Douora

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by REED CLARK ROLLINS, Editor-in-Chief

ALBERT FREDERICK HILL

STUART KIMBALL HARRIS

RALPH CARLETON BEAN Associate Editors RICHARD ALDEN HOWARD

CARROLL EMORY WOOD, JR.

VOLUME 57

1955

Che New England Botanical Club, Ine.

8 and 10 West King St.. Lancaster, Pa. Botanical Museum, Oxford St., Cambridge 38, Mass.

Douora

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by REED CLARK ROLLINS, Editor-in-Chief

ALBERT FREDERICK HILL

STUART KIMBALL HARRIS

RALPH CARLETON BEAN Associate Editors RICHARD ALDEN HOWARD

CARROLL EMORY WOOD, JR.

Vol. 57 January, 1955 No. 673

CONTENTS:

A Revision of the North American Genus Sabatia (Gentianaceae). Boben E e Soa Oe i a a 0s ee 1

Ilex glabra and a New Station for Kalmia latifolia in New Hamp- shire. A. R. Hodgdon...... 01 eerie eee rrr 34

A New Variety of Solidago ulmifolia. Arthur Cronquist......... 36

The New England Botanical Club, Ine.

8 and 10 West King St., Lancaster, Pa. Botanical Museum, Oxford St., Cambridge 38, Mass.

RHODORA.—A monthly journal of botany, devoted primarily to the flora of the Gray’s Manual Range and regions floristically related. Price, $4.00 per year, net, postpaid, in funds payable at par in United States currency in Boston; single copies (if available) of not more than 24 pages and with 1 plate, 40 cents, numbers of more than 24 pages or with more than 1 plate mostly at higher prices (see 3rd cover-page). Back volumes can be supplied at $4.00. Some: single numbers from these volumes can be supplied only at advanced prices (see 3rd cover-page). Somewhat reduced rates for complete sets can be obtained on application to Dr. Hill. Notes and short scien- tific papers, relating directly or indirectly to the plants of North America, will be considered for publication to the extent that the limited space of the journal permits. Illustrations can be used only if the cost of engraver’s blocks is met through the author or his institution. Forms may be closed five weeks in advance of publica- tion. Authors (of more than two pages of print) will receive 15 copies of the issue in which their contributions appear, if they re- quest them when returning proof. Extracted reprints, if ordered in advance, will be furnished at cost.

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GRAY HERBARIUM of Harvard University, Cambridge 38, Mass., U. S. A.

QTRbooora

JOURNAL OF

THE NEW ENGLAND BOTANICAL CLUB

Vol. 57 January, 1955 No. 673

A REVISION OF THE NORTH AMERICAN GENUS SABATIA (GENTIANACEAE)!

Rosert L. WILBUR

THE genus Sabatia is restricted in distribution to continental North America and the West Indies. It is frequently encoun- tered throughout much of its range, which extends from Nova Scotia south as far as the mountains of Hispaniola and the central plateau of Mexico and from the Atlantie seaboard westward to the prairies of Oklahoma and central Texas. All of the members of the genus recognized in this paper, seventeen species and two varieties, are to be found in the southeastern United States. The distribution of most of them is either entirely restricted to that area or is largely centered there. Most of the entities are either found in or are restricted entirely to the Coastal Plain, the oldest portions of which have been avallable to the occupancy of land plants only since the close of the Cretaceous.

This genus is believed to be an ancient one whose intrageneric divisions appear to have been clear-cut and well-defined for a long period of time. Even the species seem, in most instances, to be genetically so well isolated from their closest relatives as to show no indication of hybridization even when flowering in close proximity to one another. These are almost certainly indications of great age.

This group of plants has not been monographically treated since the time of Grisebach (1845) although it has been subjected

1 Part of a dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the University of Michigan. I should like to take this opportunity to express my appreciation for the friendship, encouragement and guidance of Dr. Rogers McVaugh throughout this investigation.

2 Rhodora [Vor. 57

to numerous regional interpretations. The present treatment is based upon the examination of more than 6500 specimens borrowed from over thirty herbaria? and the prized opportunity, made possible largely through the assistance of Professor H. H. Bartlett, director of the Botanical Gardens of the University of Michigan, of studying all but one of the species in the field.

HISTORY OF THE GENUS

Plants now considered to belong to Sabatza were first described as species of Gentiana in Gronovius’ “Flora Virginica” (1739). Linnaeus, having assisted Gronovius in that work, was personally familiar with the specimens upon which the two polynomials were based and later provided binomials for both of them in “Species Plantarum” (1753). Two additional species, based upon specimens collected by Kalm, were described for the first time in that work. Three of these species were placed by Linnaeus in the genus Chironia together with four very different species from the Old World while the remaining species known to him was assigned to the genus Swertia. Later he transferred one of the species that he had formerly assigned to Chironia to Chlora and thus the four different species known to Linnaeus were treated as representatives of three different genera.

Adanson (1763), resisting what seemed to him to be the ex- cessive generic “lumping” of Linnaeus, recognized many pre- Linnean genera and initiated some new ones as well. Sabatza,

2 The abbreviations employed in referring to these herbaria are those of Lanjouw and Stafleu (1952) and are as follows:

BRU Brown University NCU University of North Carolina

CHARL Charleston Museum NO Tulane University

CU Cornell University NY New York Botanical Gardens

DENN Dennison University OKL University of Oklahoma

DUKE Duke University OKLA Oklahoma A. & M. College

F Chicago Natural History PENN University of Pennsylvania Museum RUT Rutgers University

FLAS Florida Agr. Exp. Sta. SMU Southern Methodist Univ.

