis is by far the most variable, and it has the greatest geographical range. In Gleason’s (1906) revision of Vernonia, the species group Noveboracenses included two species V. noveboracensis (L.) Willd. and V. harperi Gleason. The former has a wide dis- tribution from Alabama to Massachusetts on the Atlantic Coastal Plain, Piedmont Plateau, and the mountains. Ac- cording to Gleason (1923), it gave rise to V. harperi of the Atlantic Coastal Plain in south Georgia; this species has larger heads with more flowers. As Ekman (1914) noted, however, Gleason’s species are very small, narrowly defined and, as a rule, based upon very meager material. The purpose of this phase of the study was to investigate the biological complexity of Vernonia noveboracensis and V. harperi in order to help clarify their systematics. In this study observations and conclusions were based on field work, on analysis of local population samples, on garden studies, on laboratory work, and on examination of her- barium specimens. Population samples: Local population samples (mass collections) of 25 individuals each were made from four localities in Georgia in late August and September of 1967. These provided a transect from the mountains to the Coastal Plain. Population sample 15048 was collected in a wet pasture in White County in the Blue Ridge Mountains and is designated as BR. Population sample 15117 was made from a colony growing on a low moist roadside in Greene County in the Piedmont Plateau and is called PP. A wet pasture in McDuffie County near the Fall Line provided 15144, and it is labeled FL. Population sample 15086 was collected from a wet power line right-of-way in Tattnal County, just a few miles from the type location of V. har- peri. Tattnal County is in the Atlantic Coastal Plain, and this sample is designated as AC. Characters in which V. harperi and V. noveboracensis differ, as well as characters in which they are similar, were used in scoring the speci- mens, Ten characters were measured or scored on each in- dividual from the samples. The numerical values thus ob- 156 Rhodora [Vol. 72 tained were then punched on IBM cards. The characters were as follows: (1) width of the middle stem leaves (cm) ; (2) length of the middle stem leaves (cm) ; (3) pubescence of the middle stem leaves (for scoring purposes, categories were set up from 1 — glabrous to 6 = pubescent with hairs over .25 mm long and crowded); (4) dots on lower leaf surface range from 1 — not dotted to 3 — strongly dotted; (5) bract tip shape (9 categories were established for scor- ing purposes; however, these populations were all scored either 8 or 9); (6) pappus color, range from 1-5, with 1 — off-white to 5 — brown with a purple tinge; (7) in- volucre width (mm) ; (8) involucre length (mm) ; (9) num- ber of flowers per head; (10) leaf length/width ratio. The population samples were compared using the com- puter program BMDO7M, Stepwise Discriminant Analysis, version of 1 Sept. 1965, Health Sciences Computing Facility, UCLA. The program is a type of multi-variant analysis using a stepwise fitting in which the sums of squares and products are re-arranged at each stage of the fitting so that the variable giving the largest reduction of the residuals is the next one fitted. The standard deviations and means were obtained from the print-out of this program. From these the coefficients of the variation were determined. The variable characters within and between the populations were compared using scatter diagrams and polygons. The type specimen of Vernonia harperi was similar to TABLE 3 Number of cases of Vernonia noveboracensis classified into group by BMDO7M, Stepwise Discriminant Analysis Pro- gram. Population! BR PP FL AC BR 15 6 4 0 PP 0 25 0 0 FL 2 0 22 1 AC 0 0 0 25 ‘Blue Ridge (BR); Piedmont Plateau (PP); Fall Line (FL) ; Atlantic Coastal Plain (AC). 1970] Vernonia — Jones 157 certain of the AC plants. Analysis of herbarium specimens indicated that there is variation in V. noveboracensis ; how- ever, the changes in characters are either gradual over long distances or not correlated geographically or morpholog- ically with other characters. The results from the computer transect analysis indicated that all populations were some- what distinct (see Table 3). However, PP and AC individ- uals would not have been misclassified into the other three populations. On the other hand, some BR plants would have been classified into PP and FL. Likewise, FL had some plants which fell into BR and AC. A E Figure 4. Polygons based on the sample means of the four popu- lations (BR, PP, FL, AC) of Vernonia noveboracensis. A, leaf width; B, leaf length; C, involucre width; D, involucre length; E, number of flowers per head; F, ratio of leaf length/width; G, pubescence score. 158 Rhodora [Vol. 72 On the basis of the multi-variant analysis, the value of the characters presented in Table 4, the coincidence of the polygons (Fig. 4) and the scatter diagram (Fig. 5), some differentiation is indicated, yet the populations are probably indistinguishable. Grant (1963) has indicated that most outbreeding populations are variable or polymorphic. He noted that this is due to the diversity in the allelic forms of one or more genes. Since allelic forms do exist, no single 18 I l6 E 2 I3I 22 : 3 12 i E14 I4 4| |! | E 333 334342311 | i. 4 43212 | -|2 23431 22 1 II > 2 4222 TT 34. 322122 I lO 4 34 44432222 ul 3234 a 423 344 3 3 498 > 5 «6 20 30 40 50 60 £70 NUMBER FLOWERS PER HEAD Figure 5. Scatter diagram of individuals of the four populations (Ac, 1; FL, 2; PP, 3; BR, 4) of Vernonia moveboracensis. 1970] Vernonia — Jones 159 individual will carry all the genetic variation found within the populations. The breeding population maintains and stores this genetic variation which is conserved by the re- productive process. Changes in the allelic frequencies could be caused by mutation, gene flow, natural selection and genetic drift. Chromatographic patterns: The chromatographic pat- terns of the phenolic compounds of eight individuals from each population sample (Bk, PP, FL and AC) were in- vestigated through the use of 2-dimensional paper chro- matography. Dried flower heads were extracted in 1% HCl in methanol for 30 hours, and the extracts were concen- trated by evaporation. About 1 ml of the extract was applied in one spot to Whatman 3 MM chromatographie paper and then developed by the descending method. Development employed the following solvents in the order listed: (1) tertiary butanol acetic acid: water (3:1:1 v/v for 24 hours), and (2) acetic acid:water (15:85 v/v for four hours). Upon drying the chromatograms were examined in daylight, in transmitted longwave ultra-violet light (UV) both with and without the presence of ammonia vapor, and by re- flected UV light after having been sprayed with Benedict’s solution, Each plant was chromatographed twice. Twenty-two compounds were followed in the course of this study. No attempt was made to identify the com- pounds; however, color reactions with several reagents in- dicated that most were probably phenolic compounds. Al- though some compounds were either absent or below the threshold of detection in certain of the plants from each population, no pattern was found that would serve to dif- ferentiate the populations. Hybridization and garden experiments: Crosses were made in all possible combinations between BR, PP, FL, AC and between Vernonia acaulis and V. glauca. The F, hybrids were grown to maturity and pairing of the chromosomes was observed at meiosis, Pollen grains were stained in ani- line blue in lactophenol for 24 hours. Transplants from several locations were observed in the garden and green- 160 Rhodora TABLE 4 population sample means (x), Vernonia noveboracensis: [Vol. 72 standard deviations (s), ranges, and coefficients of variation (C) of seven morphological characters that were variable within and between the populations. Population? x 8 C Range Middle leaf width em BR 3.8 0.8 0.2 2.4- 5.5 PP 3.3 0.6 0.2 2.5- 4.5 FL 2.5 0.7 0.3 1.8- 4.3 AC 3.8 1.0 0.3 2.6- 5.5 Middle leaf length em BR 19.1 3.7 02 12.1-24.8 PP 21.8 3. 0.2 14.5 - 28.0 FL 17.1 2.8 0.2 12.9-24.5 AC 16.5 2.6 0.2 12.4-21.0 Involucre width mm BR 7.0 0.7 0.1 6.0- 8.5 PP 7.7 0.8 0.1 6.5- 9.0 FL 7.3 0.8 0.1 6.0- 9.2 AC 8.7 0.8 0.1 7.5 - 10.0 Involucre length mm BR 11.7 1.9 0.2 8.0 - 15.0 PP 10.7 1.8 0.2 7.0 - 14.0 FL 11.7 1.9 0.2 9.0 - 16.0 AC 13.8 1.8 0.1 11.0- 17.0 Number of flowers per head BR 40.6 4.4 0.1 33 -48 PP 40.4 4.4 0.1 380 -49 FL 46.2 4.5 0.1 388 -54 AC 48.9 6.6 0.1 39 -65 Ratio leaf length/width BR 5.1 1.3 0.3 3.0- 9.3 PP 6.6 1.1 0.2 3.7- 8.8 FL 7.0 1.5 0.2 3.4- 9.9 AC 4.6 1.2 0.3 2.6- 7.3 1970] Vernonia — Jones 161 Pubescence score BR 2.6 0.8 0.3 1 - 4 FP 2.8 0.4 0.2 2 ° FL 3.0 0.5 0.2 2 -4 AC 2.9 0.6 0.2 2 -4 iBlue Ridge (BR); Piedmont Plateau (PP); Fall Line (FL) ; Atlantic Coastal Plain (AC). house for over two years. The F, hybrids were vigorous and had good chromosome pairing at meiosis. Pollen stain- ability was usually above 90%. All populations studied have a chromosome number of n = 17. The morphological variations seemed to be maintained in the transplant gar- den. Certain F. hybrids from the cross V. nove horacensis X V. glauca had reduced pollen stainability and poor chromo- some pairing at meiosis. This indicates that hybrid break- down may occur in later generations even though the F,’s are fertile. Vernonia noveboracensis hybridizes with V. altissima Nutt. in eastern Tennessee, in eastern. Kentucky and in Pennsylvania, This produces some specimens with smaller heads. In North Carolina V. noveboracensis hybridizes with V. acaulis. Also in that state, it may occasionally hybridize with V. angustifolia Michx. Herbarium specimens were observed that might possibly have been natural hybrids between V. noveboracensis and V. glauca. All of these hybrid combinations have been produced in the greenhouse. Vernonia noveboracensis is usually seasonally and ecolog- ically isolated from V. acaulis and V. glauca where they are sympatric. The former blooms in August and September, while the latter two species flower in July. Also V. nove- boracensis grows in much wetter soil than the other two. From an evolutionary viewpoint, the most important natu- ral hybridization is that with V. altissima as some local introgression seems to have occurred. POSSIBLE EVOLUTIONARY RELATIONSHIPS The presence of relatively few morphological and phys- iological differences is evidence of close relationships among 162 Rhodora [Vol. 72 these three taxa. Gleason (1923) suggests that they may have been derived from the more primitive Vernonia an- gustifolia or, more likely, a common ancestor of all of our present day eastern species. Present evidence indicates that this was a logical assumption. The taxa considered here apparently diverged from a common ancestor during past geological history. The species considered in this paper have several features in common, including around 40 flowers per head. In addi- tion, they all have acuminate bract tips, although the tips of Vernonia noveboracensis are often filiform and longer. The pappus of both V. glauca and V. acaulis is straw-colored to white, whereas it is brown to purple in V. noveboracensis. The latter flowers in August while the other two Species bloom in July. These characteristics are normally key characters in the species of Vernonia of eastern North America. The basal leaves of Vernonia acaulis seem to function in reducing the competition from the Sparse surrounding veg- etation of its habitat. The cauline leaves and longer stems of the other two taxa seem to be of selective advantage in overcoming their competition, for they are generally taller than the plants around them. This is especially true with V. noveboracensis which grows in moist areas with vigorous competing vegetation. All three species begin to grow early in the spring, Vernonia acaulis grows in uplands, sandy woods, well- drained flatwoods, and sandhill ecotones. Low, wet road- sides, wet meadows, and creek banks provide habitats for V. noveboracensis, whereas V. glauca is always found in well-drained, upland, oak-hickory woods. Geographical isolation of segments of the ancestral pop- ulation system, in conjunction with natural selection in response to influences of past changes in the earth's sur- face, climate and vegetation, could have produced the phys- iological and morphological differentiation of these species. Presently, the population systems appear to be separated 1970] Vernonia — Jones 163 by prezygotic seasonal and ecological isolating mechanisms and by a postzygotic hybrid breakdown in F. generations. DEPARTMENT OF BOTANY UNIVERSITY OF GEORGIA, ATHENS 30601 LITERATURE CITED Exman, E. L. 1914. West Indian Vernoniae. Arkiv fór Botanik. 13: 1-106. FERNALD, M. L. 1941. Another century of additions to the flora of Virginia (concluded). Rhodora 43: 635-657. GLEASON, H. A. 1923. Evolution and geographic distribution of the genus Vernonia in North America. Am. J. Botany 10: 187-202. GRANT, V. 1963. The origin of adaptations, Columbia Univ. Press, N. Y. 606 pp. Jones, S. B. 1964. Taxonomy of the narrow-leaved Vernonia of the Southeastern United States. Rhodora 66: 382-401. JONES, S. B. 1966. Experimental hybridizations in Vernonia (Com- positae). Brittonia 18: 39-44. Jones, S. B. 1967. Vernonia georgiana — species or hybrid? Brit- tonia 19: 161-164. STEELE, E. S. 1901. Additions to the flora of Washington. Proc. Biol. Soc. Washington. 14: 81-82. DISTRIBUTIONAL HISTORY OF EPILOBIUM HIRSUTUM (GREAT HAIRY WILLOW-HERB) IN NORTH AMERICA: RONALD L. STUCKEY The introduction, establishment, and spread of non-in- digenous aquatic and marsh plants in eastern North Amer- ica have continually been increasing since the coming of European man. Documented cases have been published for Butomus umbellatus (Core, 1941; Stuckey, 1968b), Hydro- charis morsus-ranae (Dore, 1968), Lycopus asper (Stuckey, 1969), Najas minor (Meriläinen, 1968), and Rorippa syl- vestris (Stuckey, 1966). Attempting to determine the dis- tributional history of these non-indigenous (introduced) species presents to present-day botanists new challenges and questions, such as: When did the species first become intro- duced? How far and to what parts of the country have the plants spread? Into what habitats do the plants become established? What are the various probable pathways and rates of migration that the plants have taken? What hab- itats and geographical areas can be predicted for the plants’ invasion and establishment in the future? These are basic questions that must be considered in understanding the biology and distribution of the species. It is important to put on record the time and location of the occurrences of plants that come into new areas because it gives botanists knowledge about when and to where the plants are moving, and therefore provides documentation for the changes that are continually occurring in the flora. Furthermore, the invasion of aquatic and marsh plants becomes critical for ‘Contribution from the Botany Program (Paper No. 766), the Her- barium, and The Franz Theodore Stone Laboratory, The Ohio State University, Columbus 43210. 164 1970] Epilobium hirsutum — Stuckey 165 Fig. 1. Epilobium hirsutum (Great hairy willow-herb) from along the banks at edge of an artificial pond at The Maples, North Central Branch of the Ohio Agricultural Research and Development Center, SE side of Sandusky Bay, NW corner of Erie County, Ohio, ca. 4 mi. NW of Castalia, 10 Aug 1968, Stuckey 7327, OS. Photograph by R. L. Stuckey. 166 Rhodora [Vol. 72 Fig. 2. Known distribution of Epilobium hirsutum (Great hairy willow-herb) in eastern North America based on herbarium specimens seen and other records cited in this paper. The position of each square 1970] Epilobium hirsutum — Stuckey 167 those interested in wildlife management. The invasion and establishment of plants new to a marsh can considerably alter the habitat and have effects on the future composition of the flora, the food chain of the animals, and perhaps ultimately determine the succession and total fate of the ecology of the marsh. This member of the evening-primrose family (Ona- graceae), Epilobium hirsutum L. (fig. 1) is a tall (up to 2 meters) plant with sessile, somewhat clasping, soft and hairy leaves. The flowers have reddish-purple, obcordate petals and are arranged in racemes in the axils of the leaves on the upper part of the plant. Reproduction is by short rootstalks or subterranean stolons and by seeds dispersed with the wind. When growing in water or water-saturated mud those parts of the plant in contact with the water develop arenchyma tissue (Batten, 1918). Great hairy willow-herb is a species of the marshes of Eurasia that has become established within the past 140 years in eastern North America. Here it occurs in a wide range of habitats. The earliest North American records indicate that the species grew in cultivated grounds, as a weed in gardens, or on ballast. As the species became estab- lished along the northeastern coast of the United States and then spread inland, it has come to occupy river, creek, and stream banks, roadside and drainage ditches, canals, edges of ponds and lakes, wet meadows and pastures, railroad tracks and ditches, extensive marshes, and edges of swamps. or circle represents a locality where a plant has been collected. Numbers in the squares are the last two digits for the year in which the plant was collected if before 1900, and the numbers in circles are the last two digits for the year of collection if since 1900. The oldest known collection is mapped for each locality. The arrows indicate the possible migration pattern that the species has taken through time as the plants have moved from the east coast to inland in the Great Lakes region and elsewhere. Since the preparation of this map, additional specimens have become available for locations along the southeast shore of Lake Erie in New York, between Ripley and Westfield, Chattauga County (Beth Evans s.n., OS, Sep 1969) and in Pennsylvania, southeast of the city of Erie, Erie County (Beth Evans s.n., OS, Sep 1969). 168 Rhodora [Vol. 72 Epilobium hirsutum can be common in low wet places, espe- cially where fires have burned the land (McKeever, 1961). According to several of the more recent floristic manuals of the region (Fernald, 1950; Gleason, 1952; Gleason and Cronquist, 1963) and a recent taxonomic revision of the genus for North America (Munz, 1965), E. hirsutum occurs in southern Quebec, southern Ontario, New England, south to New Jersey, Pennsylvania and westward through New York, Ohio, Michigan, and Illinois. Munz also reports it from Washington. Figure 2 shows the present known distribution in eastern North America. INTRODUCTION AND EARLY SPREAD The earliest known record of Epilobium hirsutum from North America is based on a collection from Newport, Rhode Island, July 1829 (Bridges s.n. PH), but no habitat or other information about its environment or history is given with the specimen. E. hirsutum was not recorded in the fourth edition (or earlier editions) of Gray's Manual (Gray, 1863), but it was reported in the fifth edition (Gray, 1867) as "Spontaneous in waste grounds, New Bedford, Mass. [without date] (T. A. Greene) and Roxbury [towards Dorchester, 1864] (D. Murray) ; and in a ravine near [1 mi n of] Albany, New York [1861] (C. H. Peck)."* According to A Catalogue of the Plants . . . of Nantucket, Mass. (Owen, 1888), E. hirsutum was “Raised in or about 1855 in a garden on Union St., from which it gradually spread by seed," until by 1888 it was “well established in many waste places about town and ... sometimes in the fields." In a flora of Essex County, Massachusetts, Robinson (1880) noted that the plants grew in “old gardens and waste heaps in Salem about 1860, and still continues in some places." Other early occurrences for eastern Massachusetts based on herbarium specimens seen are from New Bedford, 1865 (Robbins s.n., NEBC), Woods Hole, 1872 (Palmer s.n., US), Nantucket, 1873 (Harrington s.n., MICH), Plymouth, 1881 (Clark 898, NEBC), and Dartmouth, 1888 (Sturtevant 5037, "Data given in brackets are taken from the original specimens at GH. 1970] Epilobium hirsutum — Stuckey 169 NY). In the mid-nineteenth century the species must have become established in the Newport, Rhode Island, area, be- cause Tweedy (1881) reported it as “not uncommon in waste ground surrounding dwellings.” In Maine, the earliest known specimen was collected in 1889 at Portland and bears the notation, “weed in garden," (Fernald s.n., GH, NEBC). In New York, E. hirsutum is not mentioned by Torrey (1843), and except for the Albany locality given by Gray (1867), the next report for the state of New York appears to be Dudley (1886), who wrote that the plant was found in 1874 near the mill west of Cascade Place in the Cayuga Basin in the central part of New York state. A voucher specimen for this record has not been seen, Brcwn (1878) mentioned the willow-herb as “scarce” on ballast and on made land in the New York City area. In the following year he collected a plant on ballast ground at Gowanus, Brooklyn (Brown s.n., US). The species apparently became introduced farther into the interior in the Buffalo area about 1882, based on information by Day (1882-1886, 1883a, 1883b) who said that seeds of certain plants, new to the Buffalo area, were sown in waste places in the city and suburbs during 1882. According to Day (1888), E. hir- sutum was introduced, but perhaps not established, near Clifton, Ontario, which is north of Buffalo. Voucher speci- mens of the willow-herb from the city of Buffalo or town of Clifton have not been seen, but specimens are known from the area about Niagara Falls. The earliest known specimen seen is from Queen Victoria Park, Niagara Falls, July 1890 (Schafer s.n., PH). A specimen obtained in 1894 by R. Cameron (CAN) has the note “naturalized at Niagara Falls, introduced in garden seed." Before the turn of the century Gray (1889), Trelease (1891), and Britton and Brown (1897) each reported the species from waste grounds at various places on the Massachusetts and Rhode Island coast and in the interior in New York state and Ontario, EXPANSION OF THE RANGE The range of E. hirsutum continued to expand in the early 1900's. In the state of New York, House (1924) and Wie- 170 Rhodora [Vol. 72 gand and Eames (1926) both commented on its apparent recent introduction, and House (1924) reported it as “Be- coming abundant in many sections of the State, where not observed a few years ago.” Several specimens from the west central part of New York state document its occur- rence in 1916 at McLean, Tompkins County (Gershoy 6854a, GH), at Conquest, Cayuga County (Griscom & Wright 6856, GH), at Cortland, Cortland County (Munz & Wright 6854, GH), and in 1918 at Fayette, Seneca County (Evans 10445, GH). Zenkert (1934) wrote “The progress of this species across New York State in the past twenty or thirty years has been rather rapid. It seems to have followed the water- ways up the Mohawk and Seneca Rivers.” The willow-herb apparently was first found in Nova Scotia at Yarmouth in 1926 (Groh s.n., CAN, DAO), but evi- dently the species was not reported for the Maritime Prov- inces until 1951, based on a later collection from Point Pleasant, Halifax, Nova Scotia, 1 August 1949 (Erskine, 1951). For Quebec the species was not reported by Marie- Victorin (1935) in his Flore Laurentienne. In a supplement to Marie-Victorin's Flore, Rouleau (1947) apparently first reported E. hirsutum for Quebec, The earliest known speci- men comes from the “rivages du Saint-Laurent" at Les Cedres, between Melocheville and Coteau-du-Lac in 1940 (Marie-Victorin et al. 4292, CAN, F, GH, MO, NY, PH, US). Raymond and Kucyniak (1948) listed two additional sta- tions downstream as far as the western limits of the city of Montréal where plants (specimens not seen) had been found in 1944 and 1945. They pointed out that the species, *first introduced in the region of Lake Ontario, has come into Quebec [from the southwest] along the route of migra- tion followed by several native plants, such as Allium cana- dense and Justicia americana, . . . via the shores of the St. Lawrence river." Rousseau (1968) also confirmed Marie- Victorin's 1940 record and mapped the species' distribution for Quebec. Several specimen records from localities along or near Lake Ontario document its early spread in Ontario in 1930 1970] Epilobium hirsutum — Stuckey 171 at Toronto, York County (Brown 1058, TRT), in 1933 at Port Hope, Durham County (Pease & Bean 23686, GH), in 1936 at Aberfoyle, Wellington County (Stroud s.n., DAO, TRT), in 1939 at Bath, Lennox and Addington County (Min- shall 417, DAO), and along the east end of Lake Erie in 1935 at Turkey Point, Norfolk County (Bowden s.n., HAM), and in 1937 at Port Colborne, Welland County (Simon s.n., TRT). In a 1953 survey of the St. Lawrence Seaway area, Dore and Gillett (1955) pointed out that E. hirsutum was becoming common in many marshy places and ditches, but only near the River and its adjacent main highway. Mont- gomery (1957) reported E. hirsutum as “a common weed in low lying and swampy areas in the southern counties” of Ontario. The species appears to be on its way to becoming thoroughly established in the southern Ontario marsh flora as evidenced by many collections in the 1940’s, 1950’s, and 1960's. MIGRATION WESTWARD IN THE GREAT LAKES REGION The great hairy willow-herb apparently moved :nto north- eastern Ohio sometime before the 1930’s. The first known collection was obtained in 1932 along Lake Erie in a swamp near Conneaut, Ashtabula County (Hicks s.n., 08). Hicks, an active field botanist in Ohio in the 1930’s, wrote on the label, “New Ohio Record." A specimen was obtained in 1938 farther west along the lake at Perry in Lake County (Cro- foot s.n., 08). The species has been found in six other north- eastern Ohio counties as revealed by collections in The Ohio State University Herbarium from 1955 to 1958. Its occur- rence westward along Lake Erie is more recent, where it was first found at Lighthouse Point at the north end of Pelee Island in 1959 (Pinkava 59194 and Wells 173, os). Over the past several years further collections verifying the occurrence and spread of E. hirsutum have been ob- tained in western Lake Erie. In addition to those localities cited in Stuckey (1968a), the species has been found in northwestern Ohio in Erie County along old dike in marsh along west side of old Cedar Point Causeway at the NE 172 Rhodora [Vol. 72 limits of Sandusky (Stuckey 8500, os, 14 Sep 1969) ; in Ot- tawa County on mud flats along edge of marsh at East Harbor, East Harbor State Park (Stuckey 7359, GH, OS, 14 Aug 1968; Stuckey 8220, os, 31 Jul 1969), in cat-tail marsh at Turtle Creek along roadside, SW corner of Sec. 5, T7N, R15E, Carroll Twp., ca. 6 mi N of Oak Harbor (Stuckey 7526, CAN, 08, 26 Aug 1968), on wet sandy beach, along edge of marsh, Sand Point, Marblehead Peninsula, ca. 2.5 mi S of the town of Marblehead (Stuckey 7684, os, PH, 12 Sep 1968), and at edge of grape vineyard, south side of Fox's Marsh, North Bass Island (Stuckey 8205, os, 29 Jul 1969; Stuckey 8288, os, 28 Aug 1969) ; and in Monroe County, Michigan, where it was occasional in wet sand on beach along shore of Lake Erie, SW corner Sec. 35, Frenchtown Twp., N edge of Sterling Monroe State Park, E edge of the city of Monroe (Stuckey 7763, 0S, 14 Sep 1968). The first report for Michigan is that of Hanes (1945) based on plants found in 1943 in low ground near Portage, Kalamazoo County (Becker 4362, specimen not seen). A later collection comes from a marshy roadside, near Bay Port, Huron County in 1952 (Hermann 11778, GH, MICH). Since then E. hirsutum has been found in scattered locations in the Lower Peninsula of the state, several of which (from southeastern Michigan) are cited in Stuckey (1968a). The species has reached the Straits of Mackinac area where collections are known as early as 1953 (Voss, 1957a, 1957b). The species continues to persist in this area, based on a more recent collection, locally common along roadside in Macki- naw City, Emmet County (Voss 12950, os, UMBS, 10 Aug 1969). The farthest westward penetration for E. hirsutum in the Great Lakes region has been to northern Illinois (Stey- ermark & Swink, 1949) where plants were first found in 1948 at the water's edge along a ditch at the base of a railroad embankment in South Chicago, Cook County (Stey- ermark & Swink 65978, F, MO). In 1957 a specimen was collected along the Santa Fe Railroad south of Argonne Na- tional Laboratory at the southwest end of Rocky Glen Forest 1970] Epilobium hirsutum — Stuckey 173 Preserve, DuPage County (Bartel 7, F) Steyermark & Swink, 1959). Epilobium hirsutum apparently has not reeched other states bordering Lake Michigan and Lake Superior, for it is not reported for Indiana (Deam, 1940), Wisconsin (Ugent, 1963), or the counties of St. Louis and Lake in northeastern Minnesota (Lakela, 1965). It is evidently absent farther west because it is not recorded in recent checklists or floras for North Dakota (Stevens, 1950), Mani- toba (Scoggin, 1957), Saskatchewan (Breitung, 1957), and Alberta (Moss, 1959). EXPANSION IN THE SOUTHEAST PORTION OF THE RANGE Since 1947 the willow-herb has been found at additional localities in northern New Jersey in 1949 at Patterson, Passaic County (Schaeffer 31092, PH), in eastern Pennsyl- vania in 1946 at Glenburn, Lackawanna County (Glowenke 8573, GH), in 1947 at Treichlers, Northampton County Schaeffer 26937a, PH), in 1967 at Temple, Berks County (Wilkens 12793, PH), and in Maryland along railroads near Chesapeake Bay, in 1955 at Baltimore (Baltar 635, US) and in 1960 at Magnolia, Harford County (Baltar 2735, US). Reed (1964) cited collections from a large patch in wastes (Reed 39575, Oct 1957) and on chrome ore piles (Reed 45712, Oct 1959) both at Canton in the port of Baltimore, Maryland. E. hirsutum has not been reported farther in- land from such places as Centre and Huntingdon Counties, Pennsylvania (Westerfield, 1961), and West Virginia (Strausbaugh & Core, 1958), or in the coastal states of Vir- ginia (Massey, 1961) and North and South Carolina (Rad- ford, Ahles, & Bell, 1968). OCCURRENCE IN THE STATE OF WASHINGTON In the state of Washington, E. hirsutum was found in 1965 in wet ground at edge of railroad track near Belling- ham, Whatcom County (Sutherland 1084, CAN) and in 1966 abundant along a ditch beside a highway at Bingen, Klicki- tat County (Spellenberg & Sutherland 1191, NY). The oc- 174 Rhodora [Vol. 72 I T l | I T I I eoe oe o * use £ ° U @ $ 9 e ss I O = e o & e | 09-9 w o [ e su z X P. J E & e 6&6 9v @ LL. z E o * 9 e ən I < u < = 2 < 6 s e "x2 = Ww | Oo z & s e st ó < “< & w x< c E è * e xz ae: - ^ Q 9 6 e uu o = @ =< LI e @ w WH I O u ui > e su G e | 01-90 Oo _ E e e | soi o S `x 3 . š 2 e use O > o > A e e su = 3 G < "T @ | 06-3 z £ Ç > ° 4 oz e su < = > a r < m z 9 3 o e @ ww u S Wù < O Z Z ul e su wv OQ O m < e un? l l | L |I l L | + (9 u. ul Q O m < vddv 911518013 Fig. 3. Dates of Herbarium Specimens Verifying the Time of Spread and Establishment of Epilobium hirsutum in Eight Floristic 1970] Epilobium hirsutum — Stuckey 175 currences of this species in Washington probably represent recent introductions separate from its migration and spread in eastern North America. PRESENT STATUS IN NORTHEASTERN UNITED STATES Although the previous discussion and the map (fig. 2) give the early occurrences of E. hirsutum in various states or areas on the continent, there is no indication whether the species has become established in these areas. At Newport, Rhode Island, the locality of the oldest collection, several specimens are known from 1829 to the most recent seen ob- tained in 1959 (Eaton s.n., NEBC). These recorcs therefore confirm the establishment of the species in this area. Many specimens in the New England Botanical Club, New York Botanical Garden, and Gray Herbaria document its estab- lishment and spread on Nantucket Island. Bickaell (1914) stated that E. hirsutum was “Common in the lower parts of town and out into the suburbs, often massed along ditches and in corners of damp lots and low fields; also by the roadside in Shawkeno, the only place where I saw it far from town..." By 1958, according to Mackeever (1961), E. hir- sutum has spread throughout the island. In a check-list of the vascular plants of Maine (Ogden, Steinmetz, & Hyland, 1948), Cumberland County continues to be the only locality for its occurrnce in that state. In the New York City area Monachino (1959) reports that the great hairy willow-herb *is now frequent and locally abundant . . ." Through the years certain localities or general areas have been studied floristically rather frequently and more thor- oughly than other areas by botanists. Within the present range of E, hirsutum, eight of these “rather well-studied floristic areas" were selected. Each area is defined geo- graphically by several counties as given in table 1 and by the fact that one or more local floras or checklists of plants have been prepared for each area prior to obtaining the first known record of E. hirsutum. In figure 3 these floristic Areas of North America. Each dot represents one or more herbarium specimens collected within a five-year period for each floristic area. 176 Rhodora [Vol. 72 areas are listed on the vertical axis and time is represented on the horizontal axis. All specimens or records of speci- mens examined from these areas are each marked with a dot along the horizontal axis at the corresponding five-year interval of their collection. Figure 3 shows (1) the earliest specimen record known for each floristic area, (2) that the species has become established in each floristic area, be- cause specimens have been found and obtained, in the years following the original collection and even up to the present time in certain areas, and (3) the gradual progression and establishment of E. hirsutum from the northeast coast of the United States, through central New York state, and into the Great Lakes region. FUTURE EXPANSION OF THE RANGE Since 1929, E. hirsutum has expanded its range through- out northeastern United States and into the Great Lakes region. It therefore would be expected that E. hirsutum should continue to expand its distribution on the North American continent into localities with habitats similar to those where the plants have already become established. The present range of E. hirsutum has approximately the same general range in North America that Rorippa sylves- tris had about 1900 (Stuckey, 1966). R. sylvestris, a species that occurs in habitats (waste places, gardens, marshes, ditches, and along rivers) similar to those of E. hirsutum, has since 1900 spread farther westward into Wisconsin, Iowa, and North Dakota, and southward into Virginia, West Virginia, Ohio, Indiana, Illinois, and Missouri. Given an- other 70 years, perhaps E. hirsutum will spread to an extent that its range may be similar to that of R. sylvestris today. Where both species are native and have had considerable time for spreading in Europe, they both have similar dis- tributions, although E. hirsutum is absent in the extreme north (Valentine, 1964; Raven, 1968). Jonsell (1968, p. 97- 103) discusses the distributional history of R. sylvestris in Scandinavia and shows that its range has been expanding northward over the past 100 years. Comparative migration 1970] Epilobium hirsutum — Stuckey TIT rates and distributional patterns, as illustrated by these two species in North America, should be useful for predicting areas to where and when other non-indigenous shore and marsh species may or may not migrate. However, our knowledge for making these kinds of comparisons is meager because the distributional histories of many of the species have not yet been worked out. ACKNOWLEDGMENTS I am grateful to Mr. Richard Hendricks for assembling the data from the literature and several herbaria and pre- paring a preliminary version of the map. My thanks are extended to those herbaria (DAO, GH, MICH, MO, NEBC, NY, OS, PH, and US) whose specimens were examined. Records of herbarium specimens have been provided by Dr. James Wells (BLH), Dr. James Soper (CAN), Mr. Richard Lowden (F), Dr. James Pringle (HAM), Dr. James Cruise (TRT), and Dr. Edward G. Voss (UMBS). Mary R. Norcross and Elsa Prediga have contributed information from the Maria Mitchell Science Library, Nantucket, Massachusetts. The field study in the western Lake Erie area was conducted as part of the teaching and research program at The Franz Theodore Stone Laboratory, Put-in-Bay, Ohio, during the summers of 1967, 1968, and 1969. Additional support for field work has been provided by The Ohio Biological Survey. I am thankful to Mr. Ronnie Johnson, Mr. W. Alan Wentz, and Mr. Thomas Duncan who served as field assistants, and to Mr. William H. Anderson who printed the photograph and map for publication. 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Manual of the Botany of the Northern United States, Including the Dis- trict East of the Mississippi and North of North Carolina and Tennessee. American Book Co., New York. 760 p. + XXV pl. HANES, CLARENCE R. 1945. Additions to the flora of Michigan. V. Pap. Mich. Acad. (1944) 30: 63-66. House, Homer D. 1924. Annotated list of the ferns and flowering plants of New York state. N. Y. State Mus. Bull. 254: 1-759. JONSELL, BENGT. 1968. Studies in the north-west European species of Rorippa s. str. Symb. Bot. Upsal. 19: 1-121 + XI pl. LAKELA, OLGA. 1965. A Flora of Northeastern Minnesota. Univ. Minn. Press, Minneapolis. 541 p. MACKEEVER, FRANK. 1961. After a fire comes fireweed; an “Old Timer” on Nantucket. Inquirer & Mirror. January 13. MARIE-VICTORIN, FRÈRE. 1935. Flore Laurentienne. Les Fréres des Ecoles Chrétiennes, Montréal. 916 p. Massey, A. B. 1961. Virginia Flora. Va. Agr. Exp. Sta. Tech. Bull. 155. 258 p. MERILAINEN, JouKO. 1968. Najas minor. All. in North America. Rhodora 70: 161-175. MONACHINO, JOSEPH. 1959. Field trip reports — Jamaica Bay Sanc- tuary, Brooklyn, N. Y. Bull. Torrey Bot. Club 86: 64-66. Montcomery, F. H. 1957. The introduced plants of Ontario grow- ing outside of cultivation (Part II). Trans. Royal Can. Inst. 32: 1-34. Moss, E. H. 1959. Flora of Alberta. Univ. Toronto Fress, Toronto. 546 p. Munz, PHILIP A. 1965. Onagraceae. N. Am. Fl. Ser. II. Part 5. 1-231. OapEN, E. C., F. H. STEINMETZ, and F. HYLAND. 1948. Check-list of the vascular plants of Maine. Bull. Josselyn Bot. Soc., Maine No. 8. 70 p. OwEN, Marra L. 1888. A Catalogue of Plants Growing Without Cultivation in the County of Nantucket, Mass. Gazette Printing Co., Northampton. 87 p. RADFORD, ALBERT E., HARRY E. AHLES, and C. RITCHIE BELL. 1968. Manual of the Vascular Flora of the Carolinas. Univ. N. Carol. Press, Chapel Hill. 1183 p. RAvEN, P. H. 1968. Epilobiwm, p. 308-311. In Flora Europaea Vol. 2 Rosaceae to Umbelliferae. Univ. Press, Cambridge. 455 p. + v maps. 180 Rhodora [Vol. 72 RAYMOND, MARCEL, and JAMES KUCYNIAK. 1948. Six additions to the adventitious flora of Quebec. Rhodora 50: 176-180. REED, CLYDE F. 1964. A flora of the chrome and manganese ore piles at Canton, in the port of Baltimore, Maryland and at New- port News, Virginia, with descriptions of genera and species new to the flora of eastern United States. Phytologia 10: 321-406. ROBINSON, JOHN. 1880. The flora of Essex County, Massachusetts. Essex Institute, Salem. 200 p. ROULEAU, ERNEST. 1947. Supplément a la Flore Laurentienne. Fréres des Ecoles Chrétiennes, Montréal. 63 p. ROUSSEAU, CAMILLE. 1968. Historie, habitat et distribution de 220 plantes introduites au Québec. Nat. Canad. 95: 49-169. Scocein, H. J. 1957. Flora of Manitoba. Natl. Mus. Can. Bull. No. 140. 619 p. STEVENS, O. A. 1950. Handbook of North Dakota Plants. N. D. Agr. Coll., Fargo. 324 p. STEYERMARK, JULIAN A., and FLOYD A. SWINK. 1949. Plants new to Illinois or to the Chicago area in Illinois. Rhodora 51: 147-149. and —— s P 1959. Plants new to Illinois and to the Chicago region. Rhodora 61: 24-27. STRAUSBAUGH, P. D. and EARL L. CORE. 1958. Flora of West Vir- ginia (Part III). W. Va. Univ. Bull. Ser. 58(12-3) : 571-860. STUCKEY, RONALD L. 1966. The distribution of Rorippa sylvestris (Cruciferae) in North America. Sida 2: 361-376. 1968a. Aquatic flowering plants new to the Erie Islands. Ohio J. Sci. 68: 180-187. 1968b. Distributional history of Bu ‘omus umbellatus (flowering-rush) in the western Lake Erie and Lake St. Clair region. Mich. Bot. 7: 134-142. 1969. The introduction and spread of Lyco- pus asper (western water horehound) in the western Lake Erie and Lake St. Clair region. Mich. Bot. 8: 111-120. TORREY, JOHN. 1843. A Flora of the State of New-York, Compris- ing Full Descriptions of All the Indigenous and Naturalized Plants Hitherto Discovered in the State; with Remarks on their Economical and Medical Properties. Vol. I. D. Appleton & Co., New York. i-xii + 1-484 p. TRELEASE, WILLIAM. 1891. A revision of the American species of Epilobium occurring north of Mexico. 2nd. Ann, Rep. Missouri Bot. Gard. 1891: 69-117 + 47 pl. TWEEDY, FRANK. 1882. Notes on the flora of Newport, R. I. Ball. Torrey Bot. Club 9: 23-24. UGENT, DONALD. 1963. Preliminary reports on the flora of Wiscon- sin. No. 47. The orders Thymelaeales, Myrtales, and Cactales. Trans. Wis. Acad. 51: 83-134. 1970] Epilobium hirsutum — Stuckey 181 VALENTINE, D. H. 1964. Rorippa, p. 283-284. In Flora Europaea Vol. 1 Lycopodiaceae to Platanaceae. Univ. Press, Cambridge. 464 p. + v maps. Voss, EDWARD G. 1957a. New records of vascular plants from the Douglas Lake region (Emmet and Cheboygan Counties), Michi- gan. Pap. Mich. Acad. (1956) 42: 3-34. 1957b. Observations on the Michigan flora — VI. Distribution records of some Angiosperms new, rare, or mis- interpreted in the state. Brittonia 9: 83-101. WESTERFIELD, WALTER F. 1961. An annotated list of vascular plants of Centre and Huntingdon Counties, Pennsylvania. Castanea 26: 1-80. WIEGAND, KARL M., and ARTHUR J. EAMES. 1926. The flora of the Cayuga Lake Basin, New York. Cornell Univ. Agr. Exp. Sta. Mem, 92. 491 p. ZENKERT, CHARLES A. 1934. The flora of the Niagara Frontier re- gion. Buffalo Soc. Nat. Sci. Bull. 16. 328 p. Table 1. Well-Studied Floristic Areas Within the Present Range of E. hirsutum. Eastern Massachusetts Area Barnstable, Bristol, Dukes, Essex, Middlesex, Nantucket, Norfolk, Plymouth, and Suffolk Counties, Massachusetts. New York City Area Bronx, Kings, Nassau, New York, Queens, Richmond, Suffolk, and Westchester Counties, New York; Bergen and Passaic Counties, New Jersey. Niagara Falls Area Erie, Genesse, Niagara, and Orleans Counties, New York; Halton, Lincoln, Peel, Welland, Wentworth, and York Counties, Ontario. Central New York Area Cayuga, Chemung, Cortland, Madison, Monroe, Onondaga, Seneca, Tompkins, and Wayne Counties, New York. Western Lake Erie Area Erie, Lucas, Ottawa, and Sandusky Counties, Ohio; Monroe and Wayne Counties, Michigan; Essex and Kent Counties, Ontario. Southern End of Lake Michigan Area Cook, DuPage, and Lake Counties, Illinois; Lake, Laporte, and Porter Counties, Indiana; Berrien and Van Buren Counties, Michigan. Mackinac Straits Area Cheboygan, Emmet, and Mackinaw Counties, Michigan. State of Washington Klickitat and Whatcom Counties. A RE-EVALUATION OF POLYGONUM MEISNERIANUM IN NORTH AMERICA’ RICHARD S. MITCHELL In eastern North America the two widespread species of Polygonum, section Echinocaulon, are P. sagittatum and P. arifolium. These are known by their retrorse barbs, sprawling habit and sagittate to hastate leaves. A third entity, Polygonum meisnerianum C. & S. var. beyrichianum (C. & S.) Meisn., has gone almost unnoticed, although it was first collected in the United States over 140 years ago. It is one of the widest ranging aquatic angiosperms in the western hemisphere, and is unusual in that it simulates members of section Persicaria in both leaf shape and erect- emergent habit. Retrorse barbs and inflorescence type link P. meisnerianum with other ‘“Tear-thumbs” (Echinocau- lon) ; however, its center of distribution and nearest phylo- genetic relationships are in eastern South America, not in Asia, as is the case for both P. sagittatum and P. arifolium. The purpose of this paper is to bring knowledge of P. meisnerianum out of its present state of obscurity, so that the species may now be recognized by manuals and floristic treatments in the southern United States. Two collections from Florida and one from South Carolina are also reported here, and a variety based upon phenotypic modification is placed in synonymy. Historical Account Polygonum meisnerianum and P. beyrichianum were first described by Chamisso and Schlechtendal (1828) from ma- terials collected on a Brazilian expedition. A few years later, Chamisso (1833) described two additional related species, P. rubricaule and P. stelligerum. 'Thus, by 1833 there were four published species descriptions with char- "This research was supported in part by U. S. Department of the Interior Office of Water Resources Research Grant 373204-5, admin- istered by the V.P.I. Water Research Center. 182 1970] Polygonum meisnerianum — Mitchell 183 acters distributed among members of a group of closely inter-related taxa. This species-complex has its center of distribution in Brazil and Argentina. A number of names have been applied in Latin America since 1833, but most are synonyms whose descriptions add no new diagnostic information, A list of pertinent synonymy appears in the treatment to follow. Meisner, in Martius’ flora of Brazil (1855), reduced P. beyrichianum to varietal status, listing the name P. meis- nerianum C. & S. var. beyrichianum (C. & S.) Meisn., which is now the most widely accepted combination. There seems to be sufficient intermediacy in some Brazilian and Argen- tinian specimens to warrant this change, and the problem is still in need of study; however, plants designated “var. beyrichianum” do occur in typical morphological form throughout Brazil and Argentina, extending :o Central America, Mexico, Cuba and at least three states in the United States, The first person to collect P. meisnerianum in the United States was T. Drummond, whose specimen number 177 is cited, without date, by Meisner (1855). This specimen, supposedly from Texas, has not been found by the author, but Drummond's number 277, dated 1832, from New Or- leans, was found at both the Gray Herbarium of Harvard University and the U. S. National Herbarium. Specimens collected by J. L. Riddell in Louisiana are partially without dates, but the earliest seems to be August 13, 1838, from Madisonville, Louisiana. He noted in paren- theses that the species was undescribed, but labeled it P. geniculatum Ridd., a name which he then published in de Candolle's Prodromus. The species was omitted from Chapman's Flora of the Southeastern States (1889). A. B. Langlois collected the plant in 1885 and again in 1892, both times in Louisiana. He noted that it “is pro- nounced a rare plant." J. K. Small, in his monograph of North American Poly- gonum (1895), illustrated and re-described specimens under 184 Rhodora [Vol. 72 the correct name of P. meisnerianum var. beyrichianum. It is surprising that this taxon was omitted from his Manual of the Southeastern Flora (1933). Collections of this species have been extremely infrequent in North America since the turn of the century. There are no sheets of it in the herbaria of the University of Texas, Southern Methodist University or Texas Research Founda- tion, where the aquatic flora of the southwestern states is being studied extensively. There are also no specimens at Louisiana State University or Southeastern Louisiana State University. In the summer of 1968, I found P. meisnerianum var. beyrichianum growing at the south end of Lake Miccosukee on the Leon-Jefferson line in Florida. It had been collected once before in northern Florida (D. B, Ward, personal com- munication) in 1935 by Erdman West, at Newnan Lake in Alachua County. These two stations establish firmly its natural occurrence in Florida, and it has now been included in the Florida checklist. Recently a state record for South Carolina was collected, but went unnoticed. It was found in Jasper County (S. W. Leonard #2231, and A. E. Radford) in a wet ditch on High- way 601, 1.4 miles north of County road 27-169. The speci- mens are labelled "P. sagittatum L.; unusually narrow leaves". With the occurrence of P. meisnerianum in Florida, Cuba and South Carolina established, the North American dis- tribution of the species may best be described as sporadic around the Gulf of Mexico. The migration route from South America was probably along the expanded coastal plain during the Pleistocene pluvials, The direction of migration is now in doubt, however, since the occurrence of the species in the West Indies, Florida and South Caro- lina opens up the possibility of an eastern route. Morphological Transformation Polygonum meisnerianum var. beyrichianum differs from the typical variety of the species as well as its other South 1970] Polygonum meisnerianum — Mitchell 185 Fig. 1. Polygonum meisnerianum C. & S. var. beyrichianum (C. & S.) Meisn, a wide-ranging South American aquatic which reaches its northern limits of distribution in Florida, Louisiana and South Carolina. A. habit, showing dichotomously branching inflores- cence. B. inflorescence, with glandular petioles. C. nodal region with sheathing stipule (ocrea), clasping leaf base and retrorse barbs. D. trigonous achene. American relatives in having sessile, slightly auriculate leaves rather than petiolate, sagittate ones. A sagittate- leaved variety, P. meisnerianum C. & S. var. jalapense Robins., was described in 1900 from Mexico. On examina- tion, the type specimen shows a dense covering of glandular hairs on leaves, petioles and stems, and no barbs or other pubescence. 186 Rhodora [Vol. 72 Fig. 2. Leaf silhouettes as evidence that Polygonum meisnerianum var. jalapense is a name based upon a phenotypic modification of var. beyrichianum, A. leaves from P. meisnerianum var. beyrichianum collected in Florida. B. juvenile leaves from rhizomes of the same plants after being cut back and grown in the greenhouse. These leaf types persisted for months. C. leaves from the type specimen of P. meisnerianum var. jalapense. Mass collections of var. beyrichianum sent from Florida by R. K. Godfrey in 1969 showed typical leaf morphology throughout (Fig. 1); however, rhizomes of these plants, cut back and shipped live, gave surprising results when grown in a Virginia greenhouse. Young shoots produced only sagittate leaves with substantial petioles, The leaves and stems were covered with glandular hairs and lacked barbs or other pubescence. Thus, var. beyrichianum simu- lated the type specimen of var. jalapense almost exactly in its juvenile growth (Fig. 2), and continued to do so for many months after its disturbance and etiolation. It is now clear that “var. jalapense" can be induced environmentally, and should not retain taxonomic status. This atavistic mani- festation also gives good evidence that the leaf type of var. beyrichianum is a derived condition, and that this variety has evolved from plants with sagittate leaves. 1970] Polygonum meisnerianum — Mitchell 187 Pertinent Synonymy Polygonum meisnerianum var. beyrichianum appears, at present, to be the only species and variety of the South American Tear-thumb group which occurs in North Amer- ica. It is listed here with its Brazilian relatives and the most frequently used synonyms for the group. 1. Polygonum meisnerianum C. & S. var. meisnerianum, Linnaea, 3: 42. 1828. 2. Polygonum meisnerianum C. & S. var. beyrichianum (C. & S.) Meisn., in Martius’ Fl. Bras., 5: 19. 1855. P. beyrichianum C. & S., Linnaea 3: 42. 1828. P. multiangulare H. & A., Comp. Bot. Mag. 2: 62. 1836. . geniculatum Ridd. ex Meisn., DC. Prodromus, 14: 42. . chamissoanum Wedd. (non Steud.), Ama. Sci. Nat. 3(8) : 254. . türkheimii Vatke ex Small, Bull. Torr. Bot. Club 19: 370. 1892. . meisnerianum C. & S. var. jalapense Robins., Proc. Amer. Acad. 3: 324. 1900. 3. P. meisnerianum C. & S. var. usterianum Mattos, Loef- graenia 22: 2. 1965. 4. Polygonum rubricaule Cham., Linnaea 8: 130. 1833. 5. Polygonum stelligerum Cham., Linnaea 8: 131. 1833. P. acanthophyllum Lindau, Botan. Jahrb. 19(48) : 12. 1894. P. meisnerianum C. & S. var. setosum Chod. & Hass. Bull. Herb. Boiss. ser. 2, 3: 392, 1903. In addition to the preceding list there are also combina- tions under the genus T'racaulon. T V Wo ow DEPARTMENT OF BIOLOGY VIRGINIA POLYTECHNIC INSTITUTE BLACKSBURG, VIRGINIA 24061 LITERATURE CITED CHAMISSO, A, DE. 1833. Pflanzen-Missbildungen gesammelt von D. F. L. von Schlechtendal. Polygoneae. Linnaea 8: 150, 131. 188 Rhodora [Vol. 72 CHAMISSO, A. DE. and D. vON SCHLECHTENDAL. 1828. De plantis in expiditione speculatorio Romanzoffianna observatis pergunt Ad. de Chamisso et D. de Schlechtendal. Linnaea 3(3): 40-43. CHAPMAN, A. W. 1889. Flora of the Southern United States. Ive- son, Blakeman, and Co. New York. MEISNER, C. F. 1855. Polygonaceae, Thymelaeaceae, Proteaceae., in Martius’ Flora Brasiliensis, 5(1): 19-20. SMALL, J. K. 1895. A monograph of the North American species of Polygonum. Mem. Dept. Bot., Columbia College, 1, 183 pp. 1933. Manual of the Southeastern Flora. Univ. of North Carolina Press, Chapel Hill, N.C. ERAGROSTIS PLANA IN SOUTH CAROLINA. — Era- grostis plana Nees has been collected twice (Ahles & Hae- sloop 52789 in 1959, NCU; Ahles & Haesloop 53840 in 1960, NCU) on “waste ground around the Santee Wool Combing Mill" at Jamestown, South Carolina. This stout, cespitose perennial, with linear, many-flowered spikelets, and gland- ular-punctate lemma nerves, is reported by Chippendahl [1955, A guide to the identification of grasses in South Africa (part I of The grasses and pastures of South Africa). Cape Times Limited, Parow., C.P., p. 316] to grow in South Africa and “tropical Africa". There are no previous records of its occurrence in North America, The grass apparently is closely related to the weedy annual Eragrostis cilianensis (All) Lutati [E. megastachya (Koel.) Link] which also has spikelets with many florets and glandular-punctate lemma nerves. Of special note on the Jamestown specimens is the occasional occurrence of 5 strong (glandular-punctate) nerves on the lemma. This characteristic was not noted by Chippendahl and is the first variation known to the writers from the remarkably con- sistent 3-nerved condition of the Eragrostis lemma. F. W. GOULD AND R. I. LONARD DEPARTMENT OF RANGE SCIENCE TEXAS A&M UNIVERSITY COLLEGE STATION, TEXAS 77843 AN UNDESCRIBED SPECIES OF ASTRAGALUS (LEGUMINOSAE) FROM UTAH STANLEY L. WELSH An examination of additional materials of Astragalus sec- tion DESPERATI subsection NATURITENSIS from San Juan County, Utah has demonstrated the need for re-evaluation of the specimens passing under the name A. monumentalis Barneby (1953, 1964). Materials included under that epi- thet can be segregated into two entities on the basis of differences in calyx size and shape, petal length, and pod size and shape. The species can be distinguished by the following key. Calyx tube cylindric, 4.8-6.7 mm. long; flowers 11-17 mm. long; pods 20-25 mm. long, conspicuously arcuate. ee A. cottamit. Calyx tube campanulate, 3-3.5 mm. long; flowers 8-9 mm, long; pods 12-21 mm. long, straight to somewhat curved. A. monumentalis. Astragalus cottamii Welsh sp. nov. A. A. monumentale differt calycis tubo longiore cylin- drico, floribus longioribus, et leguminibus longioribus et curvatis magis. Herba perennis radice palari et caudicibus ramificantibus (interdum florens tempa prima) ; caules 0.5-6 cm. longi, ascendentes; stipulae 2-6 mm. longae, membranaceae, am- plectentes sed non connatae; folia 1.2-8 cm, longa, petiolis persistentibus, foliola (5) 9-19 (21), 2-9 mm. longa, 1-4.2 mm. lata, elliptica ad ovalia vel oblanceolata, strigulosa pilis simplicibus, saepe plicata ; pedunculi 0.7-7 em. longi; racemi 3-9 floribus, rhachidibus 0.5-2 cm. longi; bracteolae 2, min- utae vel nullae; tubus calycis cylindricus, 4.8-6.7 mm. longus, strigulosus, purpureo-suffusus, dentes ]1.2-2 mm. longi, subulati; flores 11-17 mm. longi, petala purpureo- rosea, vel vexillum pallens; legumina 20-25 mm. longa, 2.5.4 mm. lata, valde curvata, oblonga ad oblongo-lanceolata, triquetra, sutura ventralis prominens, dorsalis suleata pro- 189 190 Rhodora [Vol. 72 Figure 1. Astragalus cottamii Welsh, habit. funde, bilocularia sutura dorsali ab intrusa, purpureo-mac- ulata, strigulosa. UTAH: SAN JUAN CO., ca. 4 miles east of Clay Hills Divide, S. L. Welsh 5207, 1 May 1966 (Holotype, BRY, Iso- types ISC, NY). Comb Ridge 16 miles west of Blanding, R. C. Barneby 12783, 19 May 1955 (UTC), S. L. Welsh et al 8931, 8 May 1969 (BRY). Natural Bridges National Monu- ment, S. L. Welsh 5187, 5188, 20 April 1966, B. F. Harri- son 5922, 7 May 1933, S. L. Welsh et al 8948, 9 May 1969 (BRY). Comb Wash, S. L. Welsh et al 8932, 8938, 9 May 1969 (BRY). Bluff, W. P. Cottam 2825, 29 April 1935 (BRY). 1970] Astragalus — Welsh 191 i) d | UN m iA CLIMA / cnm. ] erm. Figure 2. Astragalus cottamii Welsh, detail of flower and fruit. The Cottam milk-vetch grows in sandy soil in weathered depressions and crevices on exposed rimrock and ledges of the Cedar Mesa (Permian), Kayenta (Jurassic), and En- tioda (Turrassic) formations, and less commonly on sandy alluvium derived from those formations. It is known to occur from the vicinity west of Clay Hills Divide on the road to Halls Crossing north-eastward to Natural Bridges National Monument and eastward to Bluff. Astragalus monumentalis occurs in similar habitats but is apparently restricted to the Cedar Mesa Formation from Fry Canyon (a tributary of White Canyon) westward along the Cedar Mesa Formation to where that formation dips beneath younger strata west of the Colorado River in Garfield Co., Utah, and northward to the Needies area of Canyonlands National Park. The species is named in honor of Dr. Walter P. Cottam, botanical collector, plant ecologist, and teacher. The altered A. monumentalis represents a more discrete entity with the exclusion of the specimens which form A. coltamii. A description and list of specimens of A. monu- mentalis is provided for comparison. Astragalus monumentalis Barneby Leafl. West. Bot. 7: 35. 1953. Perennial herb with taproot (sometimes flowering the first season) and branching caudex; stems 1-6 cm. long, ascending; stipules 2-4 mm. long, membranous, clasping but not connate around the stem; leaves 1.5-8(11) cm. long, with persistent petioles; leaflets (5) 9-17(21), 2-9 192 Rhodora [Vol. 72 Figure 3. Astragalus monumentalis Barneby, habit. mm. long, 1-4 mm. broad, oval to obovate, elliptic, or oblance- olate, strigulose with simple hairs, often folded; peduncles 1-12 cm. long; racemes 3- to 9- flowered, the rachis 0.5-7 em. long in fruit; bracts longer than the pedicels ; bracteoles lacking; calyx tube campanulate 3-3.5 mm. long, strigulose, 1970] Astragalus — Welsh 193 D cm = Fem. ECPI. Figure 4. Astragalus monumentalis Barneby, detail of flower and fruit. purplish-tinged, the teeth 0.5-1 mm. long, triangular ; flow- ers 8-9 mm. long, the petals pink-purple; pods 12-21 mm. long 2.3-3 mm. broad, straight to curved, triangular in cross- section, the ventral suture prominent, the dorsal deeply sulcate, 2-loculed by intrusion of dorsal suture, greenish or purplish-mottled, strigulose. UTAH: SAN JUAN CO. ; White Canyon, ca. 25 miles south- west of Hite, B. F. Harrison 11595, 18 May 1950 (BRY, Iso- type). Fry Canyon, S. L. Welsh 5191, 29 April 1966 (BRY, + Topotype). Ca. 8 miles west of Fry Canyon, S. L. Welsh 5212, 1 May 1966 (BRY). Canyonlands National Park; Vir- ginia Park, G. Moore 307, 14 May 1965 (BRY), Squaw Flat, G. Moore 348, 348a, 15 May 1965 (BRY), S. L. Welsh et al 2954, 1 June 1964 (BRY), Chesler Park, S. L. Welsh et al 2901, 1 June 1964 (BRY), west side of Elephant Hill, SE. Welsh et al 2888, 31 May 1964 (BRY), vicinity of Silver Stairs, S. L. Welsh 7030, 15 May 1968 (BRY). GARFIELD CO. : west side of Colorado River Bridge near Hite, Welsh 5214, 1 May 1966 (BRY), S. L. Welsh and G. Moore 7122, 28 May 1968 (BRY). DEPARTMENT OF BOTANY BRIGHAM YOUNG UNIVERSITY PROVO, UTAH 84601 LITERATURE CITED BARNEBY, R. C. 1953. Pugillus Astragalorum XV: Four new spe- cies. Leafl. West. Bot. 7: 31-38. 1964. Atlas of North American Astragalus. Mem. N. Y. Bot. Gard. 13: 1-1188. INVESTIGATIONS OF NEW ENGLAND MARINE ALGAE III COMPOSITION, SEASONAL OCCURRENCE AND REPRODUCTIVE PERIODICITY OF THE MARINE RHODOPHYCEAE IN NEW HAMPSHIRE! EDWARD J. HEHRE: & ARTHUR C. MATHIESON There are few published accounts of the marine algae in New Hampshire (Farlow, 1882; Collins, 1900, 1901, 1903, 1906; Croasdale, 1941; Wood and Straughan, 1953; Doty and Newhouse, 1954). Mathieson, Hehre and Reynolds (in press) have reviewed pertinent literature on the distribu- tion and phenology of New England marine algae and have described the seasonal occurrence and vertical distribution of 99 species of marine algae at Jaffrey Point, New Hamp- Shire. Mathieson, Reynolds and Hehre (in press) have re- corded distributional patterns of marine algae in the Great Bay Estuary System, The present paper is the culmination of a three year study of the Rhodophyceae in both coastal and estuarine en- vironments. The purpose of the study is three-fold: (1) to describe the composition of the red algal flora of New Hampshire; (2) to record the seasonal occurrence of the Rhodophyceae of New Hampshire; (3) to determine the reproductive periodicities of as many taxa as possible. Monthly collections were made from 1966-1968 at ten locations along the New Hampshire coast (Fig. 1). Prior to 1966, only irregular collections were made at each sta- tion. Sixty-two stations were established in the Estuary during the summer of 1966, eight of which were monitored approximately monthly from 1966-1968 (Fig. 1). Extreme winter conditions prohibited access during certain months. ‘Published with the approval of the Director of the University of New Hampshire Agriculture Experiment Station as Scientific Con- tribution Number 479. "Present address — Division of Natural Sciences, Southampton Col- lege, Southampton, LI, New York. 194 1970] Marine Algae — Hehre and Mathieson 195 Although monthly collections were not made at the Isles of Shoals, and the collections are somewhat rainimal, all but Duck, White and Seavey Islands are represented. The Isles of Shoals are within New Hampshire and Maine (Fig. 2), but are referred to as a single unit because of their close proximity. The methods of collection and identification of specimens are similar to those of Parts I & Il. The nomenclature of the second revised British Checklist (Parke and Dixon, 1968) has been applied whenever possible. The classifica- tion of the Acrochaetiaceae follows Papenfuss (1947). All collections not otherwise specified have been deposited in the Algal Herbarium at the University of New Hampshire (NHA). The principal individual collectors are designated as follows: E. J. Hehre (H) and A. C. Mathieson (M). The general algal collections, Algae Exsiccatae Americae- Borealis (Farlow, Anderson, and Eaton), and the volume of Phycotheca Boreali-Americana (Collins, Holden and Setchell, 1895-1919) at the Farlow Herbarium (FH) at Harvard University were examined. All previously pub- lished records on the Rhodophyceae of New Hampshire were consulted. Collecting Locations on New Hampshire Coast The New Hampshire coastline (Fig. 1) is approximately 18 miles long. The southern boundary is Seabrook Beach (42952'30" N, 10949' W) ; the boundary to the north is the mouth of the Piscataqua River at the entrance to Ports- mouth Harbour (43904/20" N, 10942/42" W). A brief de- scription of each station is summarized below. Jaffrey Point (Fig. 1, Station 1) a semi-exposed site con- sisting of massive rock outerops and many large eulittoral tidepools. The eulittoral and sublittoral fringe zones are well developed. An artificial breakwater, composed of large granitic blocks, provides excellent substrate. Fort Constitution (Fig. 1, Station 2) is a sheltered loca- tion within the entrance to Portsmouth Harbour. The substrate consists of small rock outcrops, mud and sand. The vegetation is sparse. Zostera marina L. var. stenophylla [Vol. 72 196 Rhodora P Sdn V \ Salmon Falls River ) (Yura Piu I l l Durch a MAIN i River line AM E 5, > šA N N Poris mouth N " o 43* 05 — i I S S a Greet Bey ^ > & ^, 4 \ . k 1 N \ 3 a \ : ` N s : 4 \ Ss H \ n° x 5 \ : Y | š Š \ ee S A EW HAMPSHIRE X Shoals = š: % “a, PI 7 N `` ÓN W 8 ATLANTIC OCEAN M 42 55 — lo Oe < ` ` `. 1 Mile MASS. Ve — "55" 10*45' 70 55 l I E Figure 1 The New Hampshire Coast and Great Bay Estuary System COASTAL STATIONS: 1. Jaffrey Point, 2. Fort Constitution, 8. Odiorne’s Point, 4. North 5 agged Neck, 7. Rye Ledge, 8. Wallis Sands, 5. Concord Point, 6. R Little Boar's Head, 9. Great Boar's Head, 10. Bound Rock. 1970] Marine Algae — Hehre and Mathieson 197 Aschers. et Graebn. grows on the New Hampshire Coast only at Fort Constitution and Rye Ledge. Odiorne’s Point (Fig. 1, Station 3) is a semi-exposed beach with two distinct regions: a cobble area that supports little vegetation, and an area of rock outcrops which has a well developed flora. North Wallis Sands (Fig. 1, Station 4) consists of massive rock outcrops on a sandy beach. Little vegetation is present. Concord Point (Fig. 1, Station 5) is a semi-exposed site with two basic types of substrate: a cobble beach area, which is almost devoid of vegetation, and an area with mas- sive rock outcrops. Except for the plants growing in oc- casional tidepools in the latter region, the vegetation is poor. Ragged Neck (Fig. 1, Station 6) is a cobble beach at Rye Harbour. No tidepools are present and the vegetation is poor. A breakwater delimits the harbour entrance and sup- ports a good flora. Rye Ledge (Fig. 1, Station 7) is an exposed site consisting of massive rock outcrops and an area of large boulders and cobbles. Tidepools are common and a well developed sub- littoral fringe is present. A rich algal flora is evident. Little Boar’s Head (Fig. 1, Station 8) is a semi-exposed cobble beach with scattered boulders and occasional rock outcrops. A few tidepools are present in the low eulittoral zone. The flora is poor. Great Boar's Head (Fig. 1, Station 9) is a cobble beach with numerous large boulders. The flora in the eulittoral zone is poorly developed, but there is a good sublittoral fringe zone. Topographically Great Boar's Head is an ex- posed site, but its eulittoral vegetation, which is conspic- uously dominated by fucoids, is indicative of a sheltered ESTUARY STATIONS: a. Shapleigh Island, b. Pierce Island, c. Piscataqua Rive: toll bridge, d. Hilton Park, e. Cedar Point, f. Durham Point, g. Fox Point, h. Adams Point, i. Crommet Creek, j. Weeks Point, k. Pierce Point, l. Fabyans Point, m. Squamscott River at Route 108, n. Nannie Island. 198 Rhodora [Vol. 72 ISLES OF SHOALS 42°58" A Figure 2 The Isles of Shoals 1970] Marine Algae — Hehre and Mathieson 199 location (Lewis, 1964). The cobble substrate is probably a major factor determining its vegetation. Bound Rock (Fig. 1, Station 10) is a sandy beach with massive rock outcrops and breakwaters. Few species are present because of extreme sand fluctuation. Vast beds of Mytilus and Fucus vesiculosus occur on the rock surfaces. Only a few tidepools are present and they are exposed to constant fluctuations of sand. The breakwaters are dom- inated by annuals (Porphyra umbilicalis and Bangia fus- copurpured). Isles of Shoals The Isles of Shoals (Fig. 2) are located approximately 9 miles SSE of the mouth of the Piscataqua River, and 6.5 miles due east of Straw Point, Rye. The Islands occupy an area three miles north-south by 1.5 miles east-west, and lie within the coordinates 42°58’ N, 70°37’20” W and 43°00’30” N, 70°36’ W. There are nine major islands: five belong to the town of Kittery, Maine (Duck, Appledore, Malaga, Smuttynose, Cedar) and four to Rye, N.H. (Star, White, Seavey, Lung- ing). Nine other rocks and ledges are present in the Island group (Square Rock, Halfway Rocks, Eastern Rocks, Mingo Rock, Shag Rock, Anderson Ledge, Southwest Ledge, Cedar Island Ledge, White Island Ledge). Great Bay Estuary System The Great Bay Estuary System includes the Piscataqua River, Little Bay, Great Bay and seven major tributaries (Fig. 1). It is located approximately within the coordinates 43°03’ N — 43°08’ N, 70°45’ W — 70°55’ W. Many tribu- taries, however, extend beyond these boundaries. A de- tailed description of the Great Bay Estuary Complex is given by Mathieson, Reynolds, and Hehre (in press). Table 1 summarizes the seasonal occurrence and reproduc- tive periodicity of the Rhodophyceae in New Hampshire. About 65% of the taxa found on the open coast are peren- nials (e.g. Ahnfeltia plicata, Petrocelis middendorfii, 200 Rhodora [Vol. 72 Plumaria elegans, Phycodrys rubens, Polysiphonia lanosa, Phyllophora brodiaei, P. membranifolia, Rhodochorton pur- pureum, Ptilota serrata, Cystoclonium purpureum var. cirrhosum, Ceramium rubrum, Corallina officinalis, Halo- saccion ramentaceum, Clathromorphum circumscriptum, Hildenbrandia prototypus, Dermatolithon pustulatum, Mem- branoptera alata, Chondrus crispus, Gigartina stellata, Euthora cristata, Rhodymenia palmata). Three distinct types of annuals (winter, spring and sum- mer) are distinguishable according to their season of maxi- mum growth and development. No fall annuals have been found, although many summer plants persist into the fall and early winter. The fall season appears to be a transition period between summer and winter. Most annuals which occur on the open coast grow during the winter and spring (e.g., Porphyra leucosticta, P. miniata, P. linearis, Bangia fuscopurpurea). Summer annuals, which may persist into the fall, constitute a small part of the Rhodophycean flora of the Great Bay Estuary System (e.g., Chondria baileyana, Dasya pedicellata, Lomentaria baileyana, Polysiphonia de- nudata, P. subtilissima, Ceramium strictum). Only two summer annuals are restricted to the open coast (Kylinia alariae and Nemalion helminthoides). Certain annuals (e.g., Porphyra umbilicalis) occur throughout the year and are precociously reproductive.’ Others (e.g., Dumontia incrassata) occurred throughout the year, but reproductive organs were found only in the spring and summer. Many perennials are reproductive throughout the year (e.g., Ceramium rubrum, Ahnfeltia plicata, Au- douinella membranacea, Chondrus crispus, Euthora cristata, Phycodrys rubens, Phyllophora brodiaei, Polysiphonia lanosa, P. novae-angliae, Halosaccion ramentaceum, Rhody- menia palmata, Gigartina stellata, Hildenbrandia proto- ‘Designates a plant that is reproductive throughout the year and that is usually found reproductive. 1970] Marine Algae — Hehre and Mathieson 201 typus). Others have distinct periods of reproduction which may span one or more seasons (e.g., Clathromorphum cir- cumscriptum, Polyides rotundus, Melobesia lejolisti, Mem- branoptera alata, Phyllophora membranifolia, Cystoclonium purpureum var. cirrhosum, Plumaria elegans, Petrocelis middendorfii, Rhodomela confervoides). Several species have been collected only on one or two occasions e.g., Bonnemaisonia hamifera, Trailliella intri- cata, Bangia ciliaris, Ceramium rubriforme, Erythrotrichia carnea, Pantoneura baerii, Peyssonelia rosenvingii, Lomen- taria orcadensis, Antithamnion plumula, and Callithamnion corymbosum while Gloiosiphonia capillaris, Phyllophora traillii, Antithamnion boreale, A. pylaisaei, and Poly- siphonia flexicaulis were collected once or twice by Dr. Croasdale on the Isles of Shoals during the summer of 1938. Information regarding the seasonal occurrence and repro- ductive periodicity of these 15 taxa is incomplete. ANNOTATED CHECKLIST The following checklist includes 88 taxa of Rhodophyceae from the Great Bay Estuary System and coastal environ- ments of New Hampshire (including the Isles of Shoals). It incorporates all known records of marine red algae from New Hampshire, as well as information on seasonality, reproduction and distribution (vertical and horizontal). Each taxon is cited at least once from every New Hamp- shire station at which it has been found, even though sev- eral collections may have been made. The numbers refer to the collector numbers on the specimens. Fifteen species are new records for the state of New Hampshire (Porphyra leucosticta, P. linearis, Acrochaetium polyides, Kylinia vir- gatula, Bonnemaisonia hamifera, Rhodophysema georgii, Antithamnion americanum, A. plumula, A. pylaisaei, Cal- lithamnion corymbosum, Pantoneura baerii, Chondria baileyana, Polysiphonia denudata, P. flexicaulis, P. sub- tilissima), two of which represent extensions of range on the northeast coast of North America (Acrochaetium poly- ides and Antithamnion plumula). 202 Rhodora Vol. 72 BANGIOIDEAE BANGIALES ERYTHROPELTIDACEAE Erythrotrichia carnea (Dillwyn) J. Agardh Found three times (February, September, November), veg- etative. Epiphytic on Ceramium deslongchampsii var. hooperi, Pylaiella littoralis and Polysiphonia lanosa in the lower eulittoral zone. Found in coastal and estuarine en- vironments; annual. Recorded from the tropics to Nova Scotia, St. James Bay and Baie des Chaleurs (as summarized in Taylor, 1957; Cardinal, 1967). REPRESENTATIVE SPECIMENS: Jaffrey Point, 23 February 1966 (M1637); Shaw’s Hill, Piscataqua River, 21 September 1966 (H967); Cedar Point, 23 November 1966 (H968). Porphyropsis coccinea (J. Agardh ex Areschoug) Rosen- vinge Epiphytie on Desmarestia aculeata, and probably growing in the deep sublittoral zone. The only New England records for this species are the two specimens collected by F. S. Collins ; annual (?). Recorded from New York, New Hampshire, Nova Scotia and Baie des Chaleurs (as summarized in Taylor, 1957; Edelstein and McLachlan, 1968; Cardinal, 1967). REPRESENTATIVE SPECIMEN: Hampton Beach, August 1884 (FH, J. S. Collins). BANGIACEAE Bangia ciliaris Carmichael Found once, vegetative, epiphytic on Euthora cristata in the sublittoral zone on the open coast; annual. Recorded from South Carolina to Maine, Nova Scotia, and Newfoundland (as summarized in Taylor, 1957; Edelstein and McLachlan, 1966b; Mathieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMENS: Jaffrey Point, 10 June 1967 (H225). Bangia fuscopurpurea (Dillwyn) Lyngbye Found from November to August in the mid to upper eulit- toral zone, Common on the open coast, but with restricted distribution in Great Bay Estuary; annual; monospores found from December-April. 1970] Marine Algae — Hehre and Mathieson 203 Recorded from Bermuda to Newfoundland (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Jaffrey Point, 7 January 1967 (H165); Fort Constitution, 29 May 1967 (H169); Odiorne's Point, 8 April 1967 (H170); North Wallis Sands, 20 March 1967 (H172); Concord Point, 5 March 1967 (H176); Ragged Neck, 9 March 1967 (H178); Rye Ledge, 25 May 1967 (H184); Little Boar's Head, 17 February 1967 (H185); Great Boar's Head, 24 January 1967 (H192); Bound Rock, 18 January 1967 (H194); Piscataqua River toll bridge, 29 May 1967 (H202); Hilton Park, 24 May 1967 (H204). *Porphyra leucosticta Thuret in Le Jolis* Found in January, March-July. Epiphytic on Fucus spp., Chondrus crispus, and Polysiphonia elongata in the low eulittoral and sublittoral fringe zones. Occasional on the open coast and rare in the Estuary. An- nual; a< spores found in March and April; 8 spores found March-May. Recorded from Bermuda to Maine and Baie des Chaleurs (as summarized in Taylor, 1957 ; Cardinal, 1967). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 23 July 1958 (FH I. M. Lamb A82) ; Jaffrey Point, 14 April 1968 (M5773) ; Odiorne's Point, 23 May 1968 (M2775); Hilton Park, 7 January 1967 (H2237). *Porphyra linearis Greville Found from December to May on rocks in the mid eulittoral zone on the open coast. Annual; a spores found April and May; B spores found February and March. Recorded from Northern Massachusetts, Nova Scotia, Gaspé to Newfoundland (Lamb and Zimmermann, 1964; Edelstein and McLachlan, 1966b; as summarized in Taylor, 1957 as Porphyra umbilicalis f. linearis). REPRESENTATIVE SPECIMENS: North Wallis Sands, 16 January 1967 (H2239); Concord Point, 19 February 1967 (H2240); Ragged Neck, 4 April 1967 (H2243); Rye Ledge, 10 January 1967 (H2245); Little Boar’s Head, 2 April 1967 (H2247). Porphyra miniata (C. Agardh) C. Agardh Found from March to August on rocks and epiphytic on Chondrus crispus in the upper sublittoral zone. Common on the open coast but with limited distribution in. the Estu- **New record for New Hampshire. 204 Rhodora [Vol. 72 ary. Annual; a spores found April to August; B spores found March to August. Recorded from Northern Massachusetts to the arctic (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 28 May 1968 (M5813); Jaffrey Point, 19 May 1964 (H2248); Fort Constitu- tion, 23 June 1967 (H2251) ; Odiorne's Point, 30 May 1967 (H2253) ; North Wallis Sands, 20 May 1967 (H2257); Concord Point, 27 June 1967 (H2260); Ragged Neck, 14 July 1967 (M5826); Rye Ledge, 25 May 1967 (H2261); Great Boar's Head, 18 April 1966 (M5828) ; Piscataqua River toll bridge, 29 May 1967 (H2263); Hilton Park, 23 June 1967 (H2264); Cedar Point, 24 June 1967 (H2265). Porphyra umbilicalis (Linnaeus) J. Agardh Found throughout the year on rocks in the mid to lower eulittoral zone. Common on the open coast and throughout the Estuary. Annual; a and 8 spores found January-De- cember. Recorded from New Jersey to Newfoundland (as summar- ized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Lunging Island), 22 July 1966 (M5861); Jaffrey Point, 7 January 1967 (H2268); Fort Constitution, 20 September 1966 (H2274); Odiorne's Point, 21 July 1966 (M5869); North Wallis Sands, 28 February 1967 (H22'15) ; Concord Point, 27 June 1967 (H2278); Ragged Neck, 9 March 1967 (H2280); Rye Ledge, 25 May 1967 (H2288); Little Boar's Head, 24 May 1967 (H2292); Great Boar's Head, 24 January 1967 (H2294) ; Bound Rock, 20 February 1967 (H2302) ; Piscataqua River toll bridge, 20 January 1967 (H2337); Hilton Park, 9 December 1966 (H2326); Cedar Point, 7 August 1967 (H2318); Adams Point, 22 December 1966 H2314); Weeks Point, 23 April 1966 (7245908). Porphyra umbilicalis (Linnaeus) J. Agardh forma epiphytica Collins Epiphytic on various algae (e.g. Fucus spp. and Poly- siphonia lanosa) throughout the year in the lower eulittoral and upper sublittoral zones. Common on the open coast and throughout the Estuary. Annual; a spores found Jan- uary-November; 8 spores found January-December. Recorded from Massachusetts to Maine (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 22 July 1966 (M5939); Jaffrey Point, 1 April 1967 (H2350); Fort Constitu- tion, 22 June 1966 (M5944); Odiorne’s Point, 30 May 1967 (\H2855) ; 1970] Marine Algae — Hehre and Mathieson 205 North Wallis Sands, 11 June 1967 (H2359); Concord Point, 10 August 1967 (H2363); Ragged Neck, 12 June 1967 (112367); Rye Ledge, 25 May 1967 (H2371); Little Boar’s Head, 2 April 1967 (H2372); Great Boar’s Head, 25 June 1967 (H2377); Bound Rock, 18 January 1967 (H2383); Piscataqua River toll bridge, 12 August 1967 (H2395); Hilton Park, 27 April 1967 (H2391); Cedar Point, 23 November 1966 (H2398); Durham Point, 15 November 1966 (H2396); Adams Point, 27 June 1967 (H2400). FLORIDEAE NEMALIONALES ACROCHAETIACEAE Acrochaetium polyides (Rosenvinge) Børgesen’ Found once as an endophyte in the peripheral cells of Poly- ides rotundus on the open coast. This is only the second time that the species has been recorded in North America. No reproductive structures seen; annual (7). Previously recorded from Nova Scotia (Edelstein and Mc- Lachlan, 1966a). REPRESENTATIVE SPECIMEN: North Wallis Sands, 10 September 1966 (HD). Audouinella membranaeea (Magnus) Papenfuss Endozoic in the lorica of sertularians in the mid to lower eulittoral zone. Common on the open coast, but with lim- ited distribution in the Estuary. Perennial (?) ; tetraspores found January-December. Recorded from Connecticut to Maine, Nova Scotia and Greenland (as summarized in Taylor, 1957; Ecelstein and McLachlan, 1966b; Lund, 1959). REPRESENTATIVE SPECIMENS: Jaffrey Point, 7 January 1967 (H76); Fort Constitution, 20 January 1967 (H'8); Odiorne's Point, 21 February 1967 (H83); North Wallis Sands, 20 March 1967 (H87); Concord Point, 8 January 1967 (H93); Ragged Neck, 26 December 1966 (H106); Rye Ledge, 25 April 1967 (H107); Little Boar’s Head, 23 January 1967 (H113); Great Boar’s Head, 24 January 1967 (H122); Bound Rock, 6 December 1966 (H129); Piscataqua River toll bridge, 22 June 1967 (H164); Hilton Park, 24 May 1967 (H153); Cedar Point, 3 March 1967 (H146); Durham Point, 26 June 1967 (H149); Adams Point, 17 April 1967 (H142); Weeks Pcint, 9 August 1967 (M3788). Kylinia alariae (Jónsson) Kylin Specific epiphyte on Alaria esculenta and restricted to ex- ***Range extension for the northeast coast of North America. 206 Rhodora [Vol. 72 posed coastal stations where A. esculenta occurs. Found in the sublittoral fringe zone in July and August. Annual; monospores found in July. Recorded from Northern Massachusetts to Maine and New- foundland (as summarized in Taylor, 1957; Mathieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMENS: Isles of Shoals (Lunging Island), 22 July 1966 (M4963); Rye Ledge, 18 August 1966 (M4964). Kylinia secundata (Lyngbye) Papenfuss Epiphytie on Ruppia maritima, various algae (e.g., Por- phyra umbilicalis) and epizoic on sertularians, Common on the open coast and with limited distribution in the Estu- ary. Perennial (?) ; monospores found January-December ; tetraspores rare, found once in March. Recorded from Connecticut to Maine, Nova Scotia and New- foundland (as summarized in Taylor, 1957; Edelstein and McLachlan, 1966b; Mathieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMENS: Jaffrey Point, 12 October 1966 (H1369) ; Fort Constitution, 29 May 1967 (H135); Odiorne’s Point, 21 January 1967 (H137); North Wallis Sands, 21 December 1966 (H138) ; Con- cord Point, 10 December, 1966 (H139); Ragged Neck, 18 July 1966 (M4966); Rye Ledge, 29 March 1967 (M4967); Little Boar’s Head, 25 November 1966 (H1372); Bound Rock, 7 January 1967 (H1373); Piscataqua River toll bridge, 12 August 1967 (H160); Hilton Park, 24 May 1967 (H153); Cedar Point, 21 December 1966 (H145); Durham Point, 19 September 1966 (H1374); Adams Point, 22 De- cember 1966 (H141). *Kylinia virgatula (Harvey) Papenfuss Epiphytic on specimens of Rhodymenia palmata. Found in September, October and December on the open coast. An- nual; monospores found in September, October and Decem- ber. Previously recorded from North Carolina to Southern Mas- sachusetts, Nova Scotia and Newfoundland (as summarized in Taylor, 1957; Edelstein, McLachlan and Craigie, 1967; Mathieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMENS: Rye Ledge, 15 October 1966 (H1375); Little Boar’s Head, 14 September 1966 (H1376); Great Boar’s Head, 14 October 1966 (H1377). Kylinia virgatula (Harvey) Papenfuss forma luxurians (J. Agardh) Collins 1970] Marine Algae — Hehre and Mathieson 207 Found four times (in July and September). Epiphytic on the margins of Zostera leaves. Annual; monospores found in July and September. Recorded from Connecticut to Maine and Baie des Chaleurs (as summarized in Taylor, 1957; Cardinal, 1967). REPRESENTATIVE SPECIMENS: Fort Constitution, 20 September 1966 (H1379) ; Shaw’s Hill, Piscataqua River, 21 September 1966 (H1380) ; Shapleigh Island, Piscataqua River, 21 September 1966 (H1381); Fox Point, 21 July 1966 (M4977). Rhodochorton purpureum (Lightfoot) Rosenvinge Growing in the mid and lower eulittoral zone or. rock faces covered by fucoids. Common throughout the year on the open coast, but with limited distribution in the Estuary. Perennial; tetraspores found in February, Recorded from Rhode Island to Ellesmere Island (as sum- marized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Smuttynose Island), 22 July 1966 (M6004); Jaffrey Point, 12 October 1966 (H2458) ; Fort Constitution, 28 April 1967 (46008); Odiorne’s Point, 21 January 1967 (H2469) ; North Wallis Sands, 20 March 1967 (H2479); Concord Point, 5 March 1967 (H2490); Ragged Neck, 4 April 1967 (H2491); Rye Ledge, 23 December 1966 (H2502); Little Boar’s Head, 14 August 1967 (H2509); Great Boar’s Head, 24 January 1967 (H2520); Bound Rock, 16 April 1967 (H2526); Hilton Park, 9 July 1967 (H2528). BATRACHOSPERMACEAE Batrachospermum sp. Collected at the headtide of the Oyster River (Dr. Lewis Flint, personal communication). Lemanea fucina Bory Found in rapidly flowing waters at the headtide of the Oy- ster River (Wood and Straughan, 1953). No specimens were seen from this station. L. fucina is found on the University of New Hampshire campus, where the Oyster River is en- tirely fresh water. There are specimens in the University of New Hampshire Algal Herbarium collected by Dr. A. R. Hodgdon from an estuarine habitat in Maine. REPRESENTATIVE SPECIMEN: Newcastle, Maine, 29 June 1964 (A. R. Hodgdon 14090). HELMINTHOCLADIACEAE Nemalion helminthoides (Velley in Withering) Batters 208 Rhodora [Vol. 72 Found in July and August growing on rocks and mussels (Mytilus edulis) in the extreme lower eulittoral zone. Re- stricted to the open coast. Annual; cystocarpic plants found in August. Recorded from Long Island to Nova Scotia (as summarized in Taylor, 1957 as Nemalion multifidum). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 22 July 1966 (M5120); Ragged Neck, 10 August 1967 (H1598); Rye Ledge, 12 August 1967 (M5121); Bound Rock, 18 July 1966 (H1599). BONNEMAISONIACEAE *Bonnemaisonia hamifera Hariot Collected three times (April, June and July) in drift. Veg- etative; probably growing in the deep sublittoral zone, An- nual (Chihara, 1961). Previously recorded from Long Island to Southern Massa- chusetts and Nova Scotia (as summarized in Taylor, 1957 as Asparagopsis hamifera; Edelstein and McLachlan, 1968). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), June 1965 (Kingsbury, 1965); Isles of Shoals (Cedar Island), 22 July 1966 (H75); Isles of Shoals (Smuttynose Island), 16 June 1966 (M3765). Trailliella intricata (J. Agardh) Batters Found once, vegetative, in the low eulittoral zone on the open coast. It is believed to be the tetrasporophyte of Bon- nemaisonia hamifera (Chihara, 1961, 1962). Recorded from Long Island to Newfoundland (as summar- ized in Taylor, 1957). REPRESENTATIVE SPECIMEN: Isles of Shoals (Star Island), 22 July 1966 (M6076). CRYPTONEMIALES SQUAMARIACEAE Peyssonelia rosenvingii Schmitz in Rosenvinge Found twice (January and April), vegetative, growing on rocks in the lower eulittoral and sublittoral zones on the open coast. Perennial. Recorded from Northern Massachusetts to Maine, Nova Scotia, Baie des Chaleurs, Baie de Gaspé and Greenland (as summarized in Taylor, 1957; Edelstein, McLachlan and Craigie, 1967; Cardinal, 1967; Lund, 1959). 1970] Marine Algae — Hehre and Mathieson 209 REPRESENTATIVE SPECIMENS: Jaffrey Point, 22 January 1966 (M5130) ; Jaffrey Point, 20 April 1967 (M5129). *Rhodophysema georgii Batters Found throughout most of the year as an epiphyte on Zos- tera marina, Perennial; tetraspores found in February and November. Recorded from Long Island to Maine, Nova Scotia and New- foundland (as summarized in Taylor, 1957, as Rhododermis georgii; Edelstein and McLachlan, 1966b; Mathieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMENS: Fort Constitution, 29 May 1967 (H2530) ; Rye Ledge, 11 November 1966 (41056); Cedar Point, 29 May 1967 (H2534); Adams Point, 17 April 1967 (H2532). HILDENBRANDIACEAE Hildenbrandia prototypus Nardo Growing on rocks throughout the year in the eulittoral and upper sublittoral zones. Common on the open coast and throughout the Estuary. Perennial; tetraspores found Jan- uary-December. Recorded from Florida to Baffin Island (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Lunging Island), 22 July 1966 (M4882); Jaffrey Point, 8 November 1966 (H1223); Fort Con- stitution, 23 June 1967 (H1239); Odiorne's Point, 17 October 1966 (H1240); North Wallis Sands, 11 June 1967 (H1256); Concord Point, 13 September 1966 (H1260); Ragged Neck, 9 March 1967 (H1271); Rye Ledge, 25 May 1967 (H1287); Little Boar’s Head, 23 January 1967 (H1297); Great Boar's Head, 24 April 1967 (H1806) ; Bound Rock, 6 December 1966 (H1314); Piscataqua River toll bridge, 27 February 1967 (H1361); Hilton Park, 24 May 1967 (H1353); Cedar Point, 17 April 1967 (H1325); Durham Point, 31 May 1967 (H1336); Adams Point, 17 April 1967 (H1319); Fabyan’s Point, 15 October 1966 (H1339); Pierce Point, 23 June 1967 (H1362). CORALLINACEAE Clathromorphum circumscriptum (Stromfelt) Foslie Found throughout the year in tidepools in the mid and lower eulittoral zone, and extending into the sublittoral zone. Common on the open coast and rare in the Estuary. Peren- nial; bispores found from December-May. Recorded from Northern Massachusetts to Ellesmere Island 210 Rhodora [Vol. 72 (as summarized in Taylor, 1957, as Phymatolithon compac- tum; Adey, 1965). REPRESENTATIVE SPECIMENS: Isles of Shoals (Lunging Island), 22 July 1966 (M4312); Jaffrey Point, 8 November 1966 (H586); Fort Constitution, 21 March 1967 (H594); Odiorne's Point, 26 November 1966 (H604); North Wallis Sands, 11 October 1966 (H619) ; Concord Point, 13 September 1966 (H629); Ragged Neck, 19 February 1967 (H641); Rye Ledge, 11 November 1966 (H667); Little Boar's Head, 2 April 1967 (H649); Great Boar’s Head, 24 January 1967 (H658); Hilton Park, 24 May 1967 (H617). Corallina officinalis Linnaeus Growing from the sublittoral zone to the mid eulittoral zone on the open coast. Perennial. Recorded from Bermuda to Newfoundland (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 22 July 1966 (M4363); Jaffrey Point, 7 January 1967 (H679) ; Fort Constitu- tion, 4 December 1966 (H684) ; Odiorne's Point, 26 June 1967 (H692); North Wallis Sands, 11 October 1966 (H705); Concord Point, 10 August 1967 (H713); Ragged Neck, 12 June 1967 (H725): Rye Ledge, 11 November 1966 (H741); Little Boar’s Head, 13 October 1966 (H751); Great Boar’s Head, 25 June 1967 (H759); Bound Rock, 3 September 1967 (H757). Dermatolithon pustulatum (Lamouroux) Foslie Found throughout the year as an epiphyte on various algae (e.g., Chondrus crispus, Gigartina stellata, Fucus Spp.) in the lower eulittoral and sublittoral fringe zones of the open coast. Perennial; bispores found in all months except April. Recorded from Rhode Island to Massachusetts, New Hamp- shire and Nova Scotia (as summarized in Taylor, 1957; Mathieson, Hehre and Reynolds, in press; Edelstein, Mc- Lachlan and Craigie, 1967, all as Lithophyllum macrocar- pum; Edelstein and McLachlan, 1968). REPRESENTATIVE SPECIMENS: Isles of Shoals (Lunging Island), 22 July 1966 (M5006); Jaffrey Point, 1 April 1967 (H1438); Fort Con- stitution, 27 February 1967 (H1442) ; Odiorne's Point, 10 March 1967 (H1453); North Wallis Sands, 16 January 1967 (H1463); Concord Point, 22 November 1966 (H1474); Ragged Neck, 12 June 1967 (H1483); Rye Ledge, 25 April 1967 (H1494); Little Boar's Head, 13 October 1966 (H1497); Great Boar's Head, 24 April 1967 (H1504); Bound Rock, 6 December 1966 (H1518); Hilton Park, 17 February 1968 (M5039). 1970] Marine Algae — Hehre and Mathieson 211 Lithophyllum corallinae (Crouan frat.) Heydrich Specific epiphyte on Corallina officinalis on the open coast. Perennial; bispores found in March, June and July. Recorded from Rhode Island to Maine (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Cedar Island), 22 July 1966 (H1379); Jaffrey Point, 7 January 1967 (H1384); Fort Con- stitution, 22 June 1966 (M4993); Odiorne's Point, 10 March 1967 (H1393); North Wallis Sands, 20 May 1967 (H1397); Concord Point, 5 March 1967 (H1404); Ragged Neck, 9 March 1967 (H1410); Rye Ledge, 11 November 1966 (H1417); Little Boar's Head, 8 March 1967 (H1426); Great Boar's Head, 20 March 1967 (H1432). Lithothamnium glaciale Kjellman Growing in the lower eulittoral and sublittoral zones on the open coast. Perennial; bispores found in February and April. Recorded from Massachusetts to Ellesmere Island (as sum- marized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Jaffrey Point, 1 April 1967 (H1550); Fort Constitution, 21 March 1967 (H1536); Odiorne's Point, 10 March 1967 (H1542); North Wallis Sands, 20 March 1967 (H1538) ; Concord Point, 27 June 1967 (H1524); Rye Ledge, 22 June 1967 (H1519); Little Boar's Head, 27 July 1967 (H1528); Great Boar's Head, 24 April 1967 (H1522); Bound Rock, 18 November 1966 (H1523). Melobesia lejolisii Rosanoff Found throughout the year as an epiphyte on Zostera and Phyllophora spp. in the sublittoral zone. Common in drift on the open coast and throughout the Estuary. Perennial; tetraspores found October-July. Recorded from Florida to the arctic (as summarized in Taylor, 1957, as Fosliella lejolisit). REPRESENTATIVE SPECIMENS: Jaffrey Point, 23 February 1966 (M4710); Fort Constitution, 21 March 1967 (H1040); Odiorne’s Point, 8 April 1967 (H1044); North Wallis Sands, 17 April 1967 (H1051); Concord Point, 27 June 1967 (H1052); Ragged Neck, 9 March 1967 (H1054); Rye Ledge, 15 October 1966 (111057) ; Little Boar’s Head, 25 November 1966 (H1058); Hilton Park, 3 May 1968 (M4712); Durham Point, 16 October 1966 (H1059); Pierce Point, 8 November 1966 (H1060). Phymatolithon lenormandi (Areschoug) Adey Growing on rocks and shells throughout most of the year 212 Rhodora [Vol. 72 in the sublittoral zone on the open coast, but with restricted distribution in the Estuary. No reproductive structures found; perennial. Recorded from New Jersey to the arctic (as summarized in Taylor, 1957, as Lithothamnium lenormandi; Adey, 1966). REPRESENTATIVE SPECIMENS: Isles of Shoals (Smuttynose Island), 22 July 1966 (M5312); Jaffrey Point, 6 May 1967 (M5311); Fort Constitution, 28 April 1968 (M5813); Odiorne’s Point, 17 October 1966 (H1890); North Wallis Sands, 16 February 1967 (M5815); Concord Point, 9 October 1966 (H1891); Ragged Neck, 8 October 1966 (H1892); Rye Ledge, 15 October 1966 (H1895); Little Boar's Head, 13 October 1966 (H1896); Great Boar's Head, 14 October 1966 (H1898); Hilton Park, 14 October 1966 (H1899). DUMONTIACEAE Dumontia incrassata (O. F. Müller) Lamouroux Common in the mid to lower eulittoral zone throughout the year on the open coast, and with limited distribution in the Estuary. Often found in tidepools; annual, tetraspores found March-August. Recorded from Long Island to Nova Scotia and James Bay (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Lunging Island), 22 July 1966 (M4604); Jaffrey Point, 7 January 1967 (H939); Fort Constitution, 20 January 1967 (H948); Odiorne's Point, 8 April 1967 (H954) ; North Wallis Sands, 20 March 1967 (H960) ; Concord Point, 5 March 1967 (H904); Ragged Neck, 22 May 1967 (H910); Rye Ledge, 25 May 1967 (H913); Little Boar's Head, 23 January 1967 (H920); Great Boar's Head, 24 April 1967 (H935); Bound Rock, 6 December 1966 (H938); Piscataqua River toll bridge, 4 December 1966 (H881); Hilton Park, 14 October 1966 (H870); Cedar Point, 3 March 1967 (H892); Fox Point, 17 November 1966 (H896); Nannie Island, 7 July 1966 (M4578). POLYIDEACEAE Polyides rotundus (Hudson) Greville Growing in the sublittoral zone throughout the year on the open coast and with limited distribution in the Estuary. Perennial; tetraspores found from October to January; cystocarpic nemathecia found from October to February. Recorded from Long Island to the Hudson Strait (as sum- marized in Taylor, 1957, as Polyides caprinus). 1970] Marine Algae — Hehre and Mathieson 213 REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 28 May 1968 (M5363); Jaffrey Point, 7 January 1967 (H1915); Fort Con- stitution, 24 July 1967 (M5378); Odiorne’s Point, 26 July 1967 (H1917); North Wallis Sands, 11 October 1966 (H1920); Concord Point, 8 January 1967 (H1925); Ragged Neck, 9 March 1967 (H1927); Rye Ledge, 15 October 1966 (H1928); Little Boar's Head, 24 May 1967 (H1931); Great Boar's Head, 13 November 1966 (H1938); Bound Rock, 18 January 1967 (M1945); Piscataqua River toll bridge, 27 February 1967 (H1948); Hilton Park, 27 April 1967 (H1947). GLOIOSIPHONIACEAE Gloiosiphonia capillaris (Hudson) Carmichael ex Berkeley Found once by Croasdale (July) growing on the open coast. Annual; cystocarpic in July. Recorded from Connecticut to Newfoundland (as summar- ized in Taylor, 1957). REPRESENTATIVE SPECIMEN: Isles of Shoals (Appledore Island), 25 July 1938 (Croasdale 4794). KALLYMENIACEAE Euthora cristata (C. Agardh) J. Agardh Growing in the sublittoral zone on the open coast throughout the year. Perennial; tetrasporic and cystocarpic plants found from January to December. Recorded from New Jersey to Ellesmere Island (as sum- marized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Lunging Island), 22 July 1966 (M4660) ; Jaffrey Point, 1 April 1967 (H969); Fort Con- stitution, 27 February 1967 (H974); Odiorne's Point, 30 May 1967 (H977); North Wallis Sands, 20 May 1967 (H985); Concord Point, 10 December 1966 (H992); Ragged Neck, 9 March 1957 (H1004); Rye Ledge, 10 January 1967 (H1016); Little Boar's Head, 24 May 1967 (H1023); Great Boar's Head, 25 June 1967 (H1028); Bound Rock, 6 December 1966 (H1029). CHOREOCOLACACEAE Choreocolax polysiphoniae Reinsch Parasitic on the fronds of Polysiphonia lanosa. Restricted to the open coast. Present throughout the year and peren- nial; tetraspores found from January to December; cysto- carpic plants found in July, August and November. Recorded from Connecticut to Nova Scotia and Newfound- 214 Rhodora [Vol. 72 land (as summarized in Taylor, 1957; Mathieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMENS: Jaffrey Point, 10 June 1967 (H551); Fort Constitution, 23 June 1967 (H554); Odiorne’s Point, 11 August 1967 (H561); North Wallis Sands, 17 April 1967 (H562); Concord Point, 21 May 1967 (H567); Ragged Neck, 22 May 1967 (H571); Rye Ledge, 25 May 1967 (H575); Little Boar’s Head, 14 August 1967 (H580); Great Boar’s Head, 24 April 1967 (H584). GIGARTINALES CRUORIACEAE Petrocelis middendorfii (Ruprecht) Kjellman Found throughout the year on rocks in the lower eulittoral zone. Common on the open coast and with restricted dis- tribution in the Estuary. Perennial; tetraspores found from October to June. Recorded from Long Island to Maine, Nova Scotia and the Arctic (as summarized in Taylor, 1957; Edelstein, McLach- lan and Craigie, 1967). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 22 July 1966 (M5135); Jaffrey Point, 5 December 1966 (H1604); Fort Constitution, 21 March 1967 (H1607); Odiorne’s Point, 30 May 1967 (H1617); North Wallis Sands, 11 June 1967 (H1626); Concord Point, 3 April 1967 (H1634); Ragged Neck, 20 July 1967 (H1640); Rye Ledge, 24 July 1967 (H1649); Little Boar's Head, 24 May 1967 (H1663); Great Boar's Head, 25 June 1967 (H1670); Hilton Park, 24 May 1967 (H1665). SOLIERIACEAE Agardhiella tenera (J. Agardh) Schmitz Reported by Taylor (1957) from N.H. However, none of the herbarium specimens nor references cited by Taylor in- dicate that it is found north of Gloucester, Massachusetts. Recorded from the tropics to North Carolina, north to Mas- sachusetts and New Hampshire (as summarized in Taylor, 1957). RHODOPHYLLIDACEAE Cystoclonium purpureum (Hudson) Batters var. cirrhosum Harvey Found throughout the year growing on rocks and occa- sionally on other algae in the lower eulittoral and sublittoral 1970] Marine Algae — Hehre and Mathieson 215 zones. Common on the open coast, and with restricted dis- tribution in the Estuary. Perennial; tetraspores found from April to November. Recorded from New Jersey to New Hampshire, Baie des Chaleurs and Baie de Gaspe (as summarized in Taylor, 1957; Lamb and Zimmermann, 1964; Mathieson, Hehre and Reynolds, in press; Cardinal, 1967). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 22 July 1966 (M4441); Jaffrey Point, 7 January 1967 (H772); Fort Consti- tution, 20 September 1966 (H780); Odiorne’s Point, 11 August 1967 (H781); North Wallis Sands, 20 May 1967 (H784); Concord Point, 13 September 1966 (H792); Ragged Neck, 22 May 1967 (H799); Rye Ledge, 15 October 1966 (H809); Little Boar’s Head, 24 May 1967 (H820); Great Boar's Head, 15 September 1966 (11822) ; Bound Rock, 18 November 1966 (1H829); Piscataqua River toll bridge, 27 February 1967 (H858); Hilton Park, 27 April 1967 (H&47); Durham Point, 15 November 1966 (H842); Adams Point, 22 December 1966 (H840). Rhodophyllis dichotoma (Lepeschkin) Gobi Found January-February and May-July on the open coast. The plant has been found only 4 times in drift. It was col- lected once by SCUBA in the deep sublittoral-epiphytic on Ptilota serrata. Perennial; tetraspores found in January and July; cystocarpic plants found in February, May and June. l Recorded from Northern Massachusetts to Ellesmere Island (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Appledore Island), 28 May 1968 (M6138); Hampton Beach, 27 February 1966 (H2559); Bound Rock, 24 June 1967 (H2560). GRACILARIACEAE Gracilaria foliifera (Forsskål) Børgesen Found from March to December either free-floating or at- tached to small stones in the lower eulittoral and upper sublittoral zones throughout the Estuary. Perennial; tetra- spores found from June-October; cystocarpic plants found in April and from June to November. Recorded from the tropics to New Hampshire (as summar- ized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Hilton Park, 24 May 1967 (H1205); Cedar Point, 7 August 1967 (H1193); Durham Point, 5 July 1966 216 Rhodora [Vol. 72 (M4813); Adams Point, 10 September 1966 (H1192); Fabyan’s 4813); 23 June 1967 (H1200); Pierce Point, 23 June 1967 (H1207). PHYLLOPHORACEAE Ahnfeltia plicata (Hudson) Fries Growing throughout the year in the lower eulittoral and sublittoral zones on the open coast. Restricted distribution in the Estuary. Perennial; monospores found January-De- cember. Recorded from New Jersey to Devon Island (as summar- ized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 22 July 1966 (M3669); Jaffrey Point, 27 May 1967 (H2); Fort Constitution, 24 July 1967 (M3683); Odiorne’s Pcint, 21 February 1967 (H6); North Wallis Sands, 28 February 1967 (H14); Concord Point, 8 January 1967 (H23); Ragged Neck, 9 March 1967 (H25); Rye Ledge, 11 November 1966 (H29); Little Boar’s Head, 17 February 1967 (H38); Great Boar's Head, 24 April 1967 (H45); Bound Rock, 17 March 1968 (H48); Hilton Park, 9 July 1967 (H60); Durham Point, 5 July 1966 (M3705); Adams Point, 19 July 1966 (M3704) ; Weeks Point, 14 September 1966 (H63). Ceratocolax hartzii Rosenvinge Specific parasite on Phyllophora brodiaei in the sublittoral zone on the open coast. Perennial; tetraspores found in January, February, Avril and May. Recorded from Rhode Island to the Arctic (Newroth, 1968). REPRESENTATIVE SPECIMENS: Isles of Shoals (Cedar Island), 22 July 1966 (H369); Jaffrey Point, 8 November 1966 (H370); Odiorne’s Point, 10 March 1967 (H371); North Wallis Sands, 21 December 1966 (H372); Ragged Neck, 4 April 1967 (H374); Little Boar's Head. 19 February 1966 (M4112); Bound Rock, 18 January 1967 (H375). Phyllophora brodiaei (Turner) Endlich Found throughout the year growing in the sublittoral zone on the open coast. Common in drift, Perennial; carpotetra- sporangial nemathecia found from January to December. Recorded from New Jersey to Newfoundland (as summar- ized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Lunging Island), 22 July 1966 (M5253); Jaffrey Point, 8 February 1967 (H1780); Fort Constitution, 24 July 1967 (M5265); Odiorne’s Point, 10 September 1966 (H1789); North Wallis Sands, 16 January 1967 (H1795); Concord Point, 3 April 1967 (H1804); Ragged Neck, 23 November 1970] Marine Algae — Hehre and Mathieson ZH 1966 (H1805); Rye Ledge, 25 November 1966 (H1814); Little Boar's Head, 8 March 1967 (H1818); Great Boar’s Head, 13 November 1966 (H1819); Bound Rock, 6 December 1966 (H1824). Phyllophora membranifolia (Goodenough et Woodward) J. Agardh Growing throughout the year in the lower eulittoral and sublittoral zones on the open coast, and with limited distri- bution in the Estuary. Perennial; tetraspores found from September to March ; cystocarpic plants found in January, March, April and November. Recorded from New Jersey to Baffin Island (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 22 July 1966 (M5278); Jaffrey Point, 11 July 1967 (H1827); Fort Consti- tution, 20 September 1966 (H1832) ; Odiorne’s Point, 23 June 1967 (H1836); North Wallis Sands, 20 May 1967 (H1844) ; Concord Point, 9 October 1966 (1854); Ragged Neck, 10 August 1967 (H1861); Rye Ledge, 11 November 1966 (H1864); Little Boar’s Head, 14 August 1967 (H1875); Great Boar’s Head, 24 Janvary 1957 (H1881); Bound Rock, 18 November 1966 (H1886); Hilton Park, 2 October 1968 (N. B. Reynolds 286). Phyllophora traillii Holmes ex Batters Found once by Croasdale in July growing on rocks in the low eulittoral zone on the open coast. Vegetative ; perennial. Recorded from Connecticut to Maine and Nova Scotia (as summarized in Taylor, 1957; Edelstein and McLachlan, 1968). REPRESENTATIVE SPECIMEN: Isles of Shoals (Appledore Island), 28 July 1938 (Croasdale 5297). GIGARTINACEAE Chondrus crispus Stackhouse Common throughout the year on rocks and in tidepools in the mid to lower eulittoral zone, and extending into the sublittoral zone. Found on the open coast and throughout most of the Estuary. Perennial; tetraspores found Janu- ary-December; carpospores found J anuary-May, July, Sep- tember-November. Recorded from New Jersey to Newfoundland (as summar- ized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 22 July 1966 (M4180); Jaffrey Point, 5 December 1966 (H392) ; Fort Consti- 218 Rhodora [Vol. 72 tution, 20 January 1967 (H397); Odiorne’s Point, 30 May 1967 (H412); North Wallis Sands, (16 January 1967 (H418); Concord Point, 9 October 1966 (H436); Ragged Neck, 12 June 1967 (H445); Rye Ledge, 11 November 1966 (H451); Little Boar’s Head, 8 March 1967 (H463); Great Boar’s Head, 24 April 1967 (H474); Bound Rock, 6 December 1966 (H491); Piscataqua River toll bridge, 27 February 1967 (1H535) ; Hilton Park, 14 October 1966 (H519) ; Cedar Point, 15 January 1967 (H499) ; Durham Point, 31 May 1967 (H509); Adams Point, 30 May 1967 (H493); Weeks Point, 14 September 1966 (H539). Gigartina stellata (Stackhouse) Batters Found throughout the year in the lower eulittoral and sub- littoral fringe zones on the open coast, and with limited distribution in the Estuary. Perennial; cystocarpic plants found January-December. Recorded from Rhode Island to Newfoundland (as summar- ized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 22 July 1966 (M4741) ; Jaffrey Point, 5 December 1966 (H1062) ; Fort Consti- tution, 8 November 1966 (H1077); Odiorne’s Point, 26 June 1967 (H1089); North Wallis Sands, 11 June 1967 (H1099) ; Concord Point, 19 February 1967 (H1108); Ragged Neck, 26 December 1966 (H1119); Rye Ledge, 22 June 1967 (H1129); Little Boar's Head, 14 September 1966 (H1141); Great Boar's Head, 14 October 1966 (H1152); Bound Rock, 8 August 1967 (M4774); Piscataqua River toll bridge, 27 February 1967 (H1184); Hilton Park, 24 May 1967 (H1162); Cedar Point, 3 March 1967 (H1170); Durham Point, 19 September 1966 (H1171). RHODYMENIALES RHODYMENIACEAE Halosaccion ramentaceum (Linnaeus) J. Agardh Growing throughout the year in the sublittoral fringe re- gion. Only found at three coastal stations. Perennial; tetra- spores found December-October. Recorded from Northern Massachusetts to Ellesmere Island (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Appledore Island), 25 July 1938 (Croasdale 4867) ; Jaffrey Point, 12 October 1966 (H1218) ; Rye Ledge, 26 February 1967 (H1221). 1970] Marine Algae — Hehre and Mathieson 219 Rhodymenia palmata (Linnaeus) Greville Growing throughout the year in the lower eulittoral and sublittoral zones. Common on the open coast and the Estu- ary. Perennial; tetraspores found from January to Decem- ber. Recorded from New Jersey to Ellesmere Island (as sum- marized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 28 May 1968 (M6176); Jaffrey Point, 7 January 1967 (H2565); Fort Consti- tution, 20 September 1966 (H2566) ; Odiorne’s Point, 21 February 1967 (H2575); North Wallis Sands, 20 March 1967 (H2584); Concord Point, 8 January 1967 (H2595); Ragged Neck, 10 August 1967 (H2600); Rye Ledge, 25 April 1967 (H2615); Little Boar's Head, 14 August 1967 (H2627); Great Boar's Head, 11 December 1966 (H2631); Bound Rock, 18 November 1966 (H2641); Piscataqua River toll bridge, 18 October 1966 (42685); Hilton Park, 9 July 1967 (H2650); Cedar Point, 24 June 1967 (H2671); Durham Point, 14 August 1967 (H2672); Adams Point, 22 December 1966 (H2666) ; Pierce Point, 8 November 1966 (H2679). CHAMPIACEAE Lomentaria baileyana (Harvey) Farlow Growing on shells, pebbles, epiphytic on other algae or on Zostera, or free-floating. Found from July to November ; common throughout the Estuary. Annual; tetraspores found in July and August, Recorded from the tropics to New Hampshire and Nova Scotia (as summarized in Taylor, 1957; Doty and New- house, 1954; Edelstein, McLachlan and Craigie, 1967). REPRESENTATIVE SPECIMENS: Cedar Point, 12 September 1966 (H1557); Durham Point, 19 September 1966 (H1558); Adams Point, 13 Sep- tember 1966 (H1563); Fabyan's Point, 17 September 1966 (H1565) ; Pierce Point, 13 September 1966 (H1566). Lomentaria orcadensis (Harvey) Collins ex Taylor Collected in the sublittoral zone (August and October) at two coastal and one estuarine location, Perennial; tetra- spores found in August. Recorded from Rhode Island to Maine (as suramarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Jaffrey Point, 7 August 1967 (M5085) ; Odiorne’s Point, 7 August 1967 (M5086); Hilton Park, 2 October 1968 (N. B. Reynolds 287). 220 Rhodora [Vol. 72 CERAMIALES CERAMIACEAE *Antithamnion americanum (Harvey) Farlow Two specimens seen from the open coast; vegetative; an- nual(?). Recorded from New Jersey to Labrador (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals, before 1870; (FH L. L. Thaxter 7782). Isles of Shoals (FH Cole). Antithamnion boreale (Gobi) Kjellman Found once by Croasdale (July) growing in the sublittoral zone on the open coast. Annual(?); tetraspores found in July. Recorded from Northern Massachusetts to Ellesmere Island (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMEN: Isles of Shoals (Appledore Island), 26 July 1938 (Croasdale 3728). Antithamnion cruciatum (C. Agardh) Nägeli Growing in mud or epiphytic on other algae in the lower eulittoral and sublittoral zones from May to November. Found on the open coast and throughout most of the Estu- ary. Annual; tetraspores found from June to September. Recorded from Bermuda to New Hampshire and Newfound- land (as summarized in Taylor, 1957; Mathieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 26 July 1938 (Croasdale 3729); Odiorne's Point, 7 August 1967 (M8781); Hilton Park, 9 July 1967 (H70); Cedar Point, 12 September 1966 (H69); Adams Point, 13 October 1966 (H68). Antithamnion floccosum (O. F. Müller) Kleen Growing in the sublittoral zone. Found on the open coast, and with restricted distribution in the Estuary. Collected from January to May, and August. Annual; tetraspores found in April and August. Recorded from Northern Massachusetts to Ile St. Pierre (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Jaffrey Point, 7 January 1967 (H73); Odiorne’s Point, 7 August 1967 (M8752); Rye Ledge, 20 April 1952 (M3753); Adams Point, 4 May 1968 (M3754). 1970] Marine Algae — Hehre and Mathieson 221 ** Antithamnion plumula (Ellis) Thuret in Le Jolis Found once (July) growing on mud in the low eulittoral zone in the Estuary. No reproductive structures seen; an- nual. Previously recorded from New Jersey to Southern Massa- chusetts (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMEN: Cedar Point, 27 July 1966 (H74). * Antithamnion pylaisaei (Montagne) Kjellman Growing in the sublittoral zone on the open coast. Two specimens seen (May) ; vegetative; annual (?). Recorded from Long Island to Baffin Island (as summar- ized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Appledore Island), 25 May 1938 (Croasdale 3762); Rye Beach, (FH Cole). Callithamnion baileyi Harvey Found throughout most of the year on mud, stones and epiphytic on other algae in the lower eulittoral zone. Pres- ent on the open coast and throughout the Estuary. Peren- nial(?) ; tetraspores found in March and July ; carpospores found from July to October; spermatia found in July, Sep- tember and October. Recorded from New Jersey to Prince Edward Island and Baie des Chaleurs (as summarized in Taylor, 1957; Car- dinal, 1967). REPRESENTATIVE SPECIMENS: Isles of Shoals (Appledore Island), 26 July 1938 (Croasdale 3825); Jaffrey Point, 3 May 1968 (M3826) ; Concord Point, 22 November 1966 (H210); Piscataqua River toll bridge, 12 August 1967 (H214); Hilton Park, 7 January 1967 (H218); Cedar Point, 3 March 1967 (H213); Durham Point, 16 October 1966 (H216); Adams Point, 19 July 1966 (M3827); Pierce Point, 25 July 1967 (H215). *Callithamnion corymbosum (J. E. Smith) Lyngbye Found in July in the lower eulittoral zone in the Estuary. Annual(?); tetraspores found in July. Recorded from Bermuda to Nova Scotia (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMEN: Adams Point, 23 July 1967 H219). Callithamnion roseum (Roth) Lyngbye Growing on the open coast. One specimen seen ; vegetative; annual(?). 222 Rhodora [Vol. 72 Recorded from New Jersey to Southern Massachusetts and New Hampshire (as summarized in Taylor, 1957; Doty and Newhouse, 1954). REPRESENTATIVE SPECIMEN: Isles of Shoals, before 1870 (FH L. L. Thaxter 7788). Callithamnion tetragonum (Withering) S. F. Gray Reported by Croasdale (1941) from the Isles of Shoals. No specimens seen; annual(?). Recorded from New Jersey to Maine (as summarized in Taylor, 1957). Ceramium deslongchampsii Chauvin in Duby var. hooperi (Harvev) Taylor Found from April to November growing on rocks and pil- ings under over-hanging fucoids in the mid to lower eulit- toral zone on the open coast and in the Estuary. Peren- nial(?) ; tetraspores found in August. Recorded from Connecticut to the Lower St. Lawrence (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Jaffrey Point, 9 September 1967 (H220) ; North Wallis Sands, 16 July 1967 (H221); Piscataqua River toll bridge, 12 August 1967 (H222); Cedar Point, 23 November 1966 (H223); Squamscott River, 13 September 1966 (H224). Ceramium diaphanum (Lightfoot) Roth Reported by Collins (1900) and Doty and Newhouse (1954). No specimens seen. The abundance and distribution of this plant given in the latter paper suggests possible confusion with Ceramium strictum. Recorded from Virginia to Prince Edward Island (as sum- marized in Taylor, 1957). Ceramium rubriforme Kylin prox. Found once (July) in the lower eulittoral zone on the open coast. Vegetative; perennial. Recorded from Southern Massachusetts to New Hampshire and Newfoundland (as summarized in Taylor, 1957 ; Math- ieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMEN: Isles of Shoals (Star Island), 22 July 1966 (M3905). Ceramium rubrum (Hudson) C. Agardh Found throughout the year growing on rocks and epiphytic 1970] Marine Algae — Hehre and Mathieson 223 on various algae in the lower eulittoral and sublittoral zones. Common on the open coast and throughout the Estu- ary. Perennial; tetraspores found January-December ; cys- tocarpic plants found all months except May. Recorded from the tropies to Baffin Island (as summarized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 22 July 1966 (M3940); Jaffrey Point, 7 January 1967 (!H226) ; Fort Consti- tution, 20 September 1966 (H234) ; Odiorne's Point, 21 February 1967 (H236); North Wallis Sands, 16 January 1967 (H243); Concord Point, 8 January 1967 (H253); Ragged Neck, 26 December 1966 (H264); Rye Ledge, 11 November 1966 (H271); Little Boar's Head, 24 May 1967 (H285); Great Boar’s Head, 24 January 1967 (H288); Bound Rock, 18 January 1967 (H299); Piscataqua River toll bridge, 27 February 1967 (H349); Hilton Park, 27 April 1967 (H339); Cedar Point, 3 March 1967 (H317); Durham Point, 26 June 1967 (H324); Adams Point, 13 September 1966 (H309); Faoyans Point, 8 July 1967 (H330); Pierce Point, 23 June 1967 (H346); Squam- scott River, 4 March 1967 (H356). Ceramium strictum Harvey Found from June to October growing on rocks and epiphy- tic on Zostera and certain algae, Common throughout the Estuary in the lower eulittoral and sublittoral zones. Rare on the coast. Annual; tetraspores and carpospores found from June to October. Recorded from Florida to Prince Edward Island and Baie des Chaleurs (as summarized in Taylor, 1957; Cardinal, 1967). REPRESENTATIVE SPECIMENS: Concord Point, 13 September 1966 (H357); Bound Rock, 19 July 1966 (M4079); Hilton Park, 9 July 1967 (H358); Cedar Point, 7 August 1967 (H362); Durham Point, 14 August 1967 (H363); Adams Point, 23 July 1967 (H360); Fabyans Point, 2 August 1966 (M4100); Squamscott River, 20 July 1966 (H366). Plumaria elegans (Bonnemaison) Schmitz Found throughout the year growing on vertical rock faces under over-hanging fucoids in the lower eulittoral zone on the open coast. Perennial; paraspores found from Septem- ber to January. Recorded from New Jersey to Newfoundland (as summar- ized in Taylor, 1957; Mathieson, Dawes and Humm, 1969). 224 Rhodora [Vol. 72 REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 28 July 1938 (Croasdale 5338); Jaffrey Point, 8 November 1966 (H1900); North Wallis Sands, 11 October 1966 (H1910); Rye Ledge, 11 No- vember 1966 (H1911); Great Boar’s Head, 12 August 1967 (H1913); Bound Rock, 15 March 1968 (M5343). Ptilota serrata Kiitzing Found throughout the year growing in the sublittoral zone on the open coast. Common in drift. Perennial; tetraspores found from November to June; spermatia found in Febru- ary and March; only one cystocarpic plant found (May). Recorded from New Jersey to Ellesmere Island (as sum- marized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island); 28 May 1968 (M5984); Jaffrey Point, 6 May, 1967 (M5987); Odiorne’s Point, 21 February 1967 (H2401); North Wallis Sands, 16 January 1967 (H2406); Concord Point, 10 December 1966 (H2420); Ragged Neck, 9 March 1967 (H2422); Rye Ledge, 25 April 1967 (H2430); Little Boar’s Head, 14 September 1966 (H2443); Great Boar’s Head, 24 January 1967 (H2448); Bound Rock, 18 November 1966 (H2453). Spermothamnion repens (Dillwyn) Rosenvinge Growing on mud and pebbles in the lower eulittoral zone in March and July-September, Rare on the open coast, and with limited distribution in the Estuary. Perennial; tetra- spores found in July. Recorded from Florida to Nova Scotia (as summarized in Taylor, 1957, as Spermothamnion turneri). REPRESENTATIVE SPECIMENS: North Wallis Sands, 20 March 1967 (H245); Cedar Point, 27 July 1966 (H2556); Durham Point, 19 September 1966 (H2557); Adams Point, 19 July 1966 (M6043). DELESSERIACEAE Membranoptera alata (Hudson) Stackhouse Found throughout the year epiphytic on various algae in the sublittoral zone. Frequent in drift on the open coast. Perennial; tetraspores found from November to July. Recorded from Northern Massachusetts to Baffin Island (as summarized in Taylor, 1957 ; including Membranoptera den- ticulata. See Mathieson, Hehre and Reynolds, in press). REPRESENTATIVE SPECIMENS: Isles of Shoals, before 1870 (rH L. L. Thaxter 7792); Jaffrey Point, 8 November 1966 (1H1567); Fort Constitution, 22 June 1966 (M5103); Odiorne's Point, 26 July 1967 (H1570); North Wallis Sands, 11 June 1967 (H1575); Concord Point, 1970] Marine Algae — Hehre and Mathieson 225 13 September 1966 (H1578); Ragged Neck, 18 September 1966 (H1580); Rye Ledge, 15 October 1966 (H1587); Little Boar’s Head, 2 April 1967 (H1592); Great Boar’s Head, 13 November 1966 (H1593); Bound Rock, 18 January 1967 (H1596). *Pantoneura baerii (Postels et Ruprecht) Kylin Collected twice (May and June) in the sublittoral zone on the open coast. Perennial; tetraspores found in May. Previously recorded from Northern Massachusetts, Labra- dor, Hudson Straits and Ellesmere Island (as summarized in Taylor, 1957 ; Lamb and Zimmermann, 1964). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), June 1965 (Kingsbury, 1965) ; Isles of Shoals (Appledore Island), 28 May 1968 (M5118). Phycodrys rubens (Linnaeus) Batters Growing throughout the year in the sublittoral zone of the open coast. Young plants occasionally found in deep tide- pools in the lower eulittoral zone. Perennial; tetraspores found October-July; cystocarpic plants found November- August; spermatia found once in July. Recorded from New Jersey to Ellesmere Islaad (as sum- marized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Lunging Island), 22 July 1966 (M5174); Jaffrey Point, 7 January 1967 (H1676); Fort Constitution, 23 June 1967 (H1685) ; Odiorne’s Point, 21 February 1967 (H1689); North Wallis Sands, 11 October 1966 (H1696); Con- cord Point, 13 September 1966 (H1713); Ragged Neck, 12 June 1967 (H1727); Rye Ledge, 23 December 1966 (H1728); Little Boar's Head, 24 May 1967 (H1742); Great Boar's Head, 15 September 1966 (H1752); Bound Rock, 31 May 1967 (H1761). DASYACEAE Dasya pedicellata (C. Agardh) C. Agardh Young plants grow epiphytically on various algae, while older plants occur free-floating or entangled amongst other plants. Common throughout the Estuary from July to December. Annual; tetraspores found from July-October ; spermatia and carpospores found from July to November. Recorded from the tropics to New Hampshire and Nova Scotia (as summarized in Taylor, 1957; Doty and New- house, 1954; Edelstein, McLachlan and Craigie, 1967). REPRESENTATIVE SPECIMENS: Hilton Park, 19 July 1966 (M4539) ; Cedar Point, 7 August 1967 (H864); Durham Point, 19 September 226 Rhodora [Vol. 72 1966 (H865); Adams Point, 13 October 1966 (H862); Fabyans Point, 2 August 1966 (M4535); Pierce Point, 18 October 1966 (H868). RHODOMELACEAE Bostrychia rivularis Harvey Reported by Farlow (1882). Supposedly found in the vicin- ity of the Isles of Shoals by Captain Pike. No specimens have been found to verify this species and it remains a doubtful record. Recorded from the tropics to New Hampshire (as summar- ized in Taylor, 1957). *Chondria baileyana (Montagne) Harvey Found from June-November, growing on shells, pebbles and epiphytic on other algae and on Zostera. It occurs in the lower eulittoral and sublittoral zones throughout the Estuary. Annual; tetraspores found from June to Septem- ber; cystocarpic plants found in June and September. Previously recorded from New Jersey to Northern Mass- chusetts and Nova Scotia (as summarized in Taylor, 1957; Edelstein, McLachlan and Craigie, 1967). REPRESENTATIVE SPECIMENS: Durham Point, 14 August 1967 (H377); Adams Point, 23 July 1967 (H376); Fabyans Point, 17 September 1966 (H386); Pierce Point, 13 September 1966 (H381). Chondria tenuissima (Goodenough et Woodward) C. Agardh Reported by Doty and Newhouse (1954). No specimens were available, but from their data, it would appear to be Chondria baileyana. *Polysiphonia denudata (Dillwyn) Greville ex Harvey in Hooker Found from July to December in the lower eulittoral and sublittoral zones throughout the Estuary, Annual; tetra- spores found from July to October; cystocarpic plants found from July to September. Recorded from the tropics to Prince Edward Island (as sum- marized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Hilton Park, 28 August 1968 (N. B. Reynolds 281); Cedar Point, 12 September 1966 (H1952); Durham Point, 19 September 1966 (H1953); Adams Point, 13 October 1966 (H1950); Fabyans Point, 17 September 1966 (H1955); Pierce Point, 1970] Marine Algae — Hehre and Mathieson 227 8 November 1966 (H1958); Squamscott River, 13 September 1966 (H1960). Polysiphonia elongata (Hudson) Sprengel Found throughout the year growing in tidepools in the lower eulittoral and sublittoral zones throughout the Estu- ary. Perennial; tetraspores found from June to September ; carpospores and spermatia found in July. Recorded from New York to Prince Edward Island and Newfoundland (as summarized in Taylor, 1957; Mathieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMENS: Pierce Island, Piscataqua River, 18 July 1966 (M5436); Hilton Park, 9 December 1966 (H1977); Cedar Point, 23 November 1966 (H1983); Durham Point, 15 November 1966 (H1988); Adams Point, 13 September 1966 (H1968); Fabyans Point, 15 October 1966 (H1991); Pierce Point, 8 November 1966 (H1999). *Polysiphonia flexicaulis (Harvey) Collins One record (July) on the open coast (Croasdale) and one (July) in the Estuary. Annual; cystocarpic plants found in July. Recorded from Long Island to Newfoundland (as summar- ized in Taylor, 1957 ; Mathieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMENS: Isles of Shoals (Appledore Island), 28 July 1938 (Croasdale 5471); Cedar Point, 10 July 1941 (E. Flagg 5413). Polysiphonia harveyi Bailey Reported by Collins (1900) and Doty and Newhouse (1954). No data or specimens available. Recorded from South Carolina to Newfoundland (as sum- marized in Taylor, 1957; Mathieson, Dawes and Humm, 1969). Polysiphonia lanosa (Linnaeus) Tandy Hemiparasitic on Ascophyllum nodosum. Common on the open coast, and with limited distribution in the Estuary. Present throughout the year. Perennial; tetraspores found from June to November; cystocarpic plants found from June to December; spermatia found from January to May. Recorded from New Jersey to Newfoundland (as summar- ized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 22 July 1966 (M5511); Jaffrey Point, 7 January 1967 (H2003); Fort Consti- 228 Rhodora [Vol. 72 tution, 8 November 1966 (H2018) ; Odiorne's Point, 17 October 1966 (H2028); North Wallis Sands, 11 June 1967 (H2038); Concord Point, 10 August 1967 (H2051); Ragged Neck, 22 May 1967 (H2061); Rye Ledge, 15 October 1966 (H2062); Little Boar's Head, 24 May 1967 (H2083); Great Boar's Head, 25 July 1967 (H2095); Bound Rock, 18 January 1967 (H2096); Piscataqua River toll bridge, 24 June 1966 (H2098); Hilton Park, 28 August 1968 (N. B. Reynolds 273). Polysiphonia nigra (Hudson) Batters Found in the lower eulittoral (in tidepools) and the sub- littoral zones. Occasional on the open coast, and with re- stricted distribution in the Estuary. Annual(?); tetra- spores found in March and July; cystocarpic plants found from October to December. Recorded from New Jersey to Nova Scotia and Newfound- land (as summarized in Taylor, 1957; Mathieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMENS: Jaffrey Point, 8 November 1966 (H2102); Odiorne's Point, 7 August 1967 (M5562); North Wallis Sands, 7 August 1967 (M5564); Rye Ledge, 27 February 1968 (M5565); Bound Rock, 6 December 1966 (H2103); Hilton Park, 7 January 1967 (H2108); Cedar Point, 27 July 1966 (H2105); Durham Point, 16 October 1966 (H2113); Adams Point, 13 October 1966 (H2111). Polysiphonia nigrescens (Hudson) Greville Found throughout the year on rocks in the lower eulittoral and sublittoral zones on the open coast and throughout the Estuary. Frequent in drift. Perennial; tetraspores found from May to December; cystocarpic plants found in March and from June to September; spermatia found in February. Recorded from South Carolina to Newfoundland (as sum- marized in Taylor, 1957; Mathieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMENS: Isles of Shoals (Appledore Island), 26 July 1938 (Croasdale 5586); Jaffrey Point, 8 November 1966 (H2129); Odiorne's Point, 10 March 1967 (H2119); North Wallis Sands, 10 September 1966 (H2122); Concord Point, 10 August 1967 (H2124); Ragged Neck, 23 November 1966 (H2126); Rye Ledge, 12 September 1966 (H2127); Bound Rock, 7 March 1966 (M5601) ; Hilton Park, 7 January 1967 (H2131); Cedar Point, 24 May 1967 (H2146); Durham Point, 3 March 1967 (M5632); Adams Point, 1970] Marine Algae — Hehre and Mathieson 229 13 October 1966 (H2144); Fabyans Point, 16 November 1966 (H2150); Pierce Point, 18 October 1966 (H2154). Polysiphonia novae-angliae Taylor Found throughout the year on rocks and epiphytic on other algae in the lower eulittoral and sublittoral zones. Com- mon on the open coast, and with restricted distribution in the Estuary. Perennial; tetraspores found January-Decem- ber; cystocarpic plants found from June to November; spermatia found in February, March and May. Recorded from Rhode Island to Newfoundland (as summar- ized in Taylor, 1957; Mathieson, Dawes and Humm, 1969). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), 22 July 1966 (M5656); Jaffrey Point, 6 May 1967 (M5658); Fort Consti- tution, 24 July 1967 (M5659); North Wallis Sands, 20 March 1967 (H2156); Concord Point, 8 January 1967 (H2157); Ragged Neck, 10 August 1967 (H2159); Rye Ledge, 12 September 1966 (H2160) ; Great Boar’s Head, 11 December 1966 (H2162); Bound Rock, 18 January 1967 (H2163); Piscataqua River toll bridge, 12 August 1967 (H2186); Hilton Park, 13 February 1967 (H2193); Cedar Point, 17 April 1967 (H2177); Durham Point, 15 December 1966 (H2181); Adams Point, 13 October 1966 (H2171). *Polysiphonia subtilissima Montagne Found from June to October growing on rocks and mud in the lower eulittoral and sublittoral zones. Restricted to parts of the Estuary where salinities are very low. An- nual; only one tetrasporic specimen was observed (July). Previously recorded from the tropics to Northern Massa- chusetts and with some doubt to the Baie de Gaspe (as sum- marized in Taylor, 1957; Cardinal, 1967). REPRESENTATIVE SPECIMENS: Cedar Point, 27 June 1966 (H2118); Crommet Creek, 20 June 1966 (M5573); Squamscott River, 21 June 1966 (H2115). D4'vsiphenia urceolata (Lightfoot ex Dillwyn) Greville Growing on rocks in the lower eulittoral and sublittoral zones throughout the year. Common on the open coast and with restricted distribution in the Estuary. The seasonal varieties (roseola and patens) as designated by Taylor (1957) were not distinguishable. Perennial; tetraspores found from May to August; cystocarpic plants found in July and August; spermatia found from January to May. 230 Rhodora [Vol. 72 Recorded from North Carolina to the arctic (as summar- ized in Taylor, 1957, including varieties roseola and patens). REPRESENTATIVE SPECIMENS: Isles of Shoals (Lunging Island), 22 July 1966 (M5727); Jaffrey Point, 9 August 1967 (H2204); Fort Constitution, 24 July 1967 (M5738); Odiorne's Point, 30 May 1967 (H2209); North Wallis Sands, 20 May 1967 (H2213); Concord Point, 27 June 1967 (H2215); Ragged Neck, 12 June 1967 (H2216); Rye Ledge, 24 July 1967 (H2217); Little Boar's Head, 24 May 1967 (H2222); Great Boar's Head, 24 January 1967 (H2225); Bound Rock, 20 February 1967 (H2228); Hilton Park, 14 October 1966 (H2235); Fox Point, 5 December 1966 (H2200). Rhodomela confervoides (Hudson) Silva Found throughout the year on the open coast in the low eulittoral and sublittoral zones. Perennial; tetraspores found in March and from May to August; cystocarpic plants found in June and July; spermatia found in March and April. Recorded from New Jersey to Baffin Island (as summar- ized in Taylor, 1957). REPRESENTATIVE SPECIMENS: Isles of Shoals (Star Island), June 1965 (Kingsbury, 1965); Jaffrey Point, 1 April 1967 (H2535); Fort Con- stitution, 22 June 1966 (M6031); Odiorne’s Point, 7 August 1967 (M6032); North Wallis Sands, 28 February 1967 (H2536) ; Concord Point, 5 March 1967 (H2541); Ragged Neck, 12 December 1966 (H2544); Rye Ledge, 11 November 1966 (H2550); Little Boar's Head, 22 December 1966 (H2551); Great Boar's Head, 14 October 1966 (H2552); Bound Rock, 7 March 1966 (H2554). DISCUSSION Eighty-eight taxa of red algae are listed from the coastal and estuarine environments of New Hampshire; fifteen taxa represent additions to the flora of the state. Six taxa are doubtful records (Polysiphonia harveyi, Bostrychia rivularis, Chondria tenuissima, Agardhiella tenera, Cera- mium diaphanum, Batrachospermum sp.). Many of the plants are boreal and subarctic species (e.g., Porphyra miniata, Euthora cristata, Rhodophyllis dicho- toma, Gigartina stellata, Halosaccion ramentaceum, Ptilota serrata, Phyllophora spp., Polyides rotundus, Rhodochorton purpureum). Certain components of the Estuary flora have 1970] Marine Algae — Hehre and Mathieson 231 affinities towards warm temperate regions — in particular, the summer annuals whose center of distribution is south of New England (e.g., Lomentaria baileyana, Dasya pedi- cellata, Spermothamnion repens, Polysiphonia denudata, Chondria baileyana, Gracilaria foliifera and Polysiphonia subtilissima) . The paucity of information regarding seasonality and reproductive periodicity of marine algae of northern New England is due to a lack of seasonal collections. Winter observations are essentially nonexistent and little phenolog- ical data are available. In particular, those algae which have their growth or reproduction during late fall to early spring are apt to be missed (e.g., Bangia fuscopurpurea, Plumaria elegans, Polysiphonia nigra, Porphyra linearis, Rhodochorton purpureum). Edelstein and McLachlan (1966b) studied the occurrence and reproduction of marine algae during the winter months in Nova Scotia. Data from their study and the present investigation show many sim- ilarities. The reproductive periodicities recorded for several species differ markedly from those listed in Taylor (1957). Several species reproduce throughout the year (e.g., Porphyra um- bilicalis, Audouinella membranacea, Hildenbrandia proto- typus, Euthora cristata, Choreocolax polysiphoniae, Ahn- feltia plicata, Phyllophora brodiaei, Chondrus crispus, Gigartina stellata, Rhodymenia palmata, Ptilota serrata, Ceramium rubrum, Polysiphonia lanosa, P. nigrescens, P. novae-angliae, P. urceolata), although they were reported to reproduce during only one or two seasons. Other species had more prolonged periods of reproduction than previously re- ported (e.g., Petrocelis middendorfii, Antithamnion crucia- tum, Membranoptera alata, Dasya pedicellata). Reproduc- tive specimens of Plumaria elegans were found in the fall and winter while Taylor reports reproduction from the sum- mer and earlier. He describes Porphyra miniata as being reproductive during any season, giving the impression that it occurs throughout the year. According to our observa- tions, P. miniata is an annual (spring and summer). It 232 Rhodora [Vol. 72 was found to be reproductive every month in which it was found (March-August). Our observations on P. miniata are similar to those of Lamb and Zimmermann (1964). The seasonal occurrence of most species generally agrees with those of Taylor (1957), except for Bangia fuscopur- purea and Antithamnion floccosum which differed slightly. Bangia fuscopurpurea occurred from late fall to late sum- mer. Taylor records it from early spring and summer. Antithamnion floccosum was found from winter to late summer while Taylor records it only in the spring. Both asexual and sexual structures were found. Mono- spores, bispores, paraspores and tetraspores were encoun- tered. Tetraspores were the most common type. Mono- spores were found only in Bangia fuscopurpurea, Kylinia spp. and Ahnfeltia plicata. Bispores were found only in the crustose corallines, while paraspores were found ex- clusively in Plumaria elegans. Tetraspores were found in most other species. One conspicuous exception is Gigartina stellata, where the tetrasporophyte generation is unknown. Certain plants were found only as tetrasporophytes (e.g., Antithamnion cruciatum, A. floccosum, Audouinella mem- branacea, Ceratocolax hartzii, Cystoclonium purpureum var. cirrhosum, Dumontia incrassata, Melobesia lejolisii, Halo- saccion ramentaceum, Hildenbrandia prototypus, Lomen- taria baileyana, L. orcadensis, Membranoptera alata, Petro- celis middendorfii, Rhodochorton purpureum, Rhodymenia palmata). Sexual generations have been described for most of these plants except Halosaccion ramentaceum, Hilden- brandia prototypus and Rhodymenia palmata. Sexual structures were found on most species except those which produced monospores, bispores and paraspores. Female plants were more frequent than male plants. Some species produced cystocarps and spermatia at different times of the year (e.g., Polysiphonia lanosa, P. nigrescens, P. novae-angliae, P. urceolata, Ptilota serrata, Rhodomela confervoides). Others produced both structures at the same time (e.g., Callithamnion baileyi, Dasya pedicellata, Rhy- codrys rubens, Polysiphonia denudata). 1970] Marine Algae — Hehre and Mathieson 233 A distinct overlap in the occurrence of tetraspores and carpospores was noted for several species. Callithamnion baileyi and Polysiphonia nigra showed no distinct overlap- ping. Three trends were evident: (1) Tetraspores and carpospores were found throughout the year (e.g, Ceramium rubrum, Chondrus crispus, Eu- thora cristata, Phycodrys rubens). (2) Tetraspores were found throughout the year while carpospores were found during only one or two seasons (e.g., Polysiphonia urceolata, Choreocolax polysiphoniae) . (3) Tetraspores and carpospores were presen? during one or more seasons. Both summer annuals (e.g. Ceramium strictum, Dasya pedicellata, Chondria baileyana, Polysi- phonia denudata) and perennials (e.g., Phyllophora mem- branifolia, Polyides rotundus, Polysiphonia elongata, P. lanosa, P. nigrescens) were discernable. We wish to express our sincere gratitude to the following people: Dr. A. R. Hodgdon, for his editorial advice and critical review of the manuscript; Dr. C. G. Nast, Dr. T. E. Furman, and Dr. Emery Swan, for their helpful com- ments; Mr. Norman Reynolds, for assistance with several of the sublittoral collections; Miss Joan R. Conway for assistance with collecting and with processing of herbarium specimens. DEPARTMENT OF BOTANY UNIVERSITY OF NEW HAMPSHIRE DURHAM, N. H. 03824 REFERENCES ApEY, W. H. 1965. The genus Clathromorphum (Corallinaceae) in the Gulf of Maine. Hydrobiologia 26 (3/4): 539-573. ————————, 1966. The Genera Lithothamnium, Leptophytum (nov. gen.) and Phymatolithon in the Gulf of Maine. Hydro- biologia 28(3/4) : 321-370. CARDINAL, À. 1967. Inventaire des Algues Marines Benthiques de la Baie des Chaleurs et de la Baie de Gaspe (Quebec). Natura- liste Can. 94: 233-271. 234 Rhodora [Vol. 72 CHIHARA, M. 1961. Life Cycle of the Bonnemaisoniaceous Algae of Japan (1). Science Reports of the Tokyo Kyoiku Daigaku, Sec. B. 10(153): 121-154. 1962. Life Cycle of the Bonnemaisoniaceous Algae of Japan (2). Science Reports of the Tonyo Kyoiku Daigaku, Sec. B. 11(161): 27-53. COLLINS, F. S. 1900. Preliminary Lists of New England Plants. — V. Marine Algae. Rhodora 2(14): 41-52. 1901. Notes on Algae III. Rhodora 3(29) : 132-137. 1903. Notes on Algae V. Rhodora 5(56): 204-212. 1906. Acrochaetium and Chantransia in North America. Rhodora 8(94): 189-196. I. HOLDEN and W. A. SETCHELL. 