GA University of Georgia TAES A. & M. College of Texas

GH Gray Herbarium TENN University of Tennessee

K Kew (part of collection) TEX University of Texas

KSC Kansas State College UARK University of Arkansas

MICH University of Michigan UC University of California

MO Missouri Botanical Garden Us U. S. National Herbarium

MT Université de Montréal WIS University of Wisconsin

NA U. S. National Arboretum WVA West Virginia University

NCS North Carolina State College YU Yale University

1955] Wilbur,—The North American Genus Sabatia 3

one of the latter, was entirely based upon “Gentiana Gron. virg. 29" which Linnaeus had called Chironia dodecandra. Later authors frequently ‘corrected’? Adanson’s spelling of the generic name to Sabbat?a since it was proposed in commemora- tion of an Italian botanist called Liberato Sabbati. In the "Familles des Plantes" the generic name Sabatia is invariably spelled with a single “b” while the botanist’s name always appears as Sabbati. One can therefore only conclude that Adanson deliberately chose to spell the generic name as Sabatia and, according to Article 82 of the International Code of Botan- ical Nomenclature, his choice in this matter is binding.

Pursh (1814) accepted Adanson’s genus but defined it to include all of the American species which had been placed by most earlier authors in the genus Chironia. Grisebach (1839, 1845) provided a very complete account of the eleven species and one variety that were known to him in his two monographic treatments of the Gentianaceae. These monographs were very similar in regard to the species of Sabatza, with the excep- tion of the disposition of S. gentianoides. In the first account this species was considered to form a monotypic section while six years later its distinctive characteristics were warranted to be of generie status and, furthermore, it was then placed in a different subtribe from that of its former congeners.

Gray's understanding of those species found within the United States was summarized in the ‘‘SSynoptical Flora" (1878) in which he recognized thirteen species and two varieties.

Blake (1915) provided a formalized system of classification for the genus by arranging the twenty-two species and one variety accepted by him into named subgenera, sections and subsections. This was in large part merely the application of subgenerie names and validating diagnoses to the groups of species that long had been associated together under the synoptical and indented keys of earlier authors.

GENERIC RELATIONSHIPS

Sabatia was associated in Gilg’s (1895) treatment of the family in the tribe Gentianeae, subtribe Erythraeinae together with seventeen other genera. Most of the genera of the Ery- thraeinae are composed of but few species and more than half

4 Rhodora [Vor. 57

of the genera are known only from the Old World. None of them seem particularly closely related to Sabatia. This genus has been placed most often near Centaurium Hill ( — Erythraea Borkh.) from which it may be distinguished by many characters. 'The following key employs some of their more obvious dif- ferences.

Corolla rotate, the lobes much longer than the tube; stigma deeply 2-lobed, the lobes at least one-third the length of the style, at anthesis spirally twisted about one another but uncoiling and spreading apart with time; anthers either half-twisted laterally or becoming circinnately coiled or recurved after pollen discharge... SABATIA.

Corolla salverform, the lobes equal to or shorter than the tube; stigma capitate or but barely creased, the divisions scarcely discernible, position unchanged with age; anthers becoming tightly spirally coiled in the manner of a cork- screw after pollen discharge... esee CENTAURIUM.

SYSTEMATIC TREATMENT SABATIA Adans., Fam. Pl. 2: 503. 1763.

Pleienta Raf., Fl. Tell. 3: 30. 1837.

Neurola Raf., New Fl. 4: 92. 1838.

Erect, glabrous, annual or perennial herbs. If perennial, sometimes stoloniferous and either arising from a short, erect caudex or from a short to elongate, almost always branched rhizome. Root-system of the an- nuals either a slender tap-root or a cluster of fibrous roots, that of the perennials a cluster of wiry to fleshy roots. Leaves mostly cauline but basal leaves also present in some species, decussate (or in aberrant speci- mens whorled), entire, sessile or somewhat amplexicaul, membranous to thick and turgid or even fleshy. Branches opposite or alternate. Inflorescence cymose, the cymules either compactly subeapitately clus- tered or loosely arranged, being often reduced to one or two flowers, or grouped into large, showy, compound dichasia. Flowers perfect, actino- morphic. Calyx 5-12(-14)-parted, gamosepalous, the tube campanulate to somewhat turbinate, membranous, the lobes minute to foliaceous, varying in length from shorter than the calyx-tube to many times longer. Corolla rotate, 5-12(-14)-parted, the tube urceolate to cylindrical, short but equaling or more commonly exceeding the calyx-tube; corolla- lobes usually widely spreading, dextrorsely contorted in bud. Stamens 5-12; filaments slender, inserted on the upper edge of the corolla-tube, alternating with the corolla-lobes, at first erect but strongly curving to one side after pollen discharge; anthers basally attached, linear to oblong, slender to stout, dehiscing laterally by means of longitudinal slits, after pollen discharge becoming either laterally twisted or circinnately re- curved or revolute. Ovary bicarpellate, unilocular; carpel margins slightly intruding into locule and forming 4 parietally located placental lobes each bearing numerous scattered ovules. Style slender; stigmatic

1955] Wilbur,—The North American Genus Sabatia 5

branches 2, linear to slightly spatulate, tightly spirally twisted at anthesis and bent to one side, later becoming erect, untwisting and exposing the densely papillate stigmatic surface. Capsule globose, ovoid, or cylindrical. Seeds numerous, globose or somewhat flattened, densely pitted with small depressions. TYPE SPECIES: Sabatia dodecandra (L.) BSP. (Chironia dodecandra L.)

KEY TO THE SECTIONS

Flowers usually long-pedicellate or, if short-pedicellate, the subtending bracts small and scale-like; flowers either solitary or loosely cymose; anthers becoming circinnately recurved after pollen discharge; plants, if annual, pentapetalous, if perennial, either pentapetalous or pluripetalous.

I. EUSABATIA.

Flowers sessile or very nearly so, the subtending bracts large and leaf-like; inflorescence of few to several closely associated flowers, appearing capitate; anthers slightly twisted laterally even prior to pollen discharge, never be- coming circinnately recurved; plants annual and pluripetalous.