1895-1919. Phy- cotheca Boreali-Americana. Fascicles 1-46 + A-E. Farlow Her- barium, Harvard University. CROASDALE, H. T. 1941. Additional Records of Marine Algae from New Hampshire. Rhodora 43(510): 213-216. Doty, M. and J. NEWHOUSE. 1954. The Distribution of Marine Algae into Estuarine Waters. Am. Jour. Bot. 37(6): 508-515. EDELSTEIN, T. and J. McLachlan. 1966a. Species of Acrochaetium and Kylinia New to North America. Brit. Phycol. Bull. 3(1): 37-41. 1966b. Investigations of the Ma- rine Algae of Nova Scotia. I. Winter Flora of the Atlantic Coast. Can. Jour. Bot. 44(8) : 1035-1055. 1968. Investigations of the Ma- rine Algae of Nova Scotia. V. Additional Species New or Rare to Nova Scotia. Can. Jour. Bot. 46(8): 993-1003. , and J. S. CRAIGIE. 1967. Investigations of the Marine Algae of Nova Scotia. II. Species of Rhodophyceae New or Rare to Nova Scotia. Can. Jour. Bot. 45(2) : 193-202. FARLOW, W. G. 1882. Marine Algae of New England. Report of the U.S. Commission of Fish and Fisheries for 1879. p. 1-210. KINGSBURY, J. M. 1965. Checklist of the Marine Algae of the Isles of Shoals (Star Island). Unpublished. 2 pp. LAMB, I. M. and M. H. ZIMMERMANN. 1964. Marine Vegetation of Cape Ann, Essex County, Massachusetts. Rhodora 66(767): 217-254. Lewis, J. R. 1964. The Ecology of Rocky Shores. English Univer- sities Press, London. 323 pp. LUND, SØREN. 1959. The Marine Algae of East Greenland: 1. Taxonomical Part. Medd. om Grenl., 156(1): 1-247. MATHIESON, A. C., C. J. DAWES and H. J. HUMM. 1969. Contribu- tions to the Marine Algae of Newfoundland. Rhodora 71(785) : 110-159. 1970] Marine Algae — Hehre and Mathieson 235 , E. J. HEHRE and N. B. REYNOLDS. Investigations of New England Marine Algae I. A Floristic and Descriptive Ecological Study of the Marine Algae at Jaffrey Point, New Hampshire. Nova Hedwigia (in press). , N. B. REYNOLDS and E. J. HEHRE. Investigations of New England Marine Algae II. Species Composition, Distri- bution and Zonation of Seaweeds in the Great Bay Estuary System and Adjacent Coastal Waters. Nova Hedwigia (in press). NEWROTH, P. R., and A. R. A. TAYLOR. 1968. The Distribution of Ceratocolax hartzii. Brit. Phycol. Bull. 3(3) : 543-546. PAPENFUSS, G. F. 1947. Further Contributions Toward an Under- standing of the Acrochaetium-Rhodochorton Complex of the Red Algae. Univ. Calif. Publ. Bot. 18(19) : 433-447. PARKE, M. and P. S. Drxon. 1968. Checklist of British Marine Algae — Second Revision. Jour. mar. biol. Assoc, U.K., 48(3): 783-832. TAYLOR, W. R. 1957. Marine Algae of the Northeast Coast of North America. Ann Arbor Press, 509 pp. Woop, R. D. and J. STRAUGHAN. 1953. Time-Intensity Tolerance of Lemanea fucina to Salinity. Am. Jour, Bot. 40(6): 381-384. 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DUO dO — Hoo MO les Pace s HN Pave Petts aDybun-anaou ‘qJ DOE ID Eo £010 DD OQ DIM Jg tcp suaosaubru “gq D O ə O O L == EE pur L O O 04D ‘q Job) NDS 40 LJ Oe i es eS S Se SaS DS0WD] `d Vdd AON LOO IdAS DNV ATAL ANAL AVW UdV UVW 4AA NVC VXVL 'punog jou Sjue[d — — ‘Aep aa — o ‘satods pews — g ‘satods aSae[ — V 'sa1odsei39] = L ‘eneuteds — S ‘sərodsesed — a 'so:iodsu1jo9j0d.1€2 — N ‘satodsouow — I 'sa1odsod.ie2 — 9 ‘sa1odsiq = g ein[sdureqg səy ur eveo&udopou ey} Jo Ajrorpotieq 9Anonpoidoy pue A3r[euoseog (penunuoo) I oe[q*], HYBRIDS IN RUBUS SUBGENUS EUBATUS IN NEW ENGLAND? ° FREDERIC L. STEELE AND A. R. HODGDON The background for this current study will be found in the three papers we have published on New England black- berries during the past nine years, beginning in 1962» + 5. Aside from the cited paper on hybrids of Rubus hispidus and R. setosus, we have thus far made no attempt to char- acterize hybrids in the group. In our survey of New Eng- land blackberries in 1966 we thought it would have unduly complicated our study to have done so. However, at that time we were fully aware of the significance of hybridiza- tion as we indicated in the long list of rejected binomials on pp. 510-512 where, in each instance when we suspected that a name had been given to a hybrid, we also stated what we supposed the parentage to be. The present paper then is an attempt to round out our studies on Rubus in New England, We have accumulated a great deal of data in the course of field and laboratory work, much of it serving to support our contention and that of some others before us that it is chiefly the high inci- dence of interspecific hybridization that makes classification so difficult in blackberries. Therefore, our study of New ‘Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientific Contribution No. 523. "This research has been supported by the New Hampshire Agri- cultural Experiment Station and by grants from the Central Uni- versity Research Fund of the Graduate School of the University of New Hampshire and from the Northern New England Academy of Science. "Hodgdon, A. R. & F. L. Steele. 1962. Glandularity in Rubus alle- gheniensis Porter. Rhodora 64: 161-168. ‘Steele, F. L. & A. R. Hodgdon. 1963. Hybridization of Rubus his- pidus and R. setosus. Rhodora 65: 262-270. *"Hodgdon, A. R. & F. L. Steele. 1966. Rubus subgenus Eubatus in New England. Rhodora 68: 474-518. 240 1970] Rubus Hybrids — Steele and Hodgdon 241 England Rubus would be incomplete unless we gave due consideration to hybrids. The following treatment includes those intermediates that we have observed and collected often enough to feel we have definite evidence of their hybrid nature. We are equally certain that there are some other hybrid combinations yet to be determined since we have never, for any very long period of time, failed to discover new ones. There have been found, by us and others before us, some very unusual plants that we hesitate to evaluate; presumably these may have resulted from crossing but we will ignore them in the pres- ent treatment. In what we may for convenience designate the flagellaris group of blackberries the following species, Rubus flagel- laris, R. Jaysmithii, R. arenicola and R. recurvicaulis seem to hybridize freely. At least there are overlapping ten- dencies between them and it is difficult to separate them clearly when they meet. However, it is often difficult to distinguish hybrids from unusual states of growth such as depauperate forms within the species. Also, especially on herbarium sheets, it is difficult to know which particular parents are involved. For these reasons any suspected hy- brids among these four species have been omitted from the present treatment. The cross between R. Enslenii and re- curvicaulis however, seemed to be clear enough to recognize both parents. The flagellaris group of species, which are very closely related and differ only in habit and minor aspects, would be difficult to distinguish at the species level as parents of crosses with other blackberries. Therefore, we have had to identify the parents arbitrarily on the basis of what seemed most likely at the particular site where the hybrid was ob- served. Usually this was possible to do because only Rubus flagellaris and R. recurvicaulis are widespread and common. Also, more often than not, one of these but not both occur in the immediate vicinity of a hybrid population. In ordi- nary herbarium material such hybrids would necessarily have to remain in question as to exact parentage. 242 Rhodora [Vol. 72 In the following treatment the names in parentheses that are given directly after some of the hybrid designations are binomials, in most cases rejected ones, which we consider to be equivalent to these hybrids. For convenience the generic name Rubus is omitted in the following listing of hybrids. Descriptions and discus- sions of the species involved and author's names of the binomials may be found in our 1966 study (loc. cit.). flagellaris X allegheniensis. Low doming to 2 ft or almost trailing ; leaflets 5, rather narrow, 7 to 11 cm long, slightly pubescent underneath ; primocanes 5 mm in diameter ; arma- ture of slender, hooked to straight prickles and occasional glands ; inflorescence straggling, up to 11 cm, the axis and pedicels with frequent glands. REPRESENTATIVE SPECIMENS: MAINE: LINCOLN Co., Edgecomb, Hodgdon 7231 (NHA); Boothbay, Hodgdon 10933 (NHA). NEW HAMP- SHIRE: CARROLL CO., Sandwich, Steele 3750 (STEELE); SULLIVAN CO., Claremont, Hodgdon & Steele 10198 (NHA). MASSACHUSETTS: WOR- CESTER CO., Mendon, Hodgdon & Steele 18051 (NHA). flagellaris X frondosus. Arching to doming and trailing; leaflets medium to large, 7 to 9 cm long, compound leaves 14 em across, pubescent underneath; canes 3 to 4 mm in diameter; armature of hooked prickles; inflorescence spreading, 2-6 flowered; axis pubescent but no glands pres- ent, REPRESENTATIVE SPECIMENS: NEW HAMPSHIRE: STRAFFORD CO., Durham, Hodgdon & Steele 12486 (NHA). MASSACHUSETTS: BARN- STABLE CO., Harwich, Hodgdon & Steele 3923 (STEELE). CONNECTICUT: TOLLAND CO., Tolland, Hodgdon & Steele 18052 (NHA); Stafford Springs, Steele & Hodgdon 3923 (STEELE). Jaysmithii X hispidus. REPRESENTATIVE SPECIMEN: RHODE ISLAND: WASHINGTON CO., Block Island, Champlin 7/4/67 (NEBC). Jaysmithii X elegantulus. Similar to recurvicaulis X elegantulus but with leaves slightly pubescent beneath. REPRESENTATIVE SPECIMEN: VERMONT: WINDHAM CO., Westmin- ster, Hodgdon & Steele 11372 (NHA). 1970] Rubus Hybrids — Steele and Hodgdon 243 Enslenii X recurvicaulis. Trailing to low doming or arch- ing, 1-2 ft. high; leaflets smooth, small to medium, 3-5, glabrous; prickles sparse to medium; inflorescence 2-3.5 cm long, of the recurvicaulis-type, flowers 1-4, pecicels short, slender. REPRESENTATIVE SPECIMENS: MAINE: YORK CO., Hodgdon & Steele 18050 (NHA). NEW HAMPSHIRE: STRAFFORD CO., Durham, Steele & Hodgdon 4099 (STEELE). Enslenii X pensilvanicus (K. frondosus). Arched or trail- ing; leaflets 3-5, larger than Enslenii, compound leaves up to 13 cm across, sparsely pubescent underneath; stems slender, 2.5 mm in diameter; prickles thin and slender, rather sparse; inflorescence up to 8 cm long, flowers 1-5, pedicels 2-6 cm long, glandless. REPRESENTATIVE SPECIMEN: NEW HAMPSHIRE: STRAFFORD CO., Dur- ham, Hodgdon 12197 (NHA). recurvicaulis X hispidus. Trailing; leaflets 3 to 5, charta- ceous, not lustrous; armature of short prickles, bristles and glands; inflorescence short, compact, 4 X 2 cm, 6 to 10- flowered. REPRESENTATIVE SPECIMENS: NEW HAMPSHIRE: GRAFTON CO., Wood- stock, Hodgdon & Steele 11356 (NHA); ROCKINGHAM CO., Seabrook, Hodgdon & Steele 11386 (NHA). MASSACHUSETTS: ESSEX CO., Salis- bury, Steele 3519 (STEELE). recurvicaulis X setosus (R. multiformis). Doming or trailing ; leaflets (3) 5, rather narrow, subchartaceous, glab- rous, medium to large, compound leaves up to 18 cm across; canes about 3 mm in diameter ; armature of slender prickles with few to many bristles intermixed; inflorescence of the setosus type, up to 9 cm long, axis and pedicels with glands. REPRESENTATIVE SPECIMENS: MAINE: LINCOLN CO., Boothbay, Hodg- don & Steele 7524 (NHA); SAGADAHOC CO., Arrowsic, Hodgdon 7524 (NHA). NEW HAMPSHIRE: CARROLL CO., Tamworth, Steele 3832 (STEELE); GRAFTON CO., North Woodstock, Fernald 362 (NHA). recurvicaulis X vermontanus (R. severus). Arching or doming ; leaves glabrous, often chartaceous, ovate to lanceo- late; armature of numerous sharp slender prickles, straight to hooked, up to 200 per dm. 244 Rhodora [Vol. 72 REPRESENTATIVE SPECIMENS: MAINE: CUMBERLAND CO., Harpswell, Hodgdon & Steele 12375 (NHA). NEW HAMPSHIRE: CARROLL CO., Ossipee, Steele 3477 (STEELE), Conway, Steele 2937 (STEELE) ; GRAFTON co., Thornton, Hodgdon 11347 (NHA), Waterville, Hodgdon & Steele 11306 (NHA); HILLSBORO CO., Nashua, Steele 2763 (STEELE). MASSA- CHUSETTS: ESSEX CO., Plum Island, Hodgdon & Steele 15543 (NHA). recurvicaulis X elegantulus. Arching, doming or trail- ing; leaflets large, subchartaceous, lateral petioles often short, terminal ones short to long, leaflets glabrous; canes stout; spines thin; inflorescence as in elegantulus, no glands on axis. REPRESENTATIVE SPECIMENS: MAINE: YORK CO., York, Hodgdon & Steele 12260 (NHA). NEW HAMPSHIRE: CARROLL CO., Sandwich, Steele 3783 (STEELE). recurvicaulis X allegheniensis. REPRESENTATIVE SPECIMEN: MAINE: KNOX CO., Vinal Haven, Bean 6/25/34 (NEBC). recurvicaulis < frondosus. This is very similar to flagel- laris X frondosus described above. Distinction depends on the putative parents that are present. REPRESENTATIVE SPECIMENS: MAINE: YORK CO. Wells, Fernald 13872 (NEBC). MASSACHUSETTS: BARNSTABLE CO. Harwich, Fernald 16899 (NEBC). recurvicaulis X pensilvanicus. Low-doming to 2 ft; leaf- lets 5 of medium-size, slightly pubescent beneath; primo- canes stouter than those of recurvicaulis (about 5 mm in diameter) ; armature of slender, hooked to straight prickles, without glands; inflorescence variable, axis without glands. REPRESENTATIVE SPECIMENS: MAINE: PENOBSCOT CO., Bangor, Fer- nald 13871 (NEBC). NEW HAMPSHIRE: BELKNAP CO., Laconia, Fernald 15680 (NEBC). arenicola X semisetosus. (Tentative, needs further study) ; Arching or trailing ; leaves chartaceous, soft-velvety beneath; armature of stout, hooked prickles; inflorescence of the setosus type, axis with slender prickles and. glands. REPRESENTATIVE SPECIMEN: NEW HAMPSHIRE: ROCKINGHAM CO., Seabrook, Steele 2770 (STEELE). 1970] Rubus Hybrids — Steele and Hodgdon 245 hispidus X setosus (R. adjacens, R. jacens, F. trifrons). Doming or trailing; leaflets 3 to 5, sublustrous ; armature of prickles, bristles and glands, from 2000-3000 per dm; inflorescence with some glands on axis. REPRESENTATIVE SPECIMENS: MAINE: LINCOLN CO., Monhegan Island, Hodgdon 12773 (NHA); Boothbay, Hodgdon 12272 (NHA). NEW HAMPSHIRE: CARROLL CO., Sandwich, Steele July 6, 1960 (NHA); Tamworth, Steele, Aug. 11, 1961 (NHA); GRAFTON CO., Thornton, Hodgdon & Steele 11345 (NHA); BELKNAP CO., Alton, Hodgdon & Steele 12193 (NHA); MERRIMACK CO., Newbury, Hodgdon & Steele 15484 (NHA); Pittsfield, Hodgdon & Steele 12790 (NHA); Danbury, Hodgdon & Steele 12789 (NHA); HILLSBORO CO., Litchfield, Hodgdon & Steele 16395 (NHA). MASSACHUSETTS: NORFOLK CO., Medway, Hodg- don & Steele, Sept. 2, 1964 (NHA); WORCESTER CO., Sterling, Hodgdon & Steele, July 13, 1964 (NHA). The cross between Rubus hispidus and R. setosus has been discussed earlier by us (1963) and is probably the most frequently occurring of any blackberry hybrid in New England. hispidus X vermontanus. Resembles hispidus X setosus but prickles stiffer. Trailing to arching; leaflets 3-5, larger than hispidus; compound leaves averaging 8 > 8 cm, sub- lustrous, subcoriaceous, glabrous both sides; armature of glands, bristles and definite short, stiff, hooked prickles. REPRESENTATIVE SPECIMENS: MAINE: HANCOCK CO., Somesville, Mt. Desert Island, Rand sn. (NEBC); LINCOLN CO., Southport, Hodgdon 7224 (NHA). NEW HAMPSHIRE: CARROLL CO., Tamworth, Steele 3992 (STEELE) ; GRAFTON CO., Waterville, Steele 2922 (STEELE) ; STRAFFORD co., Barrington, Hodgdon & Steele 15560 (9 collections, both putative parents present) (NHA), Rochester, Hodgdon 3461 (NHA); MERRIMACK co., Wilmot, Hodgdon & Steele 9424 (NHA); ROCKINGHAM CO., North- wood, Hodgdon & Steele 11212 (NHA); Nottingham, Hodgdon 9663 (NHA); Hampton Falls, Hodgdon & Steele 11213 (NH A). VERMONT: ESSEX CO., Ferdinand, Pease 25694 (NEBC); ADDISON CO., Leicester, Brainerd sn. (NEBC). MASSACHUSETTS: MIDDLESEX CO., Carlisle, Hodgdon & Steele 15563 (NHA). hispidus X allegheniensis (K. permixtus). Trailing or doming; leaves subcoriaceous, lustrous, pubescent under- neath, leaflets ovate or elliptic, 6-8 cm long, compound leaf averaging 13 cm X 13 cm; canes 2-4 mm in diameter; 246 Rhodora [Vol. 72 armature of short, stiff, hooked prickles and glands inter- mixed; inflorescence small to medium, often with juicy fruits. REPRESENTATIVE SPECIMENS: NEW HAMPSHIRE: CARROLL co., Tam- worth, Steele sn. (STEELE) : ROCKINGHAM co., Northwood, Hodgdon & Steele 11318 (NHA). setosus X vermontanus (R. Groutianus, Parlinii, uni- vocus). Growth habit and aspect like setosus and ver- montanus; armature of glands, bristles of various lengths and at least some acicular prickles resistant to touch and numerous (100), 1000-3000 per dm. Usually resembling setosus more than vermontanus; distinguished from setosus X elegantulus only by presence of putative parents. REPRESENTATIVE SPECIMENS: MAINE: PISCATAQUIS CO., Dover Fox- croft, Fernald (NEBC); YORK CO., York, Steele 4055 (NHA). NEW HAMPSHIRE: Coos co., Columbia, Hodgdon & Rhoades 15666 (NHA); Randolph, Hodgdon 11178 (NHA); Groveton, Hodgdon 12198 (NHA); Northumberland, Hodgdon & Steele 11179 (NHA); CARROLL CO., Sand- wich, Steele 4044 (NHA); Wakefield, Steele 4053 (NHA); Tuftonboro, Hodgdon & Steele 12268 (NHA); GRAFTON CO., Waterville, Hodgdon & Steele 11175 (NHA); Orange, Hodgdon & Steele 11268 (NHA); Graf- ton, Steele 3403 (STEELE); STRAFFORD co., Barrington, Hodgdon & Steele 15540 (NHA); Madbury, Hodgdon 14567 (NHA); BELKNAP CO., Alton, Hodgdon & Steele 12190 (NHA); MERRIMACK CO., Pittsfield, Steele 4057 (NHA); ROCKINGHAM co., Northwood, Hodgdon & Steele 11191 (NHA); Hampton Falls, Steele 3434 (STEELE) ; HILLSBORO CO., Sharon, Hodgdon 7237 (NHA). VERMONT: WINDHAM co., Westminster, Steele 3478 (STEELE) ; ESSEX CO., Victory, Pease 28414 (NEBC). CON- NECTICUT: TOLLAND CO., Tolland, Steele 3882 (STEELE) ; HARTFORD CO., Berlin, Steele 3881 (STEELE). setosus X semisetosus. Resembles setosus but with thin pubescence and small prickles on veins on under side of leaves. REPRESENTATIVE SPECIMEN: MASSACHUSETTS: NORFOLK co., Med- way, Steele 3946 (STEELE). setosus X elegantulus. Doming to arching, 2-4 ft, leaflets rather large, like elegantulus, leaflets ovate to lanceolate, subchartaceous to membranaceous ; armature of sparse to abundant thin prickles and bristles, often with glands. Very 1970] Rubus Hybrids — Steele and Hodgdon 247 close to setosus X vermontanus ; best distinguished from it by presence of putative parents. REPRESENTATIVE SPECIMENS: MAINE: LINCOLN CO., Ocean Point, Fassett 814 (NEBC). NEW HAMPSHIRE: Coos CO. Dummer, Steele 2952 (STEELE); CARROLL CO., Tuftonboro, Steele 3784 (STEELE) ; Tam- worth, Steele 2837 (STEELE); SULLIVAN CO., Sunapee, Steele & 'Hodg- don 4077 (STEELE). setosus X allegheniensis (R. frondisentis, sceleratus, aculiferus). Arching to erect, 2-4(5) ft; armature of straight strong prickles, setose bristles and glandular hairs ; leaves membranaceous to subchartaceous, resembling alle- gheniensis, with slight to abundant pubescence underneath ; axis of inflorescence pubescent and glandular; inflorescence smaller than typical allegheniensis; fruit poor and often undeveloped. REPRESENTATIVE SPECIMENS: MAINE: KNOX CO., Isle au Haut, Kid- der 8/29/19, (NEBC). NEW HAMPSHIRE: COOS CO., Northumberland, Fernald & Pease (NEBC) ; 15779 (NEBC); Randolph, Fernald & Pease 15781 (NEBC); Dummer, Hodgdon & Steele 11280 (NAA); CARROLL co., Tamworth, Steele 704 (STEELE); N. Conway, Steele 4045 (STEELE); Sandwich, Steele 4043 (STEELE); GRAFTON CO., N. Wood- stock, Hodgdon & Steele 11304 (NHA); Woodstock, Hodgdon & Steele 12374 (NHA); Lisbon, Steele 3537 (STEELE) ; MERRIMACK CO., Hooksett, Hodgdon 7213 (NHA); ROCKINGHAM CO., Rye, Steele 4067 (STEELE) ; Seabrook, Steele 2771 (STEELE); VERMONT: WINDHAM CO., Dum- merston, Brainerd 269001 (NEBC). vermontanus X semisctosus. Resembles semisetosus but pubescence thin or present only on nerves. REPRESENTATIVE SPECIMENS: MASSACHUSETTS: ESSEX CO., Newbury- port, Steele 3836 (STEELE). CONNECTICUT: Fairfield Co., New Canaan, Steele 3948 (STEELE). vermontanus X elegantulus. This hybrid undoubtedly occurs often but the 2 species are so similar at times that the cross can be identified positively only by the presence of both putative parents. REPRESENTATIVE SPECIMENS: NEW HAMPSHIRE: GRAFTON CO., North Woodstock, Steele 2949 (STEELE) ; Thornton, Steele 2929 (STEELE). 248 Rhodora [Vol. 72 vermontanus X allegheniensis. Arching, 2-4 ft; leaves medium to large, pubescent underneath; canes 3-5 mm in diameter; armature of slender prickles, often with glands intermixed, prickles shorter and thinner than typical alle- gheniensis; inflorescence like allegheniensis but somewhat smaller and poor-fruiting, axis with few to many glands. REPRESENTATIVE SPECIMENS: MAINE: LINCOLN CO. Boothbay, Hodgdon 12261 (NHA). NEW HAMPSHIRE: CARROLL CO., Sandwich, Steele 4035 (NHA); Tamworth, Steele 3804 (STEELE); Bartlett, Steele 3996 (STEELE); GRAFTON CO., Thornton, M. L. Fernald 15710 (NEBC) ; Woodstock, Steele 2927 (STEELE); STRAFFORD CO., Barrington, Hodg- don & Steele 15539 (NHA). semisetosus X frondosus. Arching, 2-4 ft; leaves of me- dium-size, compound leaf over 14 cm across, pubescent underneath; canes 3-5 mm in diameter; prickles thin, 2-4 mm long; numerous, up to 30 per dm; inflorescence of the vermontanus type, axis pubescent, no glands; fruit poor. REPRESENTATIVE SPECIMEN: MASSACHUSETTS: NORFOLK CO., Millis, Hodgdon & Steele 18290 (NHA). canadensis X elegantulus. Undoubtedly occurs often, but parents are too closely related to identify this hybrid unless both putative parents are present. REPRESENTATIVE SPECIMEN: NEW HAMPSHIRE: GRAFTON CO., Lincoln (Norcross Pond 3000 ft), Steele 3711 (STEELE). canadensis X allegheniensis. Leaves like allegheniensis, pubescent underneath; armature of sparse, short, thin prickles; axis of inflorescence with occasional glands. Re- sembles a state of pensilvanicus. REPRESENTATIVE SPECIMENS: MAINE: PISCATAQUIS CO., Dover, Fer- nald 9394 (NEBC). NEW HAMPSHIRE: CARROLL CO., Madison, Steele 3991 (STEELE); Tamworth, Steele 3967 (STEELE). canadensis X pensilvanicus. Leaves as in canadensis but with light pubescence underneath or pubescent on nerves; armature of sparse, thin prickles, 0-7 per dm; axis of in- florescence pubescent with occasional glands. REPRESENTATIVE SPECIMENS: MAINE: LINCOLN CO., Boothbay, Fas- sett 480 (NEBC). NEW HAMPSHIRE: COOS CO. Gorham, Fernold & 1970] Rubus Hybrids — Steele and Hodgdon. 249 Pease 15672 (NEBC) ; Errol, Fernald & Pease 15674 (NEEC); CARROLL co., Sandwich, Steele 2971 (STEELE); GRAFTON CO., Bath, Fernald 15692 (NEBC); Lincoln, Fernald 15730 (NEBC). elegantulus X allegheniensis. Leaves pubescent, resem- bling those of allegheniensis, subchartaceous ; prickles thin, mumerous, resembling those of elegantulus ; axis of inflores- cence with glands. REPRESENTATIVE SPECIMEN: NEW HAMPSHIRE: CARROLL CO., Sand- wich, Steele 3778 (STEELE). allegheniensis X pensilvanicus. Leaves with sparse pubescence ; armature of scattered, short, thin prickles ; axis of inflorescence with some glands. REPRESENTATIVE SPECIMENS: MAINE: ANDROSCOGGIN co., Poland, Fur- bish 1895 sn. (NEBC) ; KNOX CO., Matinicus, Long 7/1/20 sn. (NEBC) ; PISCATAQUIS CO., Dover, G. P. Fernald 115 (NEBC). NEW HAMPSHIRE: STRAFFORD CO., Lee, Hodgdon 1263 (NHA); ROCKINGHAM CO., Hampton, Steele 3516 (STEELE). allegheniensis X frondosus. Leaves pubescent beneath, of medium size, subchartaceous, with well-marked veins; armature of stout prickles; axis of inflorescence with scat- tered glands. REPRESENTATIVE SPECIMEN: RHODE ISLAND: WASHINGTON CO., Block Island, Champlin sn. (NEBC). This is probably a frequent hybrid within the range of both parents. In summary we make the following appraisal of hy- bridization-potential in Rubus subgenus Eubatus in New England. The following species hybridize frequently: Rubus hispidus, R. setosus, R. vermontanus, R. semisetosus, R. alle- gheniensis, R. pensilvanicus and R. frondosus. The following species often placed in the Series Flagel- lares presumably hybridize easily amongst themselves but less commonly with other species: Rubus flagellaris, R. recurvicaulis, R. Jaysmithii, R. arenicola and R. Enslenü. There are some species that tend to hybridize rarely: Rubus canadensis and R. elegantulus. 250 Rhodora [Vol. 72 Mostly because of their scarcity we know little about the hybridizing capacity of Rubus argutus and R. cuneifolius. ST, MARY’S-IN-THE-MOUNTAINS LITTLETON, NEW HAMPSHIRE 03561 DEPARTMENT OF BOTANY UNIVERSITY OF NEW HAMPSHIRE, DURHAM 03824 CHRYSANTHEMUM LACUSTRE BROTERO NEW TO GRAY’S MANUAL RANGE: During a collecting trip on August 15, 1968, along the western edge of Back Bay, Vir- ginia Beach, Virginia, (formerly Princess Anne County) in the southeastern corner of the state, two specimens of an unidentified composite were secured. The plants, one in flower, the other rather advanced in age, were determined by Dr. A. M. Harvill, Jr. to be Chrysanthemum lacustre Brotero, a species native to Portugal (Bailey, L. H. 1924. Manual of Cultivated Plants.) They were found by a roadside where the grassy shoulder bordered a marsh near Mill Dam Landing. Although a new record for Virginia, as well as the range of Gray’s Manual, this plant has been reported as infre- quent in old fields and ditches in Brunswick, Chowan, and Stanly Counties, North Carolina, and Abbeville County, South Carolina. (Radford, Ahles, and Bell. 1968. Manual of the Vascular Flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill, N. C.) The specimens are deposited in the herbarium of Long- wood College, Virginia. CHARLES E. STEVENS 615 PRESTON PLACE CHARLOTTESVILLE, VIRGINIA 22903 SOIL PREFERENCES AND VARIATION IN FLAVONOID PIGMENTS IN SPECIES OF ASTER WARREN G. ABRAHAMSON AND OTTO T. SOLBRIG A plant has certain environmental limits within which it can operate with greater or lesser success. The character- istics of the phenotype will determine the relative success of the plant. The environment is not static, but variable both in time and in space. For a species to occupy more than a point in time and in space, it has to have the ability to op- erate successfully under different environmental conditions. Two major mechanisms help the plant cope with a variable environment: (1) adjusting its phenotype and its metab- olism through physiological and developmental responses so that the phenotype be as close to optimum as possible for the prevailing situation; (2) adapting genetically through the production of a great array of diverse genotypes and an excess of progeny, from which presumably only the best suited will grow to maturity. The purpose of these studies is to determine the various facets involved in the adaptation of a species to con- ditions that vary in space and in time. The dilemma for the investigator is how to establish both the environmental factors that limit the distribution of a species, and also the phenotypical and genetical characteristics that make the distribution possible, given that neither the environment nor the morphological and genetical characteristics are con- stant in time and space. One classical technique is that of transplant experiments, Population samples are grown under a relatively uniform environment, and compared to one another. The differences that are found are presumably responsible for the capacity of the two populations to grow in different habitats. There are several problems in this method that will be discussed, but in general it has pro- vided interesting information. The genus Aster is widespread, common, and taxonom- ically difficult. In the northeastern United States there is 251 252 Rhodora [Vol. 72 SCALE IN MILES (i N PIN (See Collection Data on page 262) ample evidence of repeated hybridization (Wetmore and Delisle, 1939; Uttal, 1962), as well as polyploidy (Van Faasen, 1963; Solbrig, 1967). Avers (1953a, b) was able to document both hybridization and polyploidy in the hetero- phylli group, and also some degree of chromosomal homol- ogy, crossing relationships and possible phylogeny. For this reason it was felt that this group would be amenable to more in depth ecological studies, than any other group of Asters. MATERIALS AND METHODS A transect was established nearly due west from Boston to the general area of the corner of the states of Pennsyl- vania, New York, and Ohio (fig. 1). Along this transect a population was sampled every twenty to fifty miles. The sample was restricted to populations belonging to the heter- ophylli group of Aster (Fernald, 1950; Avers, 1953b). Each sample consisted of at least ten plants and not more than twenty, and these were selected at random. The plants were brought to the Botanical Gardens of the University of Mich- igan in Ann Arbor, Michigan where they were broken into two clones and both halves were then planted in a uniform garden following a randomized design. In addition a soil 1970] Asters — Abrahamson and Solbrig 253 sample was gathered in a plastic bag at most of the lo- calities and brought to the laboratory for analysis. Soil Analysis Soil samples were characterized as to pH, moisture re- tention capacity, nitrate content, and mechanical composi- tion. The hydrometer method was used for the mechanical analysis. Moisture retention was obtained using a low pres- sure ceramic plate extractor; readings at 0.8 atmospheres tension (12p.s.i.) and at 0.1 atmospheres were secured. The pH values were determined by mixing 10 grams of fresh soil and 25 ml of distilled water; readings were taken one and three hours after the mixing of the soil and the water. Soil nitrate content was determined following the phen- oldisulphonic acid method (Metson, 1956). Qualitative Studies of Pigments The fresh leaves were diced and placed into 99% MeOH and 1% HCl in the dark for 24 hours. Only mature basal leaves were used during the study. The plant material was separated from the solvent and secondary compounds by filtration through Whatman No. 1 filter paper. Two hun- dred micro-liters of this extract was spotted onto either thin-layer plates or Whatman No. 3 MM chromatography paper (46 X 57 cm sheets). Due to the fact that better separations were obtained with the paper technique the majority of the work was done in this way. The chromato- grams were developed (descending) using tertiary butyl alcohol, acetic acid, and water (3:1:1) in the first direc- tion. Caution was exercised to insure that fresh solvent was utilized since the above solvent undergoes esterification reactions rapidly. After thorough drying, the chromato- grams were again placed in the chromatocab and developed in the second dimension with 15% acetic acid and water. All the chromatography was done in an air-conditioned, dark room. The temperature was maintained as close to 21° C as possible. The compounds were tentatively identified by their char- acteristics under both short-wave and long-wave ultra- 254 Rhodora [Vol. 72 violet light, and by their behavior in the presence of ammonia vapor. Certain phenolic compounds (whitish or bluish in UV) and certain flavonoids (dark or yellowish in UV) appeared in all populations studied while other com- pounds appeared only off and on. Quantitative Studies of Anthocyanins The anthocyanin production studies were completed using leaf-dises from each plant, These were punched from the mature basal leaves with a cork borer (#6) and grown for seven days on a 0.1 M sucrose solution under constant light, at 209 C. The discs were sliced and extracted in a 1% HCl solution for a week on a shaker at approximately 69 C. The amount of anthocyanin was determined colori- metrically using the method of Thiman and Edmonson (1949). RESULTS A. Soil Analysis. 'The results of the soil ahalysis can be seen in table 1. Although the area traversed is one of varied soils, including some that are rich in silts and clays, all the populations sampled, regardless of species, grew in sandy or sandy-loam soils, with a neutral or more commonly acid pH, low organic matter and low nitrate concentration, al- though there was considerable variation in this last char- acter. The uniformity of the soil type cannot be assigned to chance and must indicate a preference for this type of soil by the species sampled. B. Secondary Compounds. Although there was variation in the kind and number of compounds present, in general there was no significant variation that could be correlated between populations. There was a great deal more physio- logical variation due to development of the plant and en- vironmental variation within the experimental plot than variation due to genetical causes. Table 2 summarizes the results of the analysis for all the populations investigated. Of the 25 compounds that were evident on the chromatographs, 6 were present in all 83 plants that were analyzed, 10 compounds were present in over 60 of the analyzed plants and 9 compounds were 1970] Asters — Abrahamson and Solbrig 255 truly variable. That is they were absent from entire popu- lations and when present in a population often only some plants would show them. No compound was species specific. Tests were run on both replicates of a clone, and in addi- tion the same plants were tested at different times of the year. The results are summarized in table 3. It can be seen that the variations are relatively great. Still greater differences can be obtained if plants are grown in the green- house. When this is done, certain spots fail to show up on the chromatogram. A further source of variation is the technique itself. If a larger or lesser amount of extract is used, the number of spots can be sometimes varied accord- ingly. | C. Anthocyanin Production. The quantitative studies were restricted to measuring the total amount of anthocyanin that the species were capable of producing. The anthocyanin was identified as cyanidin. The object was to see whether (1) different species and populations varied as to the amount of anthocyanin they can produce, and (2) the variations were species specific or if they could be cor- related with environmental factors. Preliminary studies (Solbrig, 1966) indicated that there were correlations be- tween altitude above sea level and latitude where the popu- lations grew and that there were also variations between species as to their anthocyanin production. The results of the present study are summarized in table 4. It can be seen that indeed there are variations be- tween populations and that they are not species specific. There is appreciable difference that can be ascribed to site. It should be remembered, however, that although there are differences in elevation between sites, all the populations grew at about the same latitude. DISCUSSION Uniform garden studies are designed to eliminate environ- mental differences between localities in the shaping of the phenotype. Any differences between phenotypes growing in a uniform environment can therefore be ascribed to gen- etic differences. It is conceivable, however, that dissimilar 256 Pop. No. 426 427 428 450 431 432 433 434 435 436 439 440 441 442 445 444 445 446 447 448 449 450 451 452 Nitrate N..ppm/wt of fresh soil bo e P to 89.35 111.6 99.2 5.875 41.6 31.54 2.3 80.0 17.6 79.0 14.75 212.65 227.4 140.76 b 49.8 80.5 20.0 50.6 45.6 41.65 89.225 34.0 or 35.5 Table 1. Moisture % a-d soil 1.616 Rhodora [Vol. 72 Analysis of 24 soil samples. + + E E: g: - 3. Bua B Eg S9 E së SE = “as S28 SP £ ss T$ Pon Vn n h £ n n n 8 n Bey BEy ba — Evo gy By aq o BZN Oo p^ a ® Aw oa o o . ot SSS Sok Gs aS Sk RS 67.9 48.1 21.8 3.9 28.0 26.8 47.8 26.9 13.8 3.2 25.2 22.6 240 13.9 10.8 25.8 71.8 11.4 44,2 16.1 6.6 2.0 25.1 29.5 40.9 19.5 6.4 7.3 28.8 32.7 26.9 9.6 6.2 15.1 54.0 20.2 48.0 29.2 12.0 9.6 23.8 13.8 16.9 8.6 5.0 31.8 55.4 20.6 40.6 25.9 8.9 19.5 29.4 15.6 17.6 8.4 3.7 36.6 62.0 138.4 41.3 23.6 11.8 32.6 45.2 104 33.6 22.9 8.2 8.9 39.0 25.8 34.0 18.2 3.7 14.9 22.5 25.1 34.5 20.4 8.0 18.7 27.7 21.7 37.9 19.7 7.6 52.6 22.9 20.1 37.6 22.1 8.4 27.0 19.7 16.7 40.9 29.6 9.3 8.1 5.3 9.5 44.8 24.2 8.8 33.8 16.1 14.3 30.2 16.8 9.0 26.8 37.0 15.0 37.32 2.125 8.947 19.91 33.63 19.22 0.6094 137.95 83.43 16.86 189.29 289.79 44.56 0.7806 11.745 9.134 4.106 13.76 17.023 6.675 1970] Asters — Abrahamson and Solbrig 257 Color E uu d. ur d E G i 4 quM ES. pi i FEE > 54.8 36.8 lee 5.15 10 Yr 2/2 Sandy loam/loam 47.8 42.0 10.2 4.45 10 Yr 4/2 Loam 83.2 11.0 5.8 4.72 2.5 Yr 4/2 Loamy sand 54.6 31.2 8.2 5.15 10 Yr 4/2 Sandy loam 61.5 27.0 11.2 5.72 2.5 Yr 5/2 Sandy loam 74.2 17.6 8.2 7.20 10 Yr 1/2 Sandy loam 31.6 45.8 1.66 6.20 10 Yr 5/2 Loam -— — -= 6.43 5 Yr 4/3 — — — -- 7.37 2.5 Yr 6/2 — 76.0 14.8 9.2 7.21 5 Yr 5⁄2 Sandy/sandy loam 45.0 39.8 15.2 5.46 5 Yr 4/2 Loam 75.4 12.8 11.8 7.05 5 Yr 5/2 Sandy loam 55.6 30.2 14:2 5.80 7.5 Yr 4/2 Sazdy loam 64.8 26.0 9.2 4.78 10 Yr 3/3 Sandy loam 47.6 39.6 12.8 6.92 T0 m oils Loam 49.4 29.0 21-6 6.89 10 Yr 4/3 Loam 48.0 32.8 17.2 5.82 5 Yr 4/2 boim — — — 6.58 10 Yr 3⁄1 — 36.4 40.4 23.2 6.50 10 Yr 5⁄2 Loam/clay/loam 14.8 46.8 29.2 5.52 7.5 Yr 4/2 Clay loam 30.4 46.4 23.2 TAAl JO Yr 3/2 Foam — — — 5.01 10 Yr 3/2 — E — — 5.54 10 Yr 6/2 — 52.0 31.8 16.2 6.38 10 Yr 5/2 Loam/sandy loam 52.85 32.86 14.23 6.04 296.26 133.37 40.88 0.805 17.231 11.549 6.394 0.8972 Table 2 Summary of chromatographic analysis 258 Rhodora [Vol. 72 HHH EL = O L = 6 O S y ej Se Le pepe j Oe L O = L = > gO SH SHSM SOR j oO om > HHH ++ HPS + e e= ° + HM + “> $e + Tee e s + =. + = Š ° =. + > eere e esse SOSH MH c e e c +> +++++++i+i+i++ii+ + + ++++++++i++i+ii+i+i+++++Pem +++++++++i+°S+i+++ © exer erp e e oe a S %' ° ese 00 6 C +++ +° SOH BcbLBBAV TALI LY-XA «GLO GL KXR ++++++T“% e e s L ttot 13 14 15 16 17 18 19 20 21 22 23 24 25 a + ae + ++ T+ +° ++i+TiiTi+i+ T+ + +Y Shera 0 Oe tree SSCL c< S c coco Spot No. 9 10 11 12 Vie y I + all plants of the population contained the compound ? Soper Ke SCH ee SCSt r T+++%—% +e ++++° S+ 8 — -— = + P P: T+++++i+iiTii+i++ + + + + metttrtttettett++ ttt t+ttt+tt++++ e+e et t+ettt+tet+O+ pep tere TL a ++ ++ 9 +ettt++++++ttq¢++444+ N ++% ++++++ilii+ii+ii+i+i-++i"” H ++++++iL+iUVu+Ti+i+iii+ii-T-+ some plants of the population did and others did not contain the compound 0 no plants appeared to have the compound PASIL raqunN I over 60 of the 83 plants tested showed the compound P all 83 plants showed the compound V zy t- C n G co Mi 09 Ç n CG io tO t- 00 G; C Gq x A CS c9 CQ OOM CO XB + XM MM — OMM OM XB aD aD am Go + < + HHH HHA SSG variable compound Asters — Abrahamson and Solbrig 259 1970] +° ++ + ° ++ ° Ve) N = N + ++ ++ ++ ++ + ° € uc] e P u + + + + + + + + + [an] N N ++t+++4+°+4 N - Spa yas + + Ó + 0 + + + + + + + + + + + + + 0% 6I + + + + + + + + t+ + + t + + + + + + oo m t> v + + + + + + + ° + 9I GI FI e an] sjueurdid jue[d dures oy} WOIF soA€9[ FUILI JA z 89/11/8 89/TT/8 89/11/8 + 89/61/6 + 89/9/8 4 4 + 89/11/8 4 n + Fore ea p Po k 4 o. eo c cO e o wmm ce oí a dc cct 44+ 4+ + ok oF re fo a o ~- ++tr tt + t+ 4+ + ete ot a a mp o4 B us muc. QU Qm oum qoem er DP Xl NOISE es n e i rm m © — oo eo Ves) <+ oD I uny əqd 'oN yodg JeƏ[ ur uorjerieA perodwəa} jo Aieurung € OTIdV L x9I MOY f-28F x9I MOY P-SEP SI MOY P-ZEP 9T MOY -ZEF GT MOY &-GEP juv[q 3 ‘dog 260 Rhodora [Vol. 72 TABLE 4 Total Anthocyanins produced by 10 leaf discs Pop. Species Col. Site Antho. ave. value* 426 cordifolius (1) 138 428 undulatus (3) 124 429 cordifolius (4) 352 430 cordifolius (5) 129 431 cordifolius (6) 59 433 cordifolius (8) 141 434 cordifolius (9) 148 435 ciliolatus (9) 140 436 ciliolatus (9) 62 444 lowreianus (14) 150 447 cordifolius (16) 55 448 cordifolius (17) 98 449 cordifolius (18) 169 450 cordifolius (19) 93 451 cordifolius (19) 65 454 cordifolius (20) 169 *Readings in Klett units using #54 filter and stendardized for volume. 1970] Asters — Abrahamson and Solbrig 261 genotypes will react to the same environment by producing similar phenotypes, the well known phenomenon of canal- ization. This problem to a certain degree puts into question some of the conclusions arrived at in any transplant ex- periment, The most striking feature in this study is the uniformity of the populations, considering that we are dealing with four species and three levels of ploidy (9, 13, 36). All plants in their native environment grew in very similar soils. This indicates a definite preference by these species for loose sandy soils, a fact not previously established ex- perimentally. More interesting, however, from a taxonomic point of view are the studies on the secondary compounds. The seasonal and environmental variation found should be a fair warning to those attempting to use similar methods of chromatography of secondary compounds for taxonomic purposes using random field samples or herbarium samples. At least in the heterophylli Aster secondary compounds vary greatly during the growing season, and are also af- fected by the particular environment of the plant. The en- vironment affects both the number as well as the intensity of spots produced. It appears that during the evolution of the heterophylli Aster little or no differentiation has occurred ir. soil prefer- ence, at least in the N.E. United States. There seems to have been little change in the alcohol soluble secondary compounds also. There have been no species specific changes in the amount of anthocyanin that the various species pro- duce. We would like to thank Mrs. Jennifer Ward for able laboratory assistance, and Mr. Marc Rosenstein for the identification of the anthocyanin. The work was performed with the aid of a grant from the National Science Founda- tion, which is gratefully acknowledged. DEPARTMENT OF BIOLOGY AND GRAY HERBARIUM HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS 02138 262 Rhodora [Vol. 72 LITERATURE CITED AVERS, C. 1953a. Biosystematic studies in Aster. I. Crossing re- lationships in the heterophylli. Amer. Jour. Bot. 40: 669-675. - 1953b. Biosystematic studies in Aster. II. Isolating mechanisms and some phylogenetic considerations. Evolution 7: 317-327. FERNALD, M. L. 1950. Gray’s Manual of Botany. 8th Ed. Boston METSON, A. J., 1956. Methods of Chemical Analysis for Soil Survey Samples. Dsiro, New Zealand. SoLBRIG, O. T. 1967. Some aspects of Compositae of evolutionary interest. Taxon 16: 304-307. THIMAN, K. and Y. H. EpMonson, 1949. The Biogenesis of An- thocyanins I. General nutritional conditions leading to anthocyanin formation. Arch. Biochemistry 22: 33-52. UTTAL, L. J. 1962. Synthesis of Aster herveyi. Rhodora 64: 113-117. VAN FAASEN, P. 1963. Cytotaxonomie studies in Michigan Asters. Michigan Botanist 2: 17-27. WETMORE, R. H. and A. L. DELISLE, 1939. Studies in the genetics and cytology of two species in the genus Aster and their poly- morphy in nature. Am. Jour. Bot. 26: 1-12. Collection Data Sites 1. Mass. on Rte. 111. 4.7 miles N. of Jet with Rte. 110 Harvard Township. 67-426 Aster cordifolius (Solbrig 4046) ; Aster cordi- folius (Solbrig 4047). Mass., Hwy 31, 1 mile south of Hwy 12. Plants growing under oak, beech, and maple. 67-428 Aster undulatus (Solbrig 4048). 3. Mass. 5 miles east of Hardwick, under maple forest. 67-429. Aster cordifolius (Solbrig 4049). 4. Mass., Jet of 9 and 202. Growing under a maple-oak forest. 67- 430. Aster cordifolius (Solbrig 4050). 5. Mass. Hwy 9, 1 mile east of Williamsburg, under an oak-maple forest. 67-431. Aster cordifolius (Solbrig 4051). 6. Mass., Hwy 9, 1.9 miles east of Windsor. Growing under a mixed hardwood-softwood forest, mostly at the more illuminated edges. 64-432. Aster cordifolius (Solbrig 4052). T. Mass., Hwy 20, 8.1 miles west of Pittsfield, at the edge of a maple-oak forest. 67-433. Aster cordifolius (Solbrig 4053). Do 1970] Asters — Abrahamson and Solbrig 263 8. 10. 11. 12. 13. 14. 16. y 18. 19. New York, Rte. 23, 1.2 miles east of East Windham. Plants growing in mixed maple, elm, and oak forest. 67-434. Aster cordifolius (Solbrig 4054) ; 67-435. Aster ciliolatus Lindl. (Sol- brig 4055); 67-436. Aster ciliolatus Lindl. (Solbrig 4056); 67- 437. Aster acuminatus Lindl. (Solbrig 4057); 67-438 Aster acuminatus Lindl. (Solbrig 4058). New York, Hwy 10, 4.6 miles south of Stamford. Growing on an embankment under a few maples at the edge of a field. 67- 439. Aster cordifolius L. (Solbrig 4059). New York, Hwy 28, 6.1 miles east of the Jet with 7B. 67-440. Aster cordifolius (Solbrig 4060). New York, on Hwy 7, 1.2 miles south of Riverside. Growing in a weedy area with goldenrods, brambles, rhus, etc. 67-441. Aster cordifolius (Solbrig 4061). New York, Hwy 17, 8.7 miles east of Oswego. (Jct Hwy 283) Growing in a weedy area under a few maple trees, 67-442. Aster cordifolius (Solbrig 4062). New York, Hwy 17 east of Elmira (3.2 miles east of Chemniz) Growing on the edge of a disturbed woods. 67-443. Aster cordi- folius. (Solbrig 4063) ; 67-444. Aster lowreianus (Solbrig 4064) ; 67-445. Aster laevis (Solbrig 4065). New York, Hwy 17, 1.6 miles east of Addison. Growing at the edge of a beech-oak forest. 67-446. Aster cordifolius (Solbrig 4066). New York, west outskirts of Wellsville. Growing under maples and Crataegus. 67-447. Aster cordifolius (Solbrig 4067). New York, 3 miles south of the village of Whitehouse. Growing in woodland with a border of mixed maple and oak with many introduced shrubs. 67-448. Aster cordifolius (Solbrig 4068). Pennsylvania, on Hwy 770, 4.3 miles east of the Jet with 59. Growing in a secondary forest of oak, maple, beech, and hickory. 67-449. Aster cordifolius (Solbrig 4069). Pennsylvania, Hwy 27, 2 miles west of Pittsfield. Growing at the edge of an open field, in a ditch, and at the forest edge respec- tively. 67-450. Aster cordifolius L. (Solbrig 4070); 67-451. Aster sagitifolius (Solbrig 4071); 67-455. Aster acum natus (Solbrig 4073). Pennsylvania, Hwy 77, 1 mile south-west of Jet Hwy 8. Growing in small woods at the edge of the road. 67-454. Aster cordifolius (Solbrig 4074). BIOSYSTEMATICS OF SETCREASEA BREVIFOLIA S. A. FARUQI' AND K. L. MEHRA? Setcreasea brevifolia (Torr.) Schum, & Sydow is a highly polymorphic and poorly understood species distributed in northern Mexico and southern Texas. Rose (1899, 1903, 1911) and Matuda (1955) included several taxa as syn- onyms of S, brevifolia, without indicating their similarities or differences, The similarities between certain taxa are so strong that for almost sixty years S. ovata (Coulter) Faruqi, Celarier & Mehra (Syn. Tradescantia leiandra var. ovata Coulter, Contr. U. S. Nat. Herb., 1: 50. 1890) was regarded as a synonym of S. brevifolia (Torr.) Schum. & Sydow. Floral morphology and crossing data suggested that S. ovata Faruqi et al. is related to the S. pallida Rose — S. purpurea Boom group and that it is not in the direct evolutionary line of S. brevifolia (Faruqi et al., 1962). An- other taxon, Tradescantia speciosa Buckley, considered by Rose (1899, 1911) and Matuda (1955) as a synonym of S. brevifolia and treated as S. buckleyi Johnst. (Johnston, 1944) needs taxonomic consideration since no barriers to crossability exist between these taxa and the distinction between them is ecological and morphological (Faruqi and Mehra, 1966). Furthermore, a few other taxa have never been mentioned or included within S. brevifolia in spite of their resemblance to this species. One such taxon was col- lected from the Ottine Swamp of Gonzales Co. in Texas. Its plants were very small, hardly six inches tall and with very narrow leaves, A similar plant from an unknown locality was present in our living collection and it was crossed with S. brevifolia. The F, hybrid resembled other plants from our living culture and herbarium collections. One of the herbarium sheets resembling this hybrid was annotated with a name by Bartell, but it was never pub- ‘Permanent address: Senior Lecturer, Dept. of Botany, University of Karachi, Karachi, W. Pakistan. "Permanent address: Head, Division of Plant Improvement, Indian Grassland & Fodder Research Institute, Jhansi, U.P., India. 264 1970] Setcreasea — Faruqi and Mehra 265 lished. It is important to note that all of the foresaid taxa except S. ovata are characterized by narrow connectives and pubescent ovary. Whether these taxa should be in- cluded within S. brevifolia or should be treated as different species needs consideration. Besides, the origin of these variations should also be considered. Biosystematic studies were, therefore, undertaken on living and herbarium speci- mens and four varieties were recognized within S. brevifolia (Torr.) Schum. & Sydow. The results are reported in this paper. Setereasea brevifolia (Torr.) Schum. & Sydow, in Just, Bot. Jahrb. 27(1): 452. 1899 (1901). Syn. Tradescantia leiandra var. brevifolia Torr., in Emory, U. S. and Mex. Bound. Survey, 225. 1859. Zebrina (?) leiandra Clark, in DC. Monograph, Phan. 3: 318. 1881. Tradescantia leiandra Wats., Proc. Amer. Acad. 18: 167. 1883. Tradescantia brevifolia Rose, Contr. Nat. Herb. 3: 323, pl. 16. 1895. Treleasea brevifolia Rose, Contr. Nat. Herb. 5: 207. 1899. Neotreleasea brevifolia Rose, Contr. Nat. Herb. 8: 6. 1903. Stem 8-30 em in height; sepals 3; petals 3; stamens 6, epipetalous, connectives thick and narrow; and ovary pubes- cent. Four varieties were recognized within S. brevifolia. The stem is unbranched and the leaves are 3-4 cm long in var. 1- nanella. The stems are branched and the lengths of the leaves are 5-6 cm in var. 2- pulchella, 7-10 cm in var. 3- brevifolia and 10-13 cm in var. 4- buckleyi. 1. S. brevifolia var. nanella Faruqi et Mehra, var. nov. Caule non ramoso, erecto, 10-20 cm alto; folia lanceolata vel elliptico — lanceolata, viridia, 3-4 cm longa, 1-2 cm lata. HOLOTYPE: U.S.A. TEXAS, GONZALES CO., Ottine Swamp, Cory 14581 (GH). MEXICO: D.F. MEXICO: Raynosa, Run- yon 20 (US) ; Clover 1932, cultivated (MICH). 266 Rhodora [Vol. 72 Cultivated: Setcreasea Taxon-1 (Mehra et al., 1961). Faruqi and Mehra (1967) reported that some kind of genetic barrier seems to exist between the varieties brevi- folia and nanella since only three hybrids could be obtained from forty pollinations attempted between them. Both varieties have 24 somatie chromosomes and exhibit similar cytological irregularities during meiosis (Mehra et al., 1961; Faruqi and Mehra, 1967). Binomial was a good fit to the distribution of multivalents during metaphase I of micro- sporogenesis in the var. brevifolia, indicating that all six groups of four chromosomes had an equal chance of pairing as a quadrivalent (Mehra et al., 1969). In the var. nanella, the binomial was not a good fit to the distribution of mul- tivalents, indicating differential pairing behaviour of the chromosomes during metaphase I of microsporogenesis (Mehra et al., 1969). Thus, chromosomal differentiation seems to have taken place in the evolution of the var. nanella, 2. S. brevifolia var. pulchella Faruqi & Mehra, var. nov. Caule ramoso, delicatulo, prostrato vel subascendente, 15- 25 cm longo; folia lanceolata, elliptico-ovata, viridia, 5-6 cm longa, 1.5 cm lata, HOLOTYPE: MEXICO: TAMAULIPAS, Cerro E] Palmer S. of Cruillas, Clover 13763 (MICH). D.F. MEXICO: Bartel 13763 (MICH); Clover 14920; cultivated number and the name of the collector not given (MICH). U.S.A.: TEXAS: HIDALGO CO., Lundell & Lundell 9944 (vs); Cory 20465 (GH); Corpus Christi, Meally 1894 (GH). Faruqi and Mehra (1966, 1967) reported that hybrids between varieties brevifolia and nanella resemble an un- described taxon (now called var. pulchella) represented in living culture and herbarium specimens. While the types of chromosomal irregularities were similar in the parents and hybrids, the binomial was not a good fit to the distribution of multivalents in one of the three hybrids (Mehra et al., 1969). It would suggest differential pairing behaviour of chromosomes in this hybrid. The survival of such chromo- 1970] Setcreasea — Faruqi and Mehra 267 some types with structural hybridity is enhanced owing to the vegetative mode of reproduction. 3. S. brevifolia var. brevifolia Syn. Tradescantia leiandra var. brevifolia. Torr., in Emory U. S. and Mex. Bound. Survey 225. 1859. Stem branched, prostrate or slightly ascending, about 30 em long; leaves ovate, purple-green, 7-10 cm long and 2-3 cm wide. HOLOTYPE: mountains near the mouth of the Pecos River, Oct. 19, 1852, Bigelow 1500 a (NY). MEXICO: Vic- toria, Palmer 601447 (F). U.S.A. TEXAS: Corpus Christi, Rose 1896 (GH), 839905 (GH); Nealy, 07.457 (F, GH, US) ; Heller 1427 (GH, MO, US) ; Mulley 1895 (US). CULTIVATED: Norton 1737019 (Mo) ; Rose, July 21, 1896 (GH, US). The type specimen is deposited in the N.Y. Botanical Garden Herbarium, but it is not in a condition that its floral morphology could be worked out. The type description mentions a pubescent ovary, but the sketch of the floral parts on the herbarium sheet shows glabrous ovary. Since there is no way to find out the actual position, the words from the description are taken as correct. 4. S. brevifolia var. buckleyi (Johnst.) Faruqi & Mehra, comb. nov. Syn. Tradescantia speciosa Buckl, Proc. Acad. Phila. 1862: 9. 1863, non L.f. or H.B.K. Setcreasea buckleyi Johnst, Jour. Arn. Arb. 25: 54. 1944. HOLOTYPE: U.S.A. TEXAS, Corpus Christi, Buckley, number not given on the sheet (Acad. of Natural Sciences, Philadelphia). MEXICO: TAMAULIPAS, near Santa Terasa, Johnston 5491 (UT); north of Almada, Johnston 5707 B (uT) ; Herman 13763; New Victoria, collector not known (UT). Johnston (1944) changed T'radescantia speciosa Buckl. to S. buckleyi and stated, “S. brevifolia is found from the Chisos and Davis Mts. to the lower Pecos River, and mor- 268 Rhodora [Vol. 72 phologically it is a low plant with coarse rhizomes; whereas S. buckleyi is a plant of the coastal region of Texas which has a paler corolla and a loosely branched, elongated, trail- ing stem.” The only difference betwecn this variety and var. brevi- folia is its larger more or less elliptic leaves, 10-13 cm long and 2.8-4.2 cm wide; longer internodes and many branches from the base forming mounds in its natural habitat, Under greenhouse conditions, however, the plants usually referred to as brevifolia do possess leaves as big as var. buckleyi and it is very difficult to differentiate between these varieties (Faruqi and Mehra, 1966). The inland populations of var. brevifolia exhibit shortening of leaves towards its northern area of distribution as compared to the southern area. This, however, does not seem to be the case with coastal plants of var. buckleyi (Faruqi and Mehra, 1966). Under natural conditions var. buckleyi is certainly taller than var. brevi- folia, Under greenhouse conditions it grows no taller than var, brevifolia and is slow growing and late flowering. Under laboratory conditions varieties brevifolia and buck- leyi could be crossed fairly easily and the seeds were pro- duced when either one was used as the female parent. Thus, there is no genetic barrier between these varieties and the distinction between them is ecological and morphological. But, one specimen (Johnston 5491) from north Texas re- sembled var. brevifolia in a few characters and var. buckleyi in others, indicating some gene exchange between these varieties. Since these taxa demonstrate some degree of morphological differences and have different ecological pref- erences, they only deserve varietal status. Mexico is the center of origin of the genus Setcreasea, and it is here that all of the primitive species such as S. tumida Rose, S. australis Rose, S. purpurea Boom and S. pallida Rose are distributed (Faruqi and Mehra, 1966). Primitive species of this genus have leaves larger than those of the advanced ones; by the same token a tendency towards the shortening of the shoot is also present in the advanced species. Thus, the leaves are larger and the stems are longer 1970] Setcreasea — Faruqi and Mehra 269 in the populations of S. brevifolia distributed in the south (i.e., northern Mexico and southern Texas) than those in the populations distributed in the north (i.e., mid Texas). Thus, the var. nanella seems to have originated from S. brevifolia var. brevifolia in the process of its northward migration followed by ecological adaptation. It seems that in the origin of variations in S. brevifolia ecological adapta- tion has played an important role when migration from warm climate, where growing seasons are long, to cold climate, where the seasons for its growth are short, took place. An absence of this type of variation from south to north in the coastal population (var. buckleyi) seems to be correlated with the mild climate of the coastal areas. In fact, the leaves became longer and broader in the coastal area. Experimental hybrids between varieties brevifolia and nanella resembled var. pulchella and their leaf lengths were intermediate between those of their parents. In Setcreasea brevifolia, the chromosome races with somatic chromosome numbers, 12 (Richardson, 1935), 23 (Celarier, 1955; Mehra et al., 1961), 24 (Darlington, 1929; Richardson, 1935, Mehra et al., 1961), 25 (Bose, 1962) and 56 (Anderson and Sax, 1936) have been reported. The materials of S. brevifolia used in this study had 23 (var. brevifolia) and 24 (vars. nanella, pulchella and buckleyi) somatic chromosomes. The previous reports on the chromo- some morphology indicated that changes in the chromosome structure are taking place in this species (Darlington, 1929; Richardson, 1935). The chromosome and genetic differen- tiation is not uniform in all six sets of four chromosomes in the var. nanella (Mehra et al., 1969). It is assumed that S. brevifolia is in an active stage of evolution, in which euploidy, aneuploidy, structural hybridity, and hybridiza- tion between its varieties seem to be playing a major part. The vegetative mode of reproduction and perennial habit seem to be responsible for the maintenance of varieties within this species. Through the forces of selection some of these varieties are fixed in certain ecological niches, and wherever possible self-incompatibility ensures interpopula- 270 Rhodora [Vol. 72 tion gene exchange through natural crossing. The range of morphological variation encountered is due to these factors, which in combination with each other produce an effect similar to that of facultative apomicts, exhibiting restricted gene exchange (Baker, 1953, 1959; Celarier et al., 1958; Faruqi, 1963; Faruqi and Mehra, 1968). Thanks are extended to the curators of the Gray Her- barium of Harvard University, Cambridge; United States National Herbarium, Washington, D.C.; and herbaria of the Missouri Botanical Garden, St. Louis; New York Bo- tanic Garden, N.Y.; Field Museum of Natural History, Chicago; Academy of Natural Sciences, Philadelphia; Uni- versity of Texas, Austin; and University of Michigan, Ann Arbor, for the loan of the herbarium specimens. Thanks are also due to Rev, Fr. H. Santapau, for providing the Latin diagnosis for the new varieties, and to Mr. K. C. Kanodia, for discussions. The facilities of laboratory and greenhouse were extended by the courtesy of the late Dr. W. W. Hansen, Chairman, Dept. of Botany, Oklahoma State University, Stillwater, Oklahoma, for which the authors are highly grateful. DEPARTMENT OF BOTANY AND PLANT PATHOLOGY OKLAHOMA STATE UNIVERSITY STILLWATER, OKLAHOMA 74074 LITERATURE CITED ANDERSON, E. and K. Sax. 1936. A cytological monograph of the American species of Tradescantia. Bot. Gaz. 97: 433-476. BAKER, H. G. 1953. Race formation and reproductive method in flowering plants. Symp. Soc, Exptl. Biol. 7: 114-143. 1959. Reproductive methods as factors in speciation in flowering plants. Cold Spring Harb. Symp. 24: 177-191. BosE, S. 1962. Cytotaxonomical studies in the genus Setereasea. Bull. Bot. Surv. India 4: 17-26. CELARIER, R. P. 1955. Cytology of the Tradescanticae. Bull. Torrey Bot. Club. 82: 30-38. , K. L. MEHRA and M. L. WoLF. 1958. Cytogeography of the Dichanthium annulatum complex. Brittonia 10: 59-72. 1970] Setcreasea — Faruqi and Mehra 271 DARLINGTON, C. D. 1929. Chromosome behaviour and structural hybridity in the Tradescantieae. Jour. Genet. 21: 207-286. FARUQI, S. A. 1963. The significance of apomixis in evolution and speciation of the Bothriochloininae. Biologia 9: 57-64. and K. L. MEHRA. 1966. A study of variation within Setcreasea brevifolia (Torr.) Sch. et Sydow and certain morpho- logically similar taxa. Rev. de Biol. 5: 87-294. and . 1967. Cytotaxonomical studies in the Setereasea brevifolia complex. Cytologia 32: 255-261. and . 1968. Mode of reproduction and its bearing on the genetic diversity in the genus Setcreasea. Port. Acta Biol (A.) 10: 321-326. » R. P. CELARIER and K. L. MEHRA. 1962. The tax- onomy of Setereasea ovata. Rhodora 64: 68-73. JOHNSTON, I. M. 1944. Plants of Northern Mexico, III. Jour. Arnold Arb, 25: 54. Matupa, E. 1955. Las Commelinaceas Mexicanas. Anal. Inst. Biol. Mex. 26: 303-432. MEHRA, K. L., S. A. FARUQI and R. P. CELARIER. 1961. A cytotax- onomical study of five species of Setcreasea. Phyton 17: 133-140. , P. R. SREENATH and S. A. FARUQI. 1969. Distribution of multivalents and genotypic control of chromosome pairing in Setereasea species and hybrids. Cytologia (in press). RICHARDSON, M. M. 1935. Setereasea brevifolia, a further example of polyploidy and structural hybridity in the Tradescantieae. Bot. Gaz. 97: 400-407. ROSE, J. N. 1899. Treleasea, A new genus of Commelinaceae. Contr. U. S. Nat. Herb. 5: 207-208. 1903. Studies of Mexican and Central American Plants No. 3. Contr. U. S. Nat. Herb. 8: 1-55. 1911. Studies of Mexican and Central American Plants No. 7a. Contr. U. S. Nat. Herb. 13: 291-312. NEW AND OVERLOOKED SPECIES OF THE VIR- GINIA FLORA. — Recent field and herbarium studies have turned up the following species which are apparently new to the state or were overlooked by Massey in his “Virginia Flora,” 1961. Specimens are deposited in the Longwood College Herbarium; duplicates are in herbaria indicated by symbols after collecting numbers. Najas minor Allioni. James River near Brandon, Prince George Co. Svenson & Harvill 17012. An introduction from the Old World which is local in the eastern states (Mer- ilànen, Rhodora 70: 169, 1968). Radford, Ahles and Bell list it for Wake Co., North Carolina in their “Manual of the Vascular Flora of the Carolinas,” 1968. Panicum longifolium Torrey. Boggy marsh north of Bur- gess Station, Dinwiddie Co.; sphagnous bog northwest of Dahlia, Greensville Co. Harvill 20304 & 17603. Although ranging on the coastal plain from Nova Scotia to Florida and Texas, thence north to Kentucky, it appears to be rare in Virginia. Scirpus ancistrochaetus Schuyler. High-elevation pond on the southern edge of Alleghany Co. Harvill 14559 (PH). Determined by Dr. A. E. Schuyler who states in his “Leafy Species of Scirpus,” 312, 1967, that it is known from only a few localities in Vermont, Massachusetts, New York and Pennsylvania. This is the southernmost record for the plant. Juncus validus Coville. In white sand of the Stipulicida area (Rhodora 69: 777, 1967) in Isle of Wight Co. Harvill 17062 (US). Identity confirmed by Dr. F. J. Hermann. A northern extension of the eastern arm of its distributional are which extends to Florida and Texas, thence north to Oklahoma and southwestern Missouri. The nearest known station is in Dare Co., North Carolina mapped by Radford, Ahles and Bell in their “Manual,” 279, 1968. Arenaria patula Michaux. In shallow soil on limestone flatrocks in The Cedars, Lee Co. Harvill 18530. An east- ward extension for the species which ranges from southern Ohio to Kansas, Texas, Alabama and Tennessee. 272 1970] Virginia Flora — Harvill 213 Silene gallica L. Sandy slope above a sphagnous bog north- west of Dahlia, Greensville Co. Harvill 21311. Collected by Barbara J. Harvill. An adventive from Europe which is very local in the southeastern states (Radford, Ahles and Bell, “Manual,” 447, 1968). Fraxinus quadrangulata Michaux. Limestone slope near Hilton, Scott Co. Harvill 17778. An eastward extension for the blue ash which ranges from Ontario to Oklahoma, Ala- bama and Kentucky. Mazus japonicus (Thunberg) Kuntze. Bank of a stream near Richmond, Henrico Co. Specimen sent to me by Dr. John C. Strickland of the University of Richmond. Nat- uralized from eastern Asia and very local in the southeast- ern states (Radford, Ahles and Bell, “Manual,” 942, 1968). Bellis perennis L. Weed on the campus of Randolph-Macon College, Hanover Co. Specimen sent to me by Dr, Leonard O. Morrow of Randolph-Macon College. A southeastward range extension of the introduced European plant which is rare in the southeastern states. Mapped only for Mitchell and Yancy counties, North Carolina by Radford, Ahles and Bell in their *Manual," 1071, 1968. A. M. HARVILL, JR. LONGWOOD COLLEGE FARMVILLE, VIRGINIA 23901 LISTERA OVATA (L.) R. BR. IN THE BRUCE PENINSULA, ONTARIO. — On July 11, 1968, Miss Bessie Plunkett of Port Arthur, Ontario with the senior author and a group from the Federation of Ontario Naturalists summer camp discovered a colony of about 15 orchids near the village of Red Bay in the Bruce Peninsula, Ontario. The plants were recognized as belonging to the genus Lis- tera, but were obviously not one of the five species found in northeastern North America. A single specimen was collected on July 17, 1968 by J. F. Alex of the University of Guelph with A. G. Thomas and C. R. Skelton. (J. F. Alex et al 3090 (OAC) Ontario, Bruce County, Albemarle Township, Red Bay P.O. 17 July, 1968). Alex tentatively identified the specimen as Listera ovata (L.) R. Br. and forwarded it to the Plant Research Institute of the Department of Agriculture, Ottawa. It was examined and compared with European material by C. Frankton, B. Boivin and E. W. Greenwood who confirmed Alex’s identifi- cation. F. W. Case, Jr. of Saginaw, Michigan and L. A. Garay of the Botanical Museum of Harvard University examined colored photographs and drawings of the Red Bay plants and suggested that they represented L. ovata. The orchids were found in a partly open, recently cleared area on the borders of a cedar (Thuja occidentalis L.) swamp. The soil is a sandy gravel covered with a few inches of humus and overlying dolomite. One of the plants was 60 cm in height and bore 52 blooms; six were over 40 cm high and the remainder about 30 cm. Approximately 20 plants reappeared in the same location in June, 1969 and several more specimens were discovered under eastern white cedars and black ash about 500 yards east of the original find. The field notes by J. F. Alex on the specimen label read, in part: “about 15 plants scattered through 20 m area re- cently cleared of brush, deadfall, ete, under cedar with ash, poplar and birch. Soil moderately drained organic sand. Plants ranged in height 38-61 em". 274 1970] Listera — Elliott and Cook 275 Butcher (1961) says that “it is a common plant of grass- land and woods on basic soils throughout the British Isles". Summerhayes (1951) states, “. . . . the common twayblade is probably commoner and more widely distributed in this country (Britain) than any other kind of orchid”. He gives its range as “throughout Europe from northern Scandi- navia and Russia southwards to the Mediterranean, .. . while eastwards it extends through Siberia to the region of Lake Baikal and through Asia Minor to the western end of the Himalayan Range". We are not aware of any previous occurrence of this twayblade having been recorded from North America and can only guess as to how it may have found its way to the Bruce Peninsula. A preliminary note on this finding has been published by Elliott (1969). The authors wish to thank J. F. Alex and others men- tioned for their assistance. H. VINCENT ELLIOTT 129 THOMAS STREET EXETER, ONTARIO FRANK 8. COOK DEPARTMENT OF BOTANY UNIVERSITY OF WESTERN ONTARIO LONDON, ONTARIO REFERENCES BUTCHER, R. W. 1961. A New Illustrated British Flora. Part II. Ericaceae to Gramineae. Leonard Hill (Books), London, 1080 pp. ELLIOTT, H. V. 1969. A Possible New Orchid for North America. The Ontario Naturalist 1/69: 15. SUMMERHAYES, V. S. 1951. Wild Orchids of Britain. Collins, Lon- don, SVII, 366 pp. ZONATION OF BRYOPHYTES IN THE HEADWATERS OF A NEW HAMPSHIRE STREAM JANICE M. GLIME A vertical zonation of bryophytes on the rocks of the Fox Pond inlet in Plymouth, N.H., is apparent. Here, Fontinalis novae-angliae covers the submerged portion while the ex- posed rock grades from Plagiochila asplenioides just above F. novae-angliae to Hypnaceae and Mnium on the upper- most portions, This gradation suggests the possibilities of moisture, light, or temperature gradients as causal factors, while submersion in high water seasons may also play a part. Also apparent in this stream is the dominance of Scapania undulata at upstream locations and a shift to Fontinalis novae-angliae and Plagiochila asplenioides farther down- stream. During the late winter of 1969, the inlet of Fox Pond, Plymouth, Grafton Co., N.H., was selected as a study stream for two reasons: its proximity to the campus laboratory and its lush growth of bryophytes which could be used for descriptive studies. To determine the relative abundance of the Fontinalis species and the several accompanying liverwort species, random observation stations were chosen and labelled from the source to a selected point downstream, This selected point marks the beginning of a stream drift study area and it seemed desirable to keep the two areas separate. Stations were chosen by placing the numbers 1-10 in a bucket, drawing a number, multiplying that by five, and taking that many paces to the next station. For example, station AA was placed at the most distal spring of the head- waters, and a 7 was drawn, so station BB was 35 paces downstream. Since the same researcher marked the paces each time, the distances between stations are approximately comparable and the procedure is an expedient one. By this 276 1970] Bryophytes — Glime 211 procedure, seventeen stations (AA-QQ) were chosen, Fur- ther downstream twelve stations (A-L) had previously been selected by the same procedure, giving a total of 29 stations. A station extends 2.5 m. on each side of the station label, which is painted on a nearby tree. Observations were made on September 13 and 15, 1969, at each of the 29 stations, the dominant bryophytes were identified and their locations noted from headwater down- stream. | The general trend shows a lack of aquatie bryophytes in the extreme headwaters, but it is not far (about 80 m. at the lower end of EE) before Fontinalis novae-angliae Sull. appears, along with several species of leafy liverworts. At this point also, F. antipyretica Hedw. var. gigantea (Sull.) Sull. occurs, one of two locations for the plant in this stream. As soon as submerged rocks appear in the stream, F. novae- angliae dominates the water, while Scapania undulata (L.) Dum. grows just beneath the surface and on the wet exposed surface of rocks, while higher parts of the rocks exhibit Plagiochila asplenioides (L.) Dum. or S. nemorosa (L.) Dum. Farther downstream P. asplenioides replaces S. nemo- rosa and a clear line of demarcation appears where P. asplenioides meets F. novae-angliae. lt is in this same re- gion that F. novae-angliae becomes a dominant ground cover in the stream, blanketing 50-100% of the stream bottom. There is almost no intermingling of P. asplenioides and F. novae-angliae, and on the date of observation P. asplen ioides was always out of the water. However, observations during winter and spring of 1969 showed that many areas of P. asplenioides were submerged. Farther down stream, espe- cially on the larger rocks which occur there, a 3-layered zonation appears, typically F. novae-angliae — P. aspleni- oides — Mnium & Hypnaceae, including Thuidium delicatu- ium (Hedw.) BSG. Occasional clumps of F. dalecarlica Schimp. ex. BSG occur, but never reach abundance. Several factors can be important in determining the lo- cation of these bryophytes, and the perfectly repetitive zonation pattern of F. novae-angliae to P. asplenioides in 278 Rhodora [Vol. 72 the last 14 stations is clearly not one of chance. Observa- tions of the conditions surrounding these plants during two extreme seasons (late winter and late summer) indicate that water level at critical times, coupled with summer humidity, may account for the locations. At all stations, Fontinalis was wet, although it was not always submerged during September. Furthermore, at no station was any P. asplenioides submerged in summer. But the occurrence of P. asplenioides under water for a 2-3 week period in the spring is probably one of tolerance, not of requirement, for P. asplenioides is known from soil-covered walls and slopes of open mountain woods (Watson, 1963), which are not likely to ever be submerged, On the other hand, its ability to tolerate the scouring effect of snow melted, debris-filled water could be a competitive advantage over the hypnaceous bryophytes of the upper rock surfaces. When I felt the bryophytes, a humidity gradient was apparent between these two zones on the rocks at the selected stations, and P. asplenioides was always damp. The Hypnaceae, with their compact, adnate growth form (smooth mat, Gimingham & Birse, 1957), grow on the drier upper portions of the rocks while P. asplenioides (tall turf, Gimingham & Birse, 1957) enjoys the greater humidity of the lower areas, Mr. Luna B. Witton, the owner of the property, states that the stream had much more water than usual in 1969, so the moisture stress on the exposed upper parts of the rocks would likely be much greater in other years. N evertheless, these are only speculations, and it may well be that a temperature gradient is a more important factor, where one would expect the extremes to occur on the higher parts of rocks, while the most moderate temperatures would be in the water. Or, zones could be related to light intensity, or interaction of all these factors, as suggested by Gimingham & Birse (1957) in their study of vertical zones of stream banks. Further speculation might suggest that the abundance of Scapania undulata and S. nemorosa upstream and their replacement by P. asplenioides downstream could be related to the more stable water level farther upstream. In a pre- 1970] Bryophytes — Glime 279 vious study (Glime, 1968), I indicated that Scapania un- dulata grows in Garrett Co., Maryland, on the brink of waterfalls, and in Carbon Co., Pennsylvania, on the water- air interface zones of rocks and boulders. Its areas of oc- currence appear to be areas which keep it constantly soaked, while its terrestrial counterpart, S. nemorosa, grows high enough on the rocks to be rarely submerged. Now at least we know that zonations exist, we have sev- eral hypotheses to test, and we can proceed to experimental work and detailed data to test these hypotheses. Voucher specimens are filed in the DePauw University Herbarium, Greencastle, Indiana. ACKNOWLEDGMENTS I wish to express my gratitude to Dr. Winona H. Welsh of DePauw University for help in identification of Fonti- nalis species. To Dr. Mary G. Bilheimer of Plymouth State College, I owe a debt of gratitude for criticisms of the manuscript. A special thanks goes to Mr. Luna B. Witton, who owns the property and gave us permission to use the stream for study. SCIENCE DEPARTMENT, BOYD HALL PLYMOUTH STATE COLLEGE PLYMOUTH, N. H. 02264 BIBLIOGRAPH Y GIRMINGHAM, C. H., and E. M. BIRSE. 1957. Ecological studies on erowth-form in bryophytes. J. Ecol. 45: 533-545. GLIME, J. M. 1968. Ecological observations on some bryophytes in Appalachian Mountain streams. Castanea 33: 300-325. Watson, E. V. 1963. British mosses and liverworts. Univ. Press, Cambridge. 419 p. VERNON BLACK GUM SWAMP F. R. FOSBERG AND TERRY BLUNT In the hills in the town of Vernon, Vermont, just north of the Massachusetts boundary and just west of the Con- necticut River is a small but interesting swamp, featuring a disjunct stand of Nyssa sylvatica Marsh., considerably north and inland from the usual distribution of this species. This swamp was very briefly described by H. W. Vogelmann in Vermont Natural Areas — Report 2, March 1969. We visited it on October 5, 1969, and studied it enough to pre- pare a somewhat fuller description. We also visited the owner, the distinguished architect, Mr. Jacques Delamarre, and suggested to him the desirability of preserving this area so that longer term, more adequate investigations could be carried out, and because it is a unique natural feature in this region, The swamp lies at an elevation of about 320 m on a small bench on the southwest side of a prominent knob, appar- ently unnamed. This shelf has a depression of about 2-3 hectares, rimmed by a curved ridge of hard schist up to 10 m high. At the east end is a notch, cut by a small stream that forms the outlet of the swamp. The surrounding area is a mixed Tsuga canadensis forest that has been recently severely cut. The ridge is covered by an open mixed forest from which a few hemlocks have been cut, but which has so little soil that trees of large size are scarcely to be ex- pected. Acer rubrum L., Betula papyrifera Marsh., Betula alleghaniensis Britt., Betula lenta L., Tsua canadensis (L.) Carr. and Pinus strobus L. were observed on the ridge with one large old Fagus grandifolia Ehrh. just above the level of the swamp. Ericaceous shrubs and dwarf shrubs such as Kalmia latifolia L., Vaccinium angustifolium Ait., and Gaultheria procumbens L. form a sparse shrub layer. A few saplings of Castanea dentata (Marsh.) Borkh. and Fagus grandifolia Ehrh. are scattered here and there, The depression, itself, is filled, to the level of the outlet notch, with fallen trees, humus, and Sphagnum. A very 280 1970] Black Gum — Fosberg and Blunt 281 irregular closed mixed forest about 5-8 m tall covers the swampy area with the following trees in the main closed layer: Betula alleghaniensis Britt., Nyssa sylvatica Marsh., Tsuga canadensis (L.) Carr., Picea rubens Sarg., Acer rubrum L. Scattered emergent trees, up to 25 or 30 m tall, are: Acer rubrum L., Nyssa sylvatica Marsh., Tsuga canadensis (L.) Carr., Picea rubens Sarg., In the lower end are some Pinus strobus L. A shrub layer, open, to in places quite dense, includes: Ilex verticillata (L.) Gray, Viburnum cassinoides L., Nemo- panthus mucronata (L.) Trel., Vaccinium corymbosum L., Kalmia angustifolia L. (near the lower end) and near the edges also: Viburnum alnifolium Marsh., Kalmia latifolia L., Hammamelis virginiana L. and Acer pensylvanicum L. The herb layer is closed to open, with a ground surface of litter and Sphagnum. It is composed of the following (including dwarf shrubs) : Osmunda cinnamomea L., Os- munda regalis L., Coptis groenlandica (Oeder) Fern., Aralia nudicaulis L., Gaultheria hispidula (L.) Bigel., Mitchella repens L., Vaccinium angustifolium Ait., Carex cf. rosea Schkuhr., Viburnum alnifolium Marsh., Lycopodium ob- scurum, Thelypteris dryopteris, Uvularia sessilifolia L., Aster cf. puniceus L., Kalmia angustifolia L., Gaultheria procumbens L., Pteridium aquilinum (L.) Kuhn, Quercus rubra L. (seedling). The last six listed are principally near the edges and in the lower end with Pinus strobus L. Epigaea repens L. covered a rootman of a blow-down and was seen rarely in other high spots in the swamp. The Nyssa, with their red autumn coloring, were very conspicuous. They reached 4-6 dm diameter at breast height and 20 m tall, and the large ones were all *stag-headed". The bark on leaning ones was thin and flaky on the upper sides, thick, deeply cleft and ridged on lower sides. There were the large Nyssa trees mentioned above, rather widely spaced, others 4-5 m tall, and a few seedlings, but few or no intermediates between these classes. 282 Rhodora [Vol. 72 In addition to the plants seen by us, Vogelmann mentions star flower (Trientalis borealis Raf.), wild lily-of-the-valley (Maianthemum canadense Desf.), painted trilium (Tril- lium undulatum Willd.), and interrupted fern (Osmunda claytoniana L.). He interprets the occurrences of Nyssa as a relict from a period of warmer climate. We do not dis- agree with this, since the species is now abundant only con- siderably south of this station. He says that a few scattered trees do occur elsewhere in Vermont, as near Burlington, but not such a concentration as here. It is to be hoped that this interesting pocket may be pro- tected as a nature preserve for the enjoyment and education of future generations of naturalists. SMITHSONIAN INSTITUTION WASHINGTON, D.C. 20560 CONNECTICUT RIVER WATERSHED COUNCIL GREENFIELD, MASS. 01301 RECENT PLANT RECORDS FOR NEBRASKA ROBERT J. LEMAIRE Part of my work as the biologist for the Soil Conservation Service in Nebraska includes assisting various groups and individuals throughout the state to establish nature trails. These are essentially botanical trails along which usually only vascular plants are identified and labeled. Among the plants I collected and identified along the more than 50 nature trails in the state were four species which, to my knowledge, have not previously been reported for Nebraska. One additional species was found in a very unusual habitat. Nomenclature for the following five species follows Fernald (1950). On May 18, 1966, I collected specimens of Collinsia parvi- flora Dougl. in the extreme northwest corner of the state at the Ross Raum ranch in Sioux County, about nine miles northwest of Crawford. This is a rough, ponderosa pine woodland area, part of the “Pine Ridge" in Nebraska. This annual species occurs mainly west of Nebraska. I failed to find the plant again at the same location in the spring of 1968. On September 10, 1966, I collected specimens of Leonwrus sibiricus L., at the Riverview Vacation Ranch near Com- stock in the Sandhills region, This adventive plant from Eurasia is similar to L. cardiaca L., a common naturalized weed in Nebraska. The plants were located along a fence line and in disturbed soil along a dirt road. I have not seen the species elsewhere in the state. Circaea canadensis Hill, a plant native to northeastern United States, was collected along the wooded bluffs of the Missouri River about five miles south of Nebraska City on June 29, 1966. The woodland along the Missouri River in southeastern Nebraska is the approximate western edge of the eastern deciduous forest type. In May 1966, I collected a specimen of Veronica hederae- folia, L. on the College of Agriculture campus of the Uni- 283 284 Rhodora [Vol. 72 versity of Nebraska in Lincoln. A native of Europe, this species is now reported naturalized in several eastern states at least as far west as Ohio. The specimen was growing as a weed in a Kentucky bluegrass lawn, | The roof of a mid-town Lincoln building is a very unusual location for finding a plant new to Nebraska. Part of the building is only three stories tall and part is six stories. On October 4, 1968, I collected Cyperus difformis L. growing abundantly in a “marsh” on the roof of the third floor and as scattered plants in very shallow water (up to 3/4 inch deep) on the sixth floor roof. Evidently natural rainfall and water condensed from air conditioning units maintain suitable growing conditions for this wetland plant on these flat, pebble and asphalt roofs. The “marsh” has been in existence for at least 15 years according to residents of the building. It is not known how long C. difformis has bcen growing here. The standard manuals indicate that this Asian species is adventive only in Virginia and California. Specimens of these five species have been deposited in the herbarium of the University of Nebraska at Lincoln. SOIL CONSERVATION SERVICE 194 SOUTH 12TH STREET LINCOLN, NEBRASKA 68508 LITERATURE CITED FERNALD, M. S. 1950. Gray’s Manual of Botany, 8th ed. American Book Co., New York. 1632 pp. AN ASSEMBLAGE OF HALOPHYTES IN NORTH- ERN OHIO: A remarkable group of weedy plants has been discovered on the property of the Morton Salt Company in Rittman, Wayne County, Ohio. In this area are several brine wells and a large pool in which brine is stored prior to processing. Adjacent to these wells grow Salicornia europaea L. (glasswort) and Atriplex argentea Nutt. (salt- bush), both apparently new to the Ohio flora. Salicornia, of course, is a well-known salt marsh inhabi- tant. Here at Rittman it forms great orange-red masses of thousands of individuals. The Atriplex, a western species of alkaline areas, occurs with the glasswort, but is much less common. Fernald (1950) does not include Ohio in the rango of Salicornia, but does list A. argentea as adventive in Ohio. Neither species is mentioned by Weishaupt (1968), however, and no Ohio herbarium specimens are known to the author. Company officials, unfortunately, could give no information as to when these species first appeared. Both these species occur in the most disturbed ground around the wells where no other plants grow. An exception is an occasional specimen of Coriospermum hyssopifolium L. Along the run-offs from the wells Cyperus esculentus L., Juncus effusus L., and Typha angustifolia L. are com- mon. It is hoped that a general study of this station can be developed, as this is perhaps an unique ecological situation for Ohio. All species cited are represented in the herbarium of Kent State University. Acknowledgements are gratefuily given to Dr. B. A. Foote of Kent State University for informing me of this lo- cality’s existence and to the Morton Salt Company for col- lecting privileges. ALLISON W. CUSICK DEPARTMENT OF BIOLOGICAL SCIENCES KENT STATE UNIVERSITY KENT, OHIO 44240 285 286 Rhodora [Vol. 72 LITERATURE CITED FERNALD, M.L. 1950. Gray’s Manual of Botany. American Book Company, New York. WEISHAUPT, C. G. 1968. Vascular Plants of Ohio. Wm. C. Brown Book Company, Dubuque, Iowa. A RANGE EXTENSION FOR ODONTITES SEROTINA (LAM.) DUM. (SCROPHULARIACEAE): This some- what weedy species, an adventive from Europe, has hereto- fore been reported to extend from Newfoundland to western Quebec, south to Nova Scotia, northern New England, and northern New York (Fernald, Gray's Manual, 8th Edition, 1950; Gleason and Cronquist, Manual of Vascular Plants, 1963 — but these latter exclude Newfoundland from the range). This is to report a remarkable extension of the plant's range into northeastern Wisconsin. Mr. Tom Brudnicki, a student, collected the plant in a wooded swamp in the SW part of section 19, T27N, R19E, Oconto County, Wisconsin, 14 September 1968 (Brudnicki 053). He returned to the site 6 September 1969 and collected numerous others ( Brud- nicki s.n.). Vouchers for both collections are deposited in the herbarium, Wisconsin State University-Oshkosh, and duplicates of the latter collection have been distributed to WIS, UWM, GH, and NY. Peter Salamun (1951. Preliminary Reports on the Flora of Wisconsin. XXXVI. Scrophu- lariaceae. Trans. Wis. Acad. Sci., Arts and Letters. 40(2): 111-138) does not report the plant for Wisconsin, nor are there any records for the plant in Wisconsin at UWM. Hugh Iltis (personal communication) informs me that wis has one record of this species from Door County and one from adjacent Kewaunee County, both collected since 1951. These records are sufficiently extensive to warrant including this species as a member of the Wisconsin flora. NEIL A. HARRIMAN BIOLOGY DEPARTMENT WISCONSIN STATE UNIVERSITY-OSHKOSH OSHKOSH 54901 A NEW ARGENTINE SISYRINCHIUM ROBERT C. FOSTER Among the unnamed specimens of South American Sisyr- inchium in the Gray Herbarium, there is one from Argen- tina that appears to represent an undescribed species, and is here described. Sisyrinchium papillosum R. C. Foster, spec. nov. Planta parva, plus minusve caespitosa, e rhizoma breve angustoque. Folia basalia, ad 10 cm. longa et 2-3 mm. lata, glauca, dense breveque papillata, inflorescentiam excedentia. Inflores- centia terminalis, non pseudo-lateralis, spathae pluriflorae. Spathae inaequales, exterior quam interior longior, ad 2.4 cm. longae, glaucae, plusminusve dense papillatae. Pedicelli glanduloso-villosi, pili lutei, quam spatham interiorem ad anthesin breviores vel plusminusve aequantes. Ovarium cblongum, ad 3-3.5 mm. longum, dense glanduloso-pubes- cens, pili lutei. Tepala subaequalia, lutea, venis atrobrun- neis, ovata vel obovata, acuta, ad 1 cm. longa et 4-5 mm. lata, ad basim adaxialiter plusminusve glanduloso-villosa. Filamenta coalita 2.5 mm., dense glanduloso-villosa, libera plusminusve 1.8 mm.; antherae oblongae, circa 2 mm. longae. Styli rami plusminusve 2 mm. longi, quam antherae multo breviores. Capsula glanduloso-pubescens vel glabrata. Semina matura non visa. ARGENTINA: MENDOZA: TUMUYAN: ad ripas rivulorum pr. Cuesta de los Aflijidos, iter Andinum Paso del Portillo dicto, 24 Dec. 1933, A. Ruiz Leal 1875 (type, in Gray Herb.). The partially united filaments with long, oblong anthers ally this species with the Sisyrinchium Marchio complex and to others such as S. avenaceum Klatt or even S. Wettsteinüi Hand.-Mazz. It differs from these in smaller size, glaucous leaves and spathes, these being densely short-papillate, and in the glandular villosity of the united portion of the fila- ments and basal portion of the adaxial tepal-surface. GRAY HERBARIUM, HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS 02138 287 Volume 72, No. 790 including pages 145-287, was issued July 15, 1970. NEW ENGLAND BOTANICAL CLUB LIST OF MEMBERS Officers since 1959 Presidents R. E. Schultes 1960-1962 F. C. Seymour 1963-1964 W. H. Drury, Jr. 1965-1967 S. K. Harris 1968-1970 W. D. Countryman 1970- Vice-Presidents F, C. Seymour 1960-1962 W. H. Drury, Jr. 1962-1964 S. K. Harris 1965-1967 W. D. Countryman 1968-1970 W. N. Tiffney, Sr. 1970- Corresponding Secretaries C. Schweinfurth 1949-1965 W. C. Galinat 1966-1968 T. G. Hartley 1969- Recording Secretaries S. K. Harris 1935-1964 R. M. Tryon, Jr. 1965-1967 F. A. Barkley 1968 P. D. Sorensen 1969 A. F. Tryon 1970- 'Treasurers A. F. Hill 1943-1966 H. R. Sweet 1967- Editors of Rhodora R. C. Rollins 1951-1961 A. R. Hodgdon 1962- Active Members * A]-Shebaz, Ishan Harvard University Herbaria 22 Divinity Avenue Cambridge, Mass. 02138 Ansell, Phillip W. 194 Vinton Street Melrose, Mass. 02176 Baldwin, Henry I. Hillsboro, N.H. 03244 *Banerjee, Umesh C. Harvard University Herbaria 22 Divinity Avenue Cambridge, Mass. 02138 *Baranov, Andrey Department of Biology Northeastern University Boston, Mass. 02115 Barkley, Fred A, Department of Biology Northeastern University Boston, Mass. 02115 bean, Ralph Carleton 48 Emerson Street Wakefield, Mass. 01880 Boghdan, Kalil S. 9 Fairview Avenue Watertown, Mass. 02172 Bogle, Alfred Lynn Arnold Arboretum 22 Divinity Avenue Cambridge, Mass. 02138 Bussewitz, Albert W. Rocky Knoll Nature Center 74 Maple Street Milton, Mass. 02186 *Student member Admitted March 3, 1967 December 1, 1967 April 4, 1969 January 2, 1970 December 6, 1963 November 5, 1965 December 3, 1909 April 1, 1966 December 5, 1969 June 2, 1961 *Bye, Robert Botanical Museum Oxford Street Cambridge, Mass. 02138 Canoso, Michael Anthony Harvard University Herbaria 22 Divinity Avenue Cambridge, Mass. 02138 Carr, David James Department of Biology Northeastern University Boston, Mass. 02115 Chisholm, Robert 100 Randlett Park West Newton, Mass. 02165 Colt, LeBaron C. Phillips Street Medfield, Mass. 02052 *Conway, Joan R. Marine Biological Laboratory Woods Hole, Mass. 02543 Cooley, George R. Hickory Hill Rensselaerville, N.Y. 12147 Coon, Nelson Box 1 Vineyard Haven, Mass, 02568 Countryman, William Douglas Norwich University Northfield, Vt. 05663 Creighton, Harriet B. Department of Biological Sciences Wellesley College Wellesley, Mass. 02181 DeWolf, Gordon Parker, Jr. 300 High Street Walpole, Mass. 02081 Drury, William Holland, Jr. Hatheway School of Conservation Drumlin Farm South Lincoln, Mass. 01773 December 5, 1969 March 2, 1956 January 6, 1967 June 4, 1965 January 8, 1965 March 6, 1970 December 4, 1953 November 3, 1944 December 7, 1962 January 3, 1969 December 2, 1949 November 7, 1947 Eaton, Richard Jefferson Lincoln Center, Mass. 01773 Elias, Thomas Sam Arnold Arboretum 22 Divinity Avenue Cambridge, Mass. 02138 Fordham, Alfred 896 Clapboardtree Street Day Farm Norwood, Mass. 02062 Foster, Robert Crichton Gray Herbarium 22 Divinity Avenue Cambridge, Mass. 02128 Freeman, Dana Ward 10 Dwinell Street West Roxbury, Mass, 02132 *Frolich, Michael Wick Eiological Laboratories 16 Divinity Avenue Cambridge, Mass. 02138 Galinat, David 35 Mossfield Road Waban, Mass. 02168 Galinat, Walton Clarence 33 Mossfield, Road Waban, Mass. 02168 *Gastony, Gerald C. Harvard University Herbaria 22 Divinity Avenue Cambridge, Mass. 02138 Hartley, Thomas Gordon Arnold Arboretum 22 Divinity Avenue Cambridge, Mass. 02138 Hebb, Robert Stephan 28 Bardwell Street Jamaica Plain, Mass. 02130 December 1, 1916 March 6, 1970 December 2, 1960 Ncveraber 2, 1934 November 7, 1958 January 2, 1970 January 8, 1965 January 8, 1954 January 6, 1967 June 4, 1965 April 4, 1969 Hebb, Winnifred Elliot 28 Bardwell Street Jamaica Plain, Mass. 02150 Hehre, Edward J. Southampton College Long Island University Southampton, N.Y. 11968 Hershenson, Benjamin April 4, 1969 June 4, 1965 March 1, 1963 Massachusetts College of Pharmacy 179 Longwood Avenue Bceston, Mass. 02115 Hill, Albert Frederick The Carrying Place Surry, Maine 04684 Hodedon, Albion Reed University of New Hampshire Durham, N.H. 03824 Howard, Richard Alden Arnold Arboretum 22 Divinity Avenue Cambridge, Mass. 02138 Howland, Russell Department of Botany University of New Hampshire Durham, N.H. 03824 Johnnykutty, A. Thomas Department of Biology Northeastern University Boston, Mass, 02115 Jost, Dana N. Framingham State College Framingham, Mass. 01701 Kaplan, Lawrence Department of Biology University of Massachusetts 100 Arlington Street Boston, Mass. 02116 Khudairi, A. K. Department of Biology Northeastern University Boston, Mass. 02115 Apri 7,1911 December 2, 1932 May 3, 1940 January 3, 1969 January 6, 1967 March 3, 1968 March 3, 1967 June 2, 1967 *Kistler, Larry Lynden Box 5 Monument Valley, Utah 84536 Lamb, Ivan MacKenzie Farlow Herbarium 20 Divinity Avenue Cambridge, Mass, 02138 *Lambert, Michael Department of Biology Northeastern University Boston, Mass. 02115 *Lockwood, Tommie Earl Botanical Museum Oxford Street Cambridge, Mass. 02138 Miller, Norton George Arnold Arboretum 22 Divinity Avenue Cambridge, Mass. 02138 Mish, Lawrence B. Bridgewater State College Bridgewater, Mass. 02324 Muller, Erhart Shaker Village Harvard, Mass. 01451 Nevling, Lorin Ives, Jr. Harvard University Herbaria 22 Divinity Avenue Cambridge, Mass. 02138 Nickerson, Norton H. Department of Biology Tufts University Medford, Mass, 02155 Osiecki, Walter G. 34 Park Avenue Natick, Mass. 01760 Pike, Radcliffe B. University of New Hampshire Durham, N.H. 03824 December 5, 1969 January 8, 1954 December 5, 1969 December 5, 1969 March 6, 1970 March 3, 1968 April 4, 1969 January 8, 1960 June 5, 1964 March 3, 1967 January 7, 1955 Richardson, Laurence E. Barrett’s Mill Road Concord, Mass. 01742 *Rodman, James Harvard University Herbaria 22 Divinity Avenue Cambridge, Mass. 02138 Rollins, Reed Clark Gray Herbarium 22 Divinity Avenue Cambridge, Mass. 02138 Safwat, Fuad Department of Biology University of Massachusetts 100 Arlington Street Boston, Mass, 02116 Schubert, Bernice Giduz Arnold Arboretum 22 Divinity Avenue Cambridge, Mass. 02138 Schultes, Richard Evans Botanical Museum Oxford Street Cambridge, Mass. 02138 Seeler, Edgar Viguers, Jr. 4 Berkeley Place Cambridge, Mass. 02138 Shaw, Elizabeth Anne Gray Herbarium 22 Divinity Avenue Cambridge, Mass. 02138 Sorensen, Paul Davidsen Arnold Arboretum Jamaica Plain, Mass. 02130 Spence, Willard Framingham State College Framingham, Mass. 01701 November 6, 1959 February 2, 1968 January 15, 1937 March 3, 1967 March 6, 1970 March 4, 1938 May 3, 1957 March 7, 1969 March 1, 1968 March 3, 1968 Stone, Richard Allen Department of Biology University of Massachusetts 100 Arlington Street Boston, Mass. 02116 Strong, Peter Shaker Road Harvard, Mass. 01451 Sweeney, Edward F. 20 Williams Road North Reading, Mass. 01864 Sweet, Herman Royden Department of Biology Tufts University Medford, Mass. 02155 Thomas, Aubrey 8. Department of Biology Merrimac College North Andover, Mass. 01845 *Tiffney, Bruce H. 226 Edge Hill Road Sharon, Mass. 02067 Tiffney, Wesley N., Jr. 226 Edge Hill Road Sharon, Mass. 02067 Tiffney, Wesley N., Sr. 226 Edge Hill Road Sharon, Mass. 02067 Tagani, Kamil B. El Department of Biology Northeastern University Boston, Mass. 02115 Ting, Yu-Chen 230 Bonad Road Brookline, Mass. 02146 Tryon, Alice Farber Gray Herbarium 22 Divinity Avenue Cambridge, Mass. 02158 December 6, 1968 June 7, 1963 April 6, 1962 April 3, 1959 June 2, 1967 January 6, 1967 December 1, 1961 January 8, 1960 February 7, 1969 May 3, 1957 December 6, 1968 Tryon, Rolla Milton, Jr. Gray Herbarium 22 Divinity Avenue Cambridge, Mass. 02138 Vagenas, George 314 Lafayette Street Salem, Mass. 01970 Walton, Russell 134a Sutherland Road Brookline, Mass. 02146 Wareham, Richard Thurman Route 1, Box 252 Jaffrey, N.H. 03452 Wells, Theodore Woodland Milton Academy Milton, Mass. 02186 Wetherall, Cyril 369 Ward Street Newton Center, Mass, 02159 White, Donald 356 Albion Street Wakefield, Mass. 01880 Whittier, Nathaniel Bridge Street Medfield, Mass. 02052 *Wise, David Alan Department of Botany University of New Hampshire Durham, N.H. 03824 Wood, Carroll Emory, Jr. Arnold Arboretum 22 Divinity Avenue Cambridge, Mass. 02138 January 6, 1939 March 7, 1969 February 2, 1968 October 6, 1944 February 6, 1953 December 3, 1949 April 4, 1913 June 4, 1965 December 6, 1968 November 1, 1946 Sustaining Members Adams, Franklin S. 202 Buckhout Laboratory Penn State University University Park, Pa. 16802 Andrews, Henry N., Jr. University of Connecticut Storrs, Conn. 06268 Argus, George William Canadian Forestry Service 344 Wellington Street Ottawa 4, Ontario Baldwin, John Thomas, Jr. College of William and Mary Williamsburg, Va. 23185 Barclay, Arthur Stewart New Crops Research Branch Plant Industry Station Beltsville, Md. 20705 Barneby, Rubert Charles Sterling House Box 172 Greenport, Long Island, N.Y. Barton, James Don, Jr. 15 Reynolds Street Alfred, N.Y. 14802 Beaman, John Homer Department of Botany Michigan State University East Lansing, Mich. 48823 Benninghof, William Shiffer Department of Botany University of Michigan Ann Arbor, Mich. 48104 Bergman, Herbert Floyd East Wareham, Mass. 02538 Berry, Wendell H., Jr. Lake Forest Academy Lake Forest, Ill. 60045 March 3, 1967 March 1, 1968 January 3, 1958 December 1, 1944 January 7, 1955 December 1, 1944 11101 December 2, 1960 December 4, 1953 March 1, 1940 April 3, 1931 March 4, 1966 Blohm, Henrik Apartado 69 Caracas, Venezuela Boivan, Bernard Central Experimental Farm Ottawa, Canada Brainerd, John Whiting Department of Biology Springfield College Springfield, Mass. 01109 Brown, Richard McPike Crater Lake National Park Fort Klamath Rural Station Klamath Falls, Oregon 97604 Channell, Robert Bennie Botany Department Vanderbilt University Nashville, Tenn. 37205 Chapman, Charles J, Department of Biology Concord College Athens, West Virginia 24712 Charette, Leopold Alphonse 27 Ferguson Avenue Burlington, Vt. 05401 Chase, Sherret S. State University College Oswego, N.Y. 13126 Church, George Lyle Department of Botany Brown University Providence, R.I. 02912 Clausen, Robert Theodore Weigand Herbarium Cornell University Ithaca, N.Y. 14850 Cody, William James Plant Research Institute Central Experimental Farm Ottawa, Canada May 2, 1958 May 2, 1947 November 1, 1940 December 1, 1950 March 2, 1956 February 2, 1945 March 3, 1967 February 5, 1926 December 1, 1944 February 4, 1955 Cronquist, Arthur John April 7, 1961 New York Botanical Garden Bronx, N.Y. 10458 Dahl, Anthony Orville January 6, 1939 Morris Arboretum 9414 Meadowbrook Avenue Philadelphia, Pa. 19118 Dansereau, Pierre December 1, 1944 New York Botanical Garden Bronx, N.Y. 10458 Demaree, Delzie December 1, 1944 109 South Avenue Hot Springs, Ark. 71901 Downes, Robert Marshall January 6, 1967 South Londonderry, Vt. 05155 Dressler, Robert Louis February 6, 1953 Smithsonian Tropical Research Institute P.O. Box 2072 Balboa, Canal Zone Drew, William Bradford May 13, 1932 Department of Botany Michigan State University East Lansing, Mich, 48823 Drouet, Francis March 4, 1938 Academy of Natural Sciences 19th Street & The Parkway Philadelphia, Pa. 19105 Dudley, Theodore R. December 6, 1963 U. S. National Arboretum 28th & M Streets, N.E. Washineton, D.C, 20250 Dunbar, Henry Fowler May 1, 1914 c/o Mrs, Paul Dornville box 95 Woodstock, N.Y. 12498 Duncan, Wilbur Howard March 3, 1956 Department of Botany University of Georgia Athens, Ga. 30601 Earle, Sylvia Allan Hancock Foundation University of Southern California Los Angeles, Calif. 90007 Einarsson, Eythor Museum of Natural History P.O. Box 532 Reykjavik, Iceland Eldredge, Keith Graham R.F.D. Gossville, N.H. 03239 Ernst, Wallace Roy Department of Botany Smithsonian Institution Washington, D.C. 20560 Fogg, John Milton, Jr. Barnes Arboretum Merion, Pa. 19066 Fosberg, Francis Raymond 3077 Holmes Run Road Falls Church, Va. 22042 Garton, Claude Eugene 309 Dacre Street Port Arthur, Ontario, Canada Giles, Richard Alden 118 Linden Place Ypsilanti, Mich. 48197 Gillett, John Montague March 6, 19'70 February 5, 1960 November 2, 1945 January 5, 1962 December 3, 1926 November 2, 1945 February 4, 1955 February 4, 1955 April 2, 1954 Botany & Plant Pathology Division Science Service Department of Agriculture Ottawa, Canada Glime, Janice Plymouth State College Plymouth, N.H. 03264 Goodman, George Jones University of Oklahoma Norman, Okla. 73069 December 5, 1969 January 5, 1945 Goodwin, Richard Hale December 5, 1930 Department of Botany Connecticut College New London, Conn. 06320 Gustafson, Alton Herman June 1, 1934 Bowdoin College Brunswick, Maine 04011 Habeeb, Herbert January 5, 1945 Auburn Community College Auburn, N.Y. 13021 Hand, Byron Eugene January 7, 1955 6 Caroline Avenue Caribou, Maine 04736 Hara, Hiroshi January 6, 1939 Botanical Institute University of Tokyo Tokyo, Japan Hathaway, Wilfred Adelbert May 2, 1958 Box 78 Segregansett, Mass. 02773 Hermann, Frederick Joseph December 1, 1944 Forest Service Herbarium U.S.D.A. Washington, D.C. 20250 Hinds, Harold W. January 8, 1965 Westledge School West Simsbury, Conn, 06092 Hopkins, Milton January 15, 1932 Port Washington Boulevard Roslyn, Long Island, N.Y. 11050 Hunziker, Armando Teodora April 2, 1948 Casilla de Correo 23 Cordoba, Argentina Hyde, Gregory E. January 3, 1969 41 Graden Street Red Hook, N.Y. 12571 Jablonski, Eugene December 5, 1958 Apt. A-52, Hudson View Gardens 183rd Street & Pinehurst Avenue New York, N.Y. 10023 James, Charles William Department of Botany University of Georgia Athens, Ga. 30601 Jones, George Neville Department of Botany University of Illinois Urbana, Ill. 61801 Lane, Franklin Charles Frostburg State College Frostburg, Md. 21532 Lellinger, David Bruce Department of Botany U. S. National Museum Washington, D.C. 20560 Lownes, Albert Edgar 16 Barberry Hill Providence, R.I. 02906 Maguire, Bassett New York Botanical Garden Bronx, N.Y. 10458 Manning, Wayne Eyer Department of Biology Bucknell University Lewisburg, Pa, 17837 Marderosian, Ara der January 5, 1957 January 6, 1939 March 4, 1955 November 6, 1959 March 7, 1952 December 1, 1944 February 2, 1934 December 6, 1963 Philadelphia College of Pharmacy & Science 43rd Street & Kingsessing Avenue Philadelphia, Pa. 19104 Mehlquist, Gustav A. L. Plant Science Department University of Connecticut Storrs, Conn. 06268 Moore, Albert Hanford 165 Middle Street Portsmouth, N.H. 03801 Moore, Harold Emery, Jr. Bailey Hortorium Ithaca, N.Y. 14850 June 2, 1967 May 4, 1906 March 1, 1940 Moore, John William 628 Ontario S.E. Minneapolis, Minn. 55414 Ogden, Eugene Cecil New York State Museum Albany, N.Y. 12224 Ownbey, Gerald Bruce Department of Botany University of Minnesota Minneapolis, Minn, 55455 Ownbey, Marion Department of Botany Washington State University Pullman, Wash. 99163 Partanen, Carl Richard Department of Biological Sciences University of Pittsburgh Pittsburgh, Pa. 15213 Pfeifer, Howard William University of Connecticut Storrs, Conn. 06268 Poole, James Plummer 19 E. Wheelock Street Hanover, N.H. 03755 Porter, Duncan MacNair Missouri Botanical Garden 2315 Tower Grove Avenue St. Louis, Mo. 63110 Pride, George Howard 133 Wellesley Street Weston, Mass. 02193 Pritchard, Arthur H. 42 Quarry Road Waterford, Conn. 065385 Quimby, Maynard Ward February 4, 1955 November 2, 1934 February 4, 1955 February 4, 1955 February 6, 1953 January 6, 1967 December 8, 1920 April 6, 1962 December 1, 1939 January 7, 1966 January 6, 1939 University of Mississippi School of Pharmacy University, Miss. 38677 Raup, Hugh Miller Department of Geography The Johns Hopkins University Baltimore, Md. 21218 December 6, 1929 Rudolph, Emanuel David March 2, 1956 Department of Botany & Plant Pathology Ohio State University Columbus, Ohio 43210 Schaeffer, Robert L., Jr. 32 North 8th Street Allentown, Pa. 18101 Seymour, Frank Conklin Pringle Herbarium University of Vermont Burlington, Vt. 05401 Sharp, Aaron John Department of Botany University of Tennessee Knoxville, Tenn. 37916 Shinners, Lloyd Herbert The Herbarium Southern Methodist University Dallas, Texas 75222 Smith, Ernest Chalmers Acadia University Wolfville, Nova Scotia, Canada Smith, Lyman Bradford 3941 Washington Street Kensington, Md. 20795 Smith, Wendell Phillips 911 E Street North Wilkesboro, N.C. 28659 Soper, James Herbert Museum of Natural Sciences National Museum of Canada Ottawa 4, Canada St. John, Harold Bishop Museum Honolulu, Hawaii 96819 Steele, Frederic Lincoln III St. Mary's-in-the-mountains Littleton, N.H. 053561 February 4, 1955 December 4, 1914 December 1, 1944 February 4, 1955 March 4, 1955 December 7, 1923 February 7, 1947 December 1, 1959 December 1, 1911 February 3, 1939 Whitehead, Donald Reed March 2, 1956 Department of Biology Indiana University Bloomington, Ind. 47401 Wilbur, Robert Lynch March 5, 1948 Department of Botany Duke University Durham, N.C. 27706 Wilson, Kenneth Allen January 3, 1958 San Fernando Valley State College 18111 Nordhoff Street Northridge, Calif. 91324 Winne, William Thomas February 4, 1955 2517 Troy Road Schenectady, N.Y. 12309 Youngken, Herbert Wilkinson, Jr. January 3, 1958 College of Pharmacy University of Rhode Island Kingston, R.I. 02881 OCT 2 0 1970 FARLOW HFAFEFRSENGE LIBRARY Hodova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL RALPH CARLETON BEAN ROBERT CRICHTON FOSTER ROLLA MILTON TRYON Associate Editors RADCLIFFE BARNES PIKE LORIN IVES NEVLING, JR. ELIZABETH ANNE SHAW Vol. 72 July-September, 1970 No. 791 CONTENTS: Comparative Studies of Smilax, Section Smilax, of the Southeastern United States. Ida E. Yates and Wilbur H. Duncan oeste 289 A Preliminary Checklist of the Marine Algae of Campobello Island, New Brunswick, Canada. Richard A. Stone, Edward J. Hehre, Joan M. Conway and Arthur C. Mathieson ........................ 313 Distributional History of Lycopus europaeus (European Water-Horehound) in North America. Ronald L. Stuckey and W. Louis Phstlspa- u u S a 851 (Continued on Inside Cover) The New England Botanical Club, Ine. Botanical Museum, Oxford St., Cambridge, Mass. 02188 CONTENTS: — continued Saga of the West Coast Sea-Rockets: Cakile edentula ssp. californica and C. maritima. Michael G. Barbour and James E. Rodman ........5,..« 370 Galinsoga ciliata (Compositae): Its Arrival and Spread in the Northeastern United States. Nancy N. Shontz and John P. Shontz ....... eene 386 Aster parviceps (Burgess) Mack & Bush and Aster brachy- actis Blake, New for Kansas. Kermit L. Johnson. cescccccsssercccessrsssssessencsaneccssseceesesseeeesnsansenennsees 392 Mock Bishop's Weed in the New World Tropics: Range Extensions for Ptilimnium capillaceum (Umbelliferae). W. G. D'Arey | eee eee entn entree 393 Kallstroemia in the Middle Atlantic States. Duncan M. Porter |n nennen nennen nennt 397 Nemophila microcalyx, an Incorrect Name. John W. Thieret |... rennen nnne nennen 299 Rhodora | JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 12 July-September, 1970 No. 791 COMPARATIVE STUDIES OF SMILAX, SECTION SMILAX, OF THE SOUTHEASTERN UNITED STATES IDA E. YATES AND WILBUR H. DUNCAN The species of the genus Smilax L. of the section Smilax (LILIACEAE) are unusual monocotyledons. In addition to possessing reticulate veined leaves, the taxa of this section are woody, dioecious vines with “stipular” tendrils. The combination of the above vegetative characters defines the Smilax section of Smilax as a distinctive, easily recogniz- able group. However, identification of specimens, especially sterile ones, to taxa within the section is often difficult. Also concepts of phylogenetic relationships between taxa are not clearly established and certain leaf characteristics pointed out by Duncan (1967) are not identified as to their structural nature. Therefore, attempting to alter these sit- uations, we investigated each woody species of Smilax in- digenous to the southeastern United States (Duncan, 1967), except S. pumila Walt., for epidermal and gross anatomical similarities and differences of leaves. The specimens examined in this study are of S. auricu- lata Walt., S. bona-nox L., S. glauca Walt., S. laurifolia L., S. rotundifolia L., S. smallii Morong, S. tamnoides L., and S. walteri Pursh. as interpreted by Duncan (1967). Three leaf forms of S. bona-nox (auriculate, ovate, and cordate), and two of S. tamnoides (orbicular and hastate, varieties hispida and tammoides, respectively) were treated sep- arately, making eleven kinds of Smilax studied. 289 290 Rhodora [Vol. 72 Confusion as to the delimitation of the species of Smilax dates back at least to 1753 with Linnaeus’ Species Planta- rum. As Fernald (1944) adeptly points out, Linnaeus had no clear understanding of American species, and his cita- tions cover different species. Other investigators have also disagreed regarding taxa, Morong (1894) describes S. smallii as a new species while maintaining S. lanceolata L. as a separate taxon and recognizes S. hispida Muhl. and S. beyrichii Kunth in sensu S. tamnoides L. and S. auricu- lata Walt., resnectively, Small (1933) assigns S. pseudo- china L. and S. tammoides to the species S. bona-nox and S. beyrichii, and S. smallii to the species S. lanceolata. He accepts S. hispida as S. tamnoides. Caponetti and Quimby (1956) studied anatomical struc- tures of roots, leaves, acrial stems, and rhizomes of S. tam- noides, S. auriculata, S. bona-nox, and S. glauca. They re- port the last species as the only one possessing papillae on the lower leaf epidermis and having thick-walled circular lignified cells in the region of the mesophyll beneath the midvein. They isolate S. bona-nox on the basis of the cells surrounding the midvein having thick lignified walls toward the midvein and thin non-lignified walls away from the mid- vein. Arber (1920) studied the anatomy of the tendrils of Smilax and interprets them as being equivalent in morpho- logical value to the petiole and having originated through that structure. She maintains that the blade of Smilax is not equivalent to the lamina of a dicotyledon, although quite similar in appearance internally, but is merely a “‘pseudo- lamina" representing an expansion of the upper region of the petiole. The thickened tip which characterizes the blade of some members of the genus is possibly the last relic of the unexpanded petiolavr apex. Thus each tendril on her in- terpretation is equivalent to the petiole and these to the pseudo-lamina. Coker (1944) made an extensive study of woody Smilax of the United States. He gives comprehensive descriptions and discussions of each species, pointing out important 1970] Smilax — Yates and Duncan 291 omissions and errors found in manuals and other writings. New data concerning underground parts are emphasized. Information is given on epidermal features of leaves of some taxa, but it is scanty, and mostly concerns stomates. Sixty-one fresh and dried specimens were utilized in this study. Identification by collection data and University of Georgia Herbarium acquisition number is given by Yates (1968). Two mature leaves were taken for study from each fresh collection, one larze and one small. The remainder of the collection was pressed, dried, mounted, and deposited as a voucher specimen. Additional material was obtained for study by removing one mature leaf from each of a num- ber of selected herbarium specimens previously on deposit and reconstituting it by treatment with 0.1 Normal NaOH at 609C for 24 hours. An outline drawing was made of each leaf chosen for study and a segment one mm wide cut out one-tenth of the distance of the total length beneath the tip. Two more seg- ments were taken directly below the first one. The leaf segments were placed in FAA (5 ml formalin: 5 ml glacial acetic acid: 90 ml 50% ethyl alcohol). Epidermal imprints were made from each leaf using a modification of the technique described by Sinclair and Dunn (1961). They were taken along the horizontal axis of the leaf from midvein to margin at a point approxi- mately one-third the total leaf length from the tip. Surface feature studies were first attempted from imprints made from dried material. Our stomatal counts were obtained from these preparations. Later, material preserved in FAA or reconstituted with NaOH was used to make imprints for analyzing the other surface features. Imprints were made from three leaves of each of five collections from each of the eleven kinds of Smilax. The terminology used to describe epidermal cells and venation is self-explanatory; however, those most likely to be misinterpreted will be described below. The others are described by Yates (1968). Veinal cells refer to those epi- dermal cells in areas above or below veins. Midvein and 292 Rhodora [Vol. 72 lateral veins are those arising at the base of the blade and extending longitudinally to the apex. Secondary veins are those extending horizontally from any one of the above veins to any other. Venule reticulum includes all the small veins in the leaf forming the anastomosing network char- acteristic of the genus Smilax. For the sake of brevity, the imprints of features of the epidermal cells will be referred to henceforth by the name of the feature involved. Configuration of lateral cell walls is described as straight, curved, or undulate. Straight de- notes those epidermal cells having at least three walls with- out undulations although the walls may be bowed, especially at junctions with adjacent cells, Undulate refers to those cells having at least two walls each with 2.5 or more un- dulations that are 25 ov more microns high. Curved indi- cates those epidermal cells which fall between straight and undulate. Microscopic counts and length and width measurements of the stomates were made in five fields of vision 0.093 mm? in area on imprints from three dried leaves of each collec- tion. If any part of a stomate was in the field of vision, it was included in the count. The means of these three characteristics were tested for significant differences by Duncan’s new multiple-range test (Steel and Torrie, 1960). For gross anatomical observations, the first segment cut from the large leaf in the procedure described previously was dehydrated through a tertiary butyl alcohol series, em- bedded in paraffin, and sectioned at ten microns. The sec- tions were mounted using a modification of Haupt's ad- hesive and stained with Conat's quadruple stain (Johansen, 1940). A second series of sections was stained using aniline blue and safranin as a countercheck on the former. The characteristics of the cuticle, epidermis, mesophyll, and vascular system were studied from these preparations. Two to five specimens of each of the eleven kinds of Smilax were studied anatomically. Freehand cross sections of dried, preserved, and recon- stituted material of selected species were stained with a 1970] Smilax — Yates and Duncan 293 1% solution of phloroglucin in 95% ethyl alcohol and 25% hydrochloric acid (Johansen, 1940), as a specific test for localization of lignified cell walls. Study of the epidermal imprints and cross sections re- vealed characteristics which varied from those common to all taxa of Smilax studied to those which were unique to single taxa. The interpretations presented below are neces- sarily based on characteristics of the leaf at the positions where data were obtained, namely one-tenth of the total leaf length beneath the tip for cross sections and one-third of the length for the epidermal imprints. We do not main- tain or imply that the characteristics are the same else- where. The surface features and gross anatomy of the leaves will be presented separately. Characteristies of the stomatal cells could be determined from epidermal imprints made from untreated dried ma- terial. However, imprints made from FAA preserved or NaOH reconstituted specimens were required for most characteristics of the other epidermal cells. The appearance of cells at the marginal ana veinal areas is that of parallelly aligned, rectangularly elongated struc- tures with striated surfaces and thick cell walls. On the upper epidermis, the interveinal cell type generally changes abruptly to the marginal type; whereas, on the lower epi- dermis, the change is gradual. The number of rows of distinctly marginal cells of the upper epidermis ranges from 8 to 14. Typically the number of rows of rectangularly elongated cells at the midvein is approximately three times greater on the lower epidermis than on the upper. Although there are exceptions within some taxa, the cells of the inter- veinal areas change abruptly at all veinal areas. At the midvein, the number oi rows of rectangularly elongated upper epidermal cells is 2 to 11; at the lateral veins, 2 to 8; and at secondary veins, 0 to 5. Generally, there are no rectangularly elongated cells above the venule reticulum on the upper or lower epidermis. The number of rows of whe a3 eee TX 1970] Smilax — Yates and Duncan 295 rectangularly elongated lower epidermal cells at the mid- vein is 5 to 50; at the lateral veins, 0 to 11; and at the secondary veins, 0 to 3. Although cells of interveinal regions vary considerably from species to species, there are some characteristics which are common to all. All cells of interveinal regions are ir- regularly aligned (Figs. 1, 2). Generally, undulated cell walls (Fig. 2) are thinner than straight cell walls (Fig. 1). At corresponding locations on the upper and lower epi- dermises, the cells are essentially the same size. Adjacent cells at any given interveinal location, with the exception of S. smallii, are essentially the same size. The stomatal-subsidiary cell relationship of all species in the terminology of Metcalf and Chalk (1950) is paracytic (Fig. 3). The distal ends of the stomates in all species are aligned at various angles with respect to the longi- tudinal axis of the leaf (Figs. 3, 4). Differences in micro- scopic counts and measurements of the stomates, without quantitative measurements and statistical treatment, are usually not sufficiently evident to a cursory examination with the microscope to allow quick identification of taxa. The mean stomatal frequency in a field of vision of 0.098 min? is 3.6 to 5.6; the mean stomatal length, 35.3 to 52.9 microns; and the mean stomatal width, 17.9 to 28.6 microns. Many differences in epidermal characteristics of the vari- ous kinds of Smilax studied were revealed by critical ex- amination of the imprints. Those individually unique to a given taxon will be presented first. The unique characters are as follows: S. auriculata: One of the two subsidiary cells is crescent FIGURES 1-6. Epidermal imprints, X 125. Fig. 1. S. auriculata, upper. Note indistinct features due to rough surface cell walls. Fig. 2. S. glauca, upper. Fig. 3. S. boma-noz, lower, showing large paracytic type subsidiary cells. Fig. 4. S. auriculata, lower. Note irregular outer surface of cell walls. Arrow at crescent shaped subsidiary cell. Fig. 5. S. rotundifolia, lower, showing striated subsidiary cells. Fig. 6. S. smallii, upper, showing large cells separated from each other by a row of much smaller cells. 296 Rhodora [Vol. 72 WV 4 E Ay A -, Oe MI AE sg! SERAIS X " w ` ^S .- s 2" eu "eel e Rh l s AE ° Ne STA > x X < — Ë X “ O TA a EE ANA EE E REE Ae QM PE qu. SSE T. SND A LZ Se 25 2l EA | eS Wf - M ba ` Mm < WA Ut SAR * d T^ "KT * : ays 1970] Smilax — Yates and Duncan 297 shaped (Fig. 4), rarely otherwise. In other taxa these are similar in shape to other interveinal cells (Fig. 5). S. smallii: Interveinal areas of the upper epidermis have large cells separated from each other by a row of much smaller cells, about one-sixth the size of the former (Fig. 6). Also, this is the only taxon having a combination of lateral cell walls of the upper epidermis with straight con- figuration and of the lower with undulate (Fig. 7). S. tamnoides: Rectangular cells are present and striated at the venule reticulum on the upper and lower epidermis. S. tamnoides var. hispida: Some marginal rectangular cells converge into minute marginal denticulations (Fig. 8). S. walteri: Rectangular cells are present at the venule reticulum only on the lower epidermis, Also, this taxon is the only one with both upper (Fig. 9) and lower epidermal cells having a curved configuration, and with the exposed surface of the upper epidermal cells appearing depressed in the middle (Fig. 9). Characteristics which are common to only two or a few kinds of Smilax are several: Stomates regularly present on the upper epidermis — S. rotundifolia, S. walteri (Fig. 9), frequently in S. tamnoides. These findings are essentially the same as those reported by Coker (1944) except that no stomates were found in S. tamnoides. Surface of lower epidermal cells quite irregular — S. auriculata (Fig. 4), S. laurifolia. Subsidiary cells larger than interveinal cells — S. bona- nox (Fig. 3), S. smallii (Fig. 7), S. walteri. Surface of most subsidiary cells striated — S. rotundi- folia (Fig. 5), S. tammoides (Figs. 10, 11), S. walteri. A few striations were observed in S. smallii and in the ovate leaf form of S. bona-now. Fig. 7. S. smallii, lower. Arrow at undulate cell wall. Fig. 8. S. tammoides var. hispida, upper. Fig. 9. S. walteri, upper. Arrow at stomate. Fig. 10. S. tamnoides var. hispida, lower. Fig. 11. S. tam- noides var. tammoides, lower. Fig. 12. S. glauca, lower. Arrow at stomate. [Vol. 72 Rhodora 1970] Smilax — Yates and Duncan 299 Rows of marginal rectangular cells on the upper epider- mis 10 or less — S. glauca, S. rotundifolia ; more than 20 — S. tammoides, S. walteri; 11-18 — the other four taxa. Lateral cell walls usually both thick and having a con- figuration that is straight in the upper epidermis — S. auriculata (Fig. 1) ; S. smallii (Fig. 6) ; usually in S. bona- nox; and sometimes in S. laurifolia, S. tamnoides var. tam- noides. The configuration is curved in S. walteri (Fig. 9) and in three collections of S. laurifolia from mesophytic habitats. In all other instances (S. glauca, S. rotundif olia, six collections of S. bona-nox, and two of S. tamnoides var. tamnoides) the lateral cell walls are thin and have an un- dulate configuration (Fig. 2). The specimens of S. bona- nox having straight walls in the upper epidermis are all from maritime sandy habitats, whereas those with undulate walls are from the Upper Coastal Plain, Piedmont, and Blue Ridge. Configurations of the lateral cell walls of the lower epidermis are the same as for the upper except in S. glauca (Fig. 12) which has curved; 5. smallii (Fig. 7), undulate; S. laurifolia (Fig. 13), always straight; S. tam- noides (Figs. 10, 11), always undulate; and S. boma-nox in which 6 of the 15 collections are undulate as compared to straight. Exposed cell surfaces of the upper epidermis rough — S. laurifolia (Fig. 14) ; S. auriculata (Fig. 1); sometimes rough in S. bona-nox, S. tammoides. They are smooth in other instances for the last two species and in the other taxa (Fig. 6). Exposed surfaces of the lower epidermal cells smooth (Fig. 11) — all taxa except S. auriculata and S. laurifolia. Fig. 18. S. laurifolia, lower. Note indistinct features due to ridges on surface walls. Arrow at straight cell wall. Fig. 14. S. laurifolia, upper. Note indistinct features due to rough surface walls. Fig. 15. S. tamnoides var. hispida, showing duct and marginal denticulation. Fig. 16. S. glauca, showing papillose lower epidermal cells and prominent midrih. Fig. 17. 5. laurifolia, showing large first row of palisade and lack of protruding midrib. Fig. 18. S. bona-noz, showing sclerenchyma cells in leaf margin. 300 Rhodora [Vol. 72 In the latter species the surface is so deeply ridged as to render imprints opaque when viewed with the naked eye. The ridges obscure cell wall outlines on imprints from S. laurifolia (Fig. 13) to the extent that under microscopic examination maximum illumination is usually necessary to discern the configuration of the lateral cell walls. The ratio of the number of rows of rectangular cells at the midvein of the lower epidermis to the number at the margin is lowest in S. laurifolia and S. smallii (0.3-0.7) and highest in S. bona-nox (0.6-2.0). The ratios for these and the other taxa are given by Yates (1968). The means for stomatal frequency per 0.093 mm2, and stomatal length and width measurements in microns are presented in Table 1. Means connected by one or more of the lines beneath are not significantly different at the levels indicated. Means not connected are statistically sig- nificant. Each species is indicated by the first letter of the specific epithet, and where applicable, the second letter designating the leaf form or varietal epithet. The leaf forms of S. bona-nox are indicated by Ba, Bo, and Be. The varieties of S. tamnoides are designated by Th and Tt. Sections stained with safranin-aniline blue proved to be more satisfactory for studies of the cuticle, and those stained with Conat's quedruple combination enabled a more discrete examination of the vascular system. Both staining procedures were equally revealing of other aspects of the gross anatomy. The surface of the cuticle in all species is smooth on the adaxial side and irregularly ridged on the abaxial side. In most species, it is thicker on the adaxial surface than on the abaxial surface and thicker still at the margins. At the leaf margins of most species, rib-like cuticular projec- tions extend inwardly between the radial walls of the epi- dermal cells, The continuity of the uniserate, compactly arranged epi- dermis is interrupted only by the presence of stomates and the aforementioned cuticular projections. Stomates are more distinct and more frequent in the lower epidermis of Smilax — Yates and Duncan 301 1970] X9) Əy} ur peure[dxo eie pue xnzwg jo SPU ejeorpur suormerae4qqy. "xe; ur uoeue[dxg o PAI 259 oy} 3€ * —————— — aad] AT 3€ Weoylusis JON, B — OI 98c 986 G82 GLE G96 $86 qa gI LITZ 661. BAM V od d ed S I UL d AL M [y ecg eza Teo SOG 66F 6v ver. Osh SOV 068 Ss I] gi V ed d od S A AL UL d D LOSE oec Lo eTO TAT O key LP ovy GPL $98 19'S D ed A S gi vd V od UL d AL .Cppusg JO spur uəA9[ə IOJ ,sueeur uəəajəq soUaTaYIp euj JO e»ueogruSIS ey} pu* 'suogoru Ut (TIT) U3pr^ pue (II) yu; [£3?uro3s pue (I) ¿uuu e60'0 Jed Aouonbery [ejeurojs Jo sən[6A UP I QLL 302 Rhodora [Vol. 72 all species than in the upper epidermis, where in most spe- cies they are completely absent. As seen in cross section, the epidermal cells adjacent to the guard cells do not differ markedly from other epidermal cells, In all taxa, the stomates are slightly sunken below, or are level with, the surface of the other epidermal cells. Outer and inner stomatal ledges as described by Esau (1953) project into the stomatal cavity. At corresponding locations on the adaxial and abaxial sides of the leaf, the epidermises are of approximately equal thicknesses. The cells of the epidermis of most species are rectangular and thick-walled, gradually changing into cir- cular cells with still thicker walls at the leaf margins and veinal areas. At the midrib, more cells are circular and thicker walled in the lower epidermis than in the upper epidermis; whereas, at the major lateral veins, as many cells possess these characteristics in the upper epidermis as in the lower epidermis. The epidermal cells contain no well developed plastids. The mesophyll is differentiated into palisade parenchyma on the abaxial side. The palisade parenchyma is compactly arranged and may be composed of from one to three layers of cells. The spongy parenchyma is loosely arranged and may be composed of from three to ten layers of cells, At the margins and veinal areas, the mesophyll cells are typ- ically thicker walled and more compactly arranged than mesophyll cells of interveinal regions. They may differ in size, shape, lignification of walls, and affinity for safranin. The veinal system is composed of collateral vascular bundles with the largest vein being directed along the me- dian longitudinal axis of the leaf. Sclerenchyma encloses the midvein and the major lateral veins. Surrounding the sclerenchyma is a bundle sheath composed of thin-walled parenchymatous cells with a chloroplast complement sim- ilar to the mesophyll. As previously stated, the tissue above and below major veins may differ from the mesophyll of interveinal areas. Randomly scattered throughout the mesophyll of the leaf 1970] Smilax — Yates and Duncan 303 cross sections are ducts (Fig. 15) which appear to be single celled and seem either to be void of contents or to contain compact clusters of elongated crystals. The ducts may have thick or thin walls. They may run longitudinally or at an angle. Many differences in the gross anatomy of leaves of the kinds of Smilax studied were revealed by critical examina- tion of the cross sections. Those individually unique to a given taxon will be presented first. The unique characters are as follows: S. glauca: Many lower epidermal cells are papillose and most outer walls convex (Fig. 16). The papillae are the most conspicuous in leaves of plants from the more moist habitats and least evident in those from drier habitats. These cells are flattened or rounded and lack papillae in the other species. Coker (1944) reports papillae for his “rough- ened” form of S. glauca but not for his “glabrous” form. In making no reference to papillae in the other taxa he studied, the impression is left that none were found. In our studies also, the ratio of the thicknesses of the cuticle at the margin of the blade to the thickness of the adaxial cuticle at an interveinal area 1.5 mm from the margin is always less than 2:1. Im all other taxa it is greater than 2:1. In addition, the ratio of the thickness of the cuticle of the upper epidermis to that of the lower is 4:1 or greater, whereas in other taxa the ratio is about 2:1 or less. S. tammoides: Uppermost palisade parenchyma cells are not as thick or are as thick as the upper epidermal cells (Fig. 15), whereas they are thicker than the upper epider- mal cells in all other taxa. Also mesophyll cells at the leaf margin differ from those of interveinal mesophyll by being irregularly shaped and small. S. tamnoides var. hispida: Marginal denticulations are discernible from serial cross sections (Fig. 15). S. laurifolia: The leaf thickness (about 670 microns) at interveinal areas near the midvein is greater (Fig. 17) than for the other species in which the thickness is usually between 330-370 microns (e.g. , Figs. 15, 16, 18). Upper- 304 Rhodora [Vol. 72 most palisade parenchyma cells are more than 2X as thick as upper epidermal celis (Fig. 17). These palisade cells may be nearly this large in S. rotundifolia. In all other taxa they are smaller. Also, S. lawrifolia is the only taxon in which the cuticle on the lower epidermis is thicker than on the upper. S. bona-now: More than 15 sclerid cells are in the leaf margin (Fig. 18). Other taxa have none or only one or two such cells, S. auriculata: Upper epidermal cells are seven or more microns thicker than the lower. In other taxa they are about the same thickness. Gross anatomical characteristics which are common to only two or a few kinds of Smilax are: Rib-like cuticular projections between marginal epider- mal cells — S. bona-noz, S. laurifolia, S. smallii, and S. wal- teri; often in S. auriculata and S. tamnoides. Some cell walls adjacent to sclerid cells around veins lignified, the opposite walls not — Š. bona-nox, S. tammoides. More sclerenchyma cells on the adaxial side of the major veins than on the abaxis! — S. auriculata, S. laurifolia. Cir- cular cells in the mesophyll at the leaf margin differing from cells of the interveinal mesophyll — S. laurifolia and S. smallii have smaller circular cells; S. glauca has circular cells but they are as large as adjacent interveinal cells. Conspicuously enlarged cells below the midvein — S. glauca; occasionally in S. rotundifolia and S, smallii. Midrib generally not protruding — S. laurifolia (Fig. 17) ; barely protruding in S. small/i. Smilax glauca possesses a most prominent, often ridged, midrib (Fig. 16). The midribs of the other taxa are prominent but are more rounded than in S. glauca. Although as indicated above there are differences be- tween cross sections of S. rotundifolia, S. smallii, and S. tamnoides, the general aspect of sections of the species are quite similar. Differences between some species occur in the safranin stained protoplast of the mesophyll and epidermis. The 1970] Smilax — Yates and Duncan 305 protoplast of the mesophyll of S. laurifolia and S. walteri is so abundantly stained that the sections are obviously darkened. This protoplast is stained in some of the other species, but there is no gross effect as in the above two species. Except for the two species, staining characteristics are not gross in nature and are not described here. They are described by Yates (1968). A few other minor differences in gross anatomical char- acters between taxa are given by Yates (1968). The results of the present study differ in several respects with the earlier leaf anatomical investigations of Caponetti and Quimby (1956). They report no papillae on the mid- rib of the lower epidermis. We observed some on the col- lections from mesophytic habitats. Their failure to find papillae may be due to their collections having been from xerophytic environments only. Also, neither of the staining techniques used in this study reveals the presence of lignin which they reportedly detected in the mesophyll cells abaxial to the midvein of S. glauca. No explanation for these dif- ferent observations seems possible from the information available. Their assumption that the extension of the scler- enchyma surrounding the midvein to the epidermis is char- acteristic of S. auriculata is the exception rather than the rule for specimens analyzed in the present study. Finally, S. bona-nox is not the only species, as they state, with the cells surrounding the sclerenchyma of the midvein having thick lignified inner walls and thin non-lignified outer walls. These peculiar cells are also frequently present in S. tam- noides. Because of the variation in the taxa involved, it is quite possible that the observations of Caponetti and Quim- by were limited to too few specimens to detect the vari- ability. The present study has also revealed the structural nature of the marginal ribs indicated by Duncan (1967) for S. bona-noa, S. auriculata, and S. laurifolia, Externally these ribs often seem identical to veins but were not labeled as such because no veinal connections with nearby veins were observed. In S. bona-nox the rib is largely due to 15 or 306 Rhodora [Vol. 72 more rows of sclerenchyma cells. In S. auriculata the rib seems due to the thick cuticle and compactly arranged sub- epidermal cells which also often has one or two scler- enchyma cells. In S. laurifolia a thick cuticle at the margin and compact subepidermal cells probably provide the rig- idity to form a rib when the leaf is dried. Frequently some of the sclerenchyma cells in the leaf margin of S. bona-nox are much larger than the others and look like vessels, How- ever, since our external and internal studies revealed no connection between the ribs and adjacent veins nor any phloem elements or bundle sheaths, we believe that the term “rib” should be continued in use for all three species. Analysis of the data of the present study shows that in respect to surface features of the epidermises, the nine taxa differ by unique and/or certain combinations of char- acters. The same is true for gross anatomical characters. These data, therefore, support the species concept in the section Smilax as interpreted by Duncan (1967). In ad- dition, the two varieties of S. tamnoides, as represented by the materials studied, differ in both surface features and gross anatomical characters. However, neither epidermal nor gross anatomical data were found separating the three leaf forms of S. bona-nox, Studies should be made in this species to determine whether or not specimens from mari- time habitats consistently and exclusively have straight lateral cell wall configuration in the leaf upper epidermis. Such plants may be Coker's (1944) S. bona-nox var. lit- toralis, Similar studies should be made in respect to the undulate walls in specimens from other areas. Also, more Study is needed to learn why the exposed epidermal cell walls in leaves of S. walteri appear depressed in epidermal imprints, yet as seen under microscopic examination of cross sections are not depressed. Keys presented below emphasize many of the characters most distinctive for the nine taxa and allow identification of specimens on the basis of epidermal or gross anatomical characters. Although the two varieties of S. tamnoides may be separated, in thc case of the materials we studied, 1970] Smilax — Yates and Duncan 307 we hasten to point out that only specimens distinctly one type or the other were examined. Further epidermal and gross anatomical studies are necessary to determine whether the minute denticulations on the leaf margin of var. hispida exhibit intergradation as do the leaf forms as described by Duncan (1967). Under a dissecting micro- scope both varieties appear to have denticulations, but ap- parently in var. tamnoides this is due to uneven crimping and rolling of the especially thin margins of the blade. Key to Taxa Based on Surface Features of Leaves 1. Upper epidermal cells of interveinal areas having larger cells each being separated from any other by a row of much smaller cells S. small, 1. Upper epidermal cells do not have such an arrangement of larger and smalle* cells 2 2. A crescent shaped subsidiary cell associated with the stomates on the lower epidermis .............. S. auriculata 2. No crescent shared subsidiary cell associated with the stomates on the lower epidermis 3 3. Lateral cell wall configuration of lower epidermis curved 4 4. Lateral celi wall configuration of upper epi- dermis undulated, stomates absent on upper epidermis S. glauca 4. Lateral cell wall configuration of upper epi- dermis curved, stomates present on upper epi- mis S. walteri 3. Lateral cell wall configuration of lower epidermis not curved .... 5 5. Lateral cell wall configuration on lower epi- dermis straight 6 6. Surface of lower epidermal cells of inter- veinal areas ridged iras: S. laurifolia 6. Surface of lower epidermal cells of inter- veinal areas not ridged ........... S. bona-nox 5. Lateral cell wall configuration on lower epi- dermis undulated ............. 7 308 Rhodora [Vol. 72 7. Lower epidermal cells associated with venule reticulum and striated (S. tam- noides) . .. 