II. PSEUDOCHIRONIA.

I. Section EvsABATIA Griseb., Gen. et Sp. Gent. 120. 1839.

Subg. Eusabatia (Griseb.) Blake, Rnopora 17: 56. 1915.

Annuals or perennials. Flowers either pentamerous or plurimerous (only the perennials 7-12(-14)-merous); typically long—pedicellate or at least not appearing to be sessile and capitately arranged with the clusters possessing large foliaceous bracts. Anthers becoming revolutely coiled or recurved after pollen discharge. TYPE species: Sabatia dodecandra (L.) BSP.

The fifteen species belonging to this, the larger, section of the genus show considerable diversity and may be arranged readily into several groups of species (subsections).

KEY TO THE SUBSECTIONS A. Calyx-tube very prominently 5-nerved and thinly membranous-winged ; the lateral nerves of the calyx-lobes much more strongly developed than the midnerve; flowers pentamerous; annuals......... (C) CAMPESTRES. A. Calyx-tube smooth or finely nerved and lacking membranous wings; the midnerve of the calyx-lobes almost equaling or even more strongly developed than the lateral nerves; annual or perennial and flowers either pentamerous or plurimerous. B. Perennials with elongate rhizomes; some species stoloniferous; either pentamerous or plurimerous.

C. Branches typically opposite throughout; flowers mostly 5-parted, the centermost in the cymules occasionally 6-parted; corolla white and lacking yellow “eye” at base of lobes; pedicels usually less than 1 em. long; stolon and basal rosettes never present. . . (A) DIFFORMES. C. Branches typically alternate (at least the uppermost); flowers 5-12 (-14)-parted (3 species 7-14-parted, the other species 5-6(-7)- parted); corolla typically pink or rose and with a yellow “eye” in the throat and base of the lobes; pedicels mostly over 1 em. long;

stolons and basal rosettes present in some species. (E) DODECANDRAE

6 Rhodora [Vor. 57

B. Annuals or, if perennials, the caudex short, erect, and much-branched; rhizomes and stolons never present; flowers pentamerous. D. Upper branches typically opposite; basal rosette present at some stage

of life-cycle and often persistent; annuals........ (B) ANGULAREs. D. Upper branches typically alternate; basal rosette lacking; either annuals or perennials. ..................... (D) CAMPANULATAE.

(A) Dirrormes subsect. nov., Subsect. Angulares Blake in part, RHopora 17: 56. 1915.

Robust perennials arising from a thick, gnarled, often much-branched, elongate rhizome bearing numerous fleshy roots about 2 mm. in diameter. Branches typically opposite. Leaves drying thick-chartaceous, margins revolute. Corolla white. TYPE species: Sabatia difformis (L.) Druce.

The two species associated in this subsection are strongly differentiated morphologically from the annual species of sub- section. Angulares with which they were formerly placed. In- cluding these perennials with the radically different species belonging to the Angulares would be placing undue emphasis upon the single character of an opposite-branching pattern that is typical of both groups while underestimating features such as the perennial habit and fleshy roots which seem to me to be of greater phyletie significance. The habitat oceupied by the perennials is typically that of savannas and ditches, whereas the annuals of the Angulares are generally inhabitants of drier situations.

The three entities forming subsection Difformes are a con- spicuous floristic feature of savannas or pine-barrens, or the marshy borders of low hardwood forests throughout their com- bined extensive range. "These plants are, with the exception of a very few Piedmont stations at the outer margin of that province, entirely restricted to the Coastal Plain, ranging from central New Jersey southward throughout much of peninsular Florida and westward into Louisiana. The three populations do not possess mutually exclusive areas but occupy ranges in the nature of overlapping ares somewhat radially arranged about what may have been the ancestral home of the group, the ancient Appalachian land-mass. It may be speculated that the original population, which was perhaps polymorphic,

3 Subsectio Difformes. Herbae perennes, rhizomatibus crassis, contortis, saepe multiramosis radicibus carnosis ca. 2 mm, diam. Rami plerumque oppositi. Folia in

speciminibus siccis crasse chartacea, marginibus revolutis. Corolla alba. Species typica, Sabatia difformis (L.) Druce.

1955] Wilbur,— The North American Genus Sabatia 7

migrated from Appalachia onto the Coastal Plain. The frag- ments of the original population that reached the new home continued to evolve and to spread laterally along the Coastal Plain. When they had reéstablished partial contact with their close relatives, genetical and other barriers had developed which maintained the identity of the three entities.

KEY TO THE SPECIES OF SUBSECTION DIFFORMES Leaves and upper portion of the stem not glaucous; calyx-lobes typically more than 4 mm. long and often 10 mm. long or longer; upper bractlets linear, those subtending complete (i.e. 3-flowered) cymules 4-5 mm. or more in length; corolla-lobes almost invariably longer than 7 mm. and often up to 15 mm. and occasionally longer, their color often turning saffron or orange upon drying, especially along the veins; plants rarely taller than 9 dm. high; the main stem and branches becoming strongly angled or even quadrate above from 4 internodal ridges; ranging from central New Jersey southward and westward into western Florida..................... 1. S. difformis. Leaves and upper portion of the stem glaucous; calyx-lobes rarely as long as 3 mm.; upper bractlets minutely scale-like, even those subtending complete cymules less than 3 mm. long; corolla-lobes 4-7 mm. long, color remaining white or creamy-white upon drying; plants more robust, mostly taller than 9 dm.; the main stem and branches terete throughout, not becoming angled or quadrate; ranging from central Georgia and ne. Florida westward into LOG. ou iure bEL RAE re ru Ee NE E. 2. S. macrophylla.