8 8. Marginal cells converge to form min- ute marginal dentations S. tamnoides var. hispida 8. Marginal cells do not converge to form minute marginal dentations .... a. n" S. tamnoides var. tamnoides 7. Lower epidermal cells associated with venule reticulum not different from those of interveinal areas ... s 9 9. Stomates present on upper epidermis, subsidiary cells about same size as in- terveinal cells „n. S. rotundifolia 9. Stomates absent on upper epidermis, subsidiary cells larger than interveinal cells S. bona-nox Key to Taxa Based on Gross Anatomical Characters of Leaves l. Fifteen or more sclerenchyma cells present in margin of leaf S. bona-nox l. Less than four sclerenchyma cells present in margin of leaf 2 2. Large napillose cells present in lower epidermis, ratio of thickness of the cuticle at the leaf margin to the thickness of the cuticle on the upper epidermis at an interveinal area 1.5 mm from the margin al- ways less than 2:1 S. glauca Large papillose cells not present in lower epidermis, ratio of the thickness of the cuticle at the leaf mar- gin to the thickness of the cuticle on the upper epi- dermis at an interveinal area 1.5 mm from the mar- gin always 2:1 or greater NEM 3 3. Uppermost palisade parenchyma cells not as thick as upper epidermal cells (S. tamnoides) ...... 1970] Smilax — Yates and Duncan 309 Marginal denticulations on leaf evident in serial cross sections ..... S. tamnoides var. hispida Marginal denticulations on leaf not seen in serial cross sections S. tamnoides var. tamnoides 3. Upvermost palisade parenchyma cells thicker than upper epidermal cells 5 5. wo) | Uppermost palisade parenchyma cells more than twice as thick as upper epidermal cells, cuticle on the lower epidermis thicker than on the upper S. laurifolia Uvpermost palisade parenchyma cells not more than twice as thick as upper epidermal cells, cuticle on the lower epidermis as thick as or thinner than on the upper .......5"—- 6 6. More sclerenchyma cells on adaxial side of major veins than on the abaxial side uu " S. auriculata 6. As many or fewer sclerenchyma cells on the adaxial side of major veins as on the abaxial side 7 7. Small circular cells present in the mesophyll at the leaf margin... — S. smallir 7. Small circular cells absent in the meso- phyl! at the leaf margin cscs 8 8. Mesophyll deeply stained with sa- framnin ...... a... ........... S. walteri 8. Mesophyll scarcely stained with safranin -ese S. rotundifolia We pointed out earlier that identification of sterile speci- mens to taxa is often difficult. Epidermal and gross anatom- ical characteristics revealed by this study help alleviate this problem. Gross anatomical characteristics are especially useful in the case of collections of S. bona-nox which, partly because of polymorphic leaf shapes, are often incorrectly identified as S. rotundifolia, S. tamnoides, S. auriculata, or 310 Rhodora [Vol. 72 S. smallii. Even though the character of the marginal rib of S. bona-nox leaves can give positive identification, the diagnostic value of the rib is lessened because the thin mar- gins which occur in some of the other taxa may be enrolled and appear enlarged, especially on dried specimens. Since our studies have revealed the margin to contain many sclerenchyma cells, positive identification of both dried and fresh specimens of S. bona-now may be accomplished from freehand leaf cross sections of approximately 5 mm thick- ness by staining with phloroglucin as described earlier. Within five minutes this stain will impart a purple-reddish color to lignified structures, and examination with a 10X lens will reveal the presence or absence of the unique abundance of marginal sclerenchyma cells for S. bona-nox. As is evidenced from the keys and other data presented earlier, all taxa possess both epidermal and gross anatom- ical characteristics that are distinguishing. However, ex- cept for S. bona-nox which is discussed above, epidermal imprints are more practical for identification purposes than are cross sections because of the time element and equip- ment involved. Imprint characteristics useful in checking identifications of specimens in some of the situations in which taxa are easily confused are pointed out below. Some specimens of S. smallii are similar to those of S. auriculata and S. laurifolia but may be readily distinguished from them by a unique upper epidermis where each larger cell is separated from any other by a row of smaller cells approximately one-sixth the size of the former. Smilax auriculata is easily distinguished from similar forms of S. tammoides and S. laurifolia by a crescent shaped subsidiary cell flanking each stomate. When specimens of S. glauca are dried, the glaucous na- ture of the underside of the leaf is often not apparent and the specimens are easily confused with those of S. rotundi- folia. They may be identified by the absence of stomates in the upper epidermis of the former and by their presence in the latter. Some leaf forms of S. tamnoides and of S. rotundifolia are similar. They may be identified by rectangular cells 1970] Smilax — Yates and Duncan 311 above the venule reticulum of both upper and lower epi- dermises in the former, and by a higher ratio (2.5-5.0/ 0.8-1.0) of rows of rectangular cells beneath the midvein to rows at the leaf margin in the latter. Smilax walteri, which is often similar to S. rotundifolia, is distinguished from the latter by the same ratios of rectangular cells. In addition, S. walteri has curved lateral cell walls on both epidermises while in S. rotundifolia they are undulate. Phylogenetic relationships between the Southeastern United States species of Smilax, excepting S. pumila which was not studied, seem more evident after an analysis of the data obtained in this study. A detailed presentment of presumed relationships does not seem justified here, but a summary of the more important aspects does. In coming to our conclusions, data published previously were consid- ered as well as the data we obtained. Smilax laurifolia is closely related to S. auriculata and S. smallii only, and these latter two species are much more closely related to S. laurifolia than to any other species. This is compatible with Coker’s (1944) data on under- ground parts. Because of these strong relationships the three species should be considered as a single group even though S. laurifolia and S. smallii are not strongly related. A second group of species consists of the closely related S. bona-nox, S. tamnoides, and S. rotundifolia. This group is loosely tied to the first by two similarities that S. bona- nox has with S. auriculata only and by one character it has with S. smallii only. Also, S. tamnoides has one char- acter in common with S. auriculata only. The remaining two species involved in this study, S. walteri and S. glauca, are not sufficiently related to be con- sidered as a third group. They stand instead as single taxa with relatively weak relationships to each other and to cer- tain taxa in the second group. Smilax walteri is more closely related to S. rotundifolia than any other species. This is also supported by Coker’s (1944) data on underground parts. Smilax walteri also has a character in common with S. tamnoides only. However, S. walteri has fewer ties to and more differences from the 312 Rhodora [Vol. 72 members of the S. bona-nox — S. tumnoides — S. rotundi- folia complex, than they to each other, and best should not be included with them. The several unique characteristics of S. glauca place it considerably apart from the other species. It is more closely allied to S. rotundifolia and S, walteri than any other species and therefore should be placed nearest them in any phylogenetic scheme. We wish to acknowledge the help of Robert Lane and Nancy Walker (Baker) in collecting some of the specimens from the field. DEPARTMENT OF BOTANY UNIVERSITY OF GEORGIA ATHENS 30601 LITERATURE CITED ARBER, AGNES. 1920. Tendrils of Smilax. Bot. Gaz. 69: 438-442. CAPONETTI, JAMES D. and MAYNARD W. QuiMBY. 1956. The com- parative anatomy of certain species of Smilax. J. Amer. Pharm. Ass. Sci. Ed. 45(10): 691-696. COKER, W. C. 1944. The woody smilaxes of the United States. J. of Elisha Mitchell Sci. Soc. 60: 27-69, DUNCAN, W. H. 1967. Woody vines of the southeastern states. Sida 3: 1-78. Esau, KATHERINE. 1953. Plant anatomy. John Wiley & Sons, Inc., New York. 753 p. FERNALD, M. L. 1944. Overlooked species, transfers, and novelties in the flora of eastern North America. Rhodora 46: 1-28, 32-60. JOHANSEN, D. A. 1940. P'ant microtechnique. MeGraw-Hill, New York. 523 p. METCALFE, C. R. and L. CHALK. 1950. Anatomy of the dicotyledons. Clarendon Press, Oxford. Vol. 1. MoronG, Tuomas. 1894. The Smilacaceae of North and Central America. Bull. Torr. Bot. Club 21: 419-443. SINCLAIR, C. B., and D. B. DUNN. 1961. Surface printing of plant leaves for phylogenetic studies. Stain Tech. 36: 299-309. SMALL, J. K. 1933. Manual of the southeastern flora. The author, New York. 1554 p. STEEL, ROBERT G. D. and JAMES H. ToRRIE. 1960. Principles and procedures of statistics. McGraw-Hill, New York. 481 p. Yates, IDA. E. 1968. Comparative studies of Smilax, Section Eu- smilax, of the Southeastern United States. Masters Thesis. Uni- versity of Georgia, Athens, Georgia. A PRELIMINARY CHECKLIST OF THE MARINE ALGAE OF CAMPOBELLO ISLAND, NEW BRUNSWICK, CANADA’ RICHARD A. STONE, EDWARD J. HEHRE, JOAN M. CONWAY AND ARTHUR C. MATHIESON Although several investigators have reported on the ben- thonic marine algae of the Bay of Fundy-Passamaquoddy Region of New Brunswick, Canada, none except Edelstein, Chen and McLachlan (1970) have made detailed seasonal investigations and few have interpreted the observed dis- tributional patterns. The present paper summarizes three vears of observations and collections on the benthonic ma- rine algae of Campobello Island, New Brunswick, which is in the southern portion of the Bay ot Fundy. The earliest record of marine algae from Campobello Island is given by Eaton (1873). He lists 11 species from Campobello in his account of the algae *in the vicinity of Eastport, Maine". Many other species mentioned are im- plied as being present. Farlow (1881) recorded only one species (Rhodophyllis dichotoma) from the Island. The collections of Hay (1887), Hay and MacKay (1886, 1888) and Fowler (1901, 1902) are from the environs of the Island, but they make no specific mention of Campobello Island. Klugh (1917) listed 24 species from selected sites on Campobello; again, many other species are implied as being present. Klugh was one of the first to record a variety of environments in the region and to discuss floristic dif- ferences in the Passamaquoddy-Bay of Fundy area. Bell end MacFarlane (1933), MacFarlane and Milligan (1965), Colinveaux (1966) and Edelstein, Chen and McLachian (1970) have discussed the distribution of marine algae in the Bay of Fundy and adjacent waters, The latter workers summarized a year round study of the vegetation at Digby "Published with the approval of the Director of the New Hamp- shire Agricultural Experiment Station as scientific contribution num- ber 518. 313 314 Rhodora [Vol. 72 BAY OF FUNDY | 66° 50° NEW BRUNSWICK ATLANTIC OCEAN Deer Is. Map of Campobello Island showing locations of collections. Neck, Bay of Fundy, Nova Scotia, including observations on seasonal changes, vertical distribution and reproductive structures present. Cardinal (1967a, b, c) recorded sev- eral new species from New Brunswick and Quebec, and he (Cardinal, 1968) has also given an excellent summarization of the distribution of the marine algae on the east coast of Canada. Collections and observations of marine algae were made at ten locations (Fig. 1) on Campobello Island, New Bruns- wick, Canada from May 1966 to May 1969. Table I sum- marizes the dates and sites of collections. The methods of collection and identification of specimens were similar to those of Mathieson, Hehre and Reynolds (in press) except that most collections were restricted to the littoral zone. The nomenclature of the Second Revised British Checklist 1970] Algae— Stone, Hehre, Conway & Mathieson 315 (Parke and Dixon, 1968) has been applied for most taxa, except the Acrochaetiaceae (Papenfuss, 1947). Herbarium voucher specimens of all collections (approximately 1,000) have been deposited in the Algal Herbarium of the Univer- sity of New Hampshire (NHA). Additional specimens are in the personal herbaria of the authors. The vertical dis- tribution of algae at Campobello Island is described accord- ing to the biological classification of Lewis (1964). Campobello Island is located approximately 1/3 mile off- shore from the coastal town of Lubec, Maine, near the boundary of Passamaquoddy Bay and the Bay of Fundy (Fig. 1). The maximum length and width of the Island are approximately 9 by 31/2 miles respectively. Much of the Island is covered with marshes and hills. The coast varies from steep granite cliffs to barrier beaches against marshy lowlands. The eastern shore is exposed to the greatest wave action. No major freshwater rivers are evi- dent, and the surface water salinities around the Island are probably uniform, Differences in vegetation are primarily due to physical factors such as wave action, tidal amplitude (average 15-20 feet) and substrate. A brief description of each station is summarized below. STATIONS HERRING COVE HEAD (Fig. 1, station 1): An exposed lo- cation at the north end of Herring Cove. The substrate con- sists of steep rock outcrops, which are impenetrable in some places. Many tide pools are present at all levels, and they provide a wide variety of habitats which are not found at any other station except Whiterock Cliffs. Collections were made at Herring Cove Head on most visits to the Island, and these provided a comparison for all other stations. RACCOON POINT (Fig. 1, station 2): An exposed location at the south end of Herring Cove. It is also designated as Con Robinson’s Point or Dinner Head on some maps. The substrate is similar to that at Herring Cove Head, except that the rock outcrops are lower. LIBERTY POINT (Fig. 1, station 3): An exposed location 316 Rhodora [Vol. 72 on the southern tip of the Island. The substrate is primarily composed of cobbles and small boulders. Few tide pools are present. GREATER AND LESSER DUCK PONDS (Fig. 1, station 4 and 5): Both locations are sheltered barrier beaches. The sub- strate is primarily muddy or mud-sand, but cobble and oc- casional boulders occur near low water. Freshwater stream beds empty into both of the bays. However, the freshwater outfall is limited and it only influences the vegetation in high, marshy areas. MULHOLLAND'S BEND (Fig. 1, station 6): A muddy area with occasional scattered boulders, It is located under the bridge at the southwest part of the Island. Although the area is protected from wave action it is subject to strong tidal currents. As is typical of other tidal current areas many species of seaweeds are very large in size. HARBOUR DE LOUTRE (Fig. 1, station 7): A cul-de-sac which is the most sheltered of our stations. The substrate is primarily mud, but occasional cobbles, fish weirs, and one large rock outcrop (Crabble Rock) provide a variety of habitats for seaweeds. Here, as at station 6, many species of seaweeds are very large in size. WILSON'S BEACH (Fig. 1, station 8): A sheltered shore near the town docks at Wilson's Beach — the only sizeable town on the Island. Collections were made from pier pil- ings, small scattered rocks and rock outcrops. EAST QUODDY HEAD (Fig. 1, station 9): A semi-exposed location at the easternmost tip of the Island. The substrate consists of steep rock outcrops, WHITEROCK CLIFFS (Fig. 1, station 10): An exposed location on the northeast side of the Island. The substrate consists of very steep rock outcrops, many of which are difficult to climb, A total of 114 taxa of seaweeds were collected at the ten stations, including 32 Chlorophyta, 39 Phaeophyta, 42 Rho- dophyta and 1 Xanthophyta. The distribution and seasonal occurrence of each taxa is summarized in Tables II-IV. 1970] Algae— Stone, Hehre, Conway & Mathieson 317 The following checklist includes brief comments of note- worthy features. XANTHOPHYTA Vaucheria sp.: Found on high, muddy substrates at Great Duck Pond; vegetative. CHLOROPHYTA Capsosiphon fulvescens (C. Agardh) Setchell e£ Gardner: On pebbles at Harbour de Loutre. *Chaetomorpha aerea (Dillwyn) Kützing: Found once (May) at Harbour de Loutre. Chaetomorpha melagonium (Weber et Mohr) Kützing: Common on rocks in the low eulittoral (in tide pools) and sublittoral zones of exposed locations. Cladophora flexuosa (O. F. Müller) Harvey (= Clado- phora sericeae (Hudson) Kutzing sensu C. van den Hoek, 1963) : Found once on a mud flat at Great Duck Pond. *Codiolum gregarium A. Braun: Found once (May) on rocks in the high littoral zone; mixed with C. pusillum. Codiolum gregarium may represent the “sporophyte” generation of one or more local species of Urospora (Scagel, 1966). Codiolum petrocelidis Kuckuck : Endophytic within Pet- rocelis middendorfii. It may be the “sporophyte” gen- eration of Spongomorpha spinescens or other local mem- bers of the Cladophorales (Scagel, 1966). Codiolum pusillum (Lyngbye) Kjellman in Foslie: Lo- cally abundant during the winter and spring; mixed with C. gregarium in the upper littoral zone. It may also represent the “sporophyte” generation of one or more local species of Urospora (see Scagel, 1966). Enteromorpha erecta (Lyngbye) J. Agardh: Common; free-floating or attached to small stones in the eulittoral zone. Enteromorpha groenlandica (J. Agardh) Setchell et Gard- ner: Occasional on rocks in the mid eulittoral zone. *Within known distributional range, but not previously recorded from New Brunswick. 318 Rhodora [Vol. 72 Enteromorpha intestinalis (L.) Link: Abundant on rocks throughout the eulittoral zone. Enteromorpha linza (L.) J. Agardh: Common on rocks in the lower eulittoral zone. Enteromorpha micrococca Kützing: Found once (August) on rocks in the high eulittoral zone. Enteromorpha minima Nägeli (= Blidingia minima (Nägeli ex Kützing) Kylin): Occasional on rocks in the high littoral zone. *Epicladia flustrae Reinke: Endophytie within Sertula- rians. Monostroma fuscum (Postels et Ruprecht) Wittrock: Lo- cally abundant on rocks and as an epiphyte in the lower eulittoral and sublittoral zones. Monostroma grevillei (Thuret) Wittrock: Common from late winter to early summer in the mid and lower eulit- toral zones. *Monostroma leptodermum Kjellman: A conspicuous epi- phyte on Zostera marina during the summer. Monostroma pulchrum Farlow: Locally abundant on rocks and as an epiphyte in the low eulittoral and sublittoral zones, Percursaria percursa (C. Agardh) Rosenvinge: Occa- sional on muddy, marshy substrates in the high littoral zone. Prasiola stipitata Suhr in Jessen: Occasional on high rocks (splash zone) in exposed and semi-exposed locations. *Pseudendoclonium marinum (Reinke) Aleem et Schulz: Common on rocks in the mid and lower eulittoral zones. *Rhizoclonium riparium (Roth) Harvey var, implexum (Dillwyn) Rosenvinge: Locally abundant on muddy, marshy substrates in the high littoral zone. Usually entangled among other green algae. Rhizoclonium tortuosum Kützing: Common among vari- ous plants in the lower eulittoral zone, Spongomorpha arcta (Dillwyn) Kiitzing: Abundant on rocks in the lower eulittoral zone at all stations. 1970] Algae— Stone, Hehre, Conway & Mathieson 319 *Spongomorpha hystrix Strómfelt: Found once (May) on rocks in the lower eulittoral zone. Spongomorpha spinescens Kützing: Locally abundant on rocks in the lower eulittoral (often in tide pool) and sub- littoral zones; rarely as an epiphyte. Ulothrix flacea (Dillwyn) Thuret in LeJolis: Occasional ; growing on rocks, muddy surfaces and epiphytic on vari- ous algae in the mid and high eulittoral zones. Ulva lactuca Linnaeus: Common as free-floating masses in muddy areas, or epiphytic on coarse algae in the eulit- toral zone. Only occasionally found attached to rocks or pebbles. Urospora collabens (C. Agardh) Holmes et Batters: Oc- casional on rocks in the upper littoral zone. As suggested by Mathieson, Hehre and Reynolds (in press) U. col- labens may represent a larger growth form of U. penicil- liformis. *Urospora penicilliformis (Roth) Areschoug: Occasional on rocks in the upper littoral zone. *Urospora speciosa (Carmichael ex Harvey) Leblond ef Hamel: Our material fits the description given by Edel- stein and McLachlan (1966). It occurs commonly on rocks in the mid and upper eulittoral zones, and occa- sionally epiphytic on coarse algae. Previously recorded from New Hampshire by Mathieson, Hehre and Reynolds (in press). Urospora wormskjoldii (Mertens) Rosenvinge: Uncom- mon; on rocks in the lower eulittoral zone. PHAEOPHYTA Agarum cribrosum (Mertens) Bory: Common on rocks in the lower eulittoral (tide pools) and sublittoral zones. Alaria esculenta (L.) Greville (including Alaria musae- folia (De la Pylaie) J. Agardh sensu Widdowson, 1964) : Abundant in exposed locations in the low eulittoral and sublittoral zones. During the winter the specimens are often reduced to residual portions of the stipe. Ascophyllum nodosum (L.) LeJolis: Ubiquitous through- out the eulittorai zone at all stations. 320 Rhodora [Vol. 72 Chorda filum (L.) Stackhouse: Occasional on small rocks in the lower eulittoral and sublittoral zones; also collected in drift. Chorda tomentosa Lyngbye: Common on small rocks in the lower eulittoral and sublittoral zones; also collected as free-floating specimens. Several had a larger diameter (over 4 mm) than plants recorded by Taylor (1957). Chordaria flagelliformis (O. F. Müller) C. Agardh (in- cluding var. densa Farlow) : Common on rocks, as an epi- phyte, and free-floating in the lower eulittoral zone, Fre- quently associated with Dictyosiphon foeniculaceus — particularly at muddy locations. Cladosiphon zosterae (J. Agardh) Kylin: Found once (August) growing on rocks in the lower eulittoral zone. Desmarestia aculeata (L.) Lamouroux: Common on rocks in the lower eulittoral (in tide pools) and sublittoral Zones, Desmarestia viridis (O. F. Müller) Lamouroux: Locally abundant on rocks in low tide pools and in the sublittoral zone, Dictyosiphon foeniculaceus (Hudson) Greville: Common in the lower eulittoral zone; often associated with Chor- daria flagelliformis as well as epiphytic on the latter plant. *Dictyosiphon macounii Farlow: Found once (August) at Little Duck Pond. Ectocarpus confervoides (Roth) Le Jolis: An occasional epiphyte on various coarse algae. Ectocarpus fasciculatus Harvey: Uncommon; on rocks in the lower eulittoral zone. Ectocarpus siliculosus (Dillwyn) Lyngbye: Found once (August) as an epiphyte on Laminaria. Elachista fucicola (Velley) Areschoug (including Ela- chista lubrica Ruprecht sensu Jassund, 1959) : A common epiphyte on Fucus spp. and Ascophyllum nodosum; occa- sionally found on Halosaccion ramentaceum. Fucus distichus L. subsp. distichus Powell: Abundant in high tide pools on exposed rocky shores, 1970] Algae— Stone, Hehre, Conway & Mathieson 321 Fucus distichus L. subsp. edentatus (De la Pylaie) Pow- ell: Locally abundant on rocks in the lower eulittoral zone. Fucus distichus L. subsp. evanescens (C. Agardh) Pow- ell: Distribution is similar to that of subsp. edentatus. However, it is more common in muddy habitats than the latter subspecies. *Fucus spiralis L.: Locally abundant on rocks in the upper littoral zone. Fucus vesiculosus Linnaeus: Abundant on rocks in the mid and lower eulittoral zones. *Fucus vesiculosus L. var. spiralis Farlow: Common in the mid and upper eulittoral zones of muddy areas. *Isthmoplea sphaerophora (Carmichael ex Harvey in Hooker) Kjellman: Collected once (August) as an epi- phyte on Sertularians at Raccoon Point. Laminaria digitata (Hudson) Lamouroux: Common in the lower eulittoral and sublittoral zones of exposed lo- cations. Laminaria longicruris De la Pylaie: Occasional in drift. Laminaria saccharina (L.) Lamouroux sensu Wilce, 1965: All of our specimens are the -- ecotype of Wilce (1965) or Laminaria agardhii in Taylor (1957). The plants are abundant in the lower eulittoral (in tide pools) and the sublittoral zones. Leathesia difformis (L.) Areschoug: Found once (Au- gust) as an epiphyte on Chondrus crispus. *Myrionema strangulans Greville: An occasional epiphyte on Ulva lactuca and Laminaria saccharina. Petalonia fascia (O. F. Miiller) Kuntze: Common (espe- cially during spring and summer) throughout the eulit- toral zone. *Punctaria latifolia Greville: Found twice (June and Au- gust) as an epiphyte on Zostera marina. Mixed with Monostroma leptodermum. Pilayella littoralis (L.) Kjellman: Common on rocks and epiphytic on various algae in the mid and lower eulittoral zones. 322 Rhodora [Vol. 72 *Ralfsia borneti Kuckuck: Occasional on small pebbles in the lower eulittoral zone. Ralfsia clavata (Harvey in Hooker) Crouan frat.: Found twice (February and June) on small rocks in the lower eulittoral zone. Ralfsia fungiformis (Gunner) Setchell et Gardner: Lo- cally abundant in mid and lower eulittoral tide pools of exposed stations. Plurilocular sporangia were found in February, 1969. Plurilocular sporangia have only been reported once before in North America (Edelstein, Chen and McLachlan, 1968). *Ralfsia pusilla (Stromfelt) Batters: Found once (Octo- ber) as an epiphyte on Chaetomorpha melagonium. Uni- locular sporangia were evident. Ralfsia verrucosa (Areschoug) J. Agardh: Found once (May) in a mid eulittoral tide pool. Sacchoriza dermatodea (De la Pylaie) J. Agardh: Locally abundant in low tide pools and in the sublittoral zone during late spring and summer. Scytosiphon lomentaria (Lyngbye) Link: Common throughout the eulittoral zone at all stations. Several specimens from Mulholland's Bend were larger (i.e. up to 200 cm long) than those recorded by Taylor (1957). Sphacelaria cirrosa (Roth) C. Agardh: Locally abundant on vertical rock faces which are covered with overhang- ing fucoids. *Spongonema tomentosum (Hudson) Kützing: Found once (August) as an epiphyte on the stipes of Laminaria. RHODOPHYTA Ahnfeltia plicata (Hudson) Fries: Occasional in sandy areas in the lower eulittoral and sublittoral zones. * Antithamnion cruciatum (C. Agardh) Nägeli: Found once (May) in a low eulittoral tide pool. * Antithamnion floccosum (O. F. Müller) Kleen: Occasional in tidepools in the lower eulittoral zone. * Audouinella membranacea (Magnus) Papenfuss: An oc- casional endophyte in the lorica of Sertularians. 1970] Algae— Stone, Hehre, Conway & Mathieson 323 Bangia fuscopurpurea (Dillwyn) Lyngbye: Common on rocks in the upper littoral zone at exposed locations. *Ceramium deslongchampsii Chauvin in Duby var. hooperi (Harvey) Taylor: Common throughout the year on ver- tical rock faces under overhanging fucoids. *Ceramium rubriforme Kylin, prox.: Found once (Au- gust) in the lower eulittoral zone at Little Duck Pond. Ceramium rubrum (Hudson) C. Agardh: Found three times (May, June and August) in drift. Chondrus crispus Stackhouse: Locally abundant in the lower eulittoral and sublittoral zones of rocky stations. Choreocolax polysiphoniae Reinsch: Common as a para- site on Polysiphonia lanosa. *Clathromorphum circumscriptum (Stromfelt) Foslie: Common in tide pools in the mid and lower eulittoral zones; also abundant in the sublittoral zone. Corallina officinalis Linnaeus: Common in the lower eulit- toral (in tide pools) and sublittoral zones. Cystoclonium purpureum (Hudson) Batters var. cir- rhosum Harvey: Common in tide pools and vertical rock faces in the lower eulittoral zone. *Dermatolithon pustulatum (Lamouroux) Foslie: Found once (February) as an epiphyte on Chondrus crispus. Dumontia incrassata (O. F. Müller) Lamouroux: Occa- sional on rocks in the mid eulittoral zone. Euthora cristata (C. Agardh) J. Agardh: Found once (February) in drift at Raccoon Point. Gigartina stellata (Stackhouse) Batters: Common on rocks in the lower eulittoral zone. Halosaccion ramentaceum (L.) J. Agardh: Common to locally abundant in the mid and lower eulittoral zones. Occasionally abundant in high tide pools. As described by Edelstein and McLachlan (1966) its morphology is extremely variable. Hildenbrandia prototypus Nardo: Common on rocks throughout the eulittoral zone. *Kylinia secundata (Lyngbye) Papenfuss: Occasional as 324 Rhodora [Vol. 72 an epiphyte on Spongomorpha spinescens and epizoic on Sertularians. “*Lithophyllum corallinae (Crouan frat.) Heydrich: Un- common as an epiphyte on Corallina officinalis, Previous- ly recorded from Rhode Island to Maine (Taylor, 1957). Lithothamnium glaciale Kjellman: Locally abundant in the lower eulittoral (in tide pool) and sublittoral zones. Membranoptera alata (Hudson) Stackhouse: Found oc- casionally as an epiphyte on Ptilota serrata, Petrocelis middendorfii (Ruprecht) Kjellman: Common on rocks in the lower eulittoral and sublittoral zones. Peyssonelia rosenvingii Schmitz: Found once (October) growing on Clathromorphum circumscriptum in the lower littoral zone. Phycodrys rubens (L.) Batters: Common in drift; occa- sional in tide pools in the lower eulittoral zone, and found once at -30 feet. Plumaria elegans (Bonnemaison) Schmitz: Occasional in the lower eulittoral zone at most rock stations; often on vertical rock faces under overhanging fucoids. Polyides rotundus (Hudson) Greville: Found occasionally in low tide pools. Xx **Polysiphonia arctica J. Agardh: Found once (June) at Whiterock Cliffs. Previously recorded from the arctic to Nova Scotia (as summarized in Taylor, 1957 and Edel- stein, McLachlan and Craigie, 1967). Polysiphonia flexicaulis (Harvey) Collins: Found twice May and August) as drift at Harbour de Loutre. Polysiphonia lanosa (L.) Tandy: Ubiquitous; hemipara- site on Ascophyllum nodosum throughout the mid and lower eulittoral zones, Polysiphonia urceolata (Lightfoot ex Dillwyn) Greville (Including var. roseola (C. Agardh) J. Agardh): Com- mon on rocks in the lower eulittoral and sublittoral zones. *Porphyra leucosticta Thuret. Found occasionally during the spring and summer in the lower eulittoral and sub- littoral zones. 1970] Algae— Stone, Hehre, Conway & Mathieson 325 Porphyra miniata (C. Agardh) C. Agardh: Common at most stations in the lower eulittoral and sublittoral zones during the spring and summer. Frequently free-floating and very large in size (over 10 feet in length at Harbour de Loutre). Porphyra umbilicalis (L.) J. Agardh: Abundant on rocks in the mid and upper eulittoral zones. *Porphyra umbilicalis (L.) J. Agardh forma epiphytica Collins: Common as an epiphyte on various coarse algae in the mid and lower eulittoral zones. Ptilota serrate Kützing: Common in drift; occasionally found attached in low tide pools. Rhodochorton purpureum (Lightfoot) Rosenvinge: Com- mon on vertical rock faces under overhanging fucoids; found once epiphytic on Sertularians. Rhodomela confervoides (Hudson) Silva: Common in tide pools in the mid and lower eulittoral zones. Rhodophysema georgii Batters: Found once (August) epiphytic on Zostera marina. Rhodymenia palmata (L.) Greville: Common in the lower eulittoral zone at all stations. Spermothamnium repens (Dillwyn) Rosenvinge: Found once (June) epiphytic on Polyides rotundus. Of the 114 species of marine algae found on Campobello Island, ten Chlorophyta, nine Phaeophyta and twelve Rho- dophyta are new records for New Brunswick. Two of the red algae also represent extensions of known distributional ranges: Lithophyllum corallinae was previously recorded from Rhode Island to Maine (as summarized in Taylor, 1957), while Polysiphonia arctica was previously known from the arctic to Nova Scotia (as summarized in Taylor, 1957 and Edelstein, McLachlan and Craigie, 1967). Eaton (1873) recorded Rhodophyllis dichotoma (designated as Calliblepharis ciliata) from Campobello Island, but we have not found it during our study. At present a total of 115 taxa of marine algae are recorded from Campobello Island. Several additional species (Eudesme virescens, Porphyra umiblicalis and Ceramium elegans) have been reported by 326 Rhodora [Vol. 72 Bell and McFarlane (1933) from the nearby locations of Lubec Harbor and Eastport, Maine. In addition, several other common species (e.g. Phyllophora brodiaei, P. mem- branifolia, Rhodophysema elegans, Polysiphonia nigrescens, Chaetomorpha linum and Punctaria plantaginea) which are reported from Nova Scotia by Edelstein and McLachlan (1966, 1967a, b) are conspicuously lacking from Campo- bello. Additional sublittoral observations may reveal some of these species. To date few sublittoral collections have been made, but preliminary observations have revealed a depauperate zone from 0 to 35 feet. Sea urchins (Strongylo- centrotus drobachiensis) were very conspicuous in the investigated sublittoral zones and the vegetation was re- stricted to scattered specimens of Desmarestia spp., Aga- rum cribrosum, a residual covering of crustose corallines, and a few small algae in cracks and crevices. An interesting aspect of the study has been the compara- tive composition of the flora at the various stations. A few preliminary comments can be summarized as follows: (1) The exposed, rocky shores support the greatest biomass and diversity of species. (2) Substrate is probably the most important single factor determining the floristic composition at different locations, although exposure to wave action is also an important and highly interrelated factor. (3) As described by Colinvaux (1966), there is a conspic- uous uplifting or emergence of sublittoral organisms on Campobello Island as compared to areas in central-southern New England. For example, Halosaccion ramentaceum, Alaria esculenta and Laminaria spp. are common in the mid intertidal zone (according to the physical delineation of tide levels) at Herring Cove, while they are restricted to the subtidal zone on shores of similar exposure and sub- strate in New Hampshire (Mathieson, Hehre and Reynolds, in press). (4) The relatively small number of species found on Camp- obello Island, as compared to the open Atlantic shores of 1970] Table I. Dates and locations of collections Algae — Stone, Hehre, Conway & Mathieson 696T 696T 696T 896I 896T 896T 896T 896T 896I 896I L96T LO6T LO6T 1961 196I ÁG] ABI ‘Qe “ony “ony “ony ST LI S6 cl LI OT Sny 6 ounf c ACT ABI eun f əun í eunf əunp “TRIN 6I ST LI 9I qI VI pe LO6T “FH € LO6T “OA P 996I “PO $6 9961 LLN IG 9961 LLW 06 Stations X X x x X 1. Herring Cove 2. Raccoon Point 3. Liberty Point 4, Great Duck Pond X Little Duck Pond 6. Mulholland's Bend 7. Harbour de Loutre 8. Wilson's Beach 5. x x 9. East Quoddy Head 10. Whiterock Cliffs 3 2 7 328 Rhodora [Vol. 72 Nova Scotia, may be due to its relatively sheltered location at the mouth of the Bay of Fundy. The algal flora of Campobello Island is primarily com- posed of subarctic and boreal species, but a substantial com- ponent of cosmopolitan species is also evident. A compar- ison of the marine algae from Newfoundland (Mathieson, Dawes and Humm, 1969), Campobello Island and Jaffrey Point, New Hampshire (Mathieson, Hehre and Reynolds, in press) indicates a high degree of similarity between the tnree areas. Even so there is a higher subarctic component in Newfoundland than at Campobello Island; in turn the boreal and subarctic components are higher at Campobello Island than at Jaffrey Point, New Hampshire. Thus the marine flora of Campobello Island reflects its geographical location. We wish to thank the following individuals for their help: The Franklin D. Roosevelt International Park Commission for their financial aid; Drs. A. R. Hodgdon and R. B. Pike for their encouragement and enthusiasm and for originally interesting us in the work; also to Dr. R. B. Pike and Mr. Summer Pike for their hospitality and help during our visits to Campobello Island ; and Dr. C. I. MacFarlane for supplying information concerning collections of material from Campobello Island. DEPT. OF BIOLOGY UNIVERSITY OF MASSACHUSETTS ROSTON, MASSACHUSETTS 02116 DEPT, OF BIOLOGY SOUTHAMPTON COLLEGE OF LONG ISLAND UNIVERSITY SOUTHAMPTON, NEW YORK DEPT. OF BOTANY UNIVERSITY OF MASSACHUSETTS AMHERST, MASSACHUSETTS 01003 DEPT. OF BOTANY UNIVERSITY OF NEW HAMPSHIRE DURHAM, NEW HAMPSHIRE 03824 1970] Algae-— Stone, Hehre, Conway & Mathieson 329 REFERENCES BELL, H. P. and C. MACFARLANE. 1933. The marine algae of the Maritime Provinces of Canada. I. List of species with their dis- tribution and prevalence. Can. J. Res. 9: 265-279. CARDINAL, A. 1967a. Inventaire des algues marines benthiques de la baie des Chaleurs et de la baie de Gaspé (Quebec). I. Phaéo- phycées. 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Species of Chlorophyceae new or rare to Nova Scotia. Can. J. Bot. 45: 211- 214. EDELSTEIN, T., L. CHEN and J. MCLACHLAN. 1968. Sporangia of Ralfsia fungiformis (Gunn.) Setchell and Gardner. J. Phycol. 4: 157-160. 1970. Investigations of the marine algae of Nova Scotia. VIII. The flora of Digby Neck Peninsula, Bay of Fundy. Can. J. Bot. 48: 621-629. EDELSTEIN, T., J. MCLACHLAN and J. S. CRAIGIE. 1967. Investiga- tions of the marine algae of Nova Scotia II. Species of Rho- dophyceae new or rare to Nova Scotia. Can. J. Bot. 45: 193-202. FARLOoW, W. G. 1881. The marine algae of New England. Rep. U. S. Fish and Fisheries for 1879, App. A-1, 210 p. (issued 1882) FOWLER, J. 1901. A visit to St. Andrews, N. B. with a catalogue of plants collected in its vicinity. Proc. Nat. Hist. Ass's, Mira- michi. 1: 21-28. 330 Rhodora [Vol. 72 1902. Report on the flora of St. Andrews, N. B. Contr. Can. Biol., 1901-1905: 41-48. Hay, G. U. 1887. Marine algae of the Maritime Provinces. Bull. New Brunswick Nat. Hist. Soc. 1: 62-68, Hay, G. U. and MacKay, A. H. 1886. Marine algae of Bay of Fundy. Ibid. 1: 32-33. 1888. Marine algae of New Brunswick: with an appendix containing a list of the marine algae of the Maritime Provinces of the Dominion of Canada. Trans. Roy. Soc. Canada 5: 167-174. HOEK, C. VAN DEN. 1963. Revision of the European species of Clado- phora. xi 4- 248 p. Leiden. JASSUND, E. 1959. Elachista lubrica Ruprecht and Elachista fucicola (Velley) Areschoug. Bot. Mar. 1: 101-107. KLUGH, A. R. 1917. The marine algae of the Passamaquoddy Re- gion, New Brunswick. Contrib. Can. Biol., Suppl. 6th Ann. Rep. Dept. Naval Serv., Fisheries Br., pp. 79-85. Lewis, J. R. 1964. The ecology of rocky shores. xii + 323 p. Eng- lish Univ. Press Ltd., London. MACFARLANE, C. I. and G. M. MILLIGAN. 1965. Marine algae of the Maritime Provinces of Canada, a preliminary check list. Nova Scotia Research Foundation, Halifax, 24 pp., mimeo- graphed. MATHIESON, A. C., C. J. Dawes and H. J. Humm. 1969. Contribu- tions to the marine algae of Newfoundland. Rhodora 71: 110-159. MATHIESON, A. C. E. HEHRE and N. REYNOLDs. Investigations of New England marine algae I. A floristic and descriptive ecologi- cal study of the marine algae at Jaffrey Point, New Hampshire. Nova Hedwigia (in press). PaPENFUSS, G. F. 1947. Further contributions toward an under- standing of the Acrochaetium — Rhodochorton complex of the red algae. Univ. Calif. Publ. Botany 18: 433-447. PARKE, M. and P. S. Dixon. 1968. Check-list of British marine algae — second revision. J. Mar. Biol. Ass. U. K. 48: 783-832. SCAGEL, R. F. 1966. Marine algae of British Columbia and north- ern Washington, Part I: Chlorophyceae (green algae). Nat. Mus. Canada, Biol. Ser. No. 74, Bull. No. 207; viii + 257 p. TAYLOR, W. R. 1957. Marine algae of the northeastern coast of North America. viii 4- 509 p. Univ. Michigan Press, Ann Arbor. Wippowson, T. B. 1964. A study of variation in the genus Alaria Greville. Ph.D. thesis, unpublished. University of British Colum- bia. ix + 229 p. WircE, R. T. 1965. Studies in the genus Laminaria. III, A revision of the North Atlantie species of the Simplices section of Lami- naria. Botanica Gothoburgensia III, pp. 247-256. 331 Algae — Stone, Hehre, Conway & Mathieson 1970] 9 ‘Gc G wnw42pojdo] vw042s0w0]yf 01 9 oT ‘eS SE 8 19]]4248 vwosouo W 6 ‘9 SFI I J, unasn{ mwuo042s0w0]y 6 'T on4jsnjf vippjoid y OI ‘T DUM DYALOWOLAIUA G 209909040%w, DYALOWOLAIU A "L'9 9 “g 6 PSG I Sz ‘L'I 8 DZU) Dd.2ow042]7]T Lv LY OT “6 'c 4 *6 “I I SUDUYSIJUL DydsowmosaqUg Y y I DIPUDIUGOLB DYALOWOLIIUT 6 'L ‘9 “G “2 g 6‘L‘T Z 039042 Tid40w0423u7] I I wnpsnd wmnjoio) T OT cT $2p11220429d. wnjoipoy I wninBa4AB wnjoipo) Y )Dsonzo vLoydopn) yg OT OI I *6 'c “6 “I Ç iwuniuoBnjou, Dnydsowojanyy jj Daan VDYALOUWLOLADYD L L L $uaosoa]nf uoydisosdng V f f NW Vo NW A f ‘I QEL ur Suorejs ƏY} 0} 1ogox suUIN[OD əy} ur Sioquinu ƏL 'oveoAudoio[u) Jo e»ueiinoo() Jeuoseəg pue uorjnqrtnsHT IT ?Iq*.L [Vol. 72 Rhodora 332 upjotz[su40m naodsoa y] nsowads n4odso4 7 Simsofipinuad D4odsoa y] $u9qnj]oo n.codso. N DINJIN) NAIN DIIDY L144 7] suaosouids pydcomohnods wish pydsomobunods njn DYydLomohuodsy Musonjao] wnio]oozu[or wnzojdiui “ABA MNMDALL 0006190292] WNULIDU wuniuojoopuopnosq DIDS nosni Ds.4na4od. 0408402942 WUNAYINE Duto4)8010 [y OT 'e OI “6 “c 6 fF ‘6 'c 'I L‘T 6 ‘F OT L‘T 8 6 I 6 fp 9 “G 0 8 'e fc 6 'L Z “I 6 0T ‘6 ‘LT 9P OT “I "p “Z 6 *I L OI ‘6 fL OT “6 “L OT 8 “FST -e ‘G-I *6 “L ST 6 'L ‘9 fe *T e fp 6 e fp cp 6 BT 8 “T L‘T 8z Z I 8 'c e'p c OT '6 'e-T 6 'T v f W Ei (3u09) II QLL 333 Algae-— Stone, Hehre, Conway & Mathieson 1970] L ‘On I M DIDSSDLIUL 013u0Wm(] T n371njsnd. uo 711010w42(1 6 'c 0T '6 OT '6 š 4RS02/440O “ABA Sisa] wnaimdind niwuo]2038fi 7) OI '6 Z [í '& € 'T e 0r'T 1 supuMfo DuyIDL0D I I 6 ZU UNITI SUNIN wnydiromosym p 0T ‘6 i “e “T T amuoydishjod 3102094017) 9 0T 8 'L 'e ‘T 6 'L ‘6 “I ST sndsida Snapuom) 6 p L WNLQHL nuno) e 240 huqta Wn?) OT wadooy "IBA Z *6 'e Or ‘6 PUTA usdiwm2Dwojsop wmiunaa) OT ut ‘Gui *6 ‘T no4ndandoosnfí mbung OT “6 ot 6 DIIDUDAQULAU Dnjyowwmnopny 6 “T OT 2 nsoo20y wowwm[ynqwy 0T WNIDVNMI UCUUDYIYUP 8 I »j»ond myəfuyy O V f f N V IN JA f avevAydopoyy Jo uN [euoseog pue uorjnqtisHT[ “TIT ALL [Vol. 72 N oO Hm N Rhodora 0]0]022.nm Diuoydish]og nsoun) miuowdisfijoq synvaacay miuodisfijoq 0212240 Diuowdisfijoq sSnpunjo4 sapihjog supbaja Manun) qg suəqna sñapoofin nBwiauoso4 DYAUOSShag WLOpUuappluUL $1]2204294q DjD]D 0422d0WD4QWUO JA] 91019010. wowwn101] aDUYyDL0I wmnpfiodowjv] D)»0pwunoos VDiuyhy sndfijojoud mpwupbaquappg WNIIDJUIWDL 1,00129D80]D H D]03]938 Duàj4DDir) D20]8149 DLOUINA 334 (3u05) III ə[qed 335 suada. wowaum10wods 020wQ0d. nuamfipoy q whioab nuashydopoyy Sopioa4ofuoo Djamopoyy wuno4ndand. wo340i20poqar 0]D4498 DIOP 4 paiphiydida ` f sypoyiqun Dihydsog sypoyiqun vihyd.og n30vu1uL Dihydso T 9019180279] Dfld.toq Algae — Stone, Hehre, Conway & Mathieson 1970] (3002) III QEL [ Vol. 72 Rhodora 336 L‘g 6r "y 'c T “ë “T MONINE DISIA 1 snsopnayis SNAANIOJI M L‘T snjn]noaispD| snduno0jaq FU 9 Or sapioriafuor sndano0j27, S nunosnw wowdisofijowT 6 ‘LF 6 snaonjpnamaof “I ‘L-T LT Woidtisofi]oi OL 9 I EZ I Spa vijso4puieso( 6 'c OT I ZI sg L “I ZT D)Do]non Dijsaavusacy [4 9D42j802 woidisopn,) 6 ‘LG 6 L [ PZT -G ‘G-T LT L Z “I simao finan 0i0paoo 0T ‘6 “L OT 6 -G “g “T G‘ DSOJUAWO) DPLOYD 6 LF L “I un npaouz) 6 'L I “FST OI 9 6T L ZT wnsopou unpphiydoos y 6 0T '6 ‘9 [ PZI ‘SST 6'I ZI njuəpnəsə many 6 '9 0T '6 '9 I PZI ‘§ ST 6'I ZI WUNSOLQULI nan y O S V f f W V N A f AVAOoAHdOMVHd ovooAydoyjuryx pue svaodydoovyg jo eoue.linoo() [euoseog pue uornnqrgsuq “AL QEL 337 Pa w r 1970] Algae — Stone, Hehre, Conway & Mathieson synio vilowptd nqofuv] v140j9unq MISD T1w0]019d supnjn0wDA]S nwar fo py suuuofjip visoi10o'T adJ — DULMDYIIVS DD MD] sium4owwo]. 0140100 ppp 000] poydosanyads napdouy is] sypuds “ABA snsopnaisad SNINA snsopnaisad SNIMA syouds snon,] suaosaupaa “dss suqouswp SANMA snjpjuopa “dss snyrysup SNINA snyoysip “dss snyoysyp SNINA o m ('3u09) AI IEL [Vol. 72 Rhodora 338 F I I 6z OI ‘T OT 6 ‘9-F OT ‘6 6 'e T ‘L-I ‘L'I 9 ZI ç fp I OT I ‘GI 6 “T OT G fp T V f f N ‘ds piwayonn 4 AVHIOAHJOHANVX SHnsojuowoj n wauofiuwodg p$04412 0140199741 d Ç CN L 9 D14019) uoidisojfiog Dapojnuaap DZt40990g DSOONA420 Dis {py Djjisnd. misjmor z “I SrutLofilunf Dis {Dy I DIVA] wis {paz I 192u40Q MS fima —_W W A f (77u03) AI Ə[qeL A LINDEN (TILIA) FOREST ON CAPE COD (WITH EXTENDED NOTES ON TILIA NEGLECTA, BROMUS PUBESCENS, AND RIBES HIRTELLUM). HENRY K. SVENSON An extensive area of dunes, known as Sandy Neck, ex- tends for some six miles along the north side of the Cape in the town of Barnstable. It lies between the ocean and the Great Marshes, one cf the largest salt marshes in Mas- sachusetts. About halfway down the neck and within the dunes is a swamp hollow with red maples, bordered on its southern bank by a little forest of native linden or bass- wood trees (Tilia neglecta). There are about a dozen clumps, interspersed among other trees. On the ground is a lush spring growth, a foot or two high, consisting of the grass Festuca obtusa, columbine (Aquilegia canadensis), Geranium maculatum, Smilacina stellata, Aralia quinque- folia, and Thalictrum revolutum. Less abundant are Smila- cina racemosa, meadow rue (Thalictrum polygamum ), and Circaea quadrisulcata, It is reminiscent of the dense her- baceous undergrowth of the Potomac River flats above Washington, but lacks the numerous kinds of violets on those flats. Vines of the small-fruited summer grape ( Vitis aestivalis) climb high in the trees, but only one vine was seen in fruit, and that far up out of reach. Some of the ancient maples in the swamp are uprooted and fallen. There are a few willow trees ( Salix Bebbiana) and a single bush of Salix cordata, The middle is practically bare except for dead leaves, the depression extending to a width of about 500 feet. A few bushes of [lex verticillata, with unusually long leaves and solitary fruit, extend into the open area; there are mats of marsh fern and Lycopus uniflorus and a few dwarf cinnamon ferns, but no trace of the Massachusetts fern (Thelypteris simulata), abundant in maple swamps at Osterville on the south side of the Cape. Toward the margin are some isolated clumps of sedges and rushes, including Carex seorsa, C. intumescens, 339 340 Rhodora [Vol. 72 Juncus effusus var. solutus, J. canadensis, and J. tenuis with extremely narrow leaves. The north-facing bank, some thirty feet high, is the most interesting area. It has the Tilia trees. At the lowest level are marginal and spinulese shield ferns, and some polypody fern. On this bank are sedges, Carex Swanii, C. debilis var. Rudgei, C. E'mmonsii, and C. blanda, the last-named previously known only from Osterville at a locality I can- not find. Toward the entrance to the swamp are a few sparse plants of Bromus pubescens, a grass new to south- eastern Massachusetts, Of general occurrence are Trien- talis and Canada mayflower and poison ivy, also common trees of the Cape: black and white oak, tupelo, holly, sassa- fras, and pitch pines, Witch hazel (Hammelis), scattered flowering dogwood trees (none producing flowers), and the viburnum which passes as V. venosum are on the moister parts, The only introduced plant appears to be Veronica arvensis, which grows sparingly in little openings, The linden trees are 35 to 60 feet high, each with several trunks 6 to 18 inches in diameter radiating from ground level. Only one clump was flowering. They are Tilia neg- lecta, whatever that name may represent. Spach, in Ann. Sc. Nat. ser. TI. ii: (1834) pl. 15, gave a detailed account of European and American lindens, including small figures of flower and fruit. Tilia neglecta came from “ambulacris Horti Parisiensis" and flowered at the same time as T. nigra Borkhausen, Forstbot. 2: 1220 (1803) (synonym of T. americana L.), which it resembled in stature, leaves, flowers, and bracts. Spach described the petals as “pallida lutea”, and the leaf as "subtus puberula”. Fernald, in RHODORA 412: 604-607 (1941), in a review of Tilia, said that few collections of T. neglecta could be closely matched, that leaves were green or merely grayish beneath with loosely scattered stellate hairs and simple pilosity, and that the Series was close to T. americana. In Castanea 20: 58 (1955), under the new name T., amer- icana var., neglecta, Fosberg stated that the difference lies in purple twigs and slightly stellulate-pubescent and more 1970] Tilia Forest — Svenson 341 or less glaucous under surfaces of the leaves." He remarks that Little (1953) “has reduced T. neglecta outright to T. americana, but this seems too drastic." Little was not being drastic; he used a question mark; he was just being careful. Now there are varietal synonyms that antedate var. neglecta, For example, T. nigra vestita A. Br. is listed by Rehder, Man. Cult. Trees and Shrubs, ed. 2: 625 (1940), derived from T. nigra B A. Braun in Doll, Rhein Flora, p. 674 (1843). By the Austrian J. N. Bayer, in Verhand. Zool.-bot. Ges. Wien 12: (1862) t. VIII, f. 4, Tilia neglecta was illustrated from a Spach specimen, showing leaf, foliar bract, and buds; T. nigra B was said to be sparingly stel- late-pubescent below, cited as coming from Kentucky and cultivated in Europe. It became T. americana, var. vestita (A. Br.) Victor Engler, Monographie der Gattung Tilia, Breslau (1909) p. 41. Engler described many forms and varieties, but was not too sure of var. vestita, separated with difficulty from the putative hybrid T. americana X heterophylla. This hybrid, he says, is questionably T. Mich- auxii Nuttall, in which pubescence of the lower leaf surface is greatly reduced. All of which goes to show that the European botanists knew no more about the subject than the American botanists did. Another varietal name to con- tend with is T. americana B pubescens Dippel, Handb. Laub- holzkunde 3: 63 (1893). It was described as having “‘foliis subtus pilis stellatis + pubescentes, interdum suborbicu- lata", and cited from Kentucky (Short), Pennsylvania (Gray!), and Stony Man Mt., Virginia (Steele no. 78). Differences between T. americana, T. heterophylla, and T. neglecta are pretty weak (cf. Rehder, p. 622). In pub- lishing T. glabra var. neglecta, Bush in Bull Torr. Cl. 54: 235 (1927), thought it might be only a form with straight simple firmly-attached pubescence. He cited specimens from coastal Connecticut (Guilford, July and Aug. 1921, Tre- lease). Sargent gave the range of T. neglecta (in Man. p. 133) as from Montreal to the coast of Massachusetts and New York, through the middle states to the valley of the Potomac River, then west and south, Rehder, the best de- x x * X|. |x x KIITIS — ^ FFE ae "LACE Ma aujoy Sn : LAM e s : f XALAT | d ` A AWE ia LL et LP p z vDunIIa aw D 0,02) bau Dil hydoaaya 1970] Tilia Forest — Svenson 343 scriber, says leaves are “oreenish or grayish and loosely stellate-pubescent beneath, with scattered mostly simple hairs on the veins, often tinged brownish ; petals pale yellow and 8 mm. long.” At Sandy Neck the leaves are lightly pubescent below, of firm straight hairs mixed with easily dislodged 2-4-branched flattened hairs. Bud scales and pedicels are moderately stellate (see fig. 2). The flowers are light brown and strong- smelling, and the petals reach 8 X 3 mm, about the size in Spach’s figures. In eastern Massachusetts Tilia neglecta appears as a tree of salt marsh islands or associated shores, recorded in the herbarium of the The New England Botanical Club from Mass.: Carr’s Island in salt marsh, Newburyport, Bean, Macgregor & Knowlton in 1945; abundant in seashore thicket, Hingham, Knowlton in 1943 (ripe fruit); Samp- son's Island, Orleans, Murdoch in 1913 (no flowers or fruit; Fig. 1. Tilia hair types: lower leaf surface, pedicel, and sepal. Boiled fragments, 1-5 mm in diameter, placed on slide in small drop of glue. Hairs removed with needle; some transferred to 0.5 mm-grid eyepiece on slide. T. heterophylla, Osterville, July 19, 1969 (no. 2526). Hairs in felt- like flattened layers, 120 on 1 sq. mm, mostly 8-pronged; some un- branched (see fig.). Stellate hairs of petiole and flower bracts in a dense felt-like layer. T. neglecta, Sandy Neck, July 4, 1969 (no. 2480). Count of flat hairs in a dense area 5 mra square of lower leaf surface: 4-pronged 110 (15%); 3-pronged 30 (596); 2-pronged 30 (596); 5-pronged very rare; erect type, unbranched, firmly attached to minute veins, 500 (75%); all these hairs hyaline. Axillary tufts of fascicular hairs: commonly 0.5-0.6 mm long, thickened and brownish. Pedicel: 20 scat- tered hyaline hairs on 1 sq. mm, chiefly 8-pronged; some fascicled and erect. Sepal: as on pedicel, but dense and felt-like; some erect and sinuous. On flower bract (not shown): small sparse 6-pronged on axial face, larger and 4-6-pronged on abaxial face. Minute scattered reddish glands are along veins on both surfaces of leaf. T. americana, Milton, July 9, 1969 (no. 3492). Axillary fascicular hairs as in T. heterophylla. Pedicel: 20 scattered flat, mostly 8- pronged hairs of varied size. On midrib of leaf bract: some widely scattered flat hairs, mostly 8-pronged. (not shown). On larger veins a few scattered simple hairs (not shown). 344 Rhodora [Vol. 72 noted as the only tree on the island). Conn.: “a large old tree near the coast", leaves and bark collected by Mrs. E. G. Hart. In addition to these citations, I find an old wind- blown tree without flowers or fruit at the entrance (salt marsh) to Dowse’s Point, Osterville. I fail to find Fernald and Long’s tree “about 7 feet high in 1918” at the mouth of Red River, Harwich, which has leaves loosely matted- stellate below and is maintained as T. heterophylla by Fev- nald and as T. neglecta by Sargent. As to other lindens on Cape Cod, there is a flowering tree of T. heterophylla on abandoned land on Tower Hill Road, Osterville (no. 2526). It has fragrant pale yellow flowers and thick-felted in- dumentum on the lower leaf surface. Nearby is a tangle of Populus alba, high-climbing Celastrus orbiculata, and bushes of honeysuckle (Lonicera Morrowii), three of the commonest introduced and escaped woody species on Cape Cod. T. americana. is frequently planted; T. europaea is an abundant escape from cultivation. The Sandy Neck trees are in an isolated place, and possibility of hybridization is remote. I have treated only the fringe of the problem, the great complexity of which lies in the southern Appalachians. We now come to the brome grass. Bromus pubescens Muhl. occurs sparingly (only two flowering stems observed) along with Triosteum, Galium circaezans, and G. triflorum. In no. 2481 (July 4, 1969) the leaves are more or less pubescent and sheaths glabrous. In no. 2541 (July 20, 1969) the lowest sheath is pubescent. The flange at leaf apex is in both collections about 1 mm long. The name B. pubescens is taken up with some misgivings, in conformity with Wagnon's treatment in RHODORA 52: 211-215 (1950), but I wonder if other specimens could be found in the Linnaean herbaria at Stockholm or Uppsala, (cf. Stearn's Species Plantarum Facsimile, pp. 104, 107, and 114 (1957), which might modify the indefiniteness of the Linnaean Herbarium in London. Comparative distinc- tions were not drawn as finely in 1753 as at the present, and as Stearn (p. 160) says, “The lens had not become the in- dispensible tool of Systematists in Linnaeus’ day and the 1970] Tilia Forest — Svenson 345 metric system of measurements had yet to be invented". It is interesting to note that in vol. 2. p. 72, sketches of Bromus mollis, B. lepidus, and B. squarrosus made by Lin- naeus are reproduced. These may provide insight on his ideas of the genus. Bromus purgans and B. ciliatus were actually described in Species Plantarum, and this procedure together with the new specific names, is probably associated with the relatively late acquisition of Kalm’s material. There has been a good deal of controversy about B. purgans and B. pubescens. Fernald (Man. p. 72) says the name “purgans” was given because Linnaeus erroneously identi- fied the plant with “Gramen bromoides catharticum” of Feuillée. Linnaeus did not do this; he merely cited the Peruvian Feuillée figure as questionable. We now come to the status of the Sandy Neck material. I could not find the collection cited as B. latiglumis (Rho- dora 49: 258 (1947), and Seymour in “Flora of New Eng- land" (1969) does not mention it. B. pubescens (B. pur- gans, Man. ed. 8) is known sparingly from the Boston area, but becomes common westward, and was originally de- scribed by Pursh from Nuttall's collection on the banks of the Missouri River, It was seen by Pursh growing 5 to 8 feet high in Lambert’s garden at Boyton, England. Asa Gray records an amusine visit to Lambert (Letters, vol. 1, p. 111 (1893), “the queerest old mortal I ever set eyes on". Lambert's herbarium was broken up and sold, with consequent disappearance of many of Pursh's plants. The switch of names by Wagnon is perhaps the best solution for stability, since P. latiglumis (Shear) Hitchcock appears to be untenable. It was a substitute for B. altissimus Pursh (1814), not Gilibert (1792). But McVaugh, Gentes Her- barum 8, fasc. 1 (1941) and others consider as invalid the names published in Gilibert's excessively rare “Excercitia Phytologia" (1792). The Sandy Neck material (cf. B. purgans, Rhodora 43: pl. 670 (1941) has only a slight flange at the sheath apex, as in fig. 8. Branching of the panicle and the leaves re- semble B. nottowayanus (fig. 1) from eastern Virginia, 346 Rhodora [Vol. 72 which along with B. latifolius is included by Radford, Ahles, and Bell under B. purgans in “Manual of the Vascu- lar Flora of the Carolinas", and the name B. purgans var. latiglumis Shear shows some question as to the diversity of the two plants under discussion. Is our Cape Cod plant possibly a northern extension of the southern coastal-plain Bromus nottowayanus? One interesting thing which Wag- non could not explain is the Linnaean use of the term “crispa” in reference to the panicle of B. purgans and B. ciliata. In Philosophia Botanica (1751) it refers only to leaf margins. In going through the grasses of Species Plantarum, I find only two other references: Poa alpina var. B and Poa bulbosa, and I suspect the term refers to large pulvini or similar structures. It came from Scheuch- zer, the great forerunner on Linnaeus in sedges and grasses. Bromus pubescens is cited by Seymour as “uncommon; dry open commonly rocky woods in basic soil", and in Massachusetts south to Norfolk County. It is one of the plants listed from Oak island, Revere, formerly the most interesting salt-marsh island of the Massachusetts coast (cf. the interesting account by W. P. Rich in RHODORA 4: 87-94 (1902). Probably nothing of botanical interest re- mains. Festuca obtusa Biehler. Known previously from Cape Cod only at Sandwich and Brewster and there sparingly. Carex seorsa Howe. Known previously from Monomoy (Weatherby in 1914), but it abounds in the maple swamp in the Lowell Reservation on Mashpee Pond, and Fogg cites it from Nonamesset (Elizabeth Islands. It is occasional in the Boston area south to Norton in Bristol County, and 1 have recently found it in Plymouth County at Plympton. Carex blanda Dewey. Occasional to frequent in the Bos- ton area, south to localities in Bristol County, and known from Nonamesset, At Osterville it was collected in ‘‘wet woods" by Fernald and Hunnewell in 1916. The plant is by no means smooth, as the name might imply, and the minutely serrate upper sheath apex is a good means of identification. 1970] Tilia Forest — Svenson 347 Carex intumescens Rudge. The perigynia are 14 X 6 mm; achenes average 5.0 X 3.1 mm and are not rounded at the apex. The plant is therefore typical southern C. intumes- cens. Cardamine parviflora L. var. arenicola (Britt.) O. E. Schulz. A small white-flowered cress, known previously from sandy woods in Bourne, Saconesset (W. Falmouth), and Provincetown, Aquilegia canadensis L. Fairly abundant and probably de- pendent on lime content of shells. It is known from scat- tered plants in Falmouth, Mashpee, and Osterville. Fogg in RHODORA 32: 176 (1930) cites it as one of the species on the morainal hills of the Upper Cape. It is not known from the Elizabeth Islands. “The superficial aspect of this part of the Cape is that generally associated with the Alleghe- nian flora with a slight tinge of the Canadian. This im- pression is borne out by a study of the plants which occur here, many of which are entirely lacking, or only locally known elsewhere on the Cape. There are well over 150 such plants." Ribes hirtellum Michx. var. calcicola Fernald. In RHO- DORA 72: (1970), I described the seeds. On Sandy Neck only a single flower was seen on this prickly little bush, associated here with Triosteum and Galium. It is obviously out of its environment, which is at the borders of brackish inlets, though recently I found it in a meadow in Sandwich with Habenaria psycodes. The Cape Cod plant has been somewhat questionably treated as true var. calcicola. The type was Ribes oxyacanthoides var. calcicola in RHODORA 7: 155 (1905), which came from arbor-vitae swamps at the mouth of the Bonaventure River in Quebec, and character- ized by leaves softly pubescent beneath. Inland it grows chiefly in wet calcareous soils, as noted by Wiegand and Eames in “Flora of the Cayuga Lake Basin" (1926), Mc- Vaugh in “Flora of Columbia County Area, New York" (1958), and Deam “Flora of Indiana" (1940). A detailed account was given by Fernald in “The Varieties of Ribes hirtellum”, RHODORA 13: 73-76 (1911). Perhaps the best 348 Rhodora [Vcl. 72 treatment is Alwin Bereer’s in N.Y. State Agric. Expt. Sta. (Geneva) Tech. Bull. no. 109 (1924) which says that in Grossularia hirtella the leaves of the short lateral shoots are more reniform or orbicular and with a subcordate base; the petals half as long as the sepals, and white, obovate, or with pink nervation; stamens about as long as the sepals. It has been the source of many cultivated strains of goose- berries. Gleason, IU. Fl. 2: 277 (1963) thinks the distinc- tion between R. hirtellus and R. oxyacanthoides scarcely warrants specific segregation, since intermediate forms with bracts both villous and glandular-ciliate exist. The Cape Cod speciniens show great variance with the generally accepted characteristics. The leaves are villous be- low, and vary from cuneate to cordate at the base. Flowers are 2 or 3 on a peduncle. Filaments are commonly 6 mm long and exceed the sepals in mature flowers by 1 to 1.5 mm, Sepals are oblong, green, sometimes purple-tinged, obtuse, commonly hirtellous, 3.5-4 X 2 mm. Petals are scarious white, 2.5 mm long, truncate erose at summit and narrowed to the base; styles usually about equalling the sepals, parted 1.5 mm from the base, and lanuginous for half the length. Glands are common on the apices of the branched petiolar trichomes, and sparingly on petioles, bud scales and bract margins. In addition to strong nodular spines, there are usually numerous reflexed prickles 2-7 mm long on the loose white bark of branches. Perhaps Fernald was closer to the mark when he originally described the plant as R. oxyacanthoides var. calcicola, The Cape Cod material is fairly homogeneous in the characteristics noted. The following collections are cited: among clumps of Carex prairea, at head of tide, Bumps River, Osterville (no, 2004, in flower May 27, 1967); dry bank just above tide level, Mashpee River, Mashpee (no. 2242, in flower May 10, 1969) ; swamps near old tide level, East Sandwich (no. 1988, in flower May 23, 1967). Rubus Enslenii Tratt. A trailing blackberry with solitary fruits (det. A. R. Hodgdon). He notes that “it is usually found in oak-hickory woods, and is a relatively southern 1970] Tilia Forest — Svenson 349 species of deciduous woodlands.” In their treatment of “‘Ru- bus in New England”, RHODORA 68: 491 (1966) Hodgdon and Steele cite it from Harwich. Rubus pensilvanicus Poir. (det. A. R. Hodgdon). A black- berry with tall erect canes “often looking like the common R. allegheniensis, but without the characteristic glands amongst the pubescence of the inflorescence”. It is cited from Sandwich and Falmouth. R. allegheniensis is cited by Hodgdon and Steele from Falmouth and Martha’s Vine- yard. Agrimonia gryposepala Wallr. Common in the Boston area, Known from Lakeville and Rochester in Plymouth County; and occasional in richer woodlands on Cape Cod in Pocasset, Falmouth, Centerville, and East Sandwich. Galium triflorum Michx. The leaves are six at a node, but stems vary from smooth to slightly scabrous. The plants are not sweet-scented. Known from Harwich, East Brewster, and Provincetown. Galium cireaezans Michx. var. hypomalicum Fernald. Common in northern New England, and extending south- ward into Bristol County. On Cape Cod it is known from West Barnstable and Provincetown; there are only a few plants at Sandy Neck. Triosteum aurantiacum Bicknell. The horse-gentian is occasional around Boston and extends south to Plymouth. On Cape Cod it is known from Bourne and Sandwich. Along the Sandwich Beach Road I find it growing with dog’s-tooth violet (Erythronium americanum), which is here in great abundance, and is the only locality known on Cape Cod. At one time there were scattered plants of Claytonia virginica, otherwise not known as native to eastern Massachusetts. Miss Eda M. Roos, of East Sand- wich, informs me that at the turn of the century botany classes from Sandwich High School used to go to Town Neck to see these plants. Mr. Weatherby went with her in 1932 to the locality; he collected Erythronium but they found no Claytonia. The next year Miss Roos sent me a single specimen, now in the Gray Herbarium. 350 Rhodora [Vol. 72 In the Tilia woods on Sandy Neck the Compositae are disappointing, only Aster divaricatus and Solidago caesia were noted, both of them widespread on Cape Cod. I shall give an extended account of the Amelanchier and Viburnum shortly. Since Sandy Neck is part of the Barnstable town lands, it is protected. The little saltmarsh islands along the Massachusetts coast should be searched for Tilia and asso- ciated plants, and be preserved before they, too, suffer the fate of Oak Island. Some specimens have been placed in the Cape Cod Museum of Natural History and the New England Botanical Club. OSTERVILLE, MASS, 02655 After writing this report, I talked with County Commissioner Heyworth Backus. He vaguely remembered a family living at the site, but said we should go to see Martha Dickie, of West Barnstable, who knew all about Sandy Neck. She has a good collection of arrow- heads, pestles, and axes from Sandy Neck, and mentioned the skeleton of an Indian chief, found buried there. Also, long after the Indians were gone a socially unambitious family took up quarters. They had a dull-witted daughter. She had a suitor who was not so bright. She said she did not have money for a marriage licence. “That’s all right", said the suitor, “I’ll write up a licence, and we'll take it to the county clerk, and he'll sign it, and it won't cost anything". So he wrote up the licence on a sheet of paper. It was not a success. Both were committed to an institution for the feeble-minded. It is unlikely that this family or the Indians would have planted linden trees in a remote corner of the swamp, and we have already seen that native linden trees are scattered along the Massachusetts coast. Since I wrote this footnote I have seen Spach's specimens of Tilia neglecta at Paris, and Bromus purgans in the Linnaean Herbarium in London. They will be commented upon later. DISTRIBUTIONAL HISTORY OF LYCOPUS EUROPAEUS (EUROPEAN WATER-HOREHOUND) IN NORTH AMERICA? RONALD L. STUCKEY AND W. LOUIS PHILLIPS? Within the past one hundred years, Lycopus europaeus has become naturalized in northeastern North America from Europe. Early literature and herbarium records dat- ing as far back as about 1860 indicate that the species was first found on ballast or in waste places in the eastern sea- ports of Norfolk, Camden, Philadelphia, Wilmington, New York City, and Boston, Since then the species has spread via waterways along the coast and inland where it now occurs along ditches, rivers, canals, lakes, and in marshes and waste places. Small (1933), Fernald (1950), Gleason (1952), and Gleason and Cronquist (1963) report L. euro- paeus as established in waste places and along roadsides on the east coast of the United States from Massachusetts to Virginia, Alabama, Mississippi, and Louisiana, but rarely inland. Henderson (1962) gives its distribution as only from Massachusetts to Virginia. Recent reports of addi- tional locations are North Carolina (Radford, Ahles, and Bell, 1968), Nova Scotia (Erskine, 1951), the St. Lawrence River valley in Quebec (Rousseau, 1968), Ontario (Mont- gomery, 1957), and the western basin of Lake Erie (Stuckey, 1968). The present known distribution in north- eastern North America is given in figure 1. Lycopus europaeus L. a member of the mint family (Labiatae), occurs outside of North America over most of Europe and into western Asia. Several of its common names are European water-horehound, marsh-horehound, green-archangel, bitter bugle-weed, gipsy-herb, and gipsy- wort. The plants are stout, with stems ranging 0.4-1 meter ‘Contribution from the Botany Program (Paper No. 769), the Herbarium, and the Franz Theodore Stone Laboratory, The Ohio State University, Columbus 43210. "Presently in the United States Military Service. 351 352 Rhodora [Vol. 72 ; ` @ 1860 Fig. 1. Known distribution of Lycopus ewropaeus (European water- horehound) in eastern North America based on herbarium specimens seen and other records cited in this paper. The position of each square or circle represents a locality where a plant has been collected. Num- bers in the squares are the last two digits for the year in which the plant was collected if before 1900, and the numbers in circles are the last two digits for the year of collection if since 1900. The oldest known collection is mapped for each locality. The four major areas of apparent entry and establishment for the species in North America are marked by the large black arrows with the date of the earliest known collection for that area. The smaller black arrows indicate the possible migration pattern that the species has taken in the Lake Ontario, St. Lawrence River, and Lake Erie region. 1970] Lycopus europaeus — Stuckey and Phillips 353 in height, freely branched, and usually pubescent. The leaves are 4-12 centimeters long and 1.5-5 centimeters wide, ovate, ovate-lanceolate, to narrowly lanceolate in general outline with blades sinuately lobed, but not pinnatifid. The white-petaled flowers are somewhat inconspicuous in the axils of the leaves, where later in the year four nutlets are formed each with a heavy corky crest (Henderson, 1962). The species is morphologically very similar to the wide- spread native American species, L. americanus, with which it has often been confused. Characteristics separating the two species are not always well differentiated, and the pos- sibility of hybridization should be investigated. We have neither delimited the character distinctions and possible intermediates between these two species, nor have we at- tempted to distinguish between typical L. europaeus var. europaeus and var. mollis (Kern.) Briq. Early Literature Records One of the difficulties in working out the North Amer- ican distributional history of L. europaeus is that some of the early reports in the literature are considered unreliable because the taxonomy of the genus was not known or un- derstood. For example, the earliest known North American report is by Walter (1788) in his Flora Caroliniana. This record has since been referred to L. americanus (Hender- son, 1962). Bigelow (1824) noted L. europaeus for Boston and vicinity. His description, "Lower leaves cut, upper leaves lanceoiate, serrate,” portrays L. a mericanus, a spe- cies which he did not report. Riddell (1834) in his Synopsis of the Flora of the Western States listed only L. ewropacus and L. virginicus of which the former is probably the com- mon and widespread L. americanus. Gray (1858) reported L. europaeus and two varieties, var. sinuatus and var. in- tegrifolius. These two varieties are currently recognized as L. americanus and L. rubellus, respectively. Gray's de- scription for L. europaeus could be construed to apply to either L. americanus or L. europaeus: “Stem sharply 4- angled... leaves ovate-oblong or oblong-lanceolate, sinuate- 354 Rhodora [Vol. 72 toothed or pinnatifid, more or less petioled . . , nutlets . . . equalling or exceeding the calyx tube." Gray considered typical L. europaeus to occur only in Europe, Later, Gray (1867) added a third variety, var. sessilifolius, which is currently recognized as L. sessilifolius. Gray (1870) re- vised the taxonomy of Lycopus in N orth America and for the first time recognized the presence of the European spe- cies, L. europaeus, as an adventive in North America. The reports by Walter (1788), Bigelow (1824), Riddell (1834), and Gray (1858, 1867) have created confusion for later authors. We have discarded these and other erroneous rec- ords of L. europaeus, all of which are summarized (table 1) along with the probable corrected name for each report and the author(s) who made the correction. Early Herbarium Records from Virginia, Pennsylvania, and New Jersey The time and place of introduction of L. europaeus in North America is obseure, The earliest known collection (GH, PH) was made by Elias Durand at Norfolk, Virginia, but is without a date. Durand was most active as a plant collector from about 1859 to 1868 (Pennell, 1936). This collection ig mapped with the chosen date 1860 (fig. 1). The first known Specimen with more adequate data was collected from ballast on Petty's Island in the Delaware River, New J ersey, 1 October 1867 (Parker s.n., GH). These two collections were cited by Gray ( 1870) as supposedly the first authentic records of L. ewropaeus for North Amer- ica. Later records from Virginia come from Norfolk County between Princess Anne and Berkeley, 13 July 1893 (Heller 1072, F, GH, Mo, NY, PH, US) and “from about wharf” near Suffolk, Nansemond County, 19 July 1898 (Kearney 1740, OS, US). By the late 1930’s the species apparently had be- come established and had spread locally as evidenced by collections of M. L. Fernald and Bayard Long from marshes, swales, and roadsides in Nansemond, Isle of Wight, and Surry counties of southeastern Virginia. Their records, 1970] Lycopus europaeus — Stuckey and Phillips 355 obtained from 1936 to 1941, are summarized in Fernald (1947). The early records of L. europaeus for the Philadelphia area come from ballast as indicated on the labels of plants collected at Camden, New Jersey by C. F. Parker (Petty's Island, 1 October 1867, GH ; Kaighn's Point, year 1877, GH; Camden, August 1874, MO, NY, 6 September 1879, PH) and by I. C. Martindale (Camden, August 1876, MO, September 1876, us, July 1877, P). Petty’s Island and Kaighn’s Point along the Delaware River at Camden and The Ballast Ground at the Navy Yard in Philadelphia were the most prominent locations where ballast of ships was dumped. In papers discussing the occurrence of foreign plants on ballast, Smith (1867), Burk (1877), and Martindale (1877), however, do not list L. europaeus. From ballast ground locations, L. europaeus apparently spread along the Delaware River where it was later found on the banks and in the marshes from New Castle to Port Penn, Delaware, 8 September 1869 (Commons sN., Ny). Commons also collected it three miles below New Castle (20 August 1874, PH), at Delaware City (5 September 1879, NY), in marshes and along a canal at Delaware City (22 September 1896, PH), along the river shore above Christiana river lighthouse at Wilmington (28 September 1898, PH), and along the river shore at Cherry Island marsh, Wilmington (16 Octo- ber 1900, PH). A herbarium sheet label by Commons (4 July 1901, PH) bears the note, “now well established along the river shore from Philadelphia to Port Penn and is also common around Salem, New Jersey." The species has con- tinued to spread and become a part of the flora. Forty-five years later, Tatnall (1946) wrote that L. europaeus was *common along streams, and on borders of fresh or brack- ish marshes, Coastal Plain of Cecil [Maryland] and New Castle [Delaware] counties; rare farther south : n. of Ocean City .. . naturalized from Europe." Until 1949, many col- lections from the eastern Pennsylvania-New Jersey area were deposited in the herbarium of The Academy of Nat- ural Sciences of Philadelphia. 356 Rhodora [Vol. 72 New York and Massachusetts Most of the early reports for the New York City area list L. europaeus as coming from ballast. The earliest known record is 12 July 1877 (Brown s.n., NY). It was later found at 8th Avenue and Harlem River on 4 October 1879 (Leg- gett s.n., MICH). Brown (1879) listed L. europaeus among ballast plants that he found at 8th Avenue and Gowanus in New York City, He also collected a specimen from ballast near Communipaw Ferry [now part of J ersey City], New Jersey in July 1879 (Brown s.n., US), N. L. Britton made à collection from ballast ground at 8th Avenue and 130 Street (September 1879, NY). Plants have also been ob- tained from ballast at Brooklyn in 1880 (Sehrenk s.n., NY), at Yonker's Wool Mill in 1898 (Bicknell s.n., NY), and from Cypress Hills, Long Island, in 1905 (Bicknell s.n., PH). Taylor (1915) reported the European water-horehound as occurring locally rare as a weed in waste places from New York to Virginia and considered it to be naturalized from Europe. House (1924) pointed out that the species was “locally rare as a weed in southeastern New York: natu- ralized from Europe.” In Massachusetts, John Robinson (1880) mentioned L. europaeus as "an adventitious plant and probably does not grow [ie., is not naturalized] here [Essex County]." Eu- ropean water-horehound was collected on waste ground near Parker Street, Cambridge on 6 October 1894 (B. L. Rob- inson s.n., GH). Fernald (1910) commenting upon this col- lection and locality wrote that L. europaeus ". .. has grown in waste land and neglected yards about the ‘Tin Cafion’ in Cambridge for at least sixteen years.” The plant was again obtained on Parker Street in a dump on 7 November 1903 (Moore 1537a, GH). Fernald (1947) mentioned L. euro- paeus as occasional in waste near Boston. F rom the appar- ent lack of records, it seems that L. europaeus has Spread very little in the Boston area. The most recent known col- lection for Massachusetts is cited by Seymour (1969). The Specimen comes from Provincetown, 17 September 1964 (Hinds s.n., NERC). 9 1970] Lycopus europaeus — Stuckey and Phillips 357 Inland Distribution about Lake Ontario, the St. Lawrence River, and Western Lake Erie The distribution of L. europaeus along Lake Ontario, the St. Lawrence River, and Lake Erie has come about within the past 60-70 years. The first known collection is from Lake Ontario on Toronto Island, York County, 24 August 1903 (Scott s.n., TRT). Succeeding collections have come from Toronto Island, as well as from locations at the east end of the lake by 1933 in the vicinity of Kingston and at the west end of the lake at Hamilton as early as 1957. Mont- gomery (1957) summarized the distribution of the Euro- pean water-horehound for Ontario as “Infrequently found around marshes.” The following collections are known from (or near) the shore of Lake Ontario. NEW YORK: MONROE CO.: Marsh, Irondequoit Bay, 8 August 1932 (House 19639, GH); marshy border of bay, Manitou Beach, 15 August 1933 (Howse 20772, GH, NY). ONTARIO: FRONTENAC CO.: Grass Creek Island, 7 mi below Kingston, 18-27 August 1933 (Pennell 16226, Ny, PH); Eagle Lake, 23 July 1939 (Krotokov SN., TRT); opposite grain elevator, Lake Ontario Park, 23 September 1960 (Beschel, Dore, & Hainault 11716, DAO). HASTINGS CO.: Edge of water, Belleville, 20 July 1944 (Groh 2187, DAO). LEEDS CO.: Plants numerous, swampy spot at mouth of spring among drift-debris, shore of St. Lawrence, 1000-Islands Bridge, 1 August 1956 (Dore, Soper, & Cody 16515, DAO); local along shore of St. Lawrence at base of granitic talus, 1000-Islands International Bridge near Ivy Lea, 24 September 1960 (Dore 18724, DAO). LINCOLN C0.: Edge of water, Jordan 1 N, 14 September 1966 (Davidson 364, DAO). NORTHUMBER- LAND CO.: Wooded pasture, Seymour Township, n side of the Trent River near the village of Trent River, 15 August 1952 (Gillett 6930, DAO). PRINCE EDWARD CO.: Edge of marsh, ca. 1 mi n of Huff Island, Loc 367, 4 August 1951 (Soper & Heimburger 5402, TRT). WENT- WORTH CO.: [Between the water and the railway at the extreme west end of Burlington Bay, Hamilton], 5 September 1957, 4983, 4984, 22 July 1958, 4982, 27 August 1958, 4985 (all A. Tamsalu, HAM). YORK co.: Toronto Island, September 1903 (Wilkes s.n., TRT), 18 August 1911 (White 1, GH), 20 September 1935 (Macklin s.n., HAM); Harlan's Island, Toronto, 22 August 1945 (Wage s.n., DAO). The occurrence of L. europaeus in the St. Lawrence River valley in the Province of Quebec is very recent. Rousseau 358 Rhodora [Vol. 72 (1968) mapped its distribution based on collections from Valleyfield (1964, Cinq-Mars s.n., QFA), Iles-de-la-Paix, Cté de Beauharnois (1965, Morency), Ile St. Ignace, Cté de Berthier (1965, Rouleau), and Repentigny (1966, Rouleau). These locations and recent dates strongly suggest that the plants are migrating down the St. Lawrence river. Ten years earlier Dore and Gillett (1955) did not find European water-horehound during their botanical survey of the St. Lawrence seaway area upstream in Ontario. The farthest westward inland penetration for L. euro- paeus appears to be on the islands and about the south- western shore of the western basin of Lake Erie. Here it was first collected near the camp grounds at Bay Point Peninsula, Ottawa County, on 24 July 1966 (Easterly s.n., BGSU) followed by several collections obtained in 1967 (Stuckey, 1968). The plants occur about the edges of those ponds on the Erie Islands which have connections with the water of the lake, either by a channel or by flood- ing during violent storms. Elsewhere, its occurrence is along the sandy portions of the shore of the mainland and around tne edges of ponds or in marshland directly behind these sendy beaches. The Species has not yet penetrated the extensive marshes and mud flats that are common farther inland along the Ohio and Michigan shore of Lake Erie. It is not known whether the Species has migrated into Lake Erie from older stations in Lake Ontario or whether it was separately introduced and then began to spread. The local distribution pattern strongly supports the idea that the species (1) has recently invaded western Lake Erie via a water route, rather than coming overland, and (2) is spreading in western Lake Erie by propagules being transported by water. According to Ridley (1930, p. 222), the “plant is specially adapted for dispersal by river, and is very common on English river banks, where it is rapidly dispersed. The nutlets . . . float for from 12 to 15 months (according to Praeger and Guppy)." L. europaeus is also becoming established in western Lake Erie where it is now known from 11 stations. At several 1970] Lycopus europaeus — Stuckey and Phillips 359 of these places many young plants have been seen with larger, older individuals of the same species. Many of these young plants are attached to older plants via runners and slender rhizomes. At Terwilliger’s Pond on South Bass Island some plants had 5-10 runners ranging up to two feet in length which touched the ground at various places, formed roots, and had initiated new plants. The species is spreading vegetatively at individual sites. At places where disturbance by wave action is considerable, such as at Airport Point and Peach Point on South Bass Island or at Lakeview Park in Port Clinton, L. europaeus was not found in 1969, although it was present at these sites in 1967 or 1968. The floras at these sites were drastically altered in 1969 because the unusually higher water level, and wave action had destroyed many of the species along the low rocky and sandy beaches. The following collections of L. ewropaeus from the west- ern basin of Lake Erie were taken by R. L. Stuckey, unless otherwise noted, and are in The Ohio State University Her- barium (0S) with duplicates elsewhere as stated. OHIO: ERIE co.: A few plants on wet sand and gravel dumped in the quarry from the State Park Beach, in the bottom of the north- west limestone quarry below the glacial grooves, Kelley’s Island, 14 August 1969, 8238 (CAN, GH, MICH). OTTAWA CO.: Occasional in shallow water at Smith’s Pond, s e corner of North Bass Island, 1 August 1967, 4741; Honey Point and Smith’s Pond, North Bass Island, 22 August 1969 (Phillips 181, os); South Bass Island — oc- casional at water's edge along muddy w bank of Terwilliger's Pond, 18 August 1967, 5184 (SMU), 29 August 1967, 5431, Terwilliger's Pond, 20 August 1969, (Phillips 180, os), occasional on thin, wet soil over dolomite rock at Airport Point (opposite Starve Island) on the s e shore, 27 August 1967, 5422; a few plants about boat dock on south shore of Peach Point, Fishery Bay, 24 August 1968, 7499 (DAO); occasional in wet sand behind dune at Sand Point [Bay Point], ca. 2.5 mi s of the town of Marblehead, 8 September 1967, 5716, 12 September 1968, 7685 (CAN, GH, PH); a few plants in ditch along e edge of causeway, East Harbor, East Harbor State Park, 14 August 1968, 7562; occasional on mud flat at e edge of road behind beach at East Harbor, East Harbor State Park, 23 August 1968, 7471, (DAO); East Harbor State Park, 31 July 1969, (Phillips 78, oS); a few plants in wet sand at edge of wooded area behind 360 Rhodora [Vol. 72 beach between mud flat formed in previous year, East Harbor, East Harbor State Park, 25 August 1969, 8279; portion from one of four , plants seen on wet, sandy beach (not seen here in 1967), NE 1/4 Sec. 5, T6N, R14E, Portage Twp., Lakeview Park, e edge of Port Clinton, 19 August 1968, 7451. LUCAS Co.: Occasional on mud flat along the main driveway at the north edge of Magee Marsh, Crane Creek State Park, Sec. 12, s e corner of Jerusalem Twp., ca. 10 mi e of the town of Oregon, 21 August 1967, 5307; occasional along edge of marsh behind barrier beach, west end of Crane Creek State Park, Magee Marsh, . . . , 27 August 1968, 7541. MICHIGAN: MONROE CO.: Occasional in drying cat-tail marsh near bay of Lake Erie, Sec. 33, Erie Twp., ca. 3.5 mi s e of the town of Erie, 14 September 1968, 7755; occasional in wet sand on beach along shore of Lake Erie, s w corner of Sec. 35, Frenchtown Twp., n edge of Sterling Monroe State Park, e edge of the city of Monroe, 14 Septem- ber 1968, 7760 (GH). Unlike the distributional patterns of the non-indigenous Rorippa sylvestris (Stuckey, 1966) and Epilobium hirsutum (Stuckey, 1970), in which these species apparently mi- grated inland across central New York State, L. europaeus has not been found in central New York State. It appears, therefore, that L. europaeus became separately introduced and spread in the Lake Ontario, St. Lawrence River, and Lake Erie waterway independent of its introduction and spread along the east coast of the United States. All of the known localities for the European water-horehound are along the shores of the St. Lawrence river and Lakes On- tario and Erie, except the one from the village of Trent River in Northumberland County, Ontario. Isolated Occurrences Several reliable records from isolated locations away from the areas where L. europaeus has become established ere known. Erskine (1951) reported L. europaeus as an adventive species at the edge of a ballast dump, Steele’s Pond, Point Pleasant, Halifax County, Nova Scotia, 24 August 1949 (Erskine & Erskine s.n., ACAD), Burk (1961) mentioned the species as occurring at the edge of a fresh- water pool, about one half mile from Hatteras lighthouse, Buxton Woods, Hatteras Island, Dare County, North Caro- 1970] Lycopus europaeus — Stuckey and Phillips 361 lina, 2 September 1960 (Burk b55-8, NCU). This is appar- ently the first record of the species for North Carolina. Radford, Ahles, and Bell (1968) reported it as rare in marshes which are frequently brackish. Along the Gulf of Mexico, L. europaeus is known from Cat Island, Harrison County, Mississippi, 11 August 1900 (Lloyd & Tracy s.n., NY). This record was cited by Lloyd and Tracy (1901) and Lowe (1921). Present Status In eastern North America L. europaeus has become estab- lished at several seaport cities and from there has spread locally. The species appears now to be established in six rather well-defined geographical areas. In figure 2, these geographical areas are listed on the vertical axis and time is represented on the horizontal axis, All specimens ex- amined from these areas are each marked with a dot along the horizontal axis at the corresponding five-year interval of collection. This figure shows (1) the earliest specimen record known for each geographical area, and (2) that the species has become established in each geographical area, because specimens have been found and obtained, in the years following the original collection and even up to the present time in certain areas. It can be gleaned from the information in figure 2 that no recent collections are known from the Virginia, Philadelphia, or New York City areas. It seems doubtful that L. europaeus has disappeared from these areas where the species has long been established. Rather, this gap in information suggests that collections of plants are not being made in areas that have been rather well-studied in the past. Any flora is continually changing and botanists should continue to keep records and specimens of the flora through time, Summary The probable localities and time of introduction and spread of L. europaeus in North America have been worked out from the literature and herbarium records. "These 362 Rhodora [Vol. 72 sources reveal that many of the early collections came from ballast and waste ground. Its introduction probably was from ships’ ballast at several east coast seaports, such as Norfolk, Camden, Philadelphia, Wilmington, New York City, and Boston. From these locations the species appears to have spread locally and became established. Inland, L. europaeus is not known from collections before 1900. Since then it has spread and is becoming naturalized as part of the shore flora of the upper portion of the St. Lawrence River, along Lake Ontario, and in southwestern Lake Erie. Acknowledgments Our thanks are extended to those herbaria (DAO, GH, HAM, MICH, MO, NEBC, NY, OS, PH, US) whose specimens were examined, Records of herbarium specimens have been provided by Dr. James H. Soper (CAN) and Dr. James Cruise (TRT). The field study in the western Lake Erie area was conducted as part of the teaching and research program at The Franz Theodore Stone Laboratory, Put- in-Bay, Ohio, during the summers of 1967, 1968, and 1969. Additional support for field work has been provided by The Ohio Biological Survey. We are grateful to Mr. Ronnie Johnson, Mr. W. Alan Wentz, and Mr. Thomas Duncan, who served as field assistants. COLLEGE OF BIOLOGICAL SCIENCES THE OHIO STATE UNIVERSITY COLUMBUS, OHIO 43210 REFERENCES BABCOCK, H. H. 1872. The flora of Chicago and vicinity. The Lens 1: 20-26, 65-71, 144-150, 169, 218-222. BEARDSLEE, HENRY C. 1878. Catalogue of the plants of Ohio, in- cluding flowering plants, ferns, mosses, and liverworts. 32nd. Ann. Rep. Ohio State Board Agr. 1877: 335-363. BIGELOW, JACOB. 1824. Florula Bostoniensis. A Collection of Plants of Boston and its Vicinity, . . . Cummings, Hilliard, & Co., Boston. 422 p. BILuinGs, B., JR. 1862. List of plants observed growing principally within four miles of Prescott, C. W., and for the most part in 1860. Ann. Bot. Soc. Can. 1: 114-140. 1970] Lycopus europaeus — Stuckey and Phillips 363 FLORISTIC AREA P> m; O Om l T | T j I 60-65] o e ALI) AYA MIN 2 SL13SnH2VvSSVI|AJ NHu31SV q OIlHVLNO 33v7] 3 3lu 33V] NHU31S3AA E VINIOUIA NH31SV3H1DnOG V © © e JUVMV13Q A3Su3p MaN viHa130vilHq g wo — wo en O£6l = © e e o 61-65 e 0Z6l $ 3 e l 1 1 1 1 l Fig. 2. Dates of herbarium specimens verifying the time of spread and establishment of Lycopus europaeus in six floristic areas of eastern North America. Each dot represents one or more herbarium specimens collected within a five-year period for each floristic area. 364 Rhodora [Vol. 72 BROWN, ADDISON. 1879. Ballast plants in New York City and its vicinity. Bull. Torrey Bot. Club 6: 353-360. Burk, C. JoHN. 1961. Distribution records and range extensions from the North Carolina outer banks. Castanea 26: 138-139. BURK, Isaac. 1877. List of plants recently collected on ships’ bal- last in the neighborhood of Philadelphia. Proc, Acad. Nat. Sci. Phila. 29: 105-109. COULTER, STANLEY. 1900. A catalogue of the flowering plants and the ferns and their allies indigenous to Indiana. Ann. Rep. Indiana Geol. Surv. 1899: 553-1074. DEAM, CHARLES C. 1940. Flora of Indiana. Dep. Conservation, Div. Forestry, Indianapolis. 1236 p. Dork, WILLIAM G. 1962. A centennial floristic census of Prescott, Ontario. Trans. Royal Can. Inst. (1961) 33: 49-115. . and J. M. GILLETT. 1955. Botanical Survey of the St. Lawrence Seaway Area in Ontario. 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S. 1874. The flora of Penikese Island. Am. Nat. 8: 193-197. KELLERMAN, W. A. 1899. The fourth state catalogue of Ohio plants, consisting of a serially numbered systematic check list of the Pteridophytes and Spermatcphytes. Bull. Ohio State Univ. Ser. 4, 10 (Bot. Ser. 1): 1-65. - and W. C. WERNER. “1893” [1894]. Catalogue of Ohio plants. Rep. Geol. Surv. Ohio, 7: 56-406. —, and Mns. KELLERMAN. 1900. "The non-indigenous flora of Ohio. Bull. Ohio State Univ. Ser. 4, 27 (Bot. Ser. 4): 1-28. LEWIS, I. F. 1924. The flora of Penikese, fifty years after. Rhodora 26: 181-195, 211-219, 222-229. LLovp, FRANCIS E., and S. M. Tracy. 1901. The insular flora of Mississippi and Louisiana, Bull. Torrey Bot. Club 28: 61-101 + pl. 8-11. (Reprinted as Contr. Dep. Bot. Columbia Univ. No. 174). Lowr, E. N. 1921. Plants of Mississippi. Mississippi State Geol. Surv. Bull. No. 17. 292 p. MARTINDALE, Isaac C. 1877. More about ballast plants. Bot. Gaz. 2: 127-128. MCVAUGH, ROGERS. 1935. 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Manual of the Vascular Flora of the Carolinas. Univ. North Carolina Press, Chapel Hill. 1183 p. RIDDELL, JOHN L. 1834-1835. Western Flora. Synopsis of the flora of the Western States. West. J. Med. & Phys. Sci. 8: 829-374, 489-556. Reprinted E. Deming, Cincinnati. 1835. 116 p. RIDLEY, HENRY N. 1930. The Dispersal of Plants throughout the World. L. Reeve & Co., Asford, England. 744 p. RoBINSON, JOHN. 1880. The Flora of Essex County, Massachusetts. Essex Institute, Salem. 200 p. Rousseau, CAMILLE. 1968. Historie, habitat et distribution de 220 plantes introduites au Quebec. Nat. Canad, 95: 49-169. RorHROCK, J. T. 1874. Preliminary report on the botany of central Colorado, p. 21-62. In George M. Wheeler. Catalogue of plants collected in the years 1871, 1872, and 1873, with descriptions of new species, U. S. Geogr. & Geol. Exp. & Surv. west of the one hundreth meridian. U. S. Gov. Printing Office, Washington, D.C. 62 p. SCHNECK, J. 1876. 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[Vol. 72 Rhodora 368 €coT mA P SOF cegI ma P souof Geet mA P souor ce6p “mag P seuof ZPGT ‘EQTYPPIM sdy[tyd 3 Aeyonys sdy[iud 3 Aexonys Z961 fuosiopuoH sdi[tud. 2» Aoxonjis OLST ‘Avan OL8T ‘SLID OLST ‘AVID | SNUDNLIWD “T Snyəqna UI SNUDILAWD U] SNUDNLIWD I SNUDILLIWD “I snuvawaup “I Ayqeqoard SNUDILLIWD “T “TUAW snua "[ A|qeqoid Avery snyofyissas UT yous0yy Sməqna "T "Ig Snyonus "T SNUDNLIWUD “J ZL8T '34oo»qeg LLST ‘9L8T *»euu»gs 94,8T 'uosioj1eq QL8T “uos4911Fd Te8r ^no 0981 “Ieuju PLSI "ioo 88LI 493A PEST TPPU LOST ‘Lay LOST '8egr “uD LOST ‘gegT ‘Avan snapdoina “I snopdo4no “I snuofialioqui "eA snopdo4na "T snjpnuis "IVA Snəndosnə “I snopndoa4no “I snjpnuis "IVA snapdowna `] snopndoano “I snondoamno “I snopndo4no “I Aw) smiojuissos "IBA snopndo.4mno U] Avan snyofiaboqu siga snopdo4ano "T Kua (CHA) $njonuis “IVA UT snondo4no "T AYUDA 2» OSV ‘stoulllT AoE A YSEA ‘stoui SIOUT[IT Srout[TT eulporeg unos Ņ ULOIOOr) Ayunoyg 9[189]) AON ‘QIVMV pA KeA SINJ ues *opeAo[07) SVUT[OLED s9j€]g u4əlsəAN sojejg popup, UYON saqe1g pəyYiuq u4doul(o N sojejg pəliuQq UIYHMON aouala yay | IUWEN postaey əouə1əJ9% aweN pəusrqn4q ARO] SANVN CALOAUNOO HLIM SAQMVdO/1H "I AO SLUOdAY GAHSITANd DNIGVAISIN I IS VL 369 Stuckey and Phillips Lycopus europaeus 1970] Z96T “0q sdiud 7 AexnyS sdiud 7 Aexen1s sdi[rud 7 Aexpnis cget “Ysne Ao PZT ‘SOT sdiud 7? 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CALIFORNICA AND C. MARITIMA: MICHAEL G. BARBOUR AND JAMES E. RODMAN The succulent, annual Strand plant, Cakile maritima Scop.’ is now common all along the California coast but was first seen there only 35 years ago (Rose, 1936). Prior to that, a presumed native species, C. edentula (Bigel.) Hook. Ssp. californica (Heller) Hult., was the only west coast Sea-rocket. C. maritima is a Mediterranean plant which has been introduced around the world; C. edentula is native to the Atlantic coast of North America (Pobedimova, 1963). Both species have been reported growing intermixed on the west coast strand. The subspecific rank for the presumed native taxon is not universally accepted. Pobedimova (1963) preferred Species rank (C. californica), Fernald (1922) and Schulz (1936) preferred varietal rank (C. edentula var. califor- nica), and Hitchcock et al. (1964) preferred to make no distinction at al! ("Our material is in no obvious manner different from the plant of the Atlantic coast," ) The senior author has recently investigated the autecol- ogy of C. maritima (Barbour, 1970a and 1970b) and de- sired to conduct a comparative study of C. edentula ssp. californica, The junior author is currently conducting a taxonomic revision of the genus Cakile. Both of us desired seed material of C. edentula ssp. californica, but field trips in 1969 which ranged from Ensenada, Baja California to Birch Bay, Washington convinced us that C. edentula ssp. californica has become nearly extinct in California while C. maritima has enormously increased its range and abun- dance since its introduction. Although this apparent, rapid near-extinction seems re- markable, an indirect explanation for it might be that C. "This research was supported in part by National Science Founda- tion Grant BO-14381 for the senior author; the junior author is an NSF Pre-doctoral Fellow. *Nomenclature follows Munz and Keck (1959). 370 1970] Cakile edentula — Barbour and Rodman 371 edentula ssp. californica had never been very common, thus what seems to be a real decline in its abundance has really been of minor proportions. To determine past abundance, we undertook a literature and herbarium survey. Her- baria consulted were: Australia: AD, BRI, CANB, NSW, PERTH; Britain: BM, K; Canada: DAO, UBC; Jamaica: IJ; United States: A, CAS, CU, DAV, DS, DUKE, F, FLAS, FSU, GA, GH, HSC, LA, LAF, LL, MICH, MO, MSC, NCU, NY, ORE, OSC, PH, POM, RSA, SD, SMU, TEX, UC, UCLA, Us, USF, WIS. These ab- breviations are those of Lanjouw and Stafleu (1964). The remainder of this paper documents a short but un- usual history of successive migrations: the introduction, rise, and fall of C. edentula ssp. californica ( californica) and the introduction and ascendancy of C. maritima (mari- tima). The introduction and rise of C. edentula ssp. californica At the time maritima was first seen on the west coast, some 35 years ago, californica was widely distributed from San Diego to Vancouver Island and behaved “. . . like a native...” in the strand community (Cooper, 1936). Eyer- dam (1938) and Hultén (1968) extended its range even further north to Kodiak Island off Alaska. Because of this wide distribution and supposed taxonomic differences from east coast C. cdentula, many floras have quietly contended that californica is native to the west (eg., Abrams, 1944, Calder and Taylor, 1968, Hultén, 1968, Munz and Keck, 1959, and Peck, 1961). However, botanical explorers of the early 19th century failed to report any Cakile on the west coast. The early North American floras of Hooker (1829-40) and Torrey and Gray (1838-40) record the range of the only Cakile listed (= C. edentula of today) as being the Atlantic coast and the shores of the Great Lakes, the west coast not being included. It is clear, though, that west coast habitats which support Cakile today had been examined, for modern associates of Cakile were listed: eg., Abronia arenaria Menz. (— A. latifolia Eschs.) was said by Hooker 372 Rhodora [Vol. 72 to occur on the “. . . sandy sea-shore, North West Amer- ica." Hooker and Arnott (1830-41) reported on the voyage of Captain Beechey, which stopped at Monterey and San Fran- cisco in 1827. Cakile was not listed in their flora, but mod- ern associates Franseria chamissonis and Abronia umbel- lata were, Torrey’s ( 1875) list of species collected on the voyage of Captain Wilkes in 1838-42 to Washington, Ore- gon, and northern California does not include Cakile but does include modern associates Abronia arenaria (= A. latifolia), Franseria bipinnatifida, F. chamissonis, Lathyrus maritimus, and Plantago maritima. A relatively detailed collection was made by the Russian Voznesenski along the coast between Bodega Head and Fort Ross, California in 1841 (Howell, 1937). In a per- sonal communication, Howell wrote that Cakile was not among the 214 species and varieties which comprised the collection, As late as 1882, Cakile was not on the west coast of Canada, for Macoun's Catalogue of Canadian Plants (1883- 1902) did not list it there even though he took care to note that west coast citations were based on his own “ .. very extensive collections . . ." on Vancouver Island and British Columbia in 1875, and on coast trips by Dawson in 1875-9 and Hill in 1880-2. There are no 19th century collections of Cakile from the west coast at either Kew or the British Museum of Natural History at which are deposited the collections of such important Pacific coast explorers as Menzies, Scouler, Douglas, Geyer, Burke, Bridges, and Lyall (Thomas, 1969). In southern California and Baja California, Bentham did not record Cakile during his voyage on the H.M.S, Sul- phur in 1836-42, but did report associates such as Abronia umbellata and Oenothera cheiranthifolia (Hinds, 1844). Parry’s collections made near San Diego during the bound- ary survey of 1855-6 (Emory, 1859) did not include Cakile; neither did the list compiled by Orcutt (1855) for San Diego County and adjacent parts of Baja California. 1970] Cakile edentula — Barbour and Rodman 373 Finally, as cited by Jepson (1936), Bolander did not in- clude Cakile in his 1870 Catalog of the plants in the San Francisco Bay region, nor did the California Geological Survey of 1876-80 include it. Jepson believed the first re- port of Cakile in California to be by Behr in 1888, of cali- fornica for a single locality, *Berkeley salt marshes." The earliest west coast collection of californica we have seen is dated July, 1882, from the“... beach at west-Berkeley . . ." collected by E. L. Greene and deposited at the Gray Her- barium. If the collections of Bolander in 1870 and the California Geological Survey of 187 6-80 are to be accepted as complete, then californica was introduced to the west coast in the San Francisco Bay area between 1880 and 1882. And if the west coast taxon is not morphologically distinct from the eastern C. edentula — as Hitchcock et al. (1964) contend — then one may guess it was inadvertently transported by ship in wet or dry ballast froin the Atlantic coast, The junior author has conducted tests on the effect of sea water on viability of C. edentula seeds collected at Cape Cod. Even after immersion in sea water for 10 weeks, washed seeds gave 50-60% germination, suggesting that the seeds could survive for long periods in wet ballast. From the place of its introduction, californica spread rapidly both north and south. Only a few years after its introduction, Greene (1891) reported it as “Common along sandy beaches about the Bay of San Francisco at West Berkeley, Alameda, etc. ; also at Half Moon Bay; doubtless not rare on the coast and probably indigenous . . : Half Moon Bay is about 25 miles south of San Francisco and faces the Pacific. Not long after that, B. L. Robinson, in Synoptical Flora of North America (Gray, 1895-7) listed two species of Cakile: C. maritima **. . . as ballast weed upon the Atlantic Coast of the Middle States . . .' and C. americana Nutt. (=C. edentula) on « . . gea beaches, Gulf of St. Lawrence to Florida; on the Pacific in central Cali- fornia (perhaps introduced), and along the shores of the Great Lakes." 374 Rhodora [Vol. 72 Going north, the earliest Oregon collection seems to be one by J. E. Kirkwood (NY) from the sand dunes at Nes- tucca in Tillamook County and dated 1901. The earliest Washington collection we have seen is one by E. A. Mc- Gregor (DS) from Long Beach in Pacific County, dated 1907. Jones (1936) has previously cited the first Washing- ton collection as being a single plant at Port Angeles, about 650 miles north of San Francisco, in 1908. Jones went on to point out that Piper and Beattie (1915) stated Cakile Was ", . , very rare...” on the northwest coast and cited only two locations for it, Port Angeles and Ucluelet on Vancouver Island. We have seen a collection of californica by Macoun (GH, NY) from Ucluelet, British Columbia, dated 1909. Henry (1915) also cited the only location for Cakile along the British Columbia coast as being Ucluelet. A few years later, however, Kermode (1921) reported califor- nica as being “. . . locally distributed, coast of Vancouver Island." These herbarium records suggest a progressive north- ward migration: California, 1882; Oregon, 1901; Wash- ington, 1907 ; British Columbia, 1909. However, Cakile was probably not migrating up the coast as a continuous front; it was probably jumping north considerable distances, then gradually filling in locations south of the northernmost extension. For example, early collections for Oregon are very few in number. The earliest collection for central or southern Oregon we have seen is of californica by H. H. Smith (F, NY) from Charleston Bay in Coos Co., dated 1911— yet it had reached Port Angeles at the Canadian border three years before that. Finally, Hultén (1936) and Eyerdam (1938) reported californica on Kodiak Island in 1931 — some 2000 miles north of San Francisco and implying a phenomenal rate of migration of 40 miles per year for 50 years. At the present time, californica “. . . is a very common species all along the coast of British Columbia . . .” (personal com- munication, Dr. Roy L. Taylor, University of British Co- lumbia). 1970] Cakile edentula — Barbour and Rodman 375 To the south, californica had reached Corona Del Mar, Orange County, California by 1908 (collection by A. David- son, ORE) and Ocean Beach, San Diego County by 1906 (collection by K. Brandegee, NY, UC, us). Munz (1935) reported it as present from Santa Barbara to San Diego. Smith (1952) reported it for the Channel Islands of Santa Rosa and San Miguel. According to papers by Greene (1886- 7) for Santa Cruz Island, by Brandegee (1888) for Santa Rosa Island, and by Lyon (1886) for Santa Catalina and San Clemente Islands, the Channel Islands did not support californica in the late 19th century. In fact, collections by Millspaugh and Nuttall (1923) showed it to be absent from Santa Catalina Island even in the early 20th century. A 1936 collection by Frank F. Gander (sp) showed it to be present at the United States — Mexico border in San Diego County, but apparently it has not penetrated Baja Califor- nia (Cooper, 1936, Eastwood, 1929, Goldman, 1916, John- son, 1958). Within this range, how abundant was californica? In- formation of this sort is scarce in the taxonomic literature, but the following comments have been noted: “. .. occa- sional...” from Santa Barbara to San Diego (Munz, 1936) ; “ — . scattered . . ." around Santa Barbara and on the islands of Santa Rosa and San Miguel (Smith, 1952) “... at the present day no botanist could collect on the ocean strand of San Francisco County without finding Cakile [eden- tula] ..." (Jepson, 1936) ; * — abundant..." at the base of Table Bluff in Humboldt County (collection by J. P. Tracy, 1941, UC); “... frequent...” at Trinidad in Hum- boldt County (collection by J. P. Tracy, 1948, uc); “... this plant is not uncommon on the Washington sea beaches, and in many localities is abundant...” (Jones, 1936) ; “... scat- tered ...” in Pacific County, Washington (collection by A. N. Steward, 1953, osc); “... widely distributed . . .” from San Diego to Vancouver Island (Cooper, 1936) ; *. . . com- mon...” near Prince Rupert, British Columbia (collection by J. A. Calder, 1954, Osc). One may conclude that californica at the time maritima 376 Rhodora [Vol. 72 was first seen in 1935 and for some years after that was very patchy in local distribution, sometimes scattered and sometimes abundant. It was not rare, however, and per- haps the best term to describe its past overall abundance is occasional. The rise of C. maritima and the fall of C. edentula ssp. californica The first collection of maritima on the west coast was made by Rose (1936) at Stinson Beach, Marin County in May of 1935. Stinson Beach is about 15 miles north of San Francisco. Rose stated the two Cakile species were growing intermingled, but he did not comment on their relative abundance. Morphological differences between the two species (having to do with leaf shape, fruit shape, and petal size) are marked and consistent ; in our survey we have found few misidentifications, and no misidentifications which produced a maritima collection on the west coast prior to 1935. After 1935, maritima appeared to have spread almost as fast as californica had before it. In 1937 it was at Salmon Creek, some 40 miles north of Stinson Beach (Baker, 1940) ; Roxana Ferris described it there as “. . . not un- common . . ." (1937 collection, GH). Collections by J. P. Tracy revealed that maritima reached Humboldt County, 200 or more miles north of Stinson Beach, by 1940: ". . . scattered plants, among other Cakile . . ." at Samoa (1940, GH); “. . . a single plant noticed . . .” at the foot of Table Bluff (1941, uc); “ . . introduced there since 1941 . . ." at Centerville Beach (1946, uc); “. . . a very recent introduction . . ." at Trinidad (1948, UC). Peck did not report maritima in his first edition of Oregon Flora (1941), but did include it in the second edition (1961) with the comment, “On beaches and young dunes at various points along the coast." The earliest Oregon collection of maritima we have seen is one by M. S. Doty from Sunset Bay in Coos County, dated 1942, with the comment “. . . scat- tered . . ." (ps). The earliest Washington collection we 1970] ` Cakile edentula — Barbour and Rodman 377 have seen is one by M. Ownbey and F. G. Meyer from Muk- kaw Bay, Clallam County, dated 1940 (DS, GH, UC), but more recently Gilkey and Dennis (1967) have reported maritima for sea beaches from Lane County, Oregon to northern Washington. Calder and Taylor (1968) found it on the Queen Charlotte Islands off British Columbia in 1957 — the first published account of it in British Colum- bia — but a collection by W. Newton in 1951 showed it to be present near Ucluelet some years before that (DAO, UBC). In a personal communication, Taylor added that maritima was 0 rare... al along the British Columbia coast, but judging from herbarium records, it can be locally com- mon (eg, collection by J. A. Calder and D. B. O. Savile and R. L. Taylor from one locality in Queen Charlotte Islands in 1957 had the note “. .. common . . "^, DAO). In the south, Roxana Ferris reported maritima at Mon- terey *... in large clumps . .. growing with C. edentula var. californica...” (1940 collection, GH). Smith (1952) noted maritima to be “. .. locally common . . ” around Santa Barbara but absent frcm the Channel Islands. Thorne (1967) found maritima to be“... well-established . . ." on Santa Catalina Island, and Roman Gankin, Botanist for the Davis Botanic Garden, reported it to be ". . . abun- dant..." at the west end of Santa Cruz Island in 1969 (per- sonal communication). Peter H. Raven found it “. . . abun- dant..." on San Clemente Island (1962 collection, sp), and present on San Nicolas Island (1966 collection, SD). Reid Moran noted it as “. .. Scarce. . ” as far south as Cédros Island off Baja California (1963 collection, DS, RSA, SD, UC, us). Im less than 30 years, then, maritima extended its range about 1000 miles north and south of Stinson Beach — an average migratory rate of 33 miles per year. Although this rate is slower than the 40 miles per year northward rate of californica it is still considerably faster than migra- tory rates of other California weeds. Using the careful records of weed introduction dates and areas of infestation compiled by Robbins (1940, 1952), it is possible to calculate 378 Rhodora [Vol. 72 maximum migration rates for some California weeds. Hy- pericum perfoliatum, for example, migrated 24 miles per year; Cirsium arvense, l7 miles per year; Convolvulus ar- vense, T miles per year; Cotula coronopifolia and Salsola kali var, tenuifolia, 6 miles per year. Also, the study of Taeiniatherum asperum (= Elymus caput-medusae) by McKell et al. (1962) indicated a migratory rate for that Species of 7 miles per year. These weeds are not strand plants, of course, and the more rapid rate of Cakile species migration may be due to transport of fruit and seed by ocean currents or by shipping. The junior author has found that upper joints of fruits of C. edentula can remain afloat and viable in Sea water for 11 days, Suggesting that transport by tidal and shore currents is possible. Barbour (1970b) has shown that exposure of maritima seeds to even 3% NaCl will not impair subsequent germination if the seeds are washed in distilled water. Concommitant with the spread of maritima has been the decline in abundance of californica. The disappearance of californica along the California coast has rarely been men- tioned in the literature or in herbarium collections. A 1941 collection by R. Bacigalupi (UC) from the San Fran- cisco — San Mateo County Line had the note, “Cakile mari- tima very abundant, C. edentula not in evidence.” A 1946 collection by J. P. Tracy (UC) from Centerville Beach in Humboldt County had the note, “Introduced there since 1941, but now more abundant than C. edentula var. cali- fornica.” Of californica for the San Francisco region, How- ell et al. (1958) wrote: “No recent collection of this beach plant has been seen." The most convincing evidence of replacement of califor- nica by maritima along the California coast comes from recent field visits to sites at which past herbarium collec- tions of one or both species have been made. Our own trips, and recent trips of other Davis botanists allowed us to compile the paired observations summarized in Table 1. All the 17 sites listed in Table 1 presently support mari- tima, and the abundance of maritima. ranges from occa- 1970] Cakile edentula — Barbour and Rodman 379 sional or sporadic (Samoa, San Diego) to abundant (Trini- dad, Santa Cruz Island). It must be assumed that mari- tima was absent at all 17 sites prior to 1935, based on our literature and herbarium survey. Cakile maritima, then, has enormously increased its range and can generally be described as common along the entire California coast. Cakile edentula ssp. californica, however, has reduced its range, It is probable that californica was present at all 17 sites in 1935, but there are definite records of its presence at only 13 sites. Of these 13 sites, it was still present in 1969 at only 2 sites — and in both cases (Trinidad and Table Bluff), only a single plant of californica was seen. The change in abundance of the two species is especially dramatic and well documented at Table Bluff: within 28 years, the two exactly reversed their relative abundance. This replacement of californiea by maritima is appar- ently not taking place further north. From Ensenada, Baja California, to Fort Bragg, California, we have found only maritima, usually abundant and fruiting heavily. In north- ern California and central Oregon we began seeing scat- tered individuals of californica and noted that maritima was less abundant. In northern Oregon and Washington californica was predominant, often quite abundant, with a few maritima scattered throughout the populations. Plants with hybrid characters are very rare, both in the field and in past herbarium collections, so at this point we |. hypothesize that hybridization is not occurring; the junior author, however, is at present attempting laboratory crosses. Conclusions Neither Cakile edentula ssp. californica nor C. maritima are native to the west coast of North America. Cakile eden- tula ssp. californica (perhaps better, C. edentula) was introduced between 1880 and 1882 to the San Francisco Bay area, probably by ship from the east coast. It spread as fast as 40 miles per year north to Kodiak Island and south to San Diego. Cakile maritima was introduced in [Vol. 72 Rhodora 380 (HD ‘wowpoy ‘aon *4noq4ng ) «[PUOISBI90,, pvu pue ,3uosqv,, fypo (don ‘unoqung) «[BUOISBII0,, pvu pue ,.juasqe,, {ype (doa ‘unoqung) 4[€UOISU200, punu pue «1U98q*9, fno (aoa *4noq4ng ) ,, UOUIUIOO,, Punu pue ,oe1e1 0} [fuOIS?200, imo (aoa *4noqung) «UOU “WU0d,, 7104 pu* „uəəs juv[d əuo,, fynə (HD ‘upwpoy ‘apn ‘unoq.ng ) «[BUOISBI90,, punu pue «juasqe,, fiə (aoa “enogang) ,3uvp -Unq?,, 314014 pue ,uoos juv[d ouo, fumo (qon ‘unoqung) «[BUOISBI90,, jevu pue «USEL, fimo SS eee U $9}0U uonoə[[o5 (6961) quasaig tay pur *iougjne 'uorjoo[[oo 4sed JO 93*p 0} Jajar sisəuquəred ur durpaoooe (mmu) vui 'Ə pu? (fumo) D21440/1709. *dss Dm1uo (on PUDLI) PUD JUDLD '9pG[) JUasaad 314m *(uonoe[[o ou) yuUasard Ajqissod fumo (92 “NƏH ‘OFGT) «[BUOISBOI0,, panu (qas ‘Land ‘ZEgl) juəsəad fimo (H9 “81449. ‘LEGTI) ,UOUmuooun jou, 724044 * (uonoə[[oə ou) yUasard Ajqissod fyno (oso 'jjnjdwaH, 961) 3uesead pamu *(uonoo[[oo ou) q4uesoid A[qissod fina (on ADLE *Ty Gr) «Pe Tou juv[d o[Surs e A[UO, penu (on “hovel 'gpg[) ,3uepunqe, pivu (on "opa ‘OF6T) «PNAN Tayo Suoure p2.1933905,, panu í (On Roney ‘OF6L) 3uesoud Hb (609 ‘ADe, “SFGI) 4Qu€punqe, ju ‘ (OA “ion. 'gpgr) 4AJuenbodg, fimo (uses uorjoo[[oo ou) usq; Juasqe Apue1edde punu ‘(aon "uS PUD SION 'Gpg[) 3uosoid fimo $9jou uonoəəloo jseq o T.,8€ “OO NDIVK 'Qovog uo[(q ,91,88 “oo VWONOS 'peoH v3opog ,08,8g “02 VINONOS *q9947) uouieg Jo ynom 06.68 “00 ONIDOGNAIV TOATY 9[IJ Uo], ro ymo OP OF “09 LATINO 'ga[g o[qv,, 09, OF ^09 LaToaWwaH ‘vourrs S0.1P “OO LalIoaWwaH *peprur. m ST.TP "oo LATOAWAH 'uooZeT ouojg an, Nin, J uonjoo[[O9 Jo uorjvoo' AYG — GON :930N 'pejrsodop yorya ur umrieq E SUJI '$93o0u uorj»o[[oo 3uesoid pue ysed 04 pa ayn Jo oouepunqe 9AI?e[ogp (1 ƏlqeL Cakile edentula — Barbour and Rodman 381 1970] olperods,, pnw pue auepunqe,, pvu pue „UOWW0,, 42409 pue ,juepunge,, 7pm pue UOWOD,, pvu pue ,UOUIUIOD, 41401 (HD *umupog panu pue pue ,,UOUIQO2,, 418Uu0IS9220,, panu pue 4I&uorswooo, pnw pue (HD ‘upumpoy) Quesqe, fo (qon furwpz)) | gh hte were — Ca WE Y qu A s Z oth A? rat ti, , Pea [qr dd ` < , PT Ln , ` Bariut CARNEOUE MUSEUM PITTEBukGe, pA TYPE - Fig. 1. Type specimens of Pyrola compacta illustrating the general habit and leaf shape of P. elliptica (A) and the abnormal flowers with extra petals and conspicuously mucronate anthers (B). 482 Rhodora [Vol. 72 A Fig. 2. Type specimen of Pyrola oxypetala (A & B) and an ab- normal specimen of P. asarifolia var. asarifolia (C & D). The two mutant specimens are similar in their erect flowers and extrose stamens. 1970] Pyrola — Haber 483 that the flower color must have been purplish or rose. It is more likely that the flowers were white as in typical P. elliptica since all of the specimens on the sheet are scorched to a deep brown color. The appearance of these petals is much like those on similar sheets of P. elliptica. Jennings also reported that the type specimens were very fragrant; a similar observation has also been made for P. elliptica by Lakela (1965). P. compacta appears to have been over- looked in the literature, for no mention of the name has been found applied either to a distinct species or in syn- onymy with P. elliptica. Pyrola oxypetala Austin, maintained as a distinct species in the eighth edition of Gray’s Manual, is more appropri- ately placed in synonymy with P. virens Schweigg as has been done by Gleason (1963). The type specimen (fig. 2, A & B) collected near Deposit, N.Y. exhibits the ovate leaves of P. virens, although somewhat more lanceolate than normal, as well as the short, merely acute, triangular-ovate sepal lobes which do not overlap at the bases. The flowers are abnormal in that they are held erect and that the petals taper either gradually or quite abruptly to a sharp point; as well, the stamens are extrose with the tubes facing the base of the ovary rather than being inverted (with the tubes pointing away from the ovary) as in normal Pyrola flowers during anthesis. The stamens in Pyrola buds are always extrose but become introse by the time the flowers are fully opened. The short, barely exserted, straight style of P. oxypetala with the stigmatic lobes not yet visible at the tip also appears as in many young Pyrola flowers. The erect flowers, extrose stamens, and short, straight style are not features unique to this plant alone. One speci- men of P. asarifolia from the Thunder Bay District of Ontario (LKHD 9257; fig. 2, C & D), typical in all key features such as the ovate-cordate leaves, the shape and length of the sepals, and the pink flowers, exhibits these same features. The inflorescence with its flowers borne on long, erect pedicels has the appearance of an ornate can- delabrum. Although the flowers were apparently almost 484 Rhodora [Vol. 72 fully open when the plant was collected, the stamens are still completely extrose and the short style barely exserted. The morphological changes brought on by mutations are not always quite so conspicuous ; mutations producing minor changes are more frequently found. One collection of P. asarifolia (CAN 86307) was found in which the sepals in each flower were fused into two groups — one with 3 sepals, the other with 2. Variations in the numbers of floral parts may sometimes be found as in P. asarifolia (CAN 237255) in which one flower on the plant had its sepals, petals, car- pels, and stigmas in 6’s and as well had 12 stamens. Some- what similar variations in the number of floral parts have also been observed in specimens of P. minor and P. grandi- flora. Without a doubt mutations play their role in the process of speciation through the introduction of additional varia- tion. However, such gross alterations of the basic floral structures of individual plants or clones from the population norm as seen in P. compacta and P. oxypetala must surely offer little adaptive advantage to the plants. The monstrous origin of such aberrations should be noted, but such individ- uals deserve no specific recognition. Specimens Cited Pyrola compacta Jennings (Type): Wet, open pasture near Six- mile Lake (Louise Lake), Thunder Cape, northwestern Ontario, Aug. 20, 1912, Otto E. & Grace K. Jennings (cM). Pyrola oxypetala Austin (Type): Wooded hill near Deposit, Delaware Co., N.Y., June 1, 1860, C. F. Austin (GH). Pyrola asarifolia Michx. var. asarifolia: Rocky woods along lakeshore, north shore of Little Pigeon Bay, Crooks Tp., Thunder Bay District, Ontario, July 8, 1952, C. E. Garton 1785 (LKHD 9257). Pyrola asarifolia Michx. var. purpurea (Bunge) Fern.: Near Mud Lake, Oliver Tp., Thunder Bay District, Ontario, June 30, 1937, C. E. Garton 511 (CAN 86307) ; Larch bog, vicinity of Knob Lake, north of John Lake, Quebec, 54°48’ N., 66°49 W. July 11, 1955, L. A. Viereck 618 (CAN 237255). LITERATURE CITED GLEASON, H. A. 1963. Illustrated Flora of the Northeastern United States and Adjacent Canada. New York Botanical Garden, New York. 1970] Pyrola — Haber 485 JENNINGS, O. E. 1920. New or noteworthy plants from northwest- ern Ontario. Journ. Wash. Acad. Sci. 10: 453-460. LAKELA, O. 1965. A Flora of Northeastern Minnesota. Univ. of Minnesota Press, Minneapolis. DEPARTMENT OF BOTANY UNIVERSITY OF TORONTO TORONTO 181, ONTARIO JUNCUS SLWOOKOÓRUM, A NEW SPECIES FROM ARCTIC ALASKA STEVEN B. YOUNG On July 23, 1966, I collected specimens of a small Juncus near Boxer Bay (63° 22’ N. Lat., 171° 35’ W. Long.) south- western Saint Lawrence Island, in the northern Bering Sea. These specimens cannot be identified as any known species of Juncus. They show some similarity to depauperate speci- mens of J. Fawriensis Buch., an East Asian species not known to occur in Alaska. However, the equitant leaves and unique, two flowered inflorescences (Fig. 1) of the new species are distinctive. Each of the approximately 10 in- florescences examined were identical in that they consisted of two flowers. The lower flower is essentially sessile; the upper is borne on a pedicel about 5 mm long. Each of the flowers probably represents a reduced head. The specimens were collected at the edge of a small tun- dra pond. They grew in thick, saturated moss, in association with Saxifraga nudicaulis D. Don. and S. rivularis L. The new species is named after the Slwooko family of Gambell, Saint Lawrence Island. Several members of this family contributed greatly to the success of my field work on the island, Boxer Bay, the type locality for J. Slwookoó- rum, is the ancestral hunting and trapping ground of the Slwookos. Juncus Slwookoorum S. B. Young Perennis, rhizomate brevi horizontali, radicibus filiformi- bus elongatibus. Caulis erectus, 5-15 cm altus, teres. Folia basalaria 10-15 em alta, equitantia. Vagina in auriculis duabus obtusis castaneis producta. Lamina foliosa, septis transversis completis externe plus minusve prominentibus interceptis, Inflorescentia terminalis, 2-florae, superata bractea foliacea longa. Flos inferior sessilis; flos superior portatus in pedicello plus minusve 5 mm longo. Flores 4-5 mm longi. Tepala membranacea, castanea, uninervia, lan- ceolata, acuta, aequilonga vel externa paullo longiora. 486 1970] Juncus slwookoórum — Young 487 Fig. 1. Juncas Slwookoórum: A — typical habit, B — typical in- florescence, consisting of two flowers, one sessile, the other borne on a pedicel. 488 Rhodora [Vol. 72 Stamina tres, perigonium superantia. Stylus filiformis; stigmata 3, longa, exserta, torta. Fructus c 2 mm longus, obovatus, non rostratus; pericarpium membranaceum, pal- lidum. Holotypus: S. B. Young 711 (GH): Sinus Boxeri, Insula Sancti Laurentii. Perennial with short horizontal rhizome and elongated filiform roots. Stem erect, 5-15 cm tall, terete (or slightly flattened in dried specimens). Basal leaves 10-15 em long, equitant. Sheath extended in two obtuse, castaneous auriculi. Blade flat and foliaceous, at least the upper portions having more or less prominent transverse septa. Inflorescence terminal, 2-flowered, over- topped by a long leafy bract. Lower flower sessile, the upper flower borne on a pedicel c 5 em long. Flowers 4-5 mm long. Tepais membranaceous, brown, 1-nerved, lanceolate, acute, of equal length or the exterior ones slightly longer. Stamens 2, overtopping the perigon. Style filiform, stigmas 3, long, exserted, and twisted. Fruit c 2 mm long, obovate, beak- less; pericarp membranaceous, pale. Juncus Slhwookoórum joins a small but fairly distinctive group of species whose ranges are narrowly endemic to the Bering Straits region. Other examples are Carex Jacobi- peteri Hult., Papaver Walpolei Pors., Saxifraga nudicaulis D. Don. (with disjunct stations in eastern Asia), and Ar- temisia senjavenensis Bess. These species probably repre- sent remnants of a specialized flora of the northern portion of the Bering Land Bridge. Field work on Saint Lawrence Island was supported by grants from the Institute of Evolutionary Biology of Har- vard University, the Fernald Fund of the Gray Herbarium, and Sigma Xi. INSTITUTE OF POLAR STUDIES THE OHIO STATE UNIVERSITY COLUMBUS, OHIO 43210 THE SOUTHERN LIMITS OF TRIENTALIS BOREALIS WILBUR H. DUNCAN Trientalis borealis Raf. is now known to be another of those species having northern affinities but extending south- ward in the mountains of the eastern United States into Georgia. On the basis of known stations this species is in one sense doubly disjunct. There are a number of localities where the species is known to occur in the mountains as far south as southern West Virginia and adjacent Virginia, in the Kanawha Plateau, Allegheny Mountains, and Ridge and Valley Province. Southward there are only two widely separated localities known for the species. The next station south of those in Virginia and West Virginia was reported by James (1955) from northeastern Tennessee in the hills of Sullivan County near Bristol. This locality is about 90 miles from the nearest known population to the north, that in Mercer County, West Virginia. Sharp and Baker (1964) state that in Tennessee T. borealis “has been collected only in Sullivan County.” On 12 May 1963 I collected Trientalis borealis at about 3,700 feet elevation in a north-facing ravine in the southern part of Union County, Georgia. This locality is about 160 miles from the nearest known station, in Tennessee. It is on the basis of the 90 mile and 160 mile intervals that I suggest the species might be described as doubly disjunct in the southeastern mountains. That the station in Georgia represents a significant ex- tension in range was established by analyzing distributional data from the University of Georgia and other herbaria.’ These data are recorded on Map 1. The westernmost records shown in Minnesota and the southernmost records in the states from Illinois to Georgia and Delaware are at or near the known western and southern limits of the species in the United States. However, since I sought primarily to "The records generously provided by curators from the several herbaria are gratefully acknowledged. 489 490 Rhodora [Vol. 72 Map 1. Southern distribution of Trientalis borealis. establish the southern limits of T. borealis, the records shown northward on Map 1 do not reflect its abundance there. Distributional data given by Anderson (1968) show that the species is more abundant northward, although fail- ing to show the southernmost records for Illinois and Ohio and the single record previously reported for northeastern Tennessee. The species has also been reported in Canada west to Alberta and northeast to Labrador (Gleason and Cronquist, 1963; Anderson, 1968). Much botanizing has been done in the Smoky Mountains and other parts of the southern mountains without Trien- talis being found, therefore making the “outpost” in Georgia of considerable interest. The factors which provide a suit- 1970] Trientalis borealis — Duncan 491 able habitat at the Georgia station, and which are appar- ently not present at intervening habitats, are not known. The ravine, however, is obviously cool in nature and is occupied mostly by species which are distinctly northern in distribution. The ravine drops from about 4,000 to 3,150 feet elevation in 0.35 mile between two ridges which run downwardly N 26° W and N 30° E respectively from a point where they diverge from a single higher ridge. Little sun reaches the forest floor in the heart of the ravine. Early and late in the day the two ridges block out the sun. In the middle part of the day the sun is too low on the horizon to shine on the floor except in summer and then the rays are at such a steep angle with the floor of the ravine that the canopy of the woody vegetation provides a considerable barrier to them. Much of the ravine is occupied by small to very large angular boulders in a jumbled arrangement. Some of the thousands of plants of Trientalis growing in the ravine occur on top of boulders; others occur between the boulders and on parts of the steep slopes of the eastern side of the ravine. When plants from different localities in the ravine were dug up it was discovered that all were rooted in weakly consolidated, partially decayed light brown humus composed largely of leaves from deciduous woody plants. The roots seemed to have little association with the mineral soil. Anderson (1968) reports that the rhizome grows just under the organic layer to within a centimeter of the surface in Wisconsin. Vegetation in the ravine is quite northern in aspect, most of the species occurring there being typically northern in their distribution. For example, Dryopteris goldiana (Hook.) A. Gray was reported from there as the southern- most record for the species (Duncan, 1948). The character of the vegetation in the ravine is indicated by the following list of some other species occurring there. WOODY PLANTS: Juglans cinerea, Betula lenta, B. lutea, Magnolia acuminata, M. fraseri, Rubus odoratus, 492 Rhodora [Vol. 72 Cladrastis lutea, Ilex montana, Euonymous obovatus, Acer pennsylvanicum, A. spicatum, and Aesculus octandra. HER- BACEOUS PLANTS: Trillium erectum, T. grandiflorum, T. vaseyi, Asarum canadense, Aquilegia ca nadensis, Diphyl- leia cymosa, Dicentra canadensis, D. cucullaria, Impatiens pallida, Aralia nudicaulis, Hydrophyllum canadense, and Monarda didyma. I overlooked Trientalis on previous visits to the ravine, and others may have overlooked the species elsewhere in the southeastern United States. Mature plants of Trientalis with their single whorls of leaves resemble Medeola virgin- iana and some species of Lysimachia. A second and careful look at plants of that appearance may reward the observer with the discovery of a third station for Trientalis borealis in the southeastern United States. A voucher specimen of Trientalis borealis from Georgia is on deposit in the University of Georgia Herbarium (Dun- can 22050). DEPARTMENT OF BOTANY UNIVERSITY OF GEORGIA ATHENS. GEORGIA 30601 LITERATURE CITED ANDERSON, ROGER C. 1968. The Autecology of Trientalis borealis. Ph.D. Thesis. Univ. Wisconsin (Libr. Congr. Card No. Mic. 69- 865). DUNCAN, WILBUR H. 1948. New Records of Two Ferns in Georgia. Amer. Fern Jour. 38: 69-70. GLEASON, H. A. and A. CRONQUIST. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. D. Van Nostrand Co., Inc. N.Y. 810 p. JAMES, ROBERT L. 1955. Some Botanical Notes from Northeast Tennessee. Castanea 20: 77-103. SHARP, A. J. and AILSIE BAKER. 1964. First and Interesting Re- ports of Flowering Plants in Tennessee. Castanea 29: 178-185. FURTHER ADDITIONS TO THE BRYOFLORA OF MT. KATAHDIN, MAINE F. J. HERMANN On August 19, 1963 I had an opportunity to make the ascent of Mt. Katahdin via the Abol Trail, returning by way of the Appalachian Trail, and as a result of this and an earlier partial climb as far as Chimney Pond was able to report the addition of 20 mosses and 2 hepatics to the 86 bryophytes previously known from the mountain (77 reported by Kennedy and Collins, 5 by Lorentz and 4 by Lowe). The following year a second ascent, on August 28, via the Appalachian and Saddle Trails, and a hike up South Turner Mt. (essentially a spur of Katahdin) on Septem- ber 1, resulted in the further addition of 21 unrecorded bryophytes (14 Musci and 7 Hepatics) which are here re- corded to bring the total now known from Mt. Katahdin up to 129. As with the previous year’s collections, a set has been deposited in the U. S. National Herbarium and dupli- cates of most have been distributed to several other major herbaria. To avoid repetition of habitat data the species are listed according to collecting sites. Tundra on plateau, Baxter Peak Cutoff, 4600 ft. : Dicranum muehlenbeckii BSG., 19594. Cephalozia ambigua Massal., 1959415. Although Frye and Clark recorded this species from only “Ellesmere Isl.; Pim Island; Southern Applachian Mts., Europe," it is probably more widely distributed than supposed, very likely being generally overlooked because of its minute size. It was found to be plentiful, for instance, at the edge of melting snowbanks on Mt. Baker, Washington. Vertical face of granite boulder, Saddle Trail, 3500 ft.: Ptilidium pulcherrimum (Web.) Hampe, 1960115. Humus on granite ledge above Katahdin Falls, 1700 ft.: Sharpiella striatella (Brid.) Iwats., 19571. Plagiothecium denticulatum (Hedw.) BSG., 19573. 493 494 Rhodora [Vol. 72 Wet, shaded bank of Katahdin Stream below Falls: Calypogeia meylanii Buch, 19568. Cephalozia bicuspidata (L.) Dumort., 19570. Pellia fabroniana Raddi, 19567. Vertical face of granite boulder in woods, Appalachian Trail, 3000 ft.: Trichostomum tenuirostre (Brid.) E. G. Britt., 19581. Boggy swale in Thuja-Betula woods on SE shore of Sandy Stream Pond, 1500 ft.: Calliergon stramineum (Brid.) Kindb., 19621. Dicranodontium denundatum (Brid.) E. G. Britt., 19623. An unusually wet habitat for this species. Drepanocladus exannulatus (BSG.) Warnst., 19618. Rhizomnium pseudopunctatum (Bruch & Schimp.) Kopon., 19620, Chiloscyphus pallescens (Ehrh.) Dumort., 1961915. Pellia epiphylla (L.) Corda, 19619. Gravelly trail through open woods, South Turner Mt.: Brachythecium starkei (Brid.) B. & G., 1600 ft., 19624. Bryum pallens (Web. & Mohr) Brid., 2200 ft., 19632. Hypnum haldanianum Grev. (Heterophyllum haldaneanum (Grev.) Kindb.), 2600 ft., 1963315. In pool along trail, South Turner Mt., 1600 ft.: Dichelyma falcatum (Hedw.) Myr., 19624. Drepanocladus fluitans (Hedw.) Warnst., 19625. Granite boulder along trail, South Turner Mt., 1800 ft.: Rhacomitrium heterostichum var. affine (Schleich.) C. Jens., 19627. FOREST SERVICE HERBARIUM U. S, DEPARTMENT OF AGRICULTURE WASHINGTON, D.C. LITERATURE CITED FRYE, T. C. & L. CLARK. 1945. Hepaticae of North America. Univ. of Wash. Publications in Biology, Vol. 6, No. 3. 1970] Bryoflora — Hermann 495 HERMANN, F. J. 1964. Additions to the bryophyte flora of Mt. Katahdin, Maine. The Bryologist 67: 204-206. KENNEDY, G. G. & J. F. COLLINS. 1901. The Bryophytes of Mt. Katahdin. Rhodora 3: 177-181. LORENTZE, ANNE. 1917. Notes on the Hepaticae of Mt. Ktaadn. The Bryologist 20: 41-46. Lowe, R. L. 1924. Collecting mosses on Mt. Ktaadn. The Bryolo- gist 27: 57. Lower, R. L. 1926. A second trip to Mt. Katahdin, Maine. The Bryologist 29: 15, 70. CONTRIBUTIONS TO THE FLORA OF WASHINGTON GEORGE W. DOUGLAS! and RONALD J. TAYLOR The flora of Washington is generally rather well known and has received considerable attention over the years, particularly with the completion of the comprehensive treat- ment, Vascular Plants of the Pacific Northwest, Volumes 1-5 (Hitchcock, et al., 1955, 1959, 1961, 1964, 1969). How- ever, floristic studies in the northwestern corner of the state have been relatively incomplete. This is particularly true of the Cascades north of Stevens Pass where the rugged mountain wilderness remains very inaccessible. Recent col- lections by the authors have provided a more complete botanical coverage of the North Cascades ; and in this paper, eight new records of vascular plants and mosses are re- ported for Washington together with a number of signifi- cant geographical and altitudinal range extensions, Most representatives were collected in subalpine or alpine hab- itats, a few are lowland Species, The specimens cited are deposited in one or more of the following herbaria: G. W. Douglas (private) (GD) ; New York Botanical Garden (NY) ; Skagit Valley Community College (sv); U. S. Forest Service (Fs) ; University of Washington (UW) ; and Western Washington State College (ww). Nomenclatural determinations were made by the authors unless otherwise noted. New species records are designated by an asterisk. VASCULAR PLANTS Polystichum munitum (Kaulf.) Presl. var. munitum. Green Mountain, Snohomish Co., Douglas 870 (GD); Sourdough Ridge, Whatcom Co., Douglas 1900 (Gp). This species is usually found from low to middle elevations (sea level to ‘Present address: Department of Botany, University of Alberta, Edmonton 7, Alberta. 496 1970] Flora of Washington — Douglass and Taylor 497 ca. 900 m) in the moist continuous forests of the region. The Green Mountain collection was taken at 1555 m eleva- tion on a steep colluvial slope beneath an isolated tree (Tsuga heterophyila (Raf.) Sarg.) in the Subalpine Zone. The Sourdough Ridge collection was established beneath krummholz (Abies lasiocarpa (Hook.) Nutt.) at an eleva- tion of 1920 m in the Alpine Zone. The Green Mountain collection was typical of the species while the Sourdough Ridge collection was depauperate and reached a height of only 15 em. Both specimens had well developed sporangia. Pteridium aquilinum (L.) Kuhn ssp. aquilinum var. pubes- cens Underw. Green Mountain, Snohomish Co., Douglas & Pearson 1632 (Gp). A common species at lower elevations throughout Washington. The Green Mountain specimen was collected on deep well-drained soil of a steep colluvial slope at 1610 m elevation. This fern is an important component of the lush herbaceous associations found on steep colluvial slopes or avalanche tracks throughout the Subalpine Zone (Douglas, 1970). Carex multicostata MacKenzie. Elephant Butte, Whatcom Co., Douglas 1735 (GD, FS). This collection was made on an exposed alpine ridge at 1920 m elevation. The previous known range of this species was the Cascade-Sierran region from central Washington south to the mountains of south- ern California, and in the mountains of eastern Oregon and adjacent Idaho (Hitchcock, et al., 1969). The identification was made by Dr. F. J. Hermann. Carex straminiformis Bailey. Green Mountain, Snohomish Co., Douglas 421, 431, 460, 551 (GD, FS). A frequent plant on mesic subalpine sites of the Green Mountain area. This species was known previously from the Sierra Nevada of California and adjacent Nevada extending north irregularly into the Cascade region to Mt. Adams in southern Wash- ington, and disjunet in the Wasatch Mountains of Utah and the mountains of central Idaho (Hitchcock et al., 1969). These specimens were identified by Dr. F. J. Hermann, 498 Rhodora [Vol. 72 "Carex subfusca W. Boott. Stehekin River, Chelan Co., Douglas and Lowery 1217 (GD, FS, UW). This collection, a new record for the state, was taken from a population of several scattered specimens established on a dry gravel floodplain. The species was previously known from the Klamath region of northwestern California and southwest- ern Oregon to the Sierra Nevadas and the mountains of southern California, extending east to the mountains of Arizona and southwest Utah and north into the Cascades to Mt. Hood in northern Oregon (Hitchcock, et al., 1969). The specimen was identified by Dr. F, J. Hermann. *Poa rupicola Nash. Chowder Ridge, Whatcom Co., Taylor 162, 763 (ww). These two collections were taken at 2120 m elevation from a dry interrupted alpine meadow dom- inated by Carex phaeocephala Piper. This is the first rec- ord of this grass in Washington. Hitchcock, et al., (1969) report the species is distributed from the Yukon, British Columbia and Alberta southward through the Rocky Moun- tains to Utah, Colorado and New Mexico, east to South Dakota, west to northeast Oregon, Nevada, the Sierra Nevadas of California, and northern Arizona. “Salix myrtillifolia Anderss. Miners Ridge, Snohomish Co., Naas & Naas (sv). This collection, a new record for the state, was taken from an alpine ridge at 1920 m elevation. Its previous range extended from Newfoundland and Que- bec to Alaska and British Columbia (east of the continental divide), south to California, southern Utah, Wyoming, and possibly northern Colorado (Hitchcock, et al., 1964). The specimen was identified by Dr. C. L. Hitchcock. Salix vestita Pursh var. erecta Anderss. Twin Lakes, What- com Co., Douglas 1950 (NY). This low shrub occurred on a wet, north facing subalpine scree slope at 1585 m elevation. It is an atypical specimen with style and stigmas too long, leaves too soon glabrous and bracts of the aments too green. The specimen is probably a hybrid between S. vestita var. erecta and possibly S. phylicifolia L. var. pennata (Ball) 1970] Flora of Washington — Douglass and Taylor 499 Cronq. (A. Cronquist, personal communication). This spe- cies was previously known in the Rocky Mountain region from southern Alberta and adjacent British Columbia to west central Montana, the Wallowa Mountains of Oregon, and in Chelan Co., Washington (Hitchcock, et al., 1964). The specimen was determined by Dr. A. Cronquist. Ranunculus ficaria L. Marietta, Whatcom Co., Taylor 2313, Sundquist 1114, 1165, 1550 (ww). These collections were all taken from a single population which forms a dense mat along the margin of a playground, previously a marsh. Ac- cording to Dr. Lyman Benson who confirmed the identifica- tion, this is the first recorded collection from the Pacific Northwest in the United States although a previous collec- tion was made in British Columbia, The species is native to Europe and sparingly introduced in North America (Fer- nald, 1950). Impatiens capensis Meerb. Lake Whatcom, Whatcom Co. Sundquist 1865, 1414, 1432, others (WW). This species is well established along the moist banks of Lake Whatcom. Its occurrence in the Pacific Northwest has been previously noted by Ornduff (1966). *Gentiana glauca Pall. Skyline Ridge, Whatcom Co., Taylor 1232 (ww) ; Skyline Ridge, Whatcom Co., Hobbs 12 (ww). These collections were taken at 1900 m elevation from an alpine meadow dominated by Carex spp. and disrupted by frost heaving. A new record for the state, this species was previously known from Alaska and Yukon extending south to British Columbia and in the Rocky Mountains to Mon- tana. It is also known in Asia (Hitchcock, et al., 1959). Mt. Garibaldi, in the southern British Columbia Coast Range, is apparently the southernmost station in British Columbia (Henry, 1915). Castilleja parviflora Bong. var. oreopola (Greenm.) Ownbey. Yellow Aster Butte, Whatcom Co., Douglas 1527 (GD, NY). This collection was taken in a moist subalpine meadow at 1585 m elevation. It represents a substantial range exten- 500 Rhodora [Vol. 72 sion, known previously from the Three Sisters Peak in Oregon to Mt. Rainier in Washington (Hitchcock, et al., 1959). The specimen was identified by Dr. N. H. Holmgren. *Centaurea montana L. N ewhalem, Skagit Co., Naas & Naas (SV). The specimen wags established with other members of the species in open fields of the area. This species, native to Europe (Hegi, 1954), has not previously been reported outside of cultivation in western North America. MOSSES *Pogonatum alpinum (Hedw.) Rohl var. septentrionale (Brid.) Brid. Green Mountain, Snohomish Co., Douglas 695 (GD). This collection was taken on a flat, poorly drained site in the Subalpine Zone dominated by a thick mat of Carex nigricans C. A. Mey. A new record for Washington, this moss was previously known from Greenland and Alaska, British Columbia and Colorado, as well as Europe and Asia (Grout, 1936). The specimen was identified by Dr. E. Lawton. “Bryum cyclophyllum ( Schwaegr.) B. S. G. Winchester Mountain, Whatcom Co., Douglas 734 (GD, UW). The speci- men was established in a moist rock crevice at 1585 m ele- vation. This moss, a new record for the state, is known from Greenland and arctic America extending south to New Jersey, Pennsylvania and Wisconsin, west to British Colum- bia, and also occurring in Europe and Asia (Grout, 1933). The specimen was identified by Dr, E. Lawton. Pohlia cardotii (Ren.) Broth. Seward Peak, Whatcom Co., Douglas 841 (GD, UW). This collection was taken in a poorly drained southeast basin in the Subalpine Zone. Carex nigri- cans was the sole dominant on the site. Very few collections of this species have been made and it was previously known only from Mt. Hood and Mt. Rainier (Grout, 1933). The specimen was identified by Dr. E. Lawton. Lescuraea saviana (De Not.) Lawt. Green Mountain, Sno- homish Co., Douglas 642 (GD, UW). This species occurs in 1970] Flora of Washington — Douglass and Taylor 501 moist subalpine meadows of the area. It is a common moss in Europe but is rare in North America (Lawton, 1957) where the only previous collections were made in the Olym- pic Mountains and on Mt. Rainier. The specimen was iden- tified by Dr. E. Lawton. COLLEGE OF FOREST RESOURCES UNIVERSITY OF WASHINGTON SEATTLE, WASHINGTON and DEPARTMENT OF BIOLOGY WESTERN WASHINGTON STATE COLLEGE BELLINGHAM, WASHINGTON LITERATURE CITED DovGLAs, GEORGE W. 1970. A vegetation study in the Subalpine Zone of the western North Cascades, Washington. M. 5. Thesis. Univ. of Washington. 293 p. FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th edition. Amer- ican Book Company, New York, N.Y. Grout, A. J. 1933. Moss Flora of North America North of Mexico. Author, Newfane, Vermont. Vol. 2. 285 p. Grout, A. J. 1936. Moss Flora of North America North of Mexico. Author, Newfane, Vermont. Vol. 1. 264 p. HEGI, Gustav. 1954. Illustrierte flora von mittel-Europa. Band VI, 2. Tiel. Carl Hanzer Verlag, Munchen. pp. 549-1380. HENRY, JosePu K. 1915. Flora of southern British Columbia and Vancouver Island. W. J. Gage and Co., Toronto, Ontario. 363 p. HircHucock, C. LEO, ARTHUR CRoNQUIST, MARION OWNBEY, & J. W. THOMPSON. 1955-1969. Vascular plants of the Pacific North- west. Vols. 1-5. Univ. of Washington Press, Seattle, Washington. LAWTON, ELVA. 1957. A revision of the genus Lescuraea in Europe and North America. Bull. Torrey Bot. Club 84: 281-307. ORNDUFF, ROBERT. 1967. Hybridization and regional variation in Pacifie Northwestern Impatiens (Balsaminaceae). Brittonia 19: 122-128. MONARDA STIPITATOGLANDULOSA, A NEW SPECIES FROM OKLAHOMA U. T. WATERFALL While collecting in southeastern Oklahoma in late June, 1966, the author noted a rather dwarf, erect, colonial mem- ber of the Labiateae on a mountain top, or ridge top, in Beaver Bend State Park, north of Broken Bow in McCur- tain County. It was past anthesis, and appeared to be either a Monarda, or a Pycnanthemum. He had seen it before, but not in flowering condition, so he dug it up and transplanted it at his home in Stillwater, In the spring of 1970, after having spread by rhizomes, the colony flowered abundantly. The material seemed dif- ferent enough from its nearest relative, Monarda fistulosa var. mollis, or from any other taxa in the genus as described by McClintock and Epling (1942), to warrant its descrip- tion. Its principal differentiating characteristics are: (1) its stipitate-glandular calyx teeth, the glands large, capitate, on stipes of varying length; in Monarda fistulosa var. mol- lis the calyx teeth vary from eglandular to sessile-gland- ular; (2) it has minute stipitate-glandular hairs on the external surface of the calyx tube: in M. fistulosa var. mol- lis the calyx tube varies from nearly glabrous to abundantly vestite with small, spreading or curled hairs, sometimes with sessile glands intermixed ; (3) the vestiture of the bracts may also include short, stipitate-glandular hairs, as there is often an outer, outwardly reflexed circle of hairs; (5) the corolla is white; in our M. jistulosa var. mollis it is lavender, Sometimes a lighter color, 502 1970] Monarda, New Species — Waterfall 503 The lower lip of the corolla is down-curved, with an abrupt, upward-turned apical tip 3-4 mm long; the style extends 2-3 mm beyond the tip of the upper lip; the fruit is oblong-ovate to oblong, 1.3-1.8 mm long, light brown in color. The original collection, from shallow, stony soil, was of rather dwarf plants, as compared with M. fistulosa var. mollis, the plants being 20-25 em tall. The garden-raised ones were 30-40 cm tall, growing larger under better con- ditions, but still much shorter than M. fistulosa var. mollis native in the same area. The plants began to flower May 18th, about 3-4 weeks before M. fistulosa var. mollis begins to flower in the same vicinity. M. stipitatoglandulosa Waterfall, sp. nov. Planta erecta, rhizomotosa, nana, 20-40 cm alta; caulibus minute curvi- pilis; foliis lanceolatis, vel angustolanceolatis, paucidenta- tis; petiolis 2-15 mm longis ; bracteis stipitatoglandulosis et trichomatibus longis; calycibus tubularibus minute stipita- toglandulosis ; corollis plus minusve faleatis, albis, 25-3 mm longis; labellis infernis ca 12-15 mm longis; labellis supernis 12-14 mm longis ; stylis exsertis 2-9 mm ; fructibus oblongis-ovatis vel oblongis, 1.3-1.8 mm longis, pallide brun- neis. Holotypus: Waterfall 17478, raised in garden from plants collected in an opening on a wooded mountain top, 7 miles north and 2.7 east of Broken Bow, McCurtain County, May 22. 1970 (OKLA). Isotypes will be distributed. A second collection referred to M. stipitatoglandulosa is Harry Love 159; rocky, oak-covered hillside, 1 mile south of Cloudy, Pushmataha County, June 19, 1966 (OKLA). The finding of this species in MeCurtain and Pushmataha counties is reminiscent of two other similarly endemic spe- cies, Streptanthus squamiformis Goodman (1956), also de- scribed from Beaver Bend State Park, and the unique Carex latebracteata Waterfall (1954), described from a few miles north of the park, but also found in the park, and later discovered in adjacent LeFlore County (Means, 1969). 504 Rhodora [Vol. 72 DEPARTMENT OF BOTANY AND PLANT PATHOLOGY OKLAHOMA STATE UNIVERSITY STILLWATER, OKLAHOMA 74074 LITERATURE CITED GOODMAN, G. J. A New Species of Streptanthus. Rhodora 58: 354, 355. 1956. McCLiNTOCK, E. & C. EPLING. A Review of the Genus Monarda. Univ. of Calif, Publ. Bot. 20: 149-194, 1942. MEANS, H. F., JR. Vascular Plants of Southeastern Oklahoma from the San Bois to the Kiamichi Mts. Unpublished Ph.D. thesis, Okla. State Univ. 1969. WATERFALL, U. T. A New Species of Carex (Section Phyllostachyae) from Oklahoma, Rhodora 56: 21-23. 1954. THE FLORA OF ISLE AU HAUT, MAINE! DAVID A. WISE Isle au Haut, in the outermost part of Penobscot Bay, Maine, covers an area of over seven square miles, or 4700 acres, 2800 of which are in Acadia National Park. Though first discovered by Champlain in 1604, who originally named it “Isle Haute", the island was not permanently set- tled until 1792. By 1864 the population had risen to around 300 settlers, most of whom engaged in fishing and small scale farming. Today the island’s population is seasonal, rising to a few hundred during the summer months, with only twenty or thirty permanent residents remaining through the winter. The general topographic aspect of the island is domi- nated by a central north-south ridge of hills. Of these the most prominent is Champlain Mountain (543 feet). Bogs are frequent in lowland areas with none of any consequence occurring above 200 feet. The interior of the island is drained by a number of small seasonal streams, and one relatively large body of freshwater, Long Pond, is located at the southeastern end of the island. The Isle au Haut shoreline is typical of much of the “drowned coast” of the Penobscot Bay region, For the most part it is highly irregular, rocky, and steep with pronounced headlands present on Eastern Head, Western Head, and around Duck Harbor. The beaches which do occur are ‘I am especially indebted to Dr. A. R. Hodgdon of the University of New Hampshire Botany Department for his continuous encourage- ment, assistance, and patience throughout the preparation of this paper and the Master's Thesis preceding it. In addition I would like to thank the administration of Acadia National Park for their kind cooperation and the National Science Foundation which provided most of the support for this work in the form of a National Science Founda- tion Graduate Traineeship. I am also most grateful to all my friends on Isle au Haut; their hospitality and friendship made my work a great deal more enjoyable. 505 506 Rhodora [Vol. 72 usually boulder-strewn and gravelly. Salt marshes occupy the shores of some of the coves, Isle au Haut lies within the central and eastern sector of the coastal climatic section of Maine as described by Fobes (1946). In many ways this area is typical of any moist, north temperate region and is characterized by warm days and cool nights during the summer months. As usual in coastal areas, the sea exerts an important mod- erating influence on the island's climate, making it gen- erally more equable than that of inland areas, This has the effect of reducing the summer and winter extremes of temperature. Consequently the growing season is prolonged. This prolongation of the growing season on the one hand creates a climate suitable for certain plants of southern affinities. On the other hand, the frequency of fogs together with the effects of cold sea water buffeting exposed shores and headlands create conditions similar to those in more northern regions, These exposed shores provide habitats for plants of more boreal and arctic areas. In the early decades of this century a number of well known botanists visited Isle au Haut. Mr. Nathaniel T. Kidder of Milton, Massachusetts, collected widely on the island roughly from 1900 to 1933. His notes and specimens are deposited in the library and herbarium of the New England Botanical Club. The New England Botanical Club Herbarium also contains specimens collected by Dr. Albert F. Hil! who made an extensive botanical survey of Isle au Haut from 1913 to 1919 in connection with the preparation of his Vascular Flora of the Penobscot Bay Region (Hill, 1919). In addition, Miss Kate Furbish of Brunswick, Maine, has left us a smaller, though valuable, collection of Isle au Haut plants dating from 1909. No comprehensive investigation of the flora of Isle au Haut was made until the writer's The Vascular Flora of Isle au Haut, Knox County, Maine (University of New Hampshire M. S. Thesis, unpubl.). This flora contained an annotated list of 724 taxa of vascular plants known to have occurred on Isle au Haut. It was based on collections made 1970] Isle Au Haut — Wise 507 by the writer in 1968 and 1969 all of which are deposited in the herbarium of the University of New Hampshire, and on previous collections if corroborated by herbarium speci- mens. The following list includes thirty-one additional taxa. These records are based on specimens recently found in a collection of Isle au Haut plants made by Nathaniel T. Kidder. This Kidder collection is a small portion of the Herbarium of the Portland Society of Natural History re- cently acquired on an extended loan basis by the University of New Hampshire Herbarium (NHA). With a few exceptions, the nomenclature of the Spermato- phytes employed in the following list essentially is that of Gray’s Manual of Botany (8th ed., 1950). Pteridophyte nomenclature, however, follows Seymour (1969). LIST OF PLANTS OF ISLE AU HAUT! *Equisetum arvense L., var. arvense E. sylvaticum L., var. pauciramosum Milde E. sylvaticum L., var. pauciramosum Milde, forma mul- tiramosum Fern. Lycopodium Selago L., var. appressum Desv. L. lucidulum Michx., var. lucidulum L. lucidulum Michx., var. occidentale (Clute) L. R. Wilson L. inundatum L. *L. annotinum L., var. annotinum L. annotinum L., var. acrifolium Fern, *L. clavatum L., var. clavatum L. clavatum L., var. megastachyon Fern. & Bissell *L. obscurum L., forma obscurum *L. obscurum L., forma dendroideum (Michx.) Blomq. & Corr. "Those taxa preceded by an asterisk (*) are known to have oc- curred on Isle au Haut in 1968-69. Those marked by two asterisks (**) were not included in Wise, 1970 (unpubl.) and represent the thirty-one additional Kidder specimens recently discovered in the Port- land Society of Natural History collection. Taxa preceded by a plus sign (+) were collected on Isle au Haut in 1968-69, and are not known to have been collected there previously. 508 Rhodora [Vol. 72 L. complanatum L., var. flabelliforme Fern. L. tristachyum Pursh Selaginella rupestris (L.) Spring Isoétes muricata Dur. Botrychium multifidum (Gmel) Rupr., var. interme- dium (D. C. Eat.) Farw. B. dissectum Spreng., forma dissectum B. dissectum Spreng., forma obliquum (Muhl.) Fern. +B. matricariaefolium A. Br. *Osmunda regalis L., var. spectabilis (Willd.) Gray *O. Claytoniana L. *O. cinnamomea L. Cystopteris fragilis (L.) Bernh., var. Mackayii Lawson *Onoclea sensibilis L. + Thelypteris palustris (Salisb.) Schott, var. pubescens (Lawson) Fern. +T. simulata. (Davenp.) Nieuwl. *T. noveboracensis (L.) Nieuwl. *T. Phegopteris (L.) Slosson *Gymnocarpium Dryopteris (L.) Newm. *Dryopteris spinulosa (O. F. Muell.) Watt. var. inter- media (Muhl.) Underw. *D. spinulosa (O. F. Muell) Watt, var. americana (Fisch.) Fern. *D. cristata (L.) Gray *D. marginalis (L.) Gray Polystichum acrostichoides (Michx.) Schott * Dennstaedtia punctilobula (Michx.) Moore Athyrium Filix-femina (L.) Roth, var. Michauzii (Spreng.) Farw. A. Filix-femina (L.) Roth, var. Michauxii (Spreng.) Farw., forma elatius (Link) Clute +A. Filix-femina (L.) Roth, var. Michauxii (Spreng.) Farw., forma rubellum (Gilbert) Farw. *Pteridium aquilinum | (L.) Kuhn, var. latiusculum (Desv.) Underw. * Polypodium virginianum L. Taxus canadensis Marsh. 1970] Isle Au Haut — Wise 509 * Abies balsamea (L.) Mill. ** Abies balsamea (L.) Mill., var. phanerolepis Fern. *Tsuga canadensis (L.) Carr. *Picea glauca (Moench) Voss *P. rubens Sarg. *P. mariana (Mill. BSP *Larix laricina (DuRoi) K. Koch * Pinus Strobus L. *P. resinosa Ait. *P. rigida Mill. *Thuia occidentalis L. * Juniperus communis L., var. depressa, Pursh *J. horizontalis Moench *Typha latifolia L. *Sparganium americanum Nutt. S. chlorocarpum Rydb., var. chlorocarpum *S. chlorocarpum Rydb., var. acaule (Beeby) Fern. S. angustifolium Michx. S. minimum (Hartm.) Fries Zostera marina L., var. stenophylla Aschers. & Graebn. *Potamogeton epihydrus Raf. var. Nuttallii (C. & S.) Fern. P. Oakesianus Robbins. *Triglochin maritima L. T. palustris L. Alisma triviale Pursh Sagittaria graminea Michx. *Festuca rubra L., var. rubra F. rubra L., var. rubra, forma megastachys (Gaudin) Holmb. F. elatior L. +F. elatior L., forma aristata Holmb. Puccinellia maritima (Huds.) Parl. *P. paupercula (Holm.) Fern. & Weath., var. alaskana (Scribn. & Merr.) Fern. & Wieg. *Glyceria canadensis (Michx.) Trin. *G. laxa Scribn. *G. striata (Lam.) Hitche., var. striata 510 Rhodora [Vol. 72 G. striata (Lam.) Hitche., var. stricta (Scribn.) Fern. *Poa annua L. *P. compressa L. *P. pratensis L. P. nemoralis L. *Dactylis glomerata L., var. glomerata D. glomerata L., var. ciliata Peterm. * Molinia caerulea. (L.) Moench Cynosurus cristatus L. + Agropyron trachycaulum (Link) Malte, var. majus (Vasey) Fern. A. trachycaulum (Link) Malte, var. novae-angliae (Scribn.) Fern. *A. repens (L.) Beauv., var. repens A. repens (L.) Beauv., var. repens, forma aristatum (Schum.) Holmb. A. repens (L.) Beauv., var. repens, forma trichorrachis Rohlena A. repens (L.) Beauv. var. repens, forma pilosum (Scribn.) Fern. ^ Lolium multiflorum Lam. Hordeum jubatum L. H. vulgare L, **Triticum aestivum L. Secale cereale L. *Elymus arenarius L., var. villosus Mey. *E. virginicus L., var. virginieus E. virginicus L., var. virginicus, forma hirsutiglumis (Seribn.) Fern. Avena. sativa L. *Deschampsia flexuosa (L.) Trin., var. flexuosa +D. flexuosa (L.) Trin., var. flexuosa, forma flavescens Sylven +D. caespitosa (L.) Beauv., var. parviflora (Thuill.) Richter “Danthonia spicata (L.) Beauv., var. spicata D. compressa Aust. 1970] Isle Au Haut — Wise 511 *Calamagrostis canadensis (Michx.) Nutt., var. canaden- sis Agrostis alba I., var. alba A. alba L., var. palustris (Huds.) Pers. * A. tenuis Sibth. * 4. scabra, Willd. A. perennans (Walt.) Tuckerm. Cinna latifolia (Trev.) Griseb. *Phleum pratense L. * Alopecurus pratensis L. * Muhlenbergia uniflora (Muhl.) Fern. + Brachyelytrum erectum. (Schreb.) Beauv., var. erectum B. erectum (Schreb.) Beauv., var. septentrionale Babel Oryzopsis asperifolia Michx. *O. canadensis (Poir.) Torr. *Spartina pectinata Link *S. alterniflora Loisel. *S. patens (Ait.) Muhl. *S. X caespitosa A. A. Eat. * Anthoxanthum odoratum L. * Leersia oryzoides (L.) Sw. Panicum capillare L., var. occidentale Rydb. *P. boreale Nash P. lanuginosum Ell., var. implicatum (Scribn.) Fern. +P. lanuginosum Ell., var. septentrionale Fern. +P. subvillosum Ashe Echinochloa crusgalli (L.) Beauv. *Setaria glauca (L.) Beauv. S. viridis (L.) Beauv. *Dulichium arundinaceum (L.) Britt. Eleocharis parvula (R. & S.) Link E. obtusa (Willd.) Schultes *E. halophila Fern. & Brack. *E. tenuis (Willd.) Schultes *?E. elliptica Kunth. + Scirpus cespitosus L., var. callosus Bigel. S. americanus Pers. S. acutus Muhl. 512 *g. *g. *g. * 1 dd Rhodora [Vol. 72 paludosus Nels., var. atlanticus Fern. rubrotinctus Fern. atrovirens Willd., var. georgianus (Harper) Fern. cyperinus (L.) Kunth, var. cyperinus cyperinus (L.) Kunth, var. pelius Fern. cyperinus (L.) Kunth, var. pelius Fern., forma con- densatus (Fern.) S. F. Blake pedicellatus Fern. . atrocinctus Fern, Eriophorum spissum Fern. *E. tencllum Nutt. *E. angustifolium Honckeny *E. virginicum L. *Rhynchospora alba (L.) Vahl Carex stipata Muhl. % P x x s x ^o eae casesassassas x 4 7 y. EE PAAAASS trisperma Dew., var. trisperma trisperma Dew., var. Billingsti Knight . Mackenziei recs, canescens L., var. disjuncta Fern. canescens L., var. subloliacea Laestad. brunnescens (Pers.) Poir, var. sphaerostachya (Tuckerm.) Kukenth. Deweyana Schwein. exilis Dew, atlantica Bailey cephalantha (Bailey) Bickn. angustior Mackenz. scoparia Schkuhr, var. scoparia Crawfordii Fern. tenera Dew. cumulata (Bailey) Mackenz. hormathodes Fern. hormathodes Fern., forma invisa (W. Boott) Fern. adusta Boott argyrantha 'Tuckerm. aenea Fern. leptalea Wahlenb, communis Bailey 1970] Isle Au Haut — Wise 513 *C. Emmonsii Dew. **C. artitecta Mackenz. C. novae-angliae Schwein. C. deflexa Hornem. +C. umbellata Schkuhr. C. paleacea Wahlenb. C. paleacea Wahlenb., forma erectiuscula Fern. *C. nigra (L.) Reichard, var. nigra C. nigra (L.) Reichard, var. strictiformis (Bailey) Fern. *C. stricta Lam., var. stricta *C. paupercula Michx., var. irrigua (Wahlenb.) Fern. C. Houghtonii Torr. *C. pallescens L., var. neogaea Fern. *C. arctata Boott *C. debilis Michx., var. Rudgei Bailey C. conoidea Schkuhr *C. panicea L. +C. laxiflora Lam. *C. ormostachya Wieg. C. leptonervia Fern. C. flava L. *C. viridula Michx. C. pauciflora Lightf. *C. lurida Wahlenb. *C. folliculata L. C. intumescens Rudge, var. Fernaldii Bailey *C. rostrata Stokes Arisaema Stewardsonii Britt. **Calla palustris L. *Symplocarpus foetidus (L.) Nutt. Lemna trisulca L. * Eriocaulon septangulare With. *Juncus bufonius L., var. bufonius *J. Gerardi Loisel., var. Gerardi *J. tenuis Willd., var. tenuis xJ. Greenei Oakes & Tuckerm. *J. effusus L., var. compactus Lej. & Court. 514 Rhodora [Vol. 72 J. effusus L., var. decipiens Buchenau *J. effusus L., var. Pylaei *J. balticus Willd., var. littoralis Engelm. *J. canadensis J. Gay, var. canadensis *J. brevicaudatus (Engelm.) Fern. J. militaris Bigel. *J. articulatus L., var. articulatus J. pelocarpus Mey. * Luzula acuminata Raf. *L. multiflora (Retz.) Lejeune, var. multiflora *L. multiflora (Retz.) Lejeune, var. fusconigra Celak. **L. multiflora (Retz.) Lejeune, var. acadiensis Fern. * Allium Schoenoprasum L., var. sibiricum (L.) Hartm. Hemerocallis fulva L. *Clintonia borealis (Ait.) Raf. Smilacina racemosa (L.) Desf. S. trifolia (L.) Desf. *Maianthemum canadense Desf. *Streptopus roseus Michx., var. perspectus Fassett *Polygonatum pubescens (Willd.) Pursh Medeola virginiana L. Trillium undulatum Willd. *Sisyrinchium montanum Greene, var. crebrum Fern. Iris Hookeri Penny *T, versicolor L. *Cypripedium acaule Ait. *Habenaria clavellata (Michx.) Spreng., var, ophioglos- soides Fern. orbiculata (Pursh) Torr. macrophylla Goldie obtusata (Pursh) Richards lacera (Michx.) R. Br, psyeodes (L.) Sw. fimbriata (Ait.) R. Br. * Pogonia ophioglossoides (L.) Ker *Calopogon pulchellus (Salisb.) R. Br. Arethusa bulbosa L. ** Spiranthes gracilis (Bigel.) Beck pe iq É q a B: 1970] Isle Au Haut — Wise 515 *S. cernua (L.) Richard, var. cernua *S. cernua (L.) Richard, var. ochroleuca (Rydb.) Ames S. Romanzoffiana Cham. *Goodyera repens (L.) R. Br., var. ophioides Fern. Corallorhiza trifida Chatelain, var. verna (Nutt.) Fern. *C. maculata Raf. * Malaxis unifolia Michx. Liparis Loeselii (L.) Richard Salix lucida Muhl., var. lucida alba L., var. vitellina (L.) Stokes rigida Muhl. Bebbiana Sarg. discolor Muhl. humilis Marsh, gracilis Anderss., var. textoris Fern. coactilis Fern. *Populus tremuloides Michx., var. tremuloides *P. tremuloides Michx., var. magnifica Vict. *P. grandidentata Michx. P. balsamifera L. *P. X gileadensis Rouleau *Myrica Gale L., var. subglabra (Chev.) Fern. *M. pensylvanica Loisel. *Comptonia peregrina (L.) Coult. *Ostrya virginiana. (Mill.) K. Koch “Betula alleghaniensis Britt. (= B. lutea Michx. f. of Gray’s Manual) *B. populifolia Marsh. B. caerulea-grandis Blanch. *B. populifolia Marsh. X B. papyrifera Marsh., var. papyrifera *B. papyrifera Marsh., var. papyrifera *B. papyrifera Marsh., var. cordifolia (Regel) Fern. * Alnus crispa (Ait.) Pursh, var. mollis Fern. A. rugosa (DuRoi) Spreng., var. rugosa *A. rugosa (DuRoi) Spreng., var. americana (Regel) Fern. *Fagus grandifolia Ehrh. ta ta tà ta TA to ta 516 Rhodora [Vol. 72 *Quercus rubra L. Humulus Lupulus L. ** Urtica, viridis Rydb. *U. dioica L. U. urens L. *Arceuthobium pusillum Peck Rumex altissimus Wood. R. pallidus Bigel. R. domesticus Hartm. R. orbiculatus Gray *R. crispus L. *R. Acetosella L., var. Acetosella *Polygonum aviculare L., var. vegetum Ledeb. *P. aviculare L., var. littorale (Link) W. D. J. Koch P. lapathifolium L. P. scabrum Moench P. Careyi Olney *P. Hydropiper L. *P. Persicaria L. P. punctatum Ell. *P. sagittatum L. *P. cilinode Michx. P. cilinode Michx., forma erectum (Peck) Fern. **P. Convolvulus L., var. Convolvulus P. Convolvulus L., var. subalatum Lej. & Court. Fagopyrum sagittatum Gilib. *Chenopodium album L. * Atriplex patula L., var. patula *A. patula L., var. hastata (L.) Gray *A. patula L., var. littoralis (L.) Gray +A. glabriuscula Edmondston * Salicornia europaea L., var. europaea S. europaea L., var. prostrata (Pall.) Fern. *Suaeda maritima (L.) Dumort. +S. linearis (Ell.) Moq. *Salsola Kali L., var. Kali Amaranthus retroflexus L. A. albus L. 1970] Isle Au Haut — Wise 517 Portulaca oleracea L. Montia lamprosperma Cham. *Spergularia rubra. (L.) J. & C. Presl. S. marina (L.) Griseb., var. leiospermo (Kindb.) Gurke *S. canadensis (Pers.) Don *Spergula arvensis L. *Sagina procumbens L. S. nodosa (L.) Fenzl., var. pubescens Mert. & Koch * Arenaria lateriflora L. A. peploides L., var. robusta Fern. * A. groenlandica (Retz.) Spreng. Stellaria media (L.) Cyrillo S. graminea. L. S. calycantha (Ledeb.) Bong., var. isophylla Fern. *S. calycantha (Ledeb.) Bong., var. floribunda Fern. *Cerastium vulgatum L. C. arvense L. +C. tomentosum L. ** Lychnis alba Mill. ** Silene Armeria L. S. noctiflora L. Saponaria officinalis L. *Dianthus deltoides L. D. plumarius L. *Nuphar variegatum Engelm. *Nymphaea odorata Ait. *Ranunculus Cymbalaria Pursh R. repens L. *R. acris L., var. acris *R. bulbosus L. Thalictrum polygamum Muhl. Clematis virginiana L. *Coptis groenlandica (Oeder) Fern. Aquilegia vulgaris L. Actaea rubra (Ait.) Willd. * A. pachypoda Ell. *Berberis Thunbergii DC *Corydalis sempervirens (L.) Pers. 518 Rhodora [Vol. 72 *Thlaspi arvense L. *Iberis umbellata L. Lepidiwm campestre (L.) R. Br. L. densiflorum Schrad. *Capsella Bursa-pastoris (L.) Medic. *Cakile edentula (Bigel.) Hook. Raphanus Raphanistrum L. **R, sativus L. Brassica Kaber (DC.) L. C. Wheeler, var. pinnatifida (Stokes) L. C. Wheeler B. juncea (L.) Coss. *B. nigra (L.) Koch B. Rapa L. *Sisymbrium officinale (L.) Scop., var. leiocarpum DC. S. altissimum L. Hesperis matronalis L. Erysimum cheiranthoides L. Rorippa sylvestris (L.) Bess. R. islandica (Oeder) Borbas, var. Fernaldiana Butt. & Abbe Armoracia lapathifolia Gilib. * Barbarea vulgaris R. Br., var. arcuata (Opiz) Fries B. vulgaris R. Br., var. brachycarpa Rouy & Foucaud Cardamine pensylvanica Muhl. **C. parviflora L., var. arenicola (Britt.) O. E. Schulz *Sarracenia purpurea. L. *Drosera intermedia Hayne *D. rotundifolia L., var. rotundifolia ** Sedum purpureum (L.) Link S. Rosea (L.) Scop. ** Sempervivum tectorum L. *Ribes hirtellum Michx., var. hirtellum *R. hirtellum Michx., var. calcicola Fern. R. lacustre (Pers.) Poir. *R. glandulosum Grauer R. triste Pall. R. sativum Syme *Hamamelis virginiana L., var. virginiana 1970] Isle Au Haut — Wise 519 * Spiraea latifolia (Ait.) Borkh. *S. tomentosa L. +S. japonica L.f., cv. Anthony Waterer *Pyrus Malus L. *P. floribunda Lindl. *P. melanocarpa (Michx.) Willd. *P. americana (Marsh.) DC. *P. decora (Sarg.) Hyland *P. Aucuparia (L.) Gaertn. Amelanchier stolonifera Wieg. ** A, canadensis (L.) Medic. A. arborea (Michx. f.) Fern. *A. laevis Wieg. ** A. laevis Wieg. X A. canadensis (L.) Medic. A. Bartramiana (Tausch) Roemer X A. laevis Wieg. + Crataegus chrysocarpa Ashe *C. Jonesae Sarg. *C. macrosperma Ashe *C. succulenta Link * Fragaria virginiana Duchesne, var. virginiana F. virginiana Duchesne, var. terrae-novae (Rydb.) Fern. & Wieg. F. X Ananassa Duchesne *Potentilla tridentata Ait. P. tridentata Ait., forma aurora Graustein P. palustris (L.) Scop., var. villosa (Pers.) Lehm. P. argentea L. P. pectinata Raf. *P. norvegica L. P. canadensis L. *P. simplex Michx., var. simplex P. simplex Michx., var. calvescens Fern. *P. Egedei Wormsk., var. groenlandica (Tratt.) Polunin * Filipendula Ulmaria (L.) Maxim. Geum aleppicum Jacq., var. strictum (Ait.) Fern. Rubus pubescens Raf. ** Rubus idaeus L., var. ¿idaeus R. idaeus L., var. strigosus (Michx.) Maxim. 520 Rhodora R. idaeus L., var. heterolasius Fern. *R. idaeus L., var. canadensis Richards. R. recurvicaulis Blanch. R. recurvicaulis Blanch. X R. setosus Bigel. *R. hispidus L., var. hispidus R. setosus Bigel. X R. allegheniensis Porter + ?R. semisetosus Blanch. R. canadensis L. *R. allegheniensis Porter, var. allegheniensis R. frondosus Bigel. R. pensilvanicus Poir. Dalibarda repens L. *Rosa nitida Willd. *R. virginiana Mill. ** R. virginiana Mill. X R. nitida Willd. *R. rugosa Thunb. ** R. cinnamomea L. Prunus maritima L. *P. pensylvanica L. f. **P. Avium L. *P. virginiana L. Lupinus polyphyllus Lindl. Trifolium arvense L. T. pratense L. +T. pratense L., forma pilosum (Griseb.) Hayek *T. repens L. T. hybridum L., var. elegans (Savi) Boiss. *T. agrarium L. *T. procumbens L. Medicago lupulina L. [Vol. 72 Vicia angustifolia Reichard, var. segetalis (Thuill.) W. D. J. Koch *V. Cracea L. *Lathyrus japonicus Willd., var. glaber (Ser.) Fern. L. japonicus Willd., var. pellitus Fern. L. palustris L., var. pilosus (Cham.) Ledeb. Linum usitatissimum L. *Oxalis montana Raf. 1970] Isle Au Haut — Wise 521 O. europaea Jord., var, europaea *O. europaea Jord., var. europaed, forma cymosu. (Small) Wieg. Geranium Bicknellii Britt. G. molle L. *G. Robertianum L. Polygala sanguinea L. *Euphorbia Cyparissias L. E. Peplus L. *Empetrum nigrum L. *Corema Conradii Torr. *Rhus typhina L. *R. radicans L., var. radicans *Ilex verticillata (L.) Gray, var. verticillata *I. glabra (L.) Gray *Nemopanthus mucronata (L.) Trel. 4- Celastrus orbiculatus Thunb. * Acer spicatum Lam. *A. pensylvanicum L. *A. saccharum Marsh. *4. rubrum L., var. rubrum +A. rubrum L., var. rubrum, forma tomentosum (Desf.) Dansereau *A. rubrum L., var. trilobum K. Koch *Impatiens capensis Meerb. Parthenocissus quinquefolia (L.) Planch. +P. inserta, (Kerner) K. Fritsch ** Malwa neglecta Wallr. *Hypericum perforatum L. H. boreale (Britt.) Bickn. H. mutilum L., var. parviflorum (Willd.) Fern. H. majus (Gray) Britt. *H. canadense L. *H. dissimulatum Bickn. *H. gentianoides (L.) BSP H. virginicum L., var. virginicum *H. virginicum L., var. Fraseri (Spach) Fern. Elatine americana (Pursh) Arn. 522 Rhodora [Vol. 72 **Hudsonia ericoides L. H. tomentosa N utt., var. intermedia Peck *Lechea intermedia Leggett, var. intermedia L. intermedia Leggett, var. juniperina (Bickn.) Robins. *Viola cucullata Ait. *V. septentrionalis Greene V. fimbriatula Sm. *V. pallens (Banks) Brainerd V. incognita Brainerd, var. incognita t V. incognita Brainerd, var. Forbesii Brainerd V. blanda Willd. *V. lanceolata L. V. primulifolia L., var. acuta (Bigel.) T. & G. Decodon verticillatus (L.) Ell., var. laevigatus T. & G. * Epilobium angustifolium L. E. strictum Muhl. E. leptophyllum Raf., forma umbrosum (Haussk.) Fern. E. palustre L., var. oliganthum (Michx.) Fern. E. coloratum Biehler +E. glandulosum Lehm., var. adenocaulon (Haussk.) Fern. “Oenothera biennis L., var. biennis O. parviflora L. *O. perennis L. *Circaea alpina L. “Aralia hispida Vent. *A. nudicaulis L. Hydrocotyle americana L. Cicuta maculata L. +Carum Carvi L. *Ligusticum scothieum L. *Coelopleurum lucidum (L.) Fern. *Heracleum maximum Bartr. “Daucus Carota L. “Cornus canadensis L. C. stolonifera Michx. C. alternifolia L. f. 1970] Isle Au Haut — Wise 523 Chimaphila umbellata (L.) Bart., var. cisatlantica Blake *Moneses uniflora. (L.) Gray Pyrola secunda L. P. virens Schweigger, forma paucifolia Fern. *P. elliptica Nutt. P. rotundifolia L., var. americana (Sweet) Fern. * Monotropa uniflora L. M. Hypopithys L. *Ledum groenlandicum Oeder * Rhododendron canadense (L.) Torr. * Kalmia angustifolia L. K. polifolia Wang. *Cassandra calyculata (L.) D. Don, var. angustifolia (Ait.) Rehd. (= Chamaedaphne calyculata (L.) Moench, var. angustifolia (Ait.) Rehd. of Gray’s Manual). *Gaultheria procumbens L. *G. hispidula (L.) Bigel. * Arctostaphylos Uva-ursi (L.) Spreng. var. coactilis Fern, & Mack. *Gaylussacia dumosa (Andr.) T. & G., var. Bigeloviana Fern. +G. baccata (Wang.) K. Koch Vaccinium myrtilloides Michx. +V. angustifolium Ait., var. angustifolium V. angustifolium Ait., var. laevifolium House V. angustifolium Ait., var. nigrum (Wood) Dole V. corymbosum L., var. albiflorum (Hook.) Fern. *V. Vitis-Idaea L., var. minus Lodd. V. Oxycoccos L. *V. macrocarpon Ait. *Lysimachia quadrifolia L. L. X producta (Gray) Fern. *L. terrestris (L.) BSP, var. terrestris *Trientalis borealis Raf. *Glaux maritima L., var. maritima G. maritima L., var. obtusifolia Fern. *Limonium Nashii Small 524 Rhodora [Vol. 72 *Fraxinus americana L., var. americana F. pennsylvanica Marsh., var. subintegerrima (Vahl) Fern. *Syringa vulgaris L. * Ligustrum amurense Carr. Bartonia virginica (L.) BSP *B. paniculata (Michx.) Muhl., var. intermedia Fern. *Nymphoides cordata (Ell.) Fern. * Apocynum androsaemifolium L. *Asclepias incarnata L., var. pulchra (Ehrh.) Pers. “Convolvulus sepium L., var. sepium *Cuscuta Gronovii Willd. Myosotis sylvatica Hoffm. M. arvensis (L.) Hill *Mertensia maritima (L.) S. F. Gray Verbena hastata L. “Teucrium occidentale Gray, var. boreale (Bickn.) Fern. Scutellaria lateriflora L. *S. epilobiifolia A. Hamilton Nepeta Cataria L. Glechoma hederacea L., var. hederacea *G. hederacea L., var. micrantha Moricand *Prunella vulgaris L., var. lanceolata (Bart.) Fern. P. vulgaris L., var. lanceolata (Bart.) Fern., forma iodocalyx Fern. Leonurus Cardiaca L. “Galeopsis Tetrahit L., var. bifida (Boenn.) Lej. & Court. “Hedeoma pulegioides (L.) Pers. * Lycopus uniflorus Michx. L. americanus Muhl. Mentha piperita L. M. cardiaca Baker M. arvensis L., var. arvensis *M. arvensis L., var. villosa (Benth.) S. R. Stewart *Solanum Dulcamara L. **S. nigrum L. S. rostratum Dunal *Verbascum Thapsus L. 1970] Isle Au Haut — Wise 525 Linaria vulgaris Hill *L. canadensis (L.) Dumont Scrophularia lanceolata Pursh Veronica serpyllifolia L. *V. officinalis L. V. peregrina L., var. xalapensis (HBK.) St. John & Warren V. arvensis L. *Melampyrum lineare Desr., var. lineare M. lineare Desr., var. americanum (Michx.) Beauverd Euphrasia Randii Robins. E. canadensis Townsend Odontites serotina (Lam.) Dum. *Rhinanthus Crista-galli L., var. Crista-galli Epifagus virginiana (L.) Bart. * Utricularia geminiscapa Benj. U. cornuta Michx. *Plantago major L., var. major P. major L., var. Pilgeri Domin *P. juncoides Lam., var. decipiens (Barneoud) Fern. *P. oliganthos R. & S. **P. lanceolata L. P. altissima L. *Galium Aparine L. *G. triflorum Michx. G. verum L. *G. palustre L. G. trifidum L. *G. tinctorium L., var. tinctorium Mitchella repens L. *Houstonia caerulea L. *Diervilla Lonicera Mill. Lonicera villosa (Michx.) R. & S., var. Solonis (Eat.) Fern. L. villosa (Michx.) R. & S., var. calvescens (Fern. & Wieg.) Fern. *L. canadensis Bartr. + Symphoricarpos albus (L.) Blake, var. albus 526 Rhodora [Vol. 72 S. albus (L.) Blake, var. laevigatus (Fern.) Blake *Linnaea borealis L., var. americana, (Forbes) Rehd. *Viburnum cassinoides L. *V. recognitum Fern. “Sambucus canadensis L. *S. pubens Michx. Echinocystis lobata (Michx.) T. & G. Campanula rapunculoides L. *C. rotundifolia L. *Lobelia. inflata L. *L. Dortmanna L. * Eupatorium perfoliatum L., var. perfoliatum *Solidago bicolor L. *S. puberula Nutt. *S. sempervirens L. S. juncea Ait. *S. uliginosa N utt., var. linoides (T. & G.) Fern. S. uliginosa N utt., var. terrae-novae (T. & G.) Fern. *S. nemoralis Ait. *S. rugosa. Ait., var. villosa (Pursh) Fern. *S. canadensis L. *S. graminifolia (L.) Salisb, var. Nuttallii (Greene) Fern. Aster macrophyllus L., var. macrophyllus A. macrophyllus L., var. velutinus Burgess A. cordifolius L., var. cordifolius A. polycephalus Porter A. radula Ait., var. radula A. radula Ait., var. strictus (Pursh) Gray A. lateriflorus (L.) Britt. A. lateriflorus (L.) Britt., var. tenuipes Wieg. A. simplex Willd., var. ramosissimus (T. & G.) Cronq. +A. foliaceus L. *A. johannensis Fern. "A. novi-belgii L., var. novi-belgii *A. nemoralis Ait. "A. X Blakei (Porter) House *A. acuminatus Michx. 1970] Isle Au Haut — Wise 527 * A. umbellatus Mill. *Erigeron strigosus Muhl., var. strigosus E. canadensis L. * Antennaria canadensis Greene A. neglecta Greene A. neodioica Greene A. munda Fern. * A. fallax Greene A. Brainerdii Fern. + Anaphalis margaritacea (L.) C. B. Clarke, var. mar- garitacea +A. margaritacea (L.) C. B. Clarke, var. intercedens Hara Gnaphalium obtusifolium L. *G. uliginosum L. + Ambrosia, artemisiifolia L., var. artemisiifolia A. artemisiifolia L., var. elatior (L.) Descourtils ** A. artemisiifolia L., var. elatior (L.) Descourtils, forma villosa Fern. and Grisc. * Rudbeckia serotina Nutt. * Bidens frondosa L. B. frondosa L., forma anomala (Porter) Fern. Galinsoga ciliata (Raf.) Blake + Achillea Millefolium L. Anthemis Cotula L. A. tinctoria L. Matricaria maritima L., var. maritima M. Chamomilla L. *M. matricarioides (Less.) Porter *Chrysanthemum Leucanthemum L., var. pinnatifidum Lecoq. & Lamotte C. Balsamita L. Tanacetum vulgare L. T. vulgare L., forma crispum (L.) Hayek Artemisia vulgare L. Petasites palmatus (Ait.) Gray *Erechtites hieracifolia (L.) Raf., var. intermedia Fern. *E. hieracifolia (L.) Raf., var. praealta (Raf.) Fern 528 Rhodora [Vol. 72 Senecio vulgaris L. *S. sylvaticus L. S. Robbinsii Oakes **Arctium nemorosum Lej. & Court. *A. minus (Hill) Bernh. Echinops sphaerocephalus L. *Cirsium vulgare (Savi) Tenore C. muticum Michx. *C. arvense (L.) Scop. ** Lapsana communis L. *Leontodon autumnalis L., var. autumnalis L. autumnalis L., var. pratensis (Link) W. D. J. Koch Taraxacum erythrospermum Andrz. *T. officinale Weber, var. officinale T. officinale Weber, var. palustre (Sm.) Blytt Sonchus arvensis L. *S. oleraceus L. *S. asper (L.) Hill Lactuca canadensis L., var. canadensis L. canadensis L., var. longifolia (Michx.) Farw. L. canadensis L., var. latifolia Ktze. L. biennis (Moench) Fern. *Prenanthes trifoliolata (Cass.) Fern., var. trifoliolata +P. trifoliolata (Cass.) Fern., var. nana (Bigel.) Fern. P. altissima L. P. altissima L., forma hispidula Fern. Hieracium Pilosella L. *H. aurantiacum L. *H. pratense Tausch H. florentinum All. H. canadense Michx., var. canadense H. paniculatum L. H. scabrum Michx. A close examination of the preceding list will reveal that the vascular flora of Isle au Haut is chiefly transitional in character and is generally typical of what has been termed the Canadian Transition Zone (Hill, 1923). This “zone” contains floristie elements from both the more northern 1970] Isle Au Haut — Wise 529 Boreal Region and the Alleghenian Region to the south. However, a few Carolinian plants of the southeastern coastal plain and a few Arctic plants are present on the island as well. These are of particular interest due to their markedly disjunct distributions. The following plants of Coastal Plain and Arctic phyto- geographic affinities were found to be present on Isle au Haut in 1968-69 and deserve special mention. Specimens are deposited in the herbarium of the University of New Hampshire (NHA). COASTAL PLAIN PLANTS Thelypteris simulata (Davenp.) Nieuwl. Massachusetts Fern. The Massachusetts Fern was found to be locally abundant in alder thickets at the edge of a bog at Moore’s Harbor on the western side of Isle au Haut. While found in south- western Nova Scotia the 1969 collections are the first Knox County records for this species and indicate a slight north- eastern extension of range in the United States from Lin- coln County, Maine. Corema Conradii Torr. Broom-Crowberry. This coastal plain relative of Empetrum was collected on Isle au Haut by Albert F. Hill in 1915. In 1968-69 it was frequently encountered on the rocky summits of Mt. Cham- plain and Black Dina. Ilex glabra (L.) Gray Inkberry. The inkberry was first collected on Isle au Haut by Nathaniel T. Kidder in 1922 (Kidder, 1923). It was found growing in the “Great Meadow" bog, and Mr. Kidder noted that it occurred in "considerable quantity, fifty plants or more". In 1969 the writer and Dr. A. R. Hodgdon found the plant still growing at the same site covering an area of about fifty by seventy-five feet. While quite abundant lo- cally, the plants were small, low growing, and depauperate in aspect. These collections from Isle au Haut represent 530 Rhodora [Vol. 72 the only known occurrence of Ilex glabra between Sea- brook, New Hampshire ( Hodgdon, 1955) and Nova Scotia (Kidder, 1923). Bartonia paniculata (Michx.) Muhl., var. intermedia Fern. In 1922 Kidder also collected this Coastal Plain plant in the same bog on Isle au Haut where he had found Ilex glabra. Though very infrequent in occurrence, it was still present in 1969. This station, therefore, remains the only one known for this variety of B. paniculata between Nova Scotia and the Blue Hills, Massachusetts ( Fernald, 1921; Kidder, 1923; Seymour, 1969). In addition to the above, Hudsonia ericoides L., a plant with a somewhat similar Coastal Plain distribution, was collected on Isle au Haut by Kidder in 1901 and 1926. The writer was unable to locate the collecting sites in 1968-69. Consequently the present occurrence of Hudsonia ericoides on Isle au Haut remains in doubt. ARCTIC PLANTS Scirpus cespitosus L., var. callosus Bigel. Not previously reported from Isle au Haut, this plant of Arctic affinity was found to be quite common in moist de- pressions on the top of Duck Harbor Mountain. Other taxa of similar phytogeographic relationship also frequent at the same site were: Arenaria groenlandica, Arctostaphylos Uva- ursi, and Empetrum nigrum. Mertensia maritima (L.) S. F. Gray This plant of Arctic affinity was collected previously on Isle au Haut by Kidder in 1926. Though still present in 1969, it was represented only by a few depauperate plants found along the shore of Barred Harbor. Three species of similar phytogeographic range: Iris Hookeri, Montia lamprosperma, and Sedum Rosea, though previously collected on Isle au Haut, were not found there in 1968-69. INTRODUCED PLANTS In addition to species having either Coastal Plain or 1970] Isle Au Haut — Wise 531 Arctic affinities, two introduced plants recently found on Isle au Haut deserve mention. Molinia caerulea (L.) Moench This European grass was first collected by Kidder on Point Lookout in 1918. Later collections were made by him at the same locality in 1919 and 1927. The writer and Dr. A. R. Hodgdon visited the site in 1969 and found the grass not only persisting but apparently spreading very success- fully. The only other known collections of Molinia in Maine were made by E. P. Bicknell at Mt. Desert, Bar Harbor, and York Harbor in 1897. Curiously all three collections were made on August 25 — an admirable though suspect feat considering that York Harbor is some 150 miles from Mt. Desert Island (Dix, 1945). Iberis umbellata L. Very close to the Molinia locality this plant was found growing in considerable abundance in a filled in cellar-hole. Apparently once cultivated it presently is spreading suc- cessfully as a weed. The only other known record of I. umbellata from New England is a specimen in the New England Botanical Club Herbarium collected by A. S. Pease in Randolph, New Hampshire, in 1916. DEPARTMENT OF BOTANY UNIVERSITY OF NEW HAMPSHIRE, DURHAM LITERATURE CITED Dix, W. L. 1943-44. Will the stowaway, Molinia caerulea, become naturalized? Bartonia, 23. FERNALD, M. L. 1921. The Gray Herbarium expedition to Nova Scotia, 1920. Rhodora 23: 284-300. 1950. Gray's Manual of Botany, Eighth Ed. Amer. Book Co., N.Y., 1632 pp. j FoBES, C. B. 1946. Climatic divisions of Maine. Bull. No. 40, Me. Tech. Expt. Sta., Univ. of Me., Orono. 44 pp. HiLL, A. F. 1919. Vascular Flora of the Penobscot Bay region. Proc. Port. Soc. Nat. Hist., vol. III, part 2. - 1923. The vegetation of the Penobscot Bay region, Maine, Proc. Port. Soc. Nat. Hist., vol. III, part 3. 532 Rhodora [Vol. 72 Hopepon, A. R. 1955. Ilex glabra and a new station for Kalmia latifolia in New Hampshire. Rhodora 57: 34-36. Kipper, N. T. 1923. Further notes on the plants of Isle au Haut. Rhodora 25: 147-148. Flora of Isle au Haut. Unpubl. notes, New Eng. Bot. Club Library. SEYMOUR, F. C. 1969. The flora of New England. Tuttle, Rutland, Vt. 596 pp. WisE, D. A. 1970. The vascular flora of Isle au Haut, Knox County, Maine. M. S. Thesis, Univ. N.H., Durham. Unpubl. MANUAL OF THE VASCULAR PLANTS OF TEXAS? This manual of the Texas flora is a monumental work and the authors, D. S. Correll and M. C. Johnston, as well as their collaborators, deserve the compliments of syste- matists on its publication. A definitive treatment of a large and complex flora (174 families, 1216 genera, 4839 species and 576 subspecies and varieties in Texas) must depend to a considerable extent upon the labors of predecessors. There has been a long and active series of collectors and publishers interested in Texas plants, and they have pro- vided an excellent basis for a comprehensive flora. How- ever, the special research on the Texas flora by the authors, and their extensive field experience with it, have brought to this publication an important element of personal knowl- edge. Fifty-six collaborators have provided the treatments of 20 families and of 38 separate genera. Both the authors and the collaborators are to be congratulated upon this productive cooperation. The book is well made up and the format and typography make it easy to use. An introduction provides a brief ac- count of the principal vegetation regions in Texas. This is followed by a key to the families and the taxonomic treat- ment of the flora, which encompasses 1700 pages. A brief appendix adds critical comments and some additional spe- cies from data obtained during the course of publication. The list of “Abbreviations of Authors’ Names" deserves special mention. It is in fact far more than the title states. It includes birth and death dates, and a variety of other information such as the principal professional position, or other occupation, major publications, groups or floras studied, and sometimes mention of other significant contri- butions to botany. The inclusion of this information will 'Correll, D. S. and M. C. Johnston. Manual of the vascular plants of Texas. xv + 1881 pp. frontispiece (the Texas Bluebonnet, Lupinus subcarnosus, in color), 3 maps, cloth. $30.00. (Contributions from the Texas Research Foundation, Edited by C. L. Lundell, vol. 6). Texas Research Foundation, Renner, Texas. 1970. 533 534 Rhodora [Vol. 72 remove (for many users of the Manual) the anonymity of the authors of plant names and it will relate the authors to their botanical work. It also provides an introduction to the historical basis of the Manual. A notable feature is the flexibility allowed in the tax- onomic treatments, This is a commendable policy and much more is gained by it than from the consistency resulting from a rigidly imposed format. Extended discussion is not possible in a large manual, but the authors effectively have utilized departures from the normal format and brief com- ments in difficult species-groups to point out uncertainties and problems that need further study. This flexibility also has allowed the collaborators to express preferences, for example, subspecies are used by Raven and varieties by Shinners ; hybrid formulas are used in Quercus by Muller; and Capparidaceae for the Caper Family by Iltis. The flora of Texas is dominated by four major floristic elements, The most important in terms of area and economy is that of the Prairies and Plains, predominantly a flora of Gramineae, Leguminosae and Compositae, Two impor- tant floras extend into Texas from the east, the Appala- chian-Ozark and the Coastal Plain. Together these provide the forests and woodlands of the eastern portion of the state that extend westward to the Cross-Timbers region. The Coastal Plain element also furnishes most of the spe- cies along the Gulf Coast. The Mexican flora is the most important in number of species and is of special interest because of its many endemics. This element dominates the mountains and arid valleys and basins of the Trans-Pecos region and is Strongly represented elsewhere along the Rio Grande and on the Edwards Plateau. Almost half of the Pteridophyta, for example, belong to this element, largely due to the many species of Pellaea, Cheilanthes and Notho- laena, Among other groups that show a similar geographic relation may be mentioned Ephedra and Muhlenbergia, and especially Acacia, Mimosa, Heliotropium and Baccharis. A review in Rhodora of a flora of another part of the United States invites Some comparison with the flora 1970] Vascular Plants — Tryon 535 treated in Gray’s Manual. In the 8th Edition by M. L. Fernald, the area covered is about four times as large as Texas (it requires several northeastern states to do this) ; there are similar numbers of genera (1133 to 1216 in Texas) and species (5525 to 4839 in Texas). The relatively small differences in numbers could easily be due to different taxonomic viewpoints, so the two floras might well be con- sidered nearly equivalent. There are, however, some sig- nificant differences. There are few large genera in Texas. Panicum with 68 species is the largest and only two others, Carex and Euphorbia have more than 50 species. Cyperus has 46 and there are eight other genera with more than 30 species. In the northeastern United States the largest genus is Carex with 267 species, and others such as Panicum, Juncus, Salix, Rubus, Crataegus, Viola, Solidago and Aster have over 50 species each. Genera with 30-50 species are Potamogeton, Cyperus, Eleocharis, Scirpus, Polygonum, Ranunculus, cte. Most of the larger genera have their pri- mary strength in aquatic or marsh habitats. These genera are naturally not as well represented in Texas. For some reason their place is not taken by equally large genera of xeric habitats, although these are not lacking in Mexico. The Manual of the Vascular Plants of Texas is a note- worthy achievement. It is a necessary guide to the Texas flora, but will be used as often beyond its geographic limits. It will certainly stimulate interest in Texas plants and further research on the rich Texas flora. ROLLA TRYON GRAY HERBARIUM HARVARD UNIVERSITY CAMBRIDGE, MASS. 02138 THE VARIETIES OF SILPHIUM INTEGRIFOLIUM WILBUR J. SETTLE AND T. RICHARD FISHER The genus Silphium ( Compositae) was described first by Linnaeus in 1753 in Species Plantarum which listed five species. Small (1933) recognized more than thirty species of Silphium. Perry (1937), who has written the only exten- Sive taxonomic treatment of the genus, recognized 23 spe- cies and several varieties, T. R. Fisher and some of his students currently recognize 16 species, All of the species of Silphium which have been studied are diploid (2n=14) (Figure 2). The plants reproduce sex- ually by outcrossing; selfing rarely occurs. Vegetative re- production occurs from the perennial rhizomes. Apomixis has not been reported for the genus. Flowering occurs from June to September. The genus occurs only in North America north of Mexico, ranging from Florida to Connecticut, northwest to the Dakotas, and southwest to Texas. Silphium is in the tribe Heliantheae which is characterized by radiate heads, her- baceous phyllaries, chaffy receptacles, and blunt anthers. Small (1933) divided the genus into five sections on the basis of phyllary pubescence, grouping of inflorescences, and leaf position. Section I, Perfoliata, contained the species with perfoliate leaves; sections II and III, Laciniata and Composita. respectively, contained the species with mostly basal leaves; sections IV and V, Dentata and Integrifolia respectively, contained the species with leafy stems. Sil- phium integrifolium is included in the section Integrifolia. As a part of a more extensive study of S. integrifolium, field collections and herbarium material from FSU, GA, GH, MO, ND, NY, PH, TENN, WIS, WVA, and US were studied. See Lanjouw and Stafleu (1964) for abbreviations. TAXONOMY Silphium integrifolium Michx. Fl. Bor. Am. 2: 146. 1803. (Holotype: Hab. in regione Illinoensi. A. Michaux, P). 536 1970] Silphium — Settle and Fisher Pulaski County Gry ridges. 4-5 ft. tall. Comnione po, Little Hock. Elevation tt 2754 Dste 7-221955 PET T BichARD tien. PLANTS OF ARKANSAS Collected by Delzie Demaree 537 Silphium integrifolium Michx. 37528 Figure 1. Photograph of Silphium integrifolium specimen. (Photo- graph by Wm. Anderson) 538 Rhodora [Vol. 72 Figure 2. Mitotic cell from root tip of Silphium integrifolium, 2n—14, Fisher 624. 1125 X. Silphium laevigatum Pursh, F. Am. Sept. 2: 578. 1814. (Lectotype: In Georgia, Fl. Am. Bor. A. Enslen, W!). Perennial, with short, thickened, sometimes creeping rhizome; stems to 1.5 m tall, pilose, scabrous, or almost glabrous; leaves lanceolate to elliptic, sessile, usually op- posite, toothed or entire, upper surface scabrous, lower pilose, scabrous, or glabrous; heads several in close corymb; peduncles short, much branched, with or without glandular- pubescence; phyllaries lance-ovate to elliptic, imbricate, pil- ose, hispid, or rarely glabrous, with or without glandular- 1970] Silphium — Settle and Fisher 539 pubescence; achenes obovate or orbicular-obovate, winged, toothed. Pursh’s description of S. laevigatum was based on plants collected by Aloysius Enslen in western Georgia. Pursh’s description and the type specimen are very similar to S. integrifolium. If the specimen was collected in western Georgia, it was not within the present known range of S. integrifolium. Because the present range extends eastward to central Alabama, a few plants might have grown in western Georgia at an earlier time, or even at the present time, escaping collection for some reason, The possibility also exists that the information on Enslen’s label is incor- rect. Silphium integrifolium is a rather homogeneous group of plants, morphologically. However, the most obvious and con- sistent differences are the kind and extent of vesture (vesti- ture) on the phyllaries and on the lower surface of the leaves. KEY TO VARIETIES A. Lower surface of leaves and phyllaries glabrous ............... Tet RA 2. var. gattingert A. Lower surface of leaves and/or phyllaries pubescent ... B B. Lower surface of leaves pilose (velvety) .................. T 3. var, deamii B. Lower surface of leaves hirsute, pubescent, or gla- brous C C. Phyllaries glandular-pubescent - 4. var. neglectum C. Phyllaries not glandular-pubescent hu aa DA a enn Le var, integrifolium 1. Silphium integrifolium Michx. var. integrifolium. Although Perry (1937) mentioned “typical S. integrifo- lium,” she naturally, at that time, did not refer to these plants as belonging to variety integrifolium. Cronquist (1945) called this group S. integrifolium var. integrifolium. C. A. Weatherby (1935, written communieation to C. C. 540 Rhodora [Vol. 72 Deam) examined the type sheet of S. integrifolium and found that the involucres are pustulate-scabrous or strigose, but not glandular. 2. Silphium integrifolium Michx. var. gattingeri Perry, Rhodora 39: 287, 288. 1937. (Holotype: Tennessee: Char- lotte Pike, Nashville, July 1886, Gattinger GH!). This variety has been found only in Davidson County, Tennessee. 3. Silphium integrifolium Michx. var. deamii Perry, Rho- dora 39: 287. 1937. (Holotype: Indiana: Vermillion County, right of way of the railroad 1 mile east of Dana. C. C. Deam 54376 GH!). Perry (1937) placed all of the plants of S. integrifolium having glandular-pubescent phyllaries into this variety. 4. Silphium integrifolium Michx. var. neglectum Settle and Fisher var. nov. (Holotype: Indiana: White County. T. R. Fisher 73, Bowling Green State University Herbarium!). Phyllaria cum pilis glandulosis, Pagina inferior folii glabra vel pubescens sed non pilosa. The plants of this variety have glandular-pubescent phyl- laries and leaves with hirsute, pubescent, or glabrous, but not pilose, lower surface, GEOGRAPHIC DISTRIBUTION The general distribution of Silphium integrifolium is ap- proximately the same as that of the tall grass prairie of the United States. The range of S. integrifolium. extends into eastern Indiana, west into Kansas, southwest into Texas, southeast into Alabama, and north into Wisconsin. Distribution of the varieties, compiled from field collections and from herbarium specimens, is shown in Figure 3. Silphium integrifolium and S. speciosum are sympatric in the Kansas-Oklahoma area, and S. integrifolium and S. scaberrimum are sympatric in the Oklahoma-Texas-Arkan- sas area. Naturally occurring putative hybrids between S. integrifolium and S. speciosum and between S. integri- folium and S. scaberrimum have been found. 1970] Silphium — Settle and Fisher 541 LE: X v? Silphium integrifolium ` af Var. integrifolium Oo Ver. deamii e \ Var. neglectum * + Ver, gattinger! Figure 3. Distribution of Silphium integrifolium and its varieties. 542 Rhodora [Vol. 72 The varieties (Figure 3) do not have distinct geographic distributions. The variety neglectum does have a restricted distribution, being restricted to the northeastern, glaciated portion of the range of the species. The variety deamii appears to be the most widespread geographically and also the most common of the varieties. CONCLUSIONS Three of the four varieties treated in this paper were recognized in Perry’s work published in 1937. However, Perry’s delimitation of varieties does not take into account a large portion of the plants which are distinctive because of the pilose pubescence of the lower surface of the leaves. This character is more striking and obvious than the glen- aular-pubescence of the phyllaries which Perry (1937) used as the principal basis for recognizing varieties. The vesture of the abaxial surface of the leaves, in addi- tion to the vesture of the phyllaries, seems to be a more satisfactory character upon which to base the varieties. Although S. integrifolium is a rather homogeneous group of plants, differences do exist which should be recognized. These differences are sufficient to merit recognition as separate varieties but not great enough to warrant Separat- ing the species into subspecies. Therefore, S. integrifolium should be considered to include four varieties: S. integrifo- hum var. integrifolium, S. integrifolium var, deamii Perry, S. integrifolium var. gattingeri Perry, and S. integrifolium var. neglectum Settle and Fisher. BOWLING GREEN STATE UNIVERSITY BOWLING GREEN, OHIO 43402 LITERATURE CITED CRONQUIST, A. 1945. Notes on the Compositae of northeastern United States. II. Heliantheae and Heleniae. Rhodora 47: 396- 404. LANJOUW, J. and F. A. STAFLEU. 1964. Index Herbariorum, Inter- national Bureau for Plant Taxonomy and Nomenclature of the International Association for Plant Taxonomy. Utrecht, Nether- lands. 1970] Silphium — Settle and Fisher 543 Perry, L. M. 1937. Notes on Silphium. Rhodora 39: 281-297. SMALL, J. K. 1933. Manual of the southeastern flora. Published by author, N.Y. 1554 p. WEATHERBY, C. A. 1935. A letter addressed to C. C. Deam. A PRIOR RECORD OF CHRYSANTHEMUM LACUS- TRE BROTERO: In a recent note in this journal (72:250, 1970) I reported Chrysanthemum lacustre Brotero, a Portuguese introduction, as being recently discovered for the first time in Gray’s Manual range. However, Camille Rousseau, of Université Laval in Quebec, has informed me of a record made twenty years previously in Canada. In Enumération des plantes du Canada, Naturaliste canadien, 93:1046 (1966) Bernard Boivin cites a collection from Ottawa, Ontario, deposited in the pao Herbarium, with label reading “Abandoned fields, 1 mi. west of RR. bridge over Rideau Canal, 1948, G. A. Mulligan." I am most appreciative of Prof. Rousseau's pointing out this earlier record and supplying me with the details. CHARLES E. STEVENS 615 PRESTON PLACE CHARLOTTESVILLE, VA. 22903 THE DISTRIBUTION OF PUBESCENT LEAVED INDIVIDUALS OF CONOCARPUS ERECTUS (COMBRETACEAE) JOHN C. SEMPLE! The buttonwood mangrove Conocarpus erectus L. grows along the coast of west tropical Africa, the Atlantic and Pacific coasts of tropical and subtropical North and South America and throughout the West Indies, Pubescent leaved individuals are restricted to the northern areas of the West Indies, southern Florida and northern Central America (Fig. 1) and have been variously named C. pubescens Schmach., C. erectus var. sericeus Fors. ex DC., C. sericeus (Fors. ex DC.) G. Don, C. erectus var. sericeus Griseb. and C. sericeus (Griseb.) Jimenez. Stearn (1958) noted the existence of intermediate leaved individuals (Stearn 760 from Portland Ridge, Clarendon, Jamaica) and recommended that specific level be dropped in favor of varietal level. (In this article Stearn cited the type-locality of y sericeus Griseb. as Antigua and St. Vin- cent, Lesser Antilles, but Grisebach (1860) listed sericeus only from the Bahamas (Swain a. y), a locality consistent with the findings of this author), Study of the variation in pubescence of material from F, GH, MO and fresh material in the field suggests division of the intermediate leaved condition into three groupings. The first condition consists of a few slightly pubescent leaves on an otherwise silky pubescent leaved plant. The converse of the first condition also occurs giving a plant with a few sparsely pubescent leaves on an otherwise gla- brous leaved plant. The third condition consists of individ- uals with pubescent and glabrous leaves in a cyclic pattern. ‘Supported by a NSF undergraduate grant, GY 5738 from Tufts University. Present address: Missouri Botanical Garden, St. Louis, Missouri 63110. The advice of Dr. Norton H. Nickerson, Dr. Louis V. Wilcox, and Dr. Walter H. Lewis is gratefully acknowledged. 544 Conocarpus — Semple 545 3 - š Li a | Mone 4 > KJ a E | | | E Ë š Ete) | : ELT 8 °; A a f ` ey 2 b / à Mai. e & / l Weg 2e 4 H ` | "PE | \ E ' ) `> | J | £ t°, a s ER “sl. (> ` | N Í ANN LAE KO 8 du “ E E ( A ` y (ISSN Xt (22 1 EA ~e S N | Ü 4 e 0 MW 7 o, p.d 4 1 Ne) s ds S$ e N EV [í ] yup ES - \ Á | : / we P, g | N $ ie P SN PO ` A 9 | ( Qu T / | » ea g I aes A / pt / { fy t. by f $95 / 9 9 0 c .n ° 5 $S > £ Š E mS: É > ü > o = ^ A c w 0 0 ‘ ° > > < ç 3 8 e 254299 £ i = z 8 ie i zç — o O p zy 9 É Ë š E óe6*292 5 a 9e SeN E š E Š ae 3 f : — Figure 1. Map of collection locations of Conocarpus erectus with some leaf pubescence. See text for distribution of glabrous plants and the entire species. 546 Rhodora [Vol. 72 In the complex phenomenon the degree of pubescence per leaf shifts from a high density of hairs to few or no hairs and back to the initial condition. The pattern of hair dis- tribution on the leaves in the intermediate range of density is similar to the pattern of Spiraea ulmaria L. as described by Yapp (1912). The cycle may end with either pubescent leaves at the time of collection (Elmore D21 Chachua Bay, Oaxaca, Mex.) or with glabrous leaves at the time of col- lection (Semple 11, Duck Cay, Jewfish Chain, Great Exuma, Bahamas). Collection dates from various specimens indi- cate that the rate of cycling and the cycling itself are not necessarily seasonal. Cyclic flushing of pubescence has been noted in other species as well, for example, Phragmites communis, Lysimachia vulgaris, Mentha aquatica, Pseudo- barleria sp. (all Yapp 1912), and Populus alba (H. John- Son, personal communication). The variation in leaf shape and degree of pubescence in Borrichia arborescens (L.) DC. show a similar phenomenon to that in C. erectus. The two can be found growing together in the Bahamas. The distribution of the glabrous leaved form of C. erectus is equal to the full area of distribution of the species. Gla- brous forms can be found growing with pubescent forms throughout the area indicated by Fig. 1. On various cays of Great Exuma, Bahamas, the two extremes may be found so close together that the roots and branches intermingle, The existence of a full continum of intermediate forms between the glabrous and pubescent extremes even on the same branch dictates that varietal status for the extremes is unwarranted. The only name that can apply is C. erectus L. Future descriptions of the species should note that a wide range of leaf pubescence is possible for plants occurring in the northern areas of the species’ distribution in the Americas. DEPARTMENT OF BIOLOGY TUFTS UNIVERSITY MEDFORD, MASSACHUSETTS 1970] Conocarpus — Semple 547 LITERATURE CITED GRISEBACH, A. H. R. Flora of the British West Indian Islands. Lon- don. 1860 (1859-1864). STEARN, W. T. 1958. A key to West Indian mangroves. Kew Bull. 33-37. 