1. Sabatia difformis (L.) Druce Swertia difformis L., Sp. Pl. 1: 226. 1753. Chirenia lanceolata Walt., Fl. Car. 95. 1788. C. cymosa Lam., Tabl. Encyc. et Méth. Bot. 1: 479. 1791, not Burm., Fl. Cap. 5: 1768. C. paniculata Michx., Fl. Bor.-Am. 1: 146. 1803. (C. venosa Muhl., Cat. Pl. Am. Sept. 24. 1813. Sabbatia paniculata (Michx.) Pursh, Fl. Am. Sept. 1: 138. 1814. S. paniculata æ latifolia Pursh, l.c. S. paniculata 8 angustifolia Pursh, l.c. S. corymbosa Baldw. ex Ell, Sk. Bot. S. C. & Ga. 1: 283. 1817. S. corymbosa var. angustifolia Ell., l.c. S. lanceol. [lanceolata Walt.] Raf., Fl. Tell. 3: 30. 1837, without basi- onym. S. cymosa [Lam.] Raf., l.c. without basionym. S. cymosa (Lam.) G. Don, Gen. Hist. 4: 207. 1838, as to basionym only. S. lanceolata (Walt.) T. & G., Man. ed. 1. 356. 1848. S. difformis (L.) Druce, Bot. Exch. Club & Soc. Brit. Is. 3: 423. 1914, as to basionym only.

Perennial herb (25-)45-80(-105) em. tall, 2-5 mm. in diameter, 1- several stems arising in a cluster from a gnarled, stout, branched rhizome 4-6 mm. in diameter. Stem stiffly erect, hollow, more or less terete below, becoming angular and sometimes almost quadrate above from the internodal ridges. Branches typically opposite, occasionally alter- nate at some nodes, ascendent at an angle of (10—)20-40(—50) degrees, generally restricted to the upper tenth or third of the stem, or less com- monly in very robust plants branching almost throughout the entire length of the stem. Principal branches typically 3—4-times branched and together forming a compact, corymbose, flat-topped to convex

8 Rhodora [Vor. 57

inflorescence. Numerous fleshy roots descending from the rhizome, these 8-12(-20) em. long, 1.5-3 mm. in diameter, usually bearing slender, fibrous, lateral roots. Leaves thick, rather succulent, drying somewhat thick-chartaceous, venation obscure except for the rather prominently elevated midvein and often the lateral pair of veins, or in the broader- leaved types with 2 lateral pairs prominent. Basal rosette none; the lowermost cauline leaves borne submerged or subterranean, at time of flowering usually represented only by scars, but when still present, often modified into appressed, obtuse, oblong bracts 2-3 em. long. Upper cauline leaves strongly ascendent, linear to lanceolate or even somewhat oblong or ovate-lanceolate to strongly ovate (especially in peninsular Florida), usually acute, rarely obtuse, somewhat revolute and both scarious-margined and -tipped, usually rather strongly clasping, or merely sessile in the narrower leaved types, (0.9—)1.8-4(-6.2) em. long, (3-) 4-14(-22) mm. wide, usually not more than twice exceeded by the in- ternodes and usually less; internodes occasionally 3-5 times or more longer than the nearby leaves. Pedicels slender, erect, finely ridged, 2-8(-15) mm. long. Calyx-tube only slightly ridged, rather shallowly campanulate, crateriform or even turbinate, 1-2(-3) mm. long; calyx- lobes linear to somewhat subulate, very slender, (2-)4-10(-14) mm. long, 2-8(-10) times the length of the tube, more or less outwardly arching at anthesis, erect in bud. Corolla-tube narrowly cylindrical, (2.5-) 3-5(-6) mm. long; corolla-lobes typically 5, (the centermost flower of the central cymules often 6-parted), oblong, weakly spatulate, elliptic, obtuse to acute, wide-spreading, (5-)7-15(-21) mm. long, 2.5-6(-8) mm. wide, entirely white even at the base, but often turning orange or saffron upon drying, especially along the veins. Filaments 2-3 mm. long, pale-yellowish to almost white, slender; anthers slender, lanceolate, bright yellow, 2-3 mm. long. Style 2-5 mm. long; stigmatic lobes slender, linear, 2-5 mm. long. Capsule oblong-cylindrical, 4-8 mm. long, 2.5-4 mm. wide. TYPE LOCALITY: "In Virginia." Type: Clayton 171 (British Museum); photograph of the type (GH!). DISTRIBUTION: Savannas and pine-barrens along the Coastal Plain from (perhaps Long Island) central New Jersey southward throughout much of peninsular Florida and westward into western Florida. Map. 1.

The only species with which S. difformis might be confused is S. macrophylla, which belongs to the same subsection and which shares with it the distinctive features of a stout, gnarled rhizome bearing numerous, thick, fleshy roots, characteristically opposite branches, thick leaves and white 5(6)-parted corollas. However, S. difformis may be readily distinguished from S. macrophylla by numerous striking differences among which are the absence of a glaucous bloom, the almost invariably longer calyx-lobes, bractlets, and corolla-lobes; the very common change of color to orange or saffron upon drying, the stem angled

1955] Wilbur,— The North American Genus Sabatia 9

or quadrate above, and the typically smaller stature. The ranges of the two species overlap in Georgia and in Florida; but throughout the main part of the area occupied, they are not compatriots. (See Maps 1 and 2)

S. difformis and another typically white-flowered, opposite- branched species, here called S. quadrangula but referred to by recent authors as S. paniculata, have been closely associated together in many treatments of the genus and often the two are misidentified. Gleason’s (1952) account of the species that has been passing as S. paniculata fails to distinguish clearly the two species. S. paniculata, according to Gleason, is “similar to S. difformis in habit, stature, and inflorescence” and is “very close to S. difformis in describable characters . . ." The two species may be easily distinguished and the differences are such that I feel that the two belong to different species-groups. The features of S. difformis mentioned in the preceding para- graph are in strong contrast to those of the other species which is a rosulate annual with a strongly 4-sided stem and membranous leaves.

The synonymy of this species is extensive and there has been much confusion and misunderstanding throughout the litera- ture regarding these names. It therefore seems advisable to discuss each synonym briefly.

Swertia difformis L. is based upon the Gronovian “Gentiana foliis linearibus acuminatis, pedunculis, longissimis nudis unifloris oppositis" which in turn rested upon Clayton’s collection 171 and also apparently two other specimens or perhaps descriptions sent by Clayton. These last two references apparently played no part in Linnaeus’ concept and it is very unlikely that they entered to any extent into that of Gronovius as they presented characteristics that were in direct contrast to those described in the principal account. They certainly both belonged to at least a different species. The specimen in Clayton’s her- barium is, as is shown by the photograph in the Gray Herbarium or by Blake’s sketch (Rhodora 17: pl. 112. 1915.), the uppermost portion of the rhizomatous perennial possessing characteristically opposite branches and a_ white, 5(6)-parted corolla. The specimen is peculiar in that the lateral branches, all six of them, are very much elongated and bear but one flower each. The

10 Rhodora [Vor. 57

inflorescence hence is very loose. The terminal flower is six- parted and this feature is shown very clearly in the photograph. The original description little suggests the plant that it is now known to depict and it is therefore not surprising that its proper disposition was but so recently accepted by most American botanists. The diagnosis, however, is excellent as a description of the somewhat atypical type specimen which is the opposite- branched perennial which was known throughout the latter half of the nineteenth century as S. lanceolata. Druce, in transferring the epithet, referred it to what has been called S. elliottii, an alternate-branched, white-flowered annual. In this he was following either the “Synoptical Flora" or the "Index Kewensis." Clayton's specimen has nothing to do with that species. The alternate-branched annual, furthermore, has not been found north of South Carolina; the opposite-branched perennial is known from southeastern Virginia, where Clayton served as a clerk of the court of Gloucester County for many years.

Card (Ann. Mo. Bot. Gard. 18: 262. 1931.) in his monograph of Frasera listed Swertia difformis as a synonym of Frasera caroliniensis Walt. This mistake has been previously and emphatically pointed out by St. John (Am. Midl. Nat. 26: 5. 1941.). Card was no doubt following Rafinesque (Med. Fl. 1: 196. 1828.) who listed this name among others as being syn- onymous with his substitute name, F. verticillata. Merrill (1949) also equates Frasera verticillata in part to Sabatia difformis.

Of the seven fragmentary specimens of Sabatia still represented in Walter's collection two match the brief description of Chironia lanceolata and are certainly referable to the opposite-branched perennial. The association of Walter's name with the perennial has been almost invariably the rule from the time of Pursh (1814), who, as stated in his Preface, had examined Walter's collection. The combination of Walter's epithet with Sabatia can perhaps first be credited to Torrey and Gray in Gray's Manual (1848) who, after describing the plant, listed S. corymbosa Ell. as a synonym. ‘The only name listed in synonymy under S. corym- bosa, when it was originally described, was Walter’s Chironia lanceolata. ‘The synonymy of this species presented by Grisebach (1839, 1845) was complete and these accounts were doubtless

1955] Wilbur,— The North American Genus Sabatia 11

the basis for Gray’s treatment. It is to be remembered that Gray also examined Walter's collection in 1839.

Rafinesque (Fl. Tell. 3: 30. 1837.) may have intended this transfer when he listed ''lanceol." in an enumeration of the genus Sabatia. Many of the names in this list were never described as the promised monograph never appeared. For S. lanceolata (as lanceol.), there were no basionym or description so even listing the name in synonymy may not be warranted.

A photograph of the type of Chironia cymosa Lam., which is located in Paris, was lent by the Gray Herbarium and it also proves to be a specimen of S. difformis. The specimen as is shown by the label was one of Fraser’s from “eastern [South] Carolina."

Rafinesque (Fl. Tell. 3: 30. 1837.) in the same list of names in which 'anceol." appears, also included cymosa and again without basionym or description. It is possible that a transfer was intended.

This epithet, cymosa, was transferred formally by G. Don (1838) with the necessary basionym. The description that accompanies the name, however, applies only to S. quadrangula, a very different species.

Chironia paniculata Michx. is discussed more fully under S. quadrangula, a species with which it has long been confused. The original description was brief and the few clues presented there have been either overlooked or misunderstood. The species was known to Michaux from Georgia and ''Carolina." The photograph at the Gray Herbarium of the collection in Michaux’s herbarium labeled ‘‘Chironia paniculata Georgia," illustrates an excellent specimen and one which is unmistakably S. difformis. Pursh transferred Michaux’s epithet to Sabatia and noted the synonymy with both Swertia difformis and Chironia lanceolata. He had seen the specimen upon which Swertia difformis is based in Bank’s herbarium and was acquainted with authentic material of Walter's species. In addition Pursh designated two varieties, which, as I interpret the descriptions, are merely the broad- and narrow-leaved variations that are common among representatives of this species. However, Gray (1878) and following him Blake (1915) considered Chironia paniculata B angustifolia to be, excluding C. lanceolata listed in synonymy by Pursh, what is here called S. quadrangula.

12 Rhodora (Vou. 57

Chironia venosa Muhl. was relegated to the synonymy of the opposite-branched, white-flowered perennial by Torrey as early as 1824. Those names published for the first time in Muhlen- berg’s Catalogue are so briefly characterized that Merrill and Hu (Bartonia No. 25: 22. 1949) stated that they should be considered as nomina nuda unless a basionym were included. The information presented with Chironia venosa was that the corolla was white and veiny and that the plant was a perennial found in “N. Caes. Ten." S. difformis is not known from Tennessee, nor is any other perennial or any other typically white-flowered species. It is found in New Jersey (N. Caes.). Otherwise there can be but little doubt that Chironia venosa is the same as S. difformis. The specific epithet aptly describes the orange discoloration especially noticeable along veins of the corolla-lobes in some dried specimens.

Sabbatia corymbosa Baldw. ex Ell. is the last of the new names proposed for the white-flowered, rhizomatose perennial. Au- thentie specimens from Elliott’s herbarium prove the correctness of treating this name as a synonym of S. difformis. This sheet apparently bears two collections, which may be taken perhaps as types; one with ovate leaves apparently representing the typical element and the other with narrower leaves, probably a representative of his S. corymbosa var. angustifolia. This specific name was generally employed in designating the peren- nial species until the combination of S. lanceolata was made in 1848.

REPRESENTATIVE SPECIMENS:—NEW YORK: Nassau Co., Hempstead Flats, Mackaness, 1937 (No). Since no other specimen of this species has been seen or has been reported from the state and since the region is rather well-known botanically, error in labeling is suspected. NEW JERSEY: Burlington Co., about 1.5 mi. se. of Atsion, Fogg 5663 (F, GH, PENN); Ocean Co., Tom's River, Mackenzie 2774 (MO, US). DELAWARE: Sussex Co., 0.5 mi. w. of Concord, Tatnall 3129 (GH). MARYLAND: Worchester Co., Stockton, Rusby, Aug. 1889 (NY). VIRGINIA: Sussex Co., sw. of Wakefield, Fernald, Long & Clement 15840 (GH, MO, US). NORTH CAROLINA: Bladen Co., 4 mi. n. of Elizabeth- town, Wilbur 2900 (micn); Harnett Co., about 3 mi. se. of Dunn, Wilbur 2897 (MicH); New Hanover Co., Carolina Beach, Godfrey, PL. EX. GRAYANAE 974 (F, GA, GH, MICH, MO, MT, NCS, NCU, NY, OKL, PENN, SMU, TENN, TEX, WIS, WVA, US). SOUTH CAROLINA: Berkeley Co., about 6 mi. ne. of Summerville, Wilbur & Webster 2854 (mMicu); Georgetown Co., 12 mi. n. of Georgetown, Godfrey & Tryon 65 (DUKE, F, GH, MO, NY, PENN, TENN, US). GEORGIA: Charlton Co., below Trader’s Hill, Small, 12-15 June 1895 (F, Ny); McIntosh Co., 1.5 mi. n. of Darien, Cronquist 5375 (Ga, us) Thomas Co., about 1 mi.

1955] Wilbur,—The North American Genus Sabatia 13

n. of Coolidge, Duncan 8468 (FLAS, GA, MO). FLORIDA: Dade Co., Biscayne Bay, Rusby (MicH) [Since otherwise not known south of Lake Okeechobee, this station seems questionable.] Highlands Co., near Sebring, Small, Small & Dewinkeler, 17 July 1924 (FLAS, GH, NY, US); Lake Co., Eustis, Nash 849 (cU, F, GH, MICH, MO, MT, NY, US); Manatee Co., Bradentown, Tracy 7541 (cU, F, GH, MO, NY, US, WIS); Okaloosa Co., 4 mi. e. of Crestview, Webster & Wilbur 3592 (micu); Polk Co., 16 mi. sw. of Kissimmee, Wilbur & Webster 2637 (micH); Santa Rosa Co., e. bank of Blackwater River across from the town of Milton, Webster & Wilbur 3579 (Micu); Walton Co., near DeFuniak Springs, Curtiss 5906 (CU, F, FLAS, GH, KSC, NCU, NY, SMU, US); Washington Co., 1 mi. e. of Caryville, Webster & Wilbur 3601 (micH).

2. Sabatia macrophylla Hook., Compan. Bot. Mag. 1: 171. 1836.

Perennial herb (50-)90-120(-140) em. high, 2-7 mm. in diameter, usually with several stems arising in a cluster from a stout, gnarled, branched rhizome up to 1 em. in diameter and often 10 cm. or more in length. Stem stiffly erect, hollow, terete throughout, without ridges or fine lines extending between the nodes, strikingly glaucous above. Numerous white to reddish-brown fleshy roots often 15 cm. or more in length and about 2 mm. in diameter descending in clusters from the rhizome with occasional slender, fibrous, lateral roots. Branches typically opposite, very rarely alternate, generally restricted to the upper tenth or quarter or, more rarely, throughout the upper half of the stem, as- cendent to rather strongly divergent forming an angle of 15-60 degrees with the stem, producing a compact, corymbose, flat-topped to somewhat convex inflorescence. Leaves thick, somewhat succulent, drying thickly chartaceous, strongly ascendent, very noticeably glaucous when fresh, ovate-lanceolate, lanceolate (sometimes very narrowly so), oblong, to ovate-oblong or even ovate, acute or rarely obtuse, scarious-mucronate tipped, margins conspicuously scarious and often slightly revolute, strongly clasping, (2.5-)3-6(-8.5) em. long, (0.5—)1-2.5(-4.5) em. wide, midvein prominently elevated beneath, 1—2 pairs lateral veins also often rather conspicuous. Basal rosette none; the lowermost cauline leaves absent at time of flowering. Internodes typically 3-5 times the length of the leaves. Inflorescence of numerous eymules corymbosely arranged forming a compact flat-topped to somewhat convex cluster. Leaves within the inflorescence abruptly reduced to acute, scale-like, usually non-chlorophyllous bracts 1-4 mm. long, the ultimate bracts less than 2 mm. long; the flowering clusters thus appearing naked. Pedicels slender, erect or ascendent, 1-5 mm. long. Calyx-tube and lobes more or less colorless, non-chlorophyllous; the tube campanulate, scarcely ridged or lined by elevated vascular traces, 1-2 mm. long; the lobes tri- angular-dentate, subulate, or linear, erect, slightly spreading, or somewhat outwardly curved at the tip, or strongly recurved, (0.1—)0.2-2.5(-3.0) mm. long, 0.1-3.5 times the length of the calyx-tube. Corolla-tube narrowly cylindrical, white, (2-)3-3.5(-4) mm. long. Corolla-lobes oblong to ob- long-spatulate, obtuse, widely spreading at anthesis, (4-)5-7 (78) mm. long, 2-3(4) mm. wide, entirely white or creamy-white, color not changing, or

14 Rhodora (Vor. 57

changing but little, upon drying. Filaments (1.5-)3-4 mm. long, white; anthers slender, lanceolate, creamy white to pale yellow, 1-2 mm. long. Style 1-3 mm. long; stigmatic lobes slender, 1-3 mm. long. Capsule globose or nearly so to ovoid, (3-)3.5-4(-4.5) mm. high, (2-)3-4 mm. wide.

Small (1933) described S. recurvans and, according to him, this species differed from S. macrophylla by the characteristies presented below.

S. macrophylla Hook. S. recurvans Small Calyx 2-3 mm. long, lobes Calyx 4-5 mm. long, lobes shorter than the tube. longer than the tube. Corolla-lobes 6-8 mm. long. Corolla-lobes 5-6.5 mm. long. Capsule 3-4 mm. long. Capsule 3.5-4.5 mm. long. Style and stigma about equal Style much shorter than in length. the stigma.

Available to Small at New York were five sheets which here are considered part of the recurvans population. Two of these, Harper 1415 and 708, were so annotated by Small. Excellent specimens of Harper's 1415 are represented in several herbaria. The specimen of this number at New York is chosen as the lectotype. It, however, lacks fruit, which was described from Harper's 708.

The specimens of S. recurvans available to Small at New York were too few to be significant in evaluating the supposedly distinguishing characters. This is especially true in regard to the overlapping size-ranges of the corolla-lobes and capsules published as additional evidence. "These overlapping ranges might be of some significance if based upon a large series of specimens but in this case furnished only “padding” to an otherwise scantily characterized entity. Even now few fruiting specimens are available but measurements of them indicate a range of 3-4.5 mm. in length of capsule in both species. The length of the corolla-lobes is equally useless as a distinguishing feature as the range of S. macrophylla was between 4-8 mm. and that of S. recurvans between 4—7 mm. The slight discrep- ancy would very likely disappear if a larger series of measure- ments were made. The length of the corolla-lobes is apparently strongly affected by environmental conditions and the season in which the flowers are produced. Flowers developing later in the season are generally much smaller than those appearing earlier in the year. Therefore, all of the characters with over-

1955] Wilbur,— The North American Genus Sabatia 15

lapping ranges presented by Small as supporting evidence of the distinetiveness of the two entities prove completely useless upon examination of more collections.

'The ratio of the style to the stigmas was indicated by Small to be a reliable distinguishing feature. It was even used by him as a secondary key-character. However, the comparative length of these structures varies enough so that one may find either the stigma or the style longer on the same plant; con- sequently this character is not of taxonomic value. The com- parative length of the stigma to the style varies considerably in different stages of maturity.

The calyx does possess features which seem to be valid; specimens can be assigned to one group or the other by the calyx characters and the two groups thus delineated occupy, as far as is now known, adjacent but distinct areas. Small’s very convenient hiatus of 1 mm. between the size-range of the calyx in the two entities disappears upon examination of a number of flowers. The overall calyx-length of S. macrophylla is 1.1-3(-4) mm., while that of S. recurvans is (2-)2.5-4.0(—4.5) mm. long. The overlap of these ranges is too great to be of much aid in identification.

A comparison of the length of the calyx-lobes has much more promise, but that, too, fails to provide a clear-cut distinction that might be mechanically applied. The macrophylla-popu- lation has lobes (0.1—)0.2-1.2(-2) mm. long. Of the more than six hundred lobes measured in this group nearly sixty per cent were less than 1 mm. long, and over ninety-six per cent of the lobes were less than 1.2 mm. long. The few flowers with calyx-lobes longer than 1.2 mm. were always the centermost flower of the inflorescence or the oldest flower of the cymules closest the center of the compound inflorescence. Their lobes are often longer than those of other flowers on the same plant. Calyx-lobes of the recurvans-group measured (1—)1.5-3 mm. long with only five flowers (less than three per cent of the total) with lobes less than 1.5 mm. long and with more than seventy per cent of the nearly two hundred lobes measured 2 mm. long or longer.

The ratio of the lobe/tube-length provides a somewhat sharper means of separation but even this provides no clear-

16 Rhodora (Vor. 57

eut division between the two populations. This ratio-range of the macrophylla-group is (0.1—-)0.2-1(-2) with all but five flowers possessing lobes equaling or less than the tube in length. These few flowers were again the centermost which are often strikingly different from the others in the inflorescence. The same ratio-range in the recurvans-population is (1—)1.3-3(-3.5) With only four flowers (2.3 per cent) equaling the tube and nearly forty per cent of the flowers possessing lobes two or more times greater than the tube.

Even the feature of the recurved lobes is not restricted to the recurvans-group; some of the longer-lobed representatives of S. macrophylla, which are found in the extreme eastern portion of its range, show at least the beginnings of this tendency by possessing lobes slightly recurved at the extreme tip.

The most useful criterion known to me for the separation of the two populations is that based upon the length of the calyx- lobes. Future study may demonstrate that the group known as S. macrophylla possesses pronounced clinal variation from west to east and that this variation is in the direction of the even more easternly population previously known as S. recurvans. The calycine lobes of the plants from western Florida seem longer on the average than those from Louisiana. The even more eastern S. difformis, the species most closely related to the above-discussed complex, is usually readily distinguished by calyx-lobes which are almost invariably longer than even those of S. recurvans. Again in regard to characters of the calyx there is no sharp separation between the two populations but the upper limits of S. recurvans overlap but slightly the relatively few calyx-lobes measured in S. difformis which were shorter than 4 mm. long.

The material at hand may be readily assigned to one popula- tion or the other with little more than a glance at several calyces and the two groups have, as far as is now known, separate ranges. The two entities, here considered to represent weakly differentiated geographical varieties, are not distinguished by strong, or even exclusively delimiting morphological characters. Nor are the two varieties widely separated geographically as they are now known to occur at least within fifty miles of one another.

1955] Wilbur,— The North American Genus Sabatia 17

KEY TO THE VARIETIES Calyx-lobes erect or only slighty outwardly curved at the tip, equaling or less than the calyx-tube in length....... 2a. S. macrophylla var. macrophylla. Calyx-lobes strongly recurved, exceeding the calyx-tube in length. 2b. S. macrophylla var. recurvans.

2a. Sabatia macrophylla Hook., var. macrophylla.

Calyx lobes (0.1-)0.2-1.5(-2) mm. long, triangular-dentate to linear, erect or only slightly outwardly curved at the tip, equaling or less than the calyx-tube in length (very rarely the centermost flower of the central eymules may have a ratio of lobe/tube greater than 1). TYPE LOCALITY: near Covington, Louisiana. Type: Drummond s. n. (K!). DISTRIBUTION. Savannas and pine-barrens and margins of low hardwoods from western Florida and Georgia westward to Louisiana. Map 2.

REPRESENTATIVE SPECIMENS:—GEORGIA: Muscogee Co., Columbus, Boykin (BRU, GH, NY); Sumter Co., Americus, Harper, 30 July 1897 (GH). FLORIDA: Bay Co., 1 mi. n. of Lynn Haven, Webster & Wilbur 3626 (mica); Calhoun Co., 2 mi. s. of Chipola, Thorne & M uenscher 8744 (cv); Columbia Co., Lake City, collector not stated (F); Escambia Co., 9 mi. w. of Pensacola, Webster & Wilbur 3575 (MicH); Franklin Co., Apalachicola, Biltmore Distrib. Chapman Herb. 4508a (an, NY); Gadsen Co., inter Quincy et Aspalga, Rugel, July 1843 (wo, ny); Gulf Co., Port St. Joe, Knight, 18 June 1936 (FLAS) ; Liberty Co., Bristol, West & Arnold, 23 July 1940 (Frias); Okaloosa Co., about 3 mi. e. of Crestview, Webster & Wilbur 3584 (micu); Wakulla Co., 1.5 mi. s. of Sopchoppy, Webster & Wilbur 3628 (micu); Walton Co., near Argyle, Curtiss 5931 (CU, FLAS, GH, KSC, MO, NCU, NY, SMU, US); Washington Co., 8 mi. s. of Chipley, Webster & Wilbur 3614 (micH). ALABAMA: Baldwin Co., 13.5 mi. w. of Foley, Webster & Wilbur 3530 (micu); Mobile Co., about 8 mi. w. of Mobile, Webster & Wilbur 3481 (mic); Washington Co., 40 mi. n. of Mobile, Sargent, 18 June 1950 (Ga, NCS). MISSISSIPPI: Forest Co., Lake Shelby State Park about 14 mi. s. of Hattiesburg, Webster & Wilbur 3402 (micu); Harrison Co., Biloxi, Tracy 7006 (F, MO, NY, US); Jackson Co., Ocean Springs, Pollard 1064 (cU, F, MO, NY, US); Perry Co., 9 mi. n. of Beaumont, Webster & Wilbur 3422 (micH); Stone Co., 3 mi. s. of Wiggins, Webster & Wilbur 3439 (Micu). LOUISIANA: St. Tammany Parish, Covington, Drummond TYPE (k).

2b. Sabatia macrophylla Hook., var recurvans (Small) comb. nov. Sabbatia recurvans Small, Man. SE Fl. 1049. 1933. i

Calyx-lobes (1—)1.5-3 mm. long, linear to subulate, strongly recurved, typically exceeding the calyx-tube in length (less than 3 per cent of the nearly 200 lobes measured had a lobe/tube ratio of 1 or 1.2). LECTOTYPE LOCALITY: Moist pine-barrens east of Ocilla, Irwin Co., Georgia. Lecto- type: Harper 1415 (NY!. DISTRIBUTION: Savannas and pine-barrens of southern Georgia and northeastern Florida. Map 2.

REPRESENTATIVE SPECIMENS:—GEORGIA: Brantley Co., 3 mi. e. of Nahunta, Wilbur & Webster 2748 (micu); Charlton Co., 8 mi. s. of Folkston, Wright 881 (cv); Coffee Co. without exact locality, Harper 708 (NY, vs); Emanuel Co., e. of Swainsboro, Duncan 2677 (micu); Irwin Co., e. of Ocilla, Harper 1415 (r, GH, MO, NY, US); Laurens Co., 10 mi. e. of Dublin, Pyron & McVaugh 750

18 Rhodora [Vor. 57

(G^, US); Worth Co., 3.5 mi. w. of Sylvester, Thorne 6363 (cU). FLORIDA: Baker Co., without exact locality, Curtiss 2224 (cU, F, Mo, NY, us, YU); Clay Co., without exact locality, T'odsen, 30 July 1939 (rtas); Duval Co., 15 mi. w. of Jacksonville, Wiegand & Manning 2566 (cu, au); Nassau Co., without exact locality, Knight, 1 July 1941 (Fuas).

(B) Subsection ANGuLARES Blake, Rhodora 17: 56. 1915.

Annuals with fibrous roots and with typically opposite branches. Leaves thin and membranous, margins not scarious. Basal rosette usually well-developed at some time during maturation of plant. TYPE SPECIES: Sabatia angularis (L.) Pursh.

The three species comprising subsection Angulares, as here defined, are readily distinguished from one another. Asa group it is strongly differentiated from all other subsections of section Eusabatia. The annual habit separates them from subsections Difformes and Dodecandrae, the opposite branches and the typically 3-flowered cymules set them apart from the Campanu- latae, and the