1958. YAPP, R. H. Spiraea ulmaria and its bearing on the problem of xeromorphy in marsh plants. Ann. Bot. (London) 26: 815-870. 19:12: Correction in article on Rubus Hybrids. On page 243, line 9, in this volume of Rhodora the name Rubus frondosus was erroneously given as being equivalent to the hybrid R. Enslenii X pensilvanicus. Actually this hybrid doesn't closely match any named blackberry of which we are aware. To conform with this correction the word most in the third line of page 242 should be changed to all. Thus the sentence as altered would read, “In the follow- ing treatment the names in parentheses that are given di- rectly after some of the hybrid designations are binomials, in all cases rejected ones, which we consider to be equivalent to these hybrids." F. L. Steele and A. R. Hodgdon SALIX STARKEANA IN NORTH AMERICA ALV DAN YOUNGBERG The name Salix depressa L. subsp. rostrata (Anderss.) Hiitonen (Mem. Soc. Faun. Fl. Fennica 25: 82, 1950) has been applied to Salix bebbiana Sarg. in North America by Hultén (1968). However, Salix depressa L. and Salix beb- biana Sarg. are not closely related taxa. The epithet depressa has been the cause of some confusion among salicologists because several authors have used the same epithet for dif- ferent species. Salix depressa L. (Fl. Suec. 2nd ed: 352, 1755) is a synonym of Salix lanata L. (Sp. Pl. 1019, 1753). Salix depressa (non L.) as used in part by Fries (1832, 1840) ; Ledebour (1850); and Seemen (1909); is a synonym of Salix starkeana Willd., the European and Asian willow spe- cies most closely related to Salix bebbiana Sarg. of North America. The epithet rostrata is also confusing because it has been applied to several different willow species. Richardson in 1823 used the epithet rostrata for a willow species he de- scribed from specimens collected somewhere in Canada west of Hudson Bay. However, he did not know that Thuiller had already used the epithet rostrata for a different taxon described by him in 1799. Sargent proposed Salix bebbiana as a new name for Salix rostrata Richardson in 1895. Thus, we see that the taxonomy of Hiitonen and Hultén places the taxon known as Salix bebbiana Sarg. as a sub- species of the taxon known as Salix lanata L. These two willows have completely different morphological characters and have been placed in different sections by most European salicologists including Skvortsov of the U.S.S.R. (1968). I consider Salix bebbiana to be a subspecies of Salix starkeana and propose the following taxonomy : 1. Salix starkeana Willd. subsp. starkeana. Salix starkeana Willd., Sp. Pl. 4(2) : 677, 1806. 548 1970] Salix — Youngberg 549 Salix livida Wahlenb., Fl. Lapp: 272, 1812. Salix vagans Anderss., Ofver. K, Vet. Akad. Fórhand 15: 121, 1858. Salix depressa auct. (non L. 1755, Fl. Suec. 2 ed.: 332) ; Fries, 1832, Mantissa 1: 56, id. 1840, Bot. Not.: 197; Ledb., 1850, Fl. Ross. 3, 2: 611; Seemen, 1909, in Aschers, et Graebn. Synops. 4: 115. TYPUS: Silesia, prope Gurau, leg. Starke (B, K, LE, S). 2. Salix starkeana Willd., subsp. bebbiana (Sarg.) Young- berg, comb. nov. Salix rostrata Richardson, in Franklin, Journey: 753, 1823; not Thuill., 1799. Salix fusca Hooker, Fl. Bor.-Am, 2: 151, 1839; non L. Salix vagans 1. einerascens b. occidentalis Andersson, in Ofv. Vet.-Akad. Fórh. 15: 122, 1858. Salix vagans 1. S. rostrata Andersson, in Svensk. Vet.- Akad. Handl. 6: 87, 1867. Salix vagans occidentalis Andersson, in Svensk. Vet.- Akad. Handl. 6: 87, 1867. Salix livida occidentalis Gray, Man. ed 5, 464, 1867. Salix vagans rostrata 3. obovata Andersson apud De Candolle, Prodr. 16 (2) : 227, 1868. Salix bebbiana Sarg., Gard. & Forest 8: 463, 1895. Salix perrostrata Rydb., Bull. N.Y. Bot. Gard. 2: 163, 1901. Salix depressa c. rostrata Seemen, in Ascherson & Graebner, Syn. Mitteleur. Fl. 4: 119, 1909. Salix bebbiana var. perrostrata (Rydb) Schn., Jour. Arnold Arb. 2: 71, 1920. TYPUS: Forest regions of Canada W. of Hudson Bay, Richardson (GH!, K, NY). Salix bebbiana is very similar to Salix starkeana, but these two taxa may be distinguished by the following char- acters. The mature leaves and twigs of the current season usually have some hairs present in Salix bebbiana while 550 Rhodora [Vol. 72 the mature leaves and twigs of the current season are glabrous in Salix starkeana. Subspecies bebbiana occurs in North America, and subspecies starkeana occurs in Fin- land, Czechoslovakia, Germany, Switzerland, Norway, Ro- mania, Sweden, and the U.S.S.R. SUITE 306 364-99TH AVENUE SE CALGARY 30, ALBERTA, CANADA LITERATURE CITED HULTÉN, E. 1968. Flora of Alaska and neighboring territories; a manual of vascular plants. Stanford University Press. SKVORTSOV, A. K. 1968. Willows of the USSR and Adjacent Re- gions. Moscow. THE CHROMOSOME NUMBER OF OBOLARIA VIRGINICA L. (GENTIANACEAE) KATSUHIKO KONDO Investigations of chromosomes are necessary to the fur- ther classification and clarification of the natural relation- ships between species. Obolaria virginica L. (Gentianaceae), which is mono- typic, is common in rich mesic woods throughout the pied- mont and mountain area in southeastern United States. The chromosome number of O. virginica (n—28) is here described for the first time. O. virginica was collected on the campus of the Univer- sity of North Carolina at Chapel Hill (coll. K. Kondo 562, March 10, 1970). The flower buds were fixed in Carnoy's solution: 6 : 3 : 1 of ethanol, chloroform and glacial acetic acid. Observations were made by the aceto-carmine squash method. The meiotic divisions in the PMC's were very regular and 28 bivalent chromosomes at metaphase I of meiosis were clearly observed (Figs. 1 and 2). The bivalents were nor- mal and consisted of two elements of equal size. Their forms were ring-shaped, conjugated at both ends, and stick- shaped, conjugated end to end. However, the author infers that the basic chromosome number of Obolaria. is 7, and O. virginica is the octoploid which is constant. Chromosome relationships of members of the Gentian- aceae have been well studied by Rork (1949), Favarger (1952), and Wada (1955, 1956, and 1960). Members of the Gentianaceae show heteroploidy, and their basic chro- mosome numbers are 5, 6, 7, 8, 9, 11, and 13. The same basic chromosome number as O. virginica (X=7) occurs in several genera of this family, such as Centaurium, Gen- tiana, Sabatia, and Swertia. According to Rork's report (1949), about 58 per cent of the species of the Gentianaceae investigated to date are polyploid. Perennial species show no greater tendency toward polyploidy than do subperennial 551 552 Rhodora [Vol. 72 Figs. 1 and 2. Chromosomes in pollen mother-cell of Obolaria virginica L. : »—28 (Fig. 1. X ca. 950; Fig. 2. X ca. 1450). 1970] Obolaria — Kondo 553 species; the subperennials have 32.8 per cent higher average chromosome number than the perennials. DEPARTMENT OF BOTANY THE UNIVERSITY OF NORTH CAROLINA CHAPEL HILL, N.C. 27514 LITERATURE CiTED FAVARGER, C. 1952. Contribution 4 l'etude caryologique et biologique des Gentianacées. Berichte Schweiz. Bot. Ges. 62: 244-257. Rork, C. L. 1949. Cytological studies in the Gentianaceae. Am. Journ. Bot. 36: 687-701. Wana, Z. 1955. Cytological studies of four species of Swertia and one species of Halenia. Jap. Journ. Genet. 30: 191. 1956. Cytological studies in Gentianaceae. Jap. Journ. Genet. 31: 315. 1960. Chromosome numbers in Gentianaceae. Chromo- some Information Service 7: 28-30. TRILLIUM CERNUUM L. AND GERANIUM MACULATUM L.: NEW FOR SOUTH DAKOTA Wild geranium (Geranium maculatum L.) and nodding wake robin (Trillium cernuum L.) are not listed for South Dakota by Over (1932) or Winter, Winter, and Van Brug- gen (1959). Geranium maculatum. (Johnson 73, NDA) and Trillium cernuum Johnson 69, NDA) were both collected on June 4, 1969, in Sieche Hollow State Park, 8 miles west and 6 miles north of Sisseton, South Dakota. Sieche Hol- low is an erosional ravine located on the glacial landform called the Coteau des Prairies. Both species were collected in dense, moist woods with an overstory consisting primarily of basswood (Tilia amer- icana), American elm (Ulmus americana), sugar maple (Acer saccharum), slippery elm (Ulmus rubra), and iron- wood (Ostrya virginiana). Many species found here, in- cluding the above state records, are approximately fifty miles west of their known range in the deciduous forests of west-central Minnesota. W. CARTER JOHNSON DEPARTMENT OF BOTANY NORTH DAKOTA STATE UNIVERSITY FARGO 58102 LITERATURE CITED OVER, WILLIAM HENRY. 1932. Flora of South Dakota. University of South Dakota Natural History Studies No. 3. Vermillion. 161 p. WINTER, JOHN M., CLARA K. WINTER, and THEODORE VAN BRUGGEN. 1959. A checklist of the vascular plants of South Dakota. De- partment of Botany, University of South Dakota, Vermillion. 176 p. A NEW MT. WASHINGTON STATION FOR GNAPHALIUM SUPINUM Gnaphalium supinum. L. is known in New England from Mt. Katahdin and Mt. Washington. It was first collected on Mt. Katahdin by Scribner in 1874 and has since been collected there several times probably from only two sta- tions. Notations on the sheets indicate it is rare on this mountain. On Mt. Washington it was first collected by William Boott in Tuckerman Ravine in 1855. Since then there have been a number of collections apparently confined to three or four stations. There appear to be two stations in Tuck- erman Ravine and one or perhaps two high on the cone on the slope toward the Alpine Garden. I have explored the cone and Tuckerman Ravine rather thoroughly but have been able to locate only one of these stations, This is part way up the ravine and nearly in the middle of the trail. In the past several years I have not been able to find this colony. The trail receives heavy use and some erosion results along with trampling of vegetation. I think it likely that this station has been destroyed or at least reduced to a few isolated plants. In view of the scarcity of this plant, I was most pleased, when last summer I came upon a colony of it in Huntington Ravine. This was a vigorous stand of one hundred or more plants growing in some gravel. It was far enough off the trail so that it was not in danger of trampling, but was not in a difficult place to reach. There are no records of it from Huntington Ravine in either the New England Botanical Club or the Gray Herbarium. As the Ravine has been rather thoroughly botanized in past years it is possible that this colony has developed rather recently. The windborn seeds presumably are readily transported over a considerable distance and could have arrived from the station on the cone. The plant seems to grow only on open gravelly sites of which there are rather few and another possibility is 555 556 Rhodora [Vol. 72 that it could have seeded in from some small unbotanized location on the precipitous cliffs of the Ravine. National Forest regulations forbid picking of plants in the alpine area so no collection was made. I did make care- ful notes of the colony for a record for my own herbarium and it is hoped that this note will serve as acceptable evi- dence of its occurrence in Huntington Ravine. FREDERIC L. STEELE ST. MARY’S-IN-THE-MOUNTAINS LITTLETON, NEW HAMPSHIRE VOLUME 72 1970 Errata for RHODORA Page 6, lines 20 and 24, for microparpum read microcarpon. Page 24, lines 2, 20, 29 and 36, for angustata. read angustatum. Page 147, line 36, for Verononia read Vernonia. Page 150, line 37, for piedmont read Piedmont. Page 154, line 36, for piedmont, coastal plain, read Piedmont, Coastal Plain. Page 167, line 16, for arenchyma read aerenchyma. Page 232, line 37, for Rhycodrys read Phycodrys. Page 243, line 6, for York Co., read York Co., York. Page 280, line 28, for Tsua read Tsuga. Page 281, line 15, for Hammamelis read Hamamelis. List of Sustaining members Barneby, for Rubert read Rupert Seymour, for Conklin read Conkling Youngkin, for Herbert read Heber Page 324, line 1, for spinescens read spinescens. Page 325, line 58, for umbiblicalis read umbilicalis. Page 338, line 10, for tomentosus read tomentosum. Page 339, line 12, for quinequefolia read nudicaulis. Page 340, line 14, for Hammelis read Hamamelis. Page 344, line 14, for orbiculata read orbiculatus. Page 346, line 29, insert ) after islands. Volume 72, No. 792 ineluding pages 401-582, was issued December 31, 1970. INDEX TO VOLUME 72 New scientific names and combinations are printed in bold face type Abies balsamea 418, 509, var. phanerolepis 509; lasiocarpa 497 Abrahamson, Warren G. and Otto T. Solbrig, Soil Preferences and Variation in Flavanoid Pig- ments in Species of Aster 251- 263 Abronia arenaria 371, 372; lati- folia 371, 372; umbellata 372 Acacia 534 Acanthospermum 106-108, Tax- onomie Changes and Generic Affinities in the Genus 106; australe 107; brachyceratum 106; consobrinum 107; donii 106, 107; hispidum 106; humile 106; lecocarpoides 106; lepto- lobium 106; microcarpum 106- 108; sect. Acanthospermum 107; sect. Ceratochlaena 106; sect. Lecocarpopsis 106; sect. Xanthoides 107; simule 107, 108 Acer, 54; pensylvanicum 281, 492, 521; rubrum 79, 280, 281, var. rubrum 521 f. tomentosum 521, var. trilobum 521; saccharum 521, 554; spicatum 492, 521 Achillea Millefolium 527 Acrochaetium polyides 201, 205, 236 Actaea pachypoda 517; rubra 517 Adventina ciliata 386 Aesculus octandra 492 Agardhiella tenera 214, 250 Agarum cribrosum 116, 319, 326, 336 Ageratum 94, 95; noides 94, 97 campulocli- Agrimonia gryposepala 349 Agropyron repens var. repens 510, f. aristatum 510, f. pilosum 510, f. trichorrachis 510; trachycaulum var. majus 510, var. novae-angliae 510 Agrostis alba var. alba 511, var. palustris 511; perennans 511; scabra 511; tenuis 511 Ahnfeltia plicata 117, 199, 200, 216, 231, 232, 236, 322, 333 Aira caryophyllea 7; praecox 7 Alaria esculenta 116, 205, 206, 319, 326, 336; musaefolia 319 Alaska, Juncus Slwookoorum, a New Species from 486 Alisma trivale 509 Allium canadense 170; Schoeno- prasum var. sibiricum 514 Alnus crispa var. mollis 515; rugosa var. americana 515, var. rugosa 515; serrulata 79 Alomia 95; fastigiata 97; longi- folia 95, 97; microcarpa 95 Alopecurus pratensis 511 Amaranthus albus 516; flexus 516 Ambrosia artemisiifolia var. ar- temisiifolia 527, var. elatior 527, f. villosa 527 Amelanchier 350; arborea 519; Bartramiana laevis 519; can- adensis 519; laevis 519, X can- adensis 519; stolonifera 519 Anamomis dicrana 23; simpsonii 23 Anaphalis margaritacea var. in- tercedens 527, var. margar- itacea 527 Anethum graveolens 394, 395 retro- 559 560 Andropogon gerardi 79; scoparius 79 Anisacanthus abditus 70, 71; Och- oterenae 70, 71; stramineus 70, 71 Antennaria Bayardi 136; Brain- erdii 527; canadensis 527; fal- lax 527; Longii 136; munda 527; neglecta 527; neodioica 527 Anthemis Cotula 527; nobilis 96, 98; tinctoria 527 Anthoxanthum odoratum 511, f. giganteum 47, f. odoratum 47 Antithamnion americanum 201, 220; boreale 201, 220, 236; cruciatum 117, 220, 231, 232, 236, 322, 333; floccosum 117, 220, 232, 236, 322, 333; plum- ula 201, 221, 236; pylaisaei 201, 221, 236 Apocynum androsaemifolium 524 Aquilegia canadensis 339, 347, 492; vulgaris 517 Aralia hispida 522; nudicaulis 281, 492, 522; quinquefolia 339 Arceuthobium pusillum 516 Arctium minus 528; nemorosum 528 Arctostaphylos Uva-ursi 530, var. coactilis 523 Arenaria groenlandica 517, 530; lateriflora 517; patula 272; peploides var. robusta 517 Arethusa bulbosa 514 Arisaema Stewardsonii 513 Armoracia lapathifolia 518 Artemisia senjavenensis 488; vul- garis 527 Asarum canadense 492 Asclepias 54; incarnata 29, var. pulchra 524 Ascophyllum nodosum 116, 227, 319, 320, 324, 336 Asparagopsis hamifera 208 Asperocoecus echnatus 116 Rhodora [Vol. 72 Aster 39, 251, 252, 565, Soil Pref- erences and Variation in Flava- noid Pigments in Species of 251; acuminatus 263, 401-403, 405, 407-414, 418-426, 429, 431, 432, 437, 526, f. discoideus 419, f. virescens 419; Blakei 401, 407, 419, 421, 431, 437, Chromo- some Numbers in 437; X Blakei 401, 404-407, 411, 412, 414, 418, 420-422, 424-426, 428, 430-432, 526, Evidence of the Hybrid Status of 401; bracei 41; brachyactis 392, New for Kansas 392; ciliatus 260, 263; concolor var. concolor 40, var. simulatus 39; cordifolius 260, 262, 263, var. cordifolius 526; divaricatus 350; dumosus 79, var. coridifolius 42, var. graci- lipes 41, var. sublaefolius 41, 42; ericoides 406; foliaceus 526; fontinalis 40; johannensis 526; laevis 263; lateriflorus 526, var. tenuipes 526; lowreianus 260, 263; macrophyllus var. macro- phyllus 526, var. velutinus 526; nemoralis 1, 401, 402, 405, 407- 414, 418-427, 431, 432, 437, 526, Chromosome Numbers in 437, f. alba 412, var. Blakei 401, 405, 408, var. major 405, 406, 408; novae-angliae 406; novi-belgii var. novi-belgii 526; parviceps 392, New for Kansas 392; patens 40, var. floridanus 40; pilosus 392; polycephalus 526; puniceus 281; radula var. rad- ula 526, var. strictus 526; sagit- tifolius 263; simmondsii 41; simplex 392, var. ramosissimus 526; simulatus 40; 41; tenuifolius 41, var. aphyllus 40, 41, var. tenuifolius 41; um- bellatus 527; undulatus 260, sulznerae 1970] 262; vimineus 392 Astragalus, An Undescribed Spe- cies of 189; cottamii 189-191; monumentalis 189, 191-193; sect. Desperati, subsect. Naturi- tensis 189 Athyrium Filix-femina var. Mich- auxii 508, f. elatius 508, f. ru- bellum 508 Atriplex argentea 285; glabrius- cula 516; patula var. hastata 516, var. littoralis 516, var. patula 516 Audouinella membranacea 117, 200, 205, 231, 232, 236, 322, 333 Avena sativa 510 Baccharidastrum 95; rivulare 97; triplinervium 97 Baccharis 95, 534; brevifolia 98; burchellii 98; cf. dracunculifolia 98; elaeaginoides 98; ligustrina 98; mesoneura 98; tarchonan- thoides 98; sp. 98 Baldwin, J. T. Jr., White Phase in Flower Development in Cy- pripedium acaule 142-143 Bangia ciliaris 201, 202, 236; fuscopurpurea 117, 199, 200, 202, 231, 232, 236, 323, 333 Barbarea vulgaris var. arcuata 518, var. brachycarpa 518 Barbour, Michael G. and James E. Rodman, Saga of the West Coast Sea-Rockets: Cakile ed- entula ssp. californica and C. maritima 370-386 Barneby, R. C., New Phanero- gams from the Arid Neotropics 66-71 Bartonia paniculata 15, var. in- termedia 524, 530; tenella 15; virginica 524 Batrachospermum sp, 207, 230 Bean, Ralph C., Stuart Kimball Index to Volume 72 561 Harris 126-129 Behen 146; noveboracensis 154; praealtum 154 Bellis perennis 273 Berberis Thunbergii 517 Betula alleghaniensis 280, 281, 515; caerulea-grandis 515; lenta 11, 280, 491; lutea 2, 11, 491, 515; papyrifera 12, 280, var. cordifolia 515, var. papyrifera 515; populifolia 515, X B. papyrifera var. papyrifera 515 Bidens frondosa 527, f. anomala 527; gardneri 98 Blainvillea latifolia 95; boidea 95, 98 Blidingia minima 318 Blunt, Terry and F. R. Fosberg, Vernon Black Gum Swamp 280- 282 Bonamia 32; 63 Bonnemaisonia hamifera 201, 208, 236 Borreria laevis 393 Borrichia arborescens 546 Bostrychia rivularis 226, 230 Botrychium dissectum f. dissec- tum 508, f. obliquum 508; lanceolatum 2; matricariaefo- lium 2, 508; multifidum var. intermedium 508; virginianum 4 Brachyelytrum erectum var. erec- tum 511, var. septentrionale 511 Brachythecium starkei 494 rhom- abdita 32; aquatica Brassica juncea 518; Kaber var. pinnatifida 518; nigra 518; Rapa 518 Braya Longii 136 Bromus altissimus 345; ciliatus 345; latifolius 346; latiglumis 345; lepidus 345; mollis 345; nottowayanus 345, 346; pubes- 562 cens 339, 340, 344-346; purgans 79, 345, 346, 350, var. latiglu- mis 346; racemosus 47; squar- rosus 345 Bryophytes, Zonation of 276 Bryum cyclophyllum 500; Longii 136; pallens 494 Bumelia 25; angustifolia 25, 26; celastrina 26, var. angustifolia 26, var. celastrina 25, 26 Butomus umbellatus 164 Cacalia 146; acaulis 148; nove- boracensis 154 Cakile 370-373, 378; americana 373; californica 370, 371, 373- 3779, 382, 383; edentula 370, 371, 373, 375, 378, 379, 518, ssp. californica 370, 371, 376, 379- 381, var. californica 370, 377- 379; maritima 370, 371, 373, 375-383 Calamagrostis canadensis var. canadensis 511 Calamovilfa 73, 75, A New Spe- cies of 72; arcuata 72, 74-79; brevipilis 73, 75-77; curtissii 78, 75-77; gigantea 73; longifolia 73; sect. Calamovilfa 75, 76; sect. Interior 75 Calea serrata 98 Calla palustris 513 Calliblepharis ciliata 325 Calliergon stramineum 494 Callithamnium baileyi 221, 232, 233, 236; corymbosum 201, 221, 236; roseum 221; tetragonum 222 Calypogeia meylanii 494 Calopogon pulchellus 514 Calycanthus floridus 55, var. laevigatus 55, 56 Campanula rapunculoides 526; rotundifolia 526 Capparidaceae 534 Rhodora [Vol. 72 Capsella Bursa-pastoris 518 Capsosiphon fulvescens 317, 331 Cardamine Longii 136; parviflora Var. arenicola 347, 518; pen- sylvanica 518 Carduus smallii 45; vittatum 45 Carelia 95 Carex 565; adusta 512; aenea 512; angustior 512; arctata 513; argyrantha 512; artiteeta 513; atlantica 512; Bayardi 136; blanda 340, 346; brunnescens var. sphaerostachya 512; canes- cens var. canescens 47, var. dis- juncta 512; var. subloliacea 512; cephalantha 512; commu- nis 512; conoidea 513; con- vulata 2, 7; Crawfordii 16; cumulata 512; debilis var. Rudgei 340, 513; deflexa 513; Deweyana 512; Emmonsii 340, 513; exilis 8, 512; flava 513; folliculata 513; Hormathodes 512, f. invisa 512; Houghtonii 513; intumescens 339, 347, var. Fernaldii 513; Jacobi- -peteri 488; latebracteata 503; laxicul- mis 2; laxiflora 2, 9, 513, var. blanda 9; leptalea 512; lepto- nervia 9, 513; Longii 136; lurida 513; Mackenziei 512; multicos- tata 497; nigra var. nigra 513, var. strictiformis 513; nigricans 500; novae-angliae 513; ormos- tachys 513; paleacea 513, var. erectiuscula 513; pallescens var. neogaea 515; panicea 513; pauciflora 513; paupercula var. irrigua 513; phaeocephala 498; prairea 8; radiata 8; rosea 8, 281; rostrata 513; scabrata 2, 8; scoparia var, Scoparia 512; Seorsa 339, 346; sp. 79, 499; stipata 512; straminiformis 497; stricta var. stricta 513; 1970] subfusca 498; Swanii 340; ten- era 512; trisperma var. Bill- ingsii 512, var. trisperma 512; umbellata 513; vestita 8; viri- dula 513 Carterothamnus 100, 103, 104 Carum Carvi 522 Carya glabra 2; ovalis 2; tomen- tosa 2 Cassandra calyculata var. angus- tifolia 523 Castanea dentata 12, 280 Castilleja parviflora var. oreopola 499 Celastrus orbiculatus 344, 521 Centaurea montana 500 Centaurium 551 Centratherum punctatum 94, 97 Cephalanthus 9 Ceramium deslongchampsii var. hooperi 117, 202, 222, 236, 323, 333; diaphanum 222, 230; ele- gans 325; rubriforme 201, 222, 236, 323, 333; rubrum 117, 200, 222, 231, 233, 236, 323, 333; strictum 200, 222, 223, 233, 236 Cephalozia ambigua 493; bicus- pidata 494 Cerastium arvense 517; tomen- tosum 517; vulgatum 517 Ceratocolax hartzii 216, 232, 236 Ceratophyllum demersum 13 Chamaedaphne calyculata var. angustifolia 523 Cheilanthes 534 Chaetomorpha aerea 317, 331; linum 326; melagonium 116, $17, 322, 831 Chamaecyparis 1, 2, 4, 8, 15 Chenopodium album 516; ambro- sioides 398 Chiloscyphus pallescens 494 Chimaphila umbellata var. cisat- lantica 523 Chrysanthemum Balsamita 527; Index to Volume 72 563 lacustre 250, 543, New to Gray’s Manual Range, 250, A Prior Record of 543; Leucan- themum var. pinnatifidum 527 Chrysocoma acaulis 146, 147; tomentosa 146, 154 Chrysopsis 42; floridana 44; hys- sopifolia 42; mariana var. flori- dana 44; subulata 43; traceyi 43 Chondria baileyana 200, 201, 226, 31, 233, 236; tenuissima 226, 230 Chondrus crispus 117, 200, 203, 210, 217, 231, 233, 237, 321-333 Chorda filum 320, 336; tomentosa 116, 320, 336 Chordaria flagelliformis 116, 320, 336, var. densa 320 Choreocolax polysiphoniae 117, 213, 231, 233, 237, 323, 333 Cicuta maculata 522 Cinna latifolia 5 Cireaea alpina 522; canadensis 283; quadrisuleata 339, var. canadensis 14 Cirsium 44; arvense 338, 528; horridulum 44, var. horridulum 45, var. vittatum 45; muticum 528; nuttallii 44; pinetorum 44; smallii 45; vittatum 44; vulgare 528 Cladophora flexuosa 317, sericeae 317 Cladosiphon zosterae 320, 336 Cladrastis lutea 492 Clathromorphum circumscriptum 117, 200, 201, 209, 237, 323, 324, 333 Claytonia virginica 349, 441 Clematis baldwinii 17, 18, var. baldwinii 18, var. latiuscula 18; crispa 18; fremontii 18; reticu- lata 18; virginiana 517 331; Clethra alnifolia 56, var. pubes- 564 cens 57, var. tomentosa 56 Clintonia borealis 514 Codiolum gregarium 116, 317, 331; petrocelidis 116, 317, 331; pusillum 116, 317, 331 Coelopleurum lucidum 522 Coleman, James R., Additional Chromosome Numbers for Bra- zilian Compositae 94-99 Collinsia parviflora 289 Compositae, Additional Chromo- some Numbers in Brazilian 94; Studies in the 100 Comptonia peregrina 515 Conocarpus erectus 544-546, The Distribution of Pubescent Leaved Individuals of 544, var. sericea 544; pubescens 544; sericeus 544 Convolvulus arvense 378; sepium var. sepium 524 Conway, Joan R., Edward J. Hehre and Richard A. Stone, Flora of the Wolf Islands, Part III, The Marine Algae 115-118; Richard A. Stone, Edward J. Hehre and Arthur C. Mathie- son, A Preliminary Checklist of the Marine Algae of Campo- bello Island, New Brunswick, Canada 313-338 Cook, Frank S. and H. Vincent Elliott, Listera ovata (L.) R. Br. in the Bruce Peninsula, On- tario 274-275 Coptis groenlandica 281, 517 Corallina officinalis 117, 200, 210, 211, 237, 323, 324, 333 Corallorhiza maculata 515; trifida 515 Corema Conradii 14, 521, 529 Coriospermum hyssopifolium 285 Cornus alternifolia 522; amomum 79; canadensis 522; florida 11; stolonifera 522 Rhodora [Vol. 72 Correll, D. S. and M. C. John- ston, Manual of the Vascular Plants of Texas (Review of) 533 Corydalis sempervirens 517 Cotula coronopifolia 378 Crassulaceae 460, 462 Crataegus 565; chrysocarpa 519; Jonesae 519; macrosperma 519; monogyna 13; succulenta 519 Croton floridanus 22; glandulosus 21, 22, var. floridanus 21, 22, var. glandulosus 21, var. sep- tentrionalis 22, var. simpsonii 21, 22 Cuscuta Gronovii 524 Cusick, Allison W., An Assem- blage of Halophytes in North- ern Ohio 285 Cynoctonum 27; angustifolium 28; mitreola 28, 30; sessilifo- lium 28, 30, var. angustifolium 28, 29, var. microphyllum 28, var. sessilifolium 28; succulen- tum 28-30 Cynosurus cristatus 510 Cyperus 535; difformis 284; escu- lentus 285 Cypripedium acaule 142, 514, White Phase in Flower Devel- opment in 142, f. albiflorum 142 Cystoclonium purpureum 117, var. cirrhosum 200, 201, 214, 232, 237, 323, 333 Cystopteris fragilis 508 Dactylis glomerata van. ciliata 510, var. glomerata 510 Dalibarda repens 520 Danthonia compressa 510; spicata var. spicata 510 D’Arcy, W. G., Mock Bishop's Weed in the New World Trop- ics: Range Extensions for 1970] Ptilimnium capillaceum (Um- belliferae) 393-396 Dasya pedicellata 200, 225, 231- 233, 237 Daucus Carota 522 Decodon verticillatus var. lacvi- gatus 522 Dennstaedtia punctilobula 508 Dermatolithon pustulatum 260, 210, 237, 323, 333 Deschampsia caespitosa var. par- viflora 510; flexuosa var. flex- uosa 510, f. flavescens 510 Desmarestia aculeata 116, 202, 320, 336; spp. 326; viridis 220, 336 Dianthus deltoides 517; plumarius 517 Dicentra canadensis 492; cucul- laria 492 Dichelyma falcatum 494 Dicranodontium denudatum 494 Dicranum muehlenbeckii 493 Dictyosiphon foeniculaceus 116, 320, 336; macounii 320, 336 Diervilla Lonicera 525 Diospyros virginiana 79 Diphylleia cymosa 492 Douglas, George W. and Ronald J. Taylor, Contributions to the Flora of Washington 496-501 Drepanocladus exannulatus 494; fluitans 494 Drosera intermedia 518; rotundi- folia 518 Dryopteris cristata 508; goldiana 491; marginalis 508; Phegop- teris 8; spinulosa var. ameri- cana 508, var. intermedia 508 Dulichium arundinaceum 511 Dumontia incrassata 200, 212, 232, 237, 323, 333 Duncan, Wilbur H., The Southern Limits of Trientalis borealis 489-492; and Ida E. Yates, Index to Volume 72 565 Comparative Studies of Smilax, sect. Smilax, of the Southeast- ern United States 289-312 Dunn, David B. and Lloyd W. Hess, Nomenclature of the Lu- pinus argenteus and L. cauda- tus Complexes 110-114 Echinochloa crusgalli 511 Echinocystis lobata 526 Echinops sphaerocephalus 528 Ectocarpus confervoides 116, 320, 336; fasciculatus 320, 336; sili- culosus 320, 336 Elachista fucicola 116, 320, 336; lubrica 320 Elatine americana 521 Eleocharis 565; elliptica 511; halophylla 511; nitida 48; ob- tusa 511; parvula 511; tenuis 48, 511; tuberculosa 9 Elliott, H. Vincent and Frank S. Cook, Listera ovata (L.) R.Br. in the Bruce Peninsula, On- tario 274-275 Ellisia microcalyx 400 Elvira 94; biflora 95, 98 Elymus arenarius var. villosus 510; caput-medusae 378; vil- losus 79; virginicus var. virgin- icus 510, f. hirsutiglumis 510 Empetrum nigrum 521, 530 Enteromorpha erecta 317, 331; groenlandica 116, 317, 331; in- testinalis 116, 318, 331; linza 116, 318, 331; marginata 116; micrococca 116, 318, 331; min- ima 116, 318, 331 Ephedra 534 Epicladia flustrae 116, 318, 331 Epifagus virginiana 525 Epilobium angustifolium — 522; coloratum 522; glandulosum var. adenocaulon 522; hirsutum 164-176, 181, 360, Distribu- 566 tional History in North Amer- ica of 164; leptophyllum f. um- brosum 522; palustris var, oli- ganthum 522; strictum 522 Equisetum arvense var. arvense 507; sylvaticum var. pauci- ramosum 507, f. multiramosum 507 Eragrostis cilianensis 188; mega- stachya 188; multicaulis 47; plana 188, in South Carolina 188 Erechtites hieracifolia var. inter- media 527, var. praealta 527 Erigeron canadensis 527; pul- chellus 2, 16; strigosus var. strigosus 527 Eriocaulon septangulare 513 Ernodia 35, 36; angusta 35; lit- toralis 35, 36, var. angusta 35, var. littoralis 35 Eriophorum angustifolium 512; spissum 512; tenellum 512; vir- ginicum 512 Erysimum cheiranthoides 518 Erythronium americanum 349 Erythrotrichia carnea 201, 202, 237 Eudesme virescens 325 Eugenia dicrana 23 Euonymus obovatus 492 Eupatorium amydalinum 97; bal- lotaefolium 95, 97; bupleurifo- lium 94, 97; fistulosum 79; laetevirens 95, 97; organense 94, 97; perfoliatum var. per- foliatum 526; tremulum 94, 97 Euphorbia 535; Cyparissias 521; Peplus 521 Euphrasia canadensis 525; Randii 525 Euthora cristata 177, 200, 202, 213, 230, 231, 233, 237, 323, 334 Fagopyrum sagittatum 516 Rhodora [Vol. 72 Fagus grandifolia 280, 515 Faruqi, S. A. and K. L. Mehra, Biosystematics of Setcreasea brevifolia 264-271 Festuca elatior 509, f. aristata 509; obtusa 6, 339, 346; rubra var. rubra 509, f. megastachys 509 Ficus brevifolia 23 Filipendula Ulmaria 519 Fisher, T. Richard and Wilbur T. Settle, The Varieties of Sil- phium integrifolium 536-543 Flaveria 42; floridana 42; lati- folia 42; X latifolia 42; linearis 42, var. latifolia 42 Florida, Additions and Nomen- clatural Changes in the Flora of Southern 17 Fogg, John M, Jr., Bayard Long (1885-1969) 130-136 Fontinalis 278; antipyretica 277, var. gigantea 277; dalecarlica 277; novaeangliae 276, 277 Fosberg, F. R. and Terry Blunt, Vernon Black Gum Swamp 280- 282 Fosliella lejolisii 211 Foster, Robert C. A New Ar- gentine Sisyrinchium 287 Fothergilla major 56; monticola 56 Fragaria X Ananassa 519; vir- giniana var. terrae-novae 519, var. virginiana 519 Franseria bipinnatifida 372; cha- missonis 372 Fraxinus americana 2, var. amer- icana 524; pennsylvanica var. subintegerrima 524; quadrang- ulata 273; sp. 79 Fucus distichus ssp. distichus 116, 320, 337, ssp. edentatus 116, 321, 337; ssp. evanescens 117, 321, 387; sp. 203, 204, 210, 1970] 320; spiralis 117, 321, 387; vesiculosus 117, 199, 321, 337, var. spiralis 321, 327 Galeopsis Tetrahit var. bifida 524 Galinsoga aristulata 386, 387; ciliata 386-391, 527, Its Arrival and Spread in the Northeast- ern United States 386; parvi- flora 386-390, var. hispida 386 Galium 347; Aparine 525; cir- caezans 344, var. hypomalacum 349; palustre 525; tinctorium var. tinctorium 525; trifidum 525; triflorum 344, 345, 525, verum 525 Gaultheria hispidula 281, procumbens 280, 281, 523 Gaylussacia baccata 523; dumosa var. Bigeloviana 523 Gentiana 551; crinita 15; glauca 499 Gentianaceae 551 Geranium Bicknellii 521; macu- latum 334, 554, New for South Dakota 554; molle 521; Rob- ertianum 521 Geum aleppicum var, 519 Gigartina stellata 117, 200, 210, 218, 230-232, 237, 323, 334 Glaux maritima var. maritima 523, var. obtusifolia 525 Glechoma hederacea var. hed- eracea 524, var. micrantha 524 523; strictum Glime, Janice M., Zonation of Bryophytes in the Headwaters of a New Hampshire Stream 276-279 Gloiosiphonia capillaris 201, 213, 237 Glyceria 509; striata var. var. stricta 510 canadensis 509; laxa striata 509, Index to Volume 72 567 Gnaphalium obtusifolium 27; supinum 555, A New Mt. Wash- ington Station for 555; uligi- nosum 527 Goodyera pubescens 11; repens 515 Gould, F. W. and R. I. Lonard, Eragrostis plana in South Carolina 188 Gracilaria foliifera 215, 231, 237 Grossularia hirtella 348 Gymnocarpium Dryopteris 508 Habenaria clavellata var. ophio- glossoides 514; fimbriata 514; lacera 514; macrophylla 514; obtusata 514; orbiculata 514; psycodes 347, 514 Haber, Erich, Some Aberrant Pyrola Collections from East- ern North America 480-485 Halosaccion ramentaceum 117, 200, 218, 230, 232, 287, 320, 323, 326, 334 Hamamelis 340; virginiana 281, 518 Harriman, Neil A., A Range Ex- tension for Odontites serotina (Lam.) Dum. (Scrophular- iaceae) 286 Harvill, A. M. Jr., New and Over- looked Species of the Virginia Flora 272-273 Hedeoma pulegioides 524 Hehre, Edward J., Joan R. Con- way and Richard A. Stone, Flora of the Wolf Islands, Part III, The Marine Algae 115-118; and Arthur C. Mathieson, In- vestigations of New England Marine Algae IIJ: Composition, Seasonal Occurrence and Re- productive Periodicity of the Marine Rhodophyceae in New Hampshire 194-235; Richard A. 568 Stone, Joan M. Conway and Arthur C. Mathieson, A Pre- liminary Checklist of the Marine Algae of Campobello Island, New Brunswick, Can- ada 313-338 Heliotropium 32, 534; angiosper- mum 32; curassavicum 33; fruticosum 34; horizontale 34; indieum 34; myosotoides 34; phyllostachyum 34; polyphyl- lum 33, var. horizontale 34, var. polyphyllum 33 Hemerocallis fulva 514 Hemidiodia ocimifolia 36 Heracleum maximum 522 Hermann, F. J., Further Addi- tions to the Bryoflora of M*. Katahdin, Maine 493-495 Hesperis matronalis 518 Hess, Lloyd W. and David B. Dunn, Nomenclature of the Lupinus argenteus and L. cau- datus Complexes 110-114 Heterophyllum haldaneanum 494 Heterotheca 42; floridana 44; graminifolia 43, var, traceyi 43; hyssopifolia 42, var. subulata 43; scabrella 44 Hieracium aurantiacum 528; can- adense var. canadense 528; flor- entinum 528; paniculatum 528; Pilosella 528; pratense 528; seabrum 528 Hildenbrandia prototypus 117. 200, 209, 231, 232, 237, 323, 334 Hill, L. Michael and O. M. Rogers, Chromosome Numbers of Aster Blakei and A. nemoralis 437- 438 Hodgdon, A. R., Important Work on the Flora of New England (Book Review) 137-141, and Frederic L. Steele, Hybrids in Rubus subgenus Eubatus in New England 240-250; Correc- Rhodora [Vol. 72 tion in article on Rubus Hy- brids 547 Hordeum jubatum 510; vulgare 510 Houstonia caerulea 525 Hudsonia ericoides 522, 530; tomentosa var. intermedia 522 Humulus Lupulus 516 Hydrangea 54 Hydrocharis morsus-ranae 164 Hydrocotyle americana 522 Hydrophyllum appendiculatum 399, 400; canadensis 492; trilo- bum 399, 400 Hypericum boreale 521; cana- dense 521; dissimulatum 521; gentianoides 521; majus 521; mutilum var. parviflorum 521; perfoliatum 878; perforatum 521; prolificum 79; virginicum var. Fraseri 521, var. virgini- cum 521 Hypnum haldanianum 494 Hypoxis sp. 398 Iberis umbellata 518, 531 Ichthyothere 94; rufa 95, 98 Ilex glabra 521, 529, 530; mon- tana 492; verticillata 79, 281, 339, 426, var. verticillata 521 Impatiens capensis 499, 521; pal- lida 492 Iris Hookeri 514, 530; versicolor 514 Isidorea 36; leonardii 36 Isoétes muricata 508; Tuckermani 3 Isthmoplea sphaerophora 117, 321, 337 Itea virginica 79 Jaliscoa 100; pringlei 100 Jasione montana 15 Johnson, Kermit L., Aster parvi- ceps (Burgess) Mack. & Bush pappifera — 100; 1970] and Aster brachyactis Blake, New for Kansas 392 Johnson, W. Carter, Trillium cer- nuum L. and Geranium macu- latum L., new for South Dakota 564 Jones, Samuel B., The Taxonomy of Vernonia acaulis, V. glauca and V. noveboracensis (Com- positae) 145-163 Juglans cinerea 491 Juncus 565; articulatus var. ar- ticulatus 514; balticus var. lit- toralis 514; brevicaudatus 514; bufonius var. bufonius 513; canadensis 340, var. canadensis 514; effusus 285, var. com- pactus 513, var. decipiens 514, var. Pylaei 514, var. solutus 340; Fauriensis 486-488; Ger- ardi var. Gerardi 513; Greenei 513; Longii 136; militaris 514; pelocarpus 514; Slwookoorum 486-488; tenuis 340, var. tenuis 513; validus 272 Juniperus communis var. de- pressa 4, 509; horizontalis 509 Justicia americana 170 Kallstroemia, in the Middle At- lantic States 397; intermedia 397; maxima 397; parviflora 397 Kalmia angustifolia 281, 523; lati- folia 14, 280, 281; polifolia 528 Kansas, Aster parviceps and A. brachyactis new for 392; Spir- anthes tuberosa new for 141 King, R. M. and H. Robinson, Studies in the Compositae-Eu- patorieae XV, Jaliscoa, Mac- vaughiella, Oaxacania and Planaltoa 100-105 Kondo, Katsuhiko, The Chromo- some Number of Obolaria vir- Index to Volume 72 569 ginica (Gentianaceae) 551-553 Kuhnia 38; eupatorioides 39, var. floridana 39, var. gracilis 39, var. pyramidalis 39; mosieri 38, 39; paniculata 39 Kylinia alariae 117, 200, 205, 237; secundata 117, 237, 523, 334; sp. 232; virgatula 201, 206, 237, f. luxurians 206, 237 Lactuca biennis 528; canadensis var. canadensis 528, var. lati- folia 528, var. longifolia 528 Laminaria 320, 322, 326; agardhii 321; digitata 117, 321, 33'; longicruris 117, 321, 337; sac- charina 117, 321, 397 Lantana 33; camara 34; depressa 34; ovatifolia 33, 35, var. ovati- folia 34, var. reclinata 34 Lapsana communis 528 Larix laricina 418, 509 Lathyrus japonicus var. glaber 520, var. pellitus 520; palustris var. pilosus 520; maritimus 372 Leathesia difformis 117, 321, 337 Lechea intermedia var. intermedia 522, var. juniperina 522 Lecocarpus 106-109 Ledum groenlandicum 523 Leersia oryzoides 511 Lemaire, Robert J., Recent Plant Records for Nebraska 283-284 Lemanea fucina 207 Lemna minor 9, 10; perpusilla 10; trisulea 513; valdiviana 10 Leontodon autumnalis var. au- tumnalis 528, var. pratensis 528 Leonurus Cardiaca 283, 524; sibiricus 283, 393 Lepidium campestre 518; densi- florum 518 Lescuraea saviana 500 Ligusticum scothicum 522 Ligustrum amurense 524 570 Limonium 24; angustatum 25; carolinianum 24, var. angusta- tum 24, var. carolinianum 24, 25, var. nashii 25, var. obtusi- lobum 25; nashii 24, 25, 523, var. angustatum 25, var. tricho- gonum 25; obtusilobum 24, 25; trichogonum 24, 25 Linaria canadensis 525; vulgaris 525 Linnaea borealis var. americana 526 Linum usitatissimum 520 Liparis Loeselii 515 Liquidambar styraciflua 79 Listera auriculata 47; ovata 274, in the Bruce Peninsula, On- tario 274 Lithophyllum corallinae 117, 211, 233, 324, 325, 334; macrocar- pum 210 Lithothamnium glaciale 117, 211, 238, 324, 334; lenormandi 212 Lobelia cardinalis 79; Dortmanna 526; inflata 526 Lolium multiflorum 510 Lomentaria baileyana 200, 219, 231, 232, 238; orcadensis 201, 219, 232, 238 Lonard, R. I. and F. W. Gould, Eragrostis plana in South Caro- lina 188 Long, Robert W., Additions and Nomenclatural Changes in the Flora of Southern Florida 17- 46 Lonicera canadensis 525; Mor- rowi 344; villosa var. calvescens 525, var. solonis 525 Lupinus 110; X alpestris 112, 114; argenteus 110-114, subsp. argenteus 111, var. argenteus 111, var. tenellus 111, subsp. rubricaulis 111, 112, subsp. spathulatus 111; caudatus 110, Rhodora [Vol. 72 112-114, subsp. argophyllus 113, 114, subsp. caudatus 112, subsp. cutleri 113, subsp. montigenus 113; cutleri 113; decumbens var. argophyllus 113; X inyoensis 114; montigenus 113; palmeri 114; parviflorus 111; polyphyl- lus 50, 520; rubricaulis 111; spathulatus 111; tenellus 111 Luzula acuminata 514; multiflora var. acadiensis 514, var. fus- conigra 8, 10, 514, var. multi- flora 514 Lychnis alba 517 Lycopersicon esculentum 393 Lycopodium annotinum var. acri- folium 507, var. annotinum 507; clavatum 1, var. clavatum 507, var. megastachyon 507; com- planatum var. flabelliforme 508; inundatum 507; lucidulum 1, var. lucidulum 507, var. occi- dentale 507; obscurum 281, f. dendroideum 507, f. obscurum 907; Selago var. appressum 507; tristachyum 508 Lycopus 354; americanus 79, 352, 358, 524, var. Longii 136; asper 164; europaeus 351-363, Distri- butional History in North America of 351, var. europaeus 353, var. integrifolius 353, var. mollis 353, var. sessilifolius 354, var. sinuatus 353; rubellus 353; sessilifolius 354; uniflorus 339, 524; virginicus 353 Lysimachia X producta 523; quadrifolia 523; spp. 492; ter- restris var. terrestris 523; vul- garis 546 Macvaughiella 100, 101, mexicana 101, 102, var. mexicana 101, var. standleyi 101, 102; stand- leyi 102 1970] Magnolia acuminata 491; fraseri 491 Magrath, Lawrence K., Spir- anthes tuberosa, New for Kan- sas 141 Maianthemum canadense 282, 514 Maine, Further Additions to the Bryoflora of Mt. Katahdin 493, The Flora of Isle au Haut 505 Malaxis Bayardi 136; unifolia 515 Malva neglecta 521 Mathieson, Arthur C. and Edward J. Hehre, Investigations of New England Marine Algae III: Composition, Seasonal Occur- rence and Reproductive Peri- odicity of the Marine Rhodo- phyceae of New Hampshire 194-235; Richard A. Stone, Ed- ward J. Hehre and Joan M. Conway, A Preliminary Check- list of the Marine Algae of Campobello Island, New Bruns- wick, Canada 313-338 Matricaria Chamomilla 527; mar- itima var. maritima 527; matri- carioides 527 Mazus japonicus 273 Medeola virginiana 493, 514 Medicago lupulina 520 Mehra K. L. and S. A. Faruqi, Biosystematics of Setcreasia brevifolia 264-271 Melampodium 107, 108; longifo- lium 108; paniculatum 98 Melobesia lejolisii 201, 211, 282, 238 Melothria 37; crassifolia 38; microcarpa 35; Nashii 38; pen- dula 37, var. aspera 37, 38; var. crassifolia 37, 38; var. micro- carpa 88; var. pendula 37, 38 Membranoptera alata 117, 200, 201, 224, 231, 232, 238, 324, 334; denticulata 224 Index to Volume 72 571 Mentha aquatica 546; arvensis var. arvensis 524, var. villosa 524; cardiaca 524 piperita 524 Melampyrum lineare var. ameri- canum 525, var. lineare 525 Mertensia maritima 524, 530 Metopium toxiferum 23 Mikania 4 Mimosa 534 Mitchell, Richard S., A Re-eval- uation of Polygonum meisneri- anum in North America 182- 188 Mitchella repens 281, 525 Mitreola sessilifolia 8 angusti- folia 28 Mnium 276, 277 Molinia caerulea 7, 510, 531 Monarda 502; didyma 492; fistu- losa var. mollis 502, 503; stipitatoglandulosa 503 Moneses uniflora 523 Monostroma fuscum 318, 331, f. blyttii 116; grevillei 116, 318, 331; leptodermum 116, 318, 321, 331; pulchrum 116, 318, 332 Monotropa Hypopithys 523 Monotropsis lehmaniae 55; odo- rata 55, var. lehmaniae 54, 55, var. odorata 55 Montia lamprosperma 517, 530 Muhlenbergia 534; sylvatica 79; uniflora 511 Myosotis arvensis 524; scorpioides 15; sylvatica 524 Myrcianthes dicrana 23; fragrans 22, 23, var. fragrans 23, var. simpsonii 22; simpsonii 23 Myrica Gale 426, var. subglabra 515; pensylvanica 515 Myrionema strangularis 117, 321, 337 Mytilus 199; edulis 208 Najus minor 164, 272 572 Nasturtium officinale 13 Nebraska, Recent Plant Records for 283 Nemalion helminthoides 200, 207, 238; multifidum 208 Nemopanthus 426; 281, 521 Nemophila microcalyx 400, An In- correct Name 399; triloba 400 Neotreleasea brevifolia 265 Nepeta Cataria 524 New Brunswick, A Preliminary Checklist of the Marine Algae of Campobello Island 313 New England, Hybrids in Rubus subgenus Eubatus in 240 New England Botanical Club, List of Members and Officers 287 seq. New Hampshire, Composition, Seasonal Occurrence and Re- productive Periodicity of the Marine Rhodophyceae in 194; Zonation of Bryophytes in the Headwaters of a Stream in 276 North America, A New Species of Calamovilfa from 72; A Re- evaluation of Polygonum meis- nerianum in 182; Distributional History of Lycopus europaeus in 351; Distributional History of Epilobium hirsutum in 164; Salix starkeana in 548; Some Aberrant Pyrola Collections from Eastern 480 Notholaena 584 Nuphar 54; variegatum 517 Nymphaea odorata 517 Nymphoides cordata 524 Nyssa sylvatica 79, 280, 281 mucronata Oaxacania 100, 103, 104; malvae- folia 108 Obolaria virginica 551, 552, The Chromosome Numbers of 551 Rhodora [Vol. 72 Odontites serotina 286, 525, A Range Extension for 286 Oenothera biennis var. biennis 522; cheiranthifolia 372; parvi- flora 522; perennis 522 Ohio, An Assemblage of Halo- phytes in Northern 285 Oklahoma, Monarda stipitato- glandulosa, A New Species from 502 Onoclea sensibilis 508 Ontario, Listera ovata Bruce Peninsula 274 Ophioglossum vulgatum var. pseudopodum 3 Orchis spectabilis f. Willeyi 48 Orobanche uniflora 15 Oryzopsis asperifolia 511; cana- densis 511 Osmunda cinnamomea 281, 508; Claytoniana 3, 282, 508; regalis 281, var. spectabilis 508 Ostrya 4; virginiana 2, 515, 554 Oxalis 387; barrelieri 395; eur- opaea var. europaea 521, f. cymosa 521; montana 521 Oxypolis rigidior 79 in the Panicum 535; amarum 5; an- nulum 6; boreale 511; capillare var. occidentale 511; clandes- tinum 79; dichotomiflorum 48, var. dichotomiflorum 48, var. geniculatum 48; lanuginosum var. implicatum 511, var. sep- tentrionalis 511; longifolium 272; mattamuskeetense 5, 6; microcarpon 6, 79; spretum 5; subvillosum 511; virgatum 5 Pantoneura baerii 201, 225, 238 Papaver Walpolei 488 Parthenocissus inserta 521; quin- quefolia 521 Passiflora edulis 396 Pellaea 534 1970] Peliia epiphylla 494; fabroniana 494 Percursaria percursa 318, 382 Petalonia fascia 117, 321, 397 Petasites palmatus 527 Petrocelis middendorfii 117, 199, 201, 214, 231, 232, 238, 324, 334 Peyssonelia rosenvingii 117, 201, 208, 238, 324, 334 Phillips, W. Louis and Ronald L. Stuckey, Distributional History of Lycopus europaeus (Euro- pean Water-Horehound) in North America 351-369 Phleum pratense 511 Phlox 54 Phragmites communis 546 Phycodrys rubens 117, 200, 225, 232, 233, 238, 324, 334 Phyllophora brodiaei 200, 216, 231, 238, 326; membranifolia 200, 201, 217, 233, 258, 326; spp. 211, 230; traillii 201, 217, 238 Phymatolithon compactum lenormandi 117, 211, 238 Picea glauca 509; mariana 509; rubens 281, 418, 509 Pike, Radcliffe B., Evidence for the Hybrid Status of Aster X Blakei (Porter) House 401-455 Pilayella littoralis 117 Pilea parietaria 345 Pinus elliottii 19; resinosa 509; rigida 509; Strobus 280, 281, 509; virginiana 79 210; Plagiochila asplenioides 276-275 Plagiothecium denticulatum 495 Planaitoa 100, 104; lychnopho- roides 104, 105; salviifolia 104 Plantago 387; altissima 525; junc- oides var. decipiens 525; lanceo- lata 525; maritima 372; major 393, var. major 525, var. Pil- geri 525; oliganthos 525 Index to Volume 72 573 Plumaria elegans 117, 200, 201, 223, 231, 232, 238, 324, 334 Poa alpina var. B 346; annua 570; bulbosa 346; compressa 510; nemoralis 510; pratensis 510; rupicola 498 Pogonatum alpinum var. septen- trionale 500 Pogonia ophioglossoides 514 Pohlia cardotii 500 Polygala boykinii 19, var. boy- kinii 19, var. sparsifolia 19, 20, var. suborbicularis 19; coral- licola 21; cumulicola 20; ery- throrrhiza 69, 70; evolvulacea 67, 68; flagellaris 20; glandu- losa 68; grandifolia 20, var. angustifolia 21, var. grandi- folia 20, 21, var. leiodes 21, var. leptophylla 21; Lind- heimeri 70; macradenia 67, 68, var. glanduloso-pilosa 67, 66; miamiensis 20; nitida 70; phoe- nicistes 68; praetervisa 20; sanguinea 521; sect. Adeno- phora 67, 68; sect. Eurhino- tropis 67, 68; sparsifolia 20; subsect. Adenophora 68; sub- sect. Eurhinotropis 69, 70; Tweedyi 69, 70 Polygonatum pubescens 514 Polygonum acanthophyllum 187; arifolium 182; aviculare var. littorale 516, var. vegetum 516; beyrichianum 182, 183, 186, 187; Careyi 516; chamissoanum 187; cilinode 516, f. erectum 516; Convolvulus var. Convolv- ulus 516, var. subalatum 516; geniculatum 183, 187; Hydro- piper 516; lapathifolium 516; meisnerianum 182-184, A Re- evaluation of 182, var. beyrichi- anum 182-187, var. jalapense 185-187, - var. meisnerianum 574 187, var. setosum 187, var. usterianum 187; multiangulare 187; Persicaria 516, var. ruder- ale 49; punctatum 516; rubri- caule 182, 187; sagittatum 182, 184, 516; scabrum 516; sect. Echinocaulon 182; stelligerum 182, 187; türkheimii 187 Polyides caprinus 212; rotundus 117, 201, 205, 212, 230, 233, 238, 324, 325, 334 Polypodium virginianum 4, 508; vulgare 4 Polysiphonia arctica 324, 825, 334; denudata 200, 201, 221, 231-233, 238; elongata 203, 227, 233, 238; flexicaulis 201, 227, 238, 324, 334; harveyi 227, 230; lanosa 117, 200, 202, 204, 213, 221, 231-233, 239, 323, 324, 334; nigra 228, 231, 233, 239; nigres- cens 228, 231-233, 239, 326; novae-angliae 200, 229, 231, 232, 239; subtillisima 200, 201, 229, 231, 239; urceolata 117, 229, 231-233, 239, 324, 334, var. patens 229, var. roseola 229, 324 Polystichum acrostichoides 4, 508; munitum var. munitum 496 Populus alba 344, 546; balsami- fera 515; X gileadensis 515; grandidentata 515; tremuloides var. magnifica 515, var. tremu- loides 515 Porphyra leucosticta 118, 200, 201, 203, 239, 324, 335; linearis 118, 200, 201, 203, 231, 239; miniata 118, 200, 203, 230-232, 239, 325, 335; umbilicalis 118, 119, 200, 204, 206, 231, 239, 325, 335, f. epiphytica 118, 204, 239, 325, 335, f. linearis 203 Porphyropsis coccinea 202, 239 Porter, Duncan M., Kallstroemia Rhodora [Vol. 72 in the Middle Atlantic States 397-398 Portulaca oleracea 517 Potamogeton Berchtoldii var. lacunatus 47; epihydrus var. Nuttallii 509; pectinatus 48; Oakesianus 509 Potentilla argentea 519; arguta 47; canadensis 519; Egedei var. groenlandica 519; fruticosa 416; norvegica 519; simplex var. calvescens 519, var. sim- plex 519; palustris var. villosa 519; pectinata 519 Prasiola stipitata 116, 318, 332 Prenanthes altissima 528, f. his- pidula 528; trifoliolata var. nana 528, var. trifoliolata 528 Prunella vulgaris var. lanceolata 524, f. iodocalyx 524 Prunus Avium 520; maritima 520; pensylvanica 520; virginiana 520 Pseudendoclonium marinum 116, 318, 332 Pseudobarleria sp. 546 Psoralea holosericea 66, 67; mex- icana 66, 67, var. Trianae 66, 67 Pteridium aquilinum 281, ssp. aquilinum var. pubescens 497, var. latiusculum 508 Ptilidium pulcherrimum 493 Ptilimnium 395; capillaceum 393, 394, Range Extension for 393 Ptilota serrata 118, 200, 215, 224, 230-232, 239, 324, 325, 335 Puccinellia maritima 509; pauper- cula var. alaskana 509 Punctaria latifolia 321, plantaginea 326 Pycnanthemum 502; pilosum 50; tenuifolium 50, var. Laurencei 50 Pylaiella littoralis 202, 321, 387 337; 1970] Pyrola 480, 483, Aberrant Collec- tions from Eastern North America of 480; asarifolia 483, 484, var. asarifolia 482, 484, var. purpurea 484; compacta 480, 481, 483, 484; elliptica 480, 481, 483, 523; grandiflora 484; minor 484; oxypetala 482-484; rotundifolia var. americana 528; secunda 523; virens 483, f. paucifolia 523 Pyrus americana 519; Aucuparia 519; decora 519; floribunda 519; Malus 519; melanocarpa 519 Quercus 534; rubra 12, 281, 516 Ralfsia borneti 322, 338; clavata 322, 338; fungiformia 117, 322, 338; pusilla 322, 338; verrucosa 322, 338 Ranunculus 565; abortivus 2; acris var. acris 517; bulbosus 517; Cymbalaria 517; ficaria 499; recurvatus 2; repens 517 Raphanus Raphanistrum 518; sat- ivus 518 Rhacomitrium heterostichum var. affine 494 Rhamnus frangula 14 Rhinanthus Crista-galli Crista-galli 525 Rhizoclonium riparium var. im- plexum 116, 318, 332; tortuo- sum 116, 318, 332 Rhodochorton purpureum 118, 200, 207, 230-232, 239, 325, 335 Rhododendron canadense 1, 14, 528; sp. 79 Rhododermis georgii 209 Rhodomela confervoides 118, 201, 230, 232, 239, 325, 335 Rhodophyceae 194-235 Rhodophyllis dichotoma 215, 230, 239, 318, 325 var. Index to Volume 72 575 Rhodophysema elegans 326; georgii 201, 209, 239, 325, 335 Rhodymenia palmata 118, 200, 206, 219, 231, 232, 239, 325, 335 Rhus radicans var. radicans 521; typhina 521 Rhynchospora alba 512; capitel- lata 9 Rhizomnium 494 Ribes glandulosum 518; hirtellum 339, 347, 948, var. calcicola 13, 347, 518, var. hirtellum 518; lacustre 518; oxyacanthoides 348, var. calcicola 347, 348; sat- ivum 518; triste 518 Robinson, H. and R. M. King, Studies in the Compositae- Eupatorieae XV: Jaliscoa, Mac- vaughiella, Oaxacania and Planaltoa 100-105 Rodman, James E. and Michael G. Barbour, Saga of the West Coast Sea-Rockets: Cakile ed- entula ssp. californica and C. maritima 370-386 Rogers, Ken E., A New Species of Calamovilfa (Gramineae) from North America 72-80 Rogers, O. M. and L. Michael Hill, Chromosome Numbers in Aster Blakei and A. nemoralis 437- 438 Rorippa islandica var. Fernald- iana 518; sylvestris 164, 360, 518 Rosa cinnamomea 520; nitida 520; palustris 79; rugosa 520; vir- giniana 520, X R. nitida 520 Rubus 240, 241, 547; aculiferus 247; adjacens 245; alleghanien- sis 242, 244, 245, 247, 249, 349, var. alleghaniensis 520; areni- cola 241, 244, 249; argutus 250; canadensis 248, 249, 520; cunei- pseudopunctatum 576 folius 250; elegantulus 242, 244, 246-249; Enslenii 243, 244, 249, 349; flagellaris 241, 242, 249; frondisentis 249, 520; frondosus 248, 244, 248, 249; Groutianus 246; hispidus 240, 242, 243, 245, 249, var. hispidus 520; idaeus var. canadensis 520, var. idaeus 519, var. heterolasius 520, var. strigosus 519; jacens 245; Jay- smithii 241, 242, 249; Longii 136; multiformis 243; odoratus 491; Parlinii 246; pensilvanicus 243, 244, 248, 249, 349, 520; per- mixtus 245; pubescens 519; recurvicaulis 241, 243, 244, 249, 520; sceleratus 247; semiseto- sus 244, 246-249, 520; setosus 240, 243, 245, 246, 249; severus 243; subgenus Eubatus 240, 249, Hybrids in 240; trifrons 245; univocus 246; vermontanus 248, 245-249 Rubus hybrids, alleghaniensis X frondosus 249, X pensilvanicus 249; arenicola X semisetosus 244; canadensis X alleghanien- sis 248, X elegantulus 248, x pensilvanicus 248; elegantulus X alleghaniensis 249; Enslenii X pensilvanicus 243, 547, X re- curvicaulis 243; flagellaris X alleghaniensis 242, X frondosus 242, 244; hispidus X alleghani- ensis 245, XX setosus 245, X vermontanus 245; Jaysmithii X elegantulus 242, X hispidus 242; recurvicaulis X alleghan- iensis 244, X frondosus 244, X elegantulus 242, 244, X his- pidus 243, X pensilvanicus 249, X setosus 243, 520, X vermont- anus 243; semisetosus X fron- dosus 248; setosus X alleghan- iensis 247, 520, X elegantulus Rhodora [Vol. 72 246, X semisetosus 246, X ver- montanus 246, 247; vermont- anus X alleghaniensis 248, X elegantulus 247, X semiseto- sus 247 Rudbeckia serotina 527 Rumex Acetosella var. Acetosella 516; altissimus 516; crispus 393, 516; domesticus 516; orbicula- tus 516; pallidus 516 Ruppia maritima 206 Sabal palmetto 19 Sabatia 551; bartramii 57; dode- candra 57, var. coriacea 57, var. dodecandra 57, var. kennedy- ana 57; foliosa 57; harperi 57; kennedyana 57 Sacchoriza dermatodea 323, 338 Sagina nodosa var. pubescens 517; procumbens 517 Sagittaria graminea 509 Salicornia europaea 285, var. eu- ropaea 516, var. prostrata 516 Salix 565; alba var. vitellina 515; Bebbiana 339, 515, 548, 549, var. perrostrata 549; coactilis 515; cordata 49, 339; depressa 548, 549, subsp. rostrata 548, c. rostrata 549; discolor 515; fusca 549; gracilis var. textoris 515; humilis 515; lanata 548; livida 549, occidentalis 549; lucida var. lucida 515; myrtilli- folia 498; perrostrata 549; phylicifolia var. pennata 498; rigida 49, 515, var. angustata 49; rostrata 548, 549; spp. 426; starkeana 548-550, in North America 548, subsp. bebbiana 549, 550, subsp. starkeana 548, 550; subsericea 49; vagans 549, l cinerascens b. occidentalis 549, occidentalis 549, 1 S. ros- trata 549, 3 obovata 549; ves- 1970] tita var. erecta 498 Salsola Kali var, Kali 516, var. tenuifolia 378 Salvia coccinea 393 Sambucus canadensis 79, 526; pubens 526 Sanchezia 119; leucerythra 119, 120 Saponaria officinalis 517 Sarracenia purpurea 518 Saurauia aequatoriensis 86, 90, 92; biserrata 86, 90, 92; brachy- botrys 86, 90, 92; bullosa 82, 84, 87, 88, 90, 92; chiliantha 87, 90, 92; Humboldtiana 86, 90, 92; isoxanthotricha 84, 86, 90, 92; omichlophila 82, 84, 90, 92; pastasana 86; peduncularis 88, 90, 92; portachuelensis 86, 87, 90, 92; Prainiana 90, 92; putamayonis 90, 92; Species and their Chromosomes $81; Stapfiana 86, 90, 92; tomentosa 86, 87, 90, 92; ursina 90, 92 Saxifraga nudicaulis 486, 488; rivularis 486 Seapania nemorosa 277-279; un- dulata 276-279 Schaetzellia 101; standleyi 102 Scirpus 565; acutus 511; ameri- canus 511; ancistrochaetus 272; atrocinctus 512; atrovirens var. georgianus 512; cespitosus var. callosus 511, 520; cyperinus var. cyperinus 512, var. pelius 512, f. condensatus 512; Longii 133, 136; paludosus var. atlanticus 512; pedicellatus 512; rubro- tinctus 512 Scrophularia lanceolata 525 mexicana 102; Scutellaria epilobiifolia 524; lat- eriflora 524; sp. 79 Scytosiphon lomentaria 117, 322, 338 Index to Volume 72 577 Secale cereale 510 Sedum 460; beyrichianum 467; glaucophyllum 460-462, 463, 465-472, 474-476, Heteroploidy in 460; nevii 460, 465-469; pulchellum 460, 462, 463, 465, 469; purpureum 518; rosea 460, 518, 530; ternatum 460, 462, 463, 465-469, 471, 474, 475 Selaginella rupestris 508 Sempervivum tectorum 518 Semple, John C., The Distribution of Pubescent Leaved Individ- uals of Conocarpus erectus (Combretaceae) 544-547 Senecio cuneifolius 99; desider- abilis 99; hemmendorffii 99; icoglossus 99; Robbinsii 528; sylvaticus 528; vulgaris 528 Serratula caroliniana 146, 154; glauca 147, 150; noveboracen- sis 146, 153; praealta 146, 153 Setaria glauca 511; viridis 511 Setcreasea 268; australis 268; brevifolia 264, 265, 267, 269, Biosystemics of 264, var. brevi- folia 265, 266, 268, 269, var. buckleyi 265, 267-269, var. nanella 265, 266, 269, var. pul- chella 265, 266, 269; buckleyi 264, 267, 268; ovata 264, 265; pallida 264, 268; purpurea 264, 268; tumida 268 Settle, Wilbur J. and T. Richard Fisher, The Varieties of Sil- phium integrifolium 536-543 Seymour, Frank C., The Flora of New England (Review of) 157; Notes from the Pringle Her- barium III. 47-50 Sharpiella striatella 493 Shontz, John P. and Nancy N., Galinsoga ciliata (Compositae): Its Arrival and Spread in the Northeastern United States 578 386-392 Shontz, Nancy N. and John P., Galinsoga ciliata (Compositae): Its Arrival and Spread in the Northeastern United States 386-392 Sida rhombifolia 393 Silene Armeria 517; caroliniana 59-62, ssp. caroliniana 59-62, var. caroliniana 62, 63, var. pensylvanica 59, 61-63, ssp. pensylvanica 64, ssp. wherryi 59-62, 64 var. wherryi 64; gal- lica 273; noctiflora 517; pensyl- vanica 62, 64; virginica 60; wherryi 60 Silphium 536; integrifolium 536- 542, The Varieties of 536, var. deamii 539, 540, 542, var. gat- tingeri 539, 540, 542, var. in- tegrifolium 539, 542, var. neg- lectum 539, 540, 542; laeviga- tum 538, 539; scaberrimum 540; speciosum 540 Simsia dombeyana 94, 98 Sisymbrium altissimum 518; of- ficinale var. leiocarpum 518 Sisyrinchium, A New Argentine 287; avanaceum 287; Marchio 287; montanum var. crebrum 514; papillosum 287; Wett- steinii 287 Smilacina racemosa 3839, 514; stellata 330; trifolia 514 Smilax 289-293, 295, 297, 301, 303, 306, 311; auriculata 289, 290, 294, 295, 297, 299, 304, 305, 307, 309-311; beyrichii 290; bona- nox 289, 290, 294, 295, 297-300, 304-312, var. littoralis 306; glauca 2, 289, 290, 294, 295, 297-299, 303-305, 307, 308, 310- 312; hispida 290; lanceolata 290; laurifolia 289, 297-300, 303-307, 309-311; pseudo-china Rhodora [Vol. 72 290; pumila 289, 311; rotundi- folia 2, 289, 294, 295, 297, 299, 304, 308-312; sect. Smilax, Comparative Studies of 289; smallii 289, 290, 294-297, 299, 300, 304, 307, 309-311; tam- noides 289, 290, 297, 299, 300, 303-306, 308-312, var. hispida 289, 296-299, 303, 307-309, var. tamnoides 289, 296, 297, 299, 307-209; walteri 289, 296, 297, 299, 304-307, 309, 311, 312 Soejarto, Djaja D., Saurauia Spe- cies and their Chromosomes 81- 93 Solanum americanum 393; Dulca- mara 524; nigrum 524; rostra- tum 524 Solbrig, Otto T. and Warren G. Abrahamson, Soil Preferences and Variation in Flavanoid Pig- ments in Species of Aster 251- 263 Solidago 565; bicolor 526; caesia 350; canadensis 526; gramini- folia var. Nuttallii 526; juncea 526; nemoralis 526; puberula 526; rugosa 79, var. villosa 526; sempervirens 526; uliginosa var. linoides 526, var. terrae- novae 526 Sonchus arvensis 528; asper 528; oleraceus 528; uliginosus 50 South Carolina, Eragrostis plana in 188 South Dakota, Trillium cernuum L. and Geranium maculatum New for 554 Sparganium americanum 509; an- gustifolium 509; chlorocarpum var. acaule 509, var. chlorocar- pum 509; minimum 509 511; X 511; alterniflora patens Spartina caespitosa 511; pectinata 511 1970] Spergula arvensis 517 Spergularia canadensis 517; mar- ina var. leiosperma 517; rubra 517 Spermacoce 36; floridana 36; key- ensis 35; portoricensis 34; tenuior 36, var. floridana 36, var. tenuior 36; tetraquetra 36, 37 Spermothamnion repens 224, 231 239, 325, 335; turneri 224 Sphacelaria cirrosa 117, 322, 338 Sphagnum 280, 281 Spilanthes acmella 95, 98; amer- icana 95 Spiraea Bumalda cv. Anthony Waterer 519; latifolia 519; tomentosa 519; ulmaria 546 Spiranthes cernua var. cernua 515, var. ochroleuca 515; graci- lis 514; Grayii 11; Romanzof- fiana 515; tuberosa 141 Spirodela polyrhiza 10 Spongomorpha 324; arcta 116, 318, 332; hystrix 319, 332; lanosa 116; spinescens 116, 319, 332 Spongonema tomentosum 322, 338 Sporobolus cryptandrus 7 Stachytarpheta jamaicensis 393 Statice brasiliensis var. angusta- tum 25 Steele, Frederic L, A New Mt. Washington Station for Gna- phalium supinum 555-556; and A. R. Hodgdon, Hybrids in Rubus subgenus Eubatus in New England 240-250; Correc- tion in article on Rubus Hy- brids 547 Stellaria calycantha var. flori- bunda 517, var. isophylla 517; graminea 517; media 517 Stevens, Charles E., Chrysanthe- mum lacustre Brotero New to Index to Volume 72 579 Gray’s Manual Range 250; A Prior Record of Chrysanthe- mum lacustre Brotero 543 Stevia 95; organensis 95, 97 Stictocardia tiliifolia 32 Stone, Richard A., Edward J. Hehre and Joan M. Conway, Flora of the Wolf Islands, Part III: The Marine Algae 115-118; Edward J. Hehre, Joan M. Con- way and Arthur C. Mathieson, A Preliminary Checklist of the Marine Algae of Campobello Island New Brunswick, Canada 313-338 Streptanthus squamiformis 503 Streptopus roseus var. perspectus 514 Strongylocentrotus drobachiensis 326 Stuckey, Ronald L., Distributional History of Epilobium hirsutum (Great Hairy Willow-Herb) in North America 164-181; and W. Louis Phillips, Distributional History of Lycopus europaeus (European Water-Horehound) in North America 351-369 Stuessy, Tod F., The Genus Acan- thospermum (Compositae-Hel- ientheae-Melampodinae): Tax- onomic Changes and Generic Affinities 106-109 Stylisma 32; abdita 32 Suaeda linearis 516; maritima 516 Suessenguthia 119, 120, A Synop- sis of the Genus 119; cuscoensis 120-123; leucerythra 119, 120; trochilophila 119, 120, 124; var- gasii 119-121, 123 Suprago glauca 150 Svenson, Henry K., A Linden (Tilia) Forest on Cape Cod (with Extended Notes on Tilia neglecta, Bromus pubescens and 580 Ribes hirtellum) 339-350; West- ern Cape Cod: Plant Notes 1-16 Swertia 551 Symphoricarpus albus var. albus 525, var. laevigatus 526 Symplocarpus foetidus 513 Syringa vulgaris 524 Taeiniatherum asperum 378 Tanacetum vulgare 527, f. cris- pum 527 Taraxacum erythrospermum 528; officinale var. officinale 528, var. palustre 528 Taxus canadensis 508 Taylor, Ronald J. and George W. Douglas, Contributions to the Flora of Washington 496-501 Teucrium occidentale var. boreale 524 Thalictrum polygamum 339, 517; ravolutum 13, 339 Thelypteris dryopteris 281; nove- boracensis 508; palustris var. pubescens 508; Phegopteris 508; simulata 339, 508, 529 Thieret, John W., Nemophila microcalyx, an Incorrect Name 399-400 Thlaspi arvense 518 Thuidium delicatulum 277 Thuja occidentalis 274, 509 Tiaridium indicum 34 Tilia 339, 340, 343, 350; ameri- cana 340-344, 554, B pubescens 341, heterophylla 341, var. neglecta 340, var. vestita 341; europaea 344; glabra var. neg- lecta 341; heterophylla 341-344; Michauxii 341; neglecta 339- 344, 350; nigra 340, ñ 341, ves- tita 341 Tracaulon 187 Tradescantia brevifolia 265; lei- andra 265, var. brevifolia 265, Rhodora [Vol. 72 267, var. ovata 264; 264, 267 Trailliella intricata 201, 208, 239 Treleasea brevifolia 265 Trichogonia 94 Trichostomum tenuirostre 494 Trientalis 340; borealis 282, 489- 492, 523, The Southern Limits of 489 Trifolium 387; agrarium 520; ar- vense 520; hybridum var. ele- gans 520; pratense 520, f. pil- osum 520; procumbens 520; repens 520 Triglochin maritima 509; palus- tris 509 Trillium 2; cernuum 11, 554, New for South Dakota 554; erectum 492; grandiflorum 492; undula- tum 282, 514; vaseyi 492 Triosteum 344, 347; aurantiacum 349 Triticum aestivum 510 Tryon, Rolla, Manual of the Vas- cular Plants of Texas (Book Review) 533-535 Tsuga canadensis 5, 280, 281, 509; heterophylla 497 Typha angustifolia 285; latifolia 509 speciosa Uhl, Charles H., Heteroploidy in Sedum glaucophyllum 460-477 Ulmus americana 554; rubra 554 Ulothrix flacca 319, 332 Ulva lactuca 116, 319, 321, 332 United States, Comparative Studies of Smilax, section Smilax, in the Southeastern 289; Galinsoga ciliata: Its Ar- rival and Spread in the North- eastern 386 Urechites 30; lutea 30, var. lutea 30, 31, var. sericea 30, 31; jaegeri 31; pinetorum 31 1970] Urospora 317; collabens 116, 319, 332; penicilliformis 116, 319, 332; speciosa 116, 319, 332; wormskjoldii 116, 319, 332 Urtica dioica 516; urens 516; vir- idis 516 Utah, An Undescribed Species of Astragalus from 189 Utricularia cornuta 525; gemini- scapa 525; inflata 58, var. minor 58; radiata 58 Uvularia sessilifolia 281 Vaccinium angustifolium 280, 281, var. angustifolium 5238, var. laevifolium 523, var. nigrum 523; corymbosum 281, var. al- biflorum 523; gaultherioides 443, 446, 449; macrocarpon 523; microphyllum 450; myrtilloides 523; occidentale 454; Oxycoc- cus 523; pubescens 448; salici- num 443, 453; uliginosum 439- 450, 452, 453, 455, 456, f. pubes- cens 448, ssp. gaultherioides 443, 447, 449-451, 455, 456, ssp. occidentale 454-457, var. occi- dentale 448, 453-455, 457, var. salicinum 448, 453-455, 457, ssp. pedris 443, 447, 448, 450, 452, 455, 457, ssp. pubescens 447- 449, 452, 455, 456, ssp. uligi- nosum 447, 450, 452, 453, 455, var. alpinum 448, f. Langeanum 448, var. gaultherioides 450, var. Langeanum 448, var. ped- ris 450, var. pubescens 448, * microphyllum 450, Taxonomy and Distribution of 439; Vitis- Idaea var. minus 523 Vaucheria sp. 517 Verbascum Thapsus 524 Verbena bonariensis 393; hastata 524 Verbesina encelioides 396 Index to Volume 72 581 Vernonia 145, 146, 155; acaulis 145-151, 159, 161, 162; altis- sima 161; angustifolia 146, 148- 150, 161, 162; flaccidifolia 152; georgiana 146; glauca 145, 147- 152, 159-162, f. longiaristata 147, 150, 152; harperi 147, 154- 156; noveboracensis 145-158, 155-158, 160, 162, f. albiflora 154, f. lilacina 154, var. lati- folia 147, 150, 154, var. praealta 154, [var.] tomentosa 147, 150, 154; oligophylla 146, 148; praealta 146, 154; rugeliana 147, 154; tomentosa 147, 154 Veronica arvensis 340, 525; hed- eraefolia 283; officinalis 525; peregrina var. xalapensis 525; serpyllifolia 525 Viburnum alnifolium 281; cassi- noides 281, 526; recognitum 526; venosum 340 Vicia angustifolia var. segetalis 520; Cracca 520 Viola 535; blanda 522; cucullata 522; fimbriatula 522; incognita var. Forbesii 522, var. incognita 522; lanceolata 522; pallens 522; primulifolia var. acuta 522; septentrionalis 522 Viorna baldwinii 18 Vrginia, New and Overlooked Species of the Flora of 272 Vitis aestivalis 339 Washington, Contributions to the Flora of 496 Wasshausen, Dieter C., A Synop- sis of the Genus Suessenguthia (Acanthaceae) 119-125 Waterfall, U. T., Monarda stipi- tatoglandulosa, a New Species from Oklahoma 502-504 Welsh, Stanley L., An Unde- scribed Species of Astragalus issouri 582 (Leguminosae) from Utah 189- 193 Wilbur, Robert L., Infraspecific Classification in the Carolina Flora 51-65 Wise, David A., The Flora of Isle au Haut, Maine 505-532 Wulffia stenoglossa 98 Xyris Bayardi 136; caroliniana 6; torta 6, 79 Yates, Ida E. and Wilbur H. Dun- can, Comparative Studies of Smilax, section Smilax, of the Southeastern United States 3 1753 00341 3942 Rhodora [Vol. 72 289-312 Young, Steven B., Juncus Slwoo- koórum, a New Species from Alaska 486-488; On the Tax- onomy and Distribution of Vac- cinium uliginosum 439-459 Youngberg, Alv Dan, Salix stark- eana in North America 548-550 Zebrina leiandra 265 Zenobia cassinefolia 56; pulveru- lenta 56 Zostera 207, 211, 219, 226; mar- ina 209, 318, 321, 325, var. stenophylla 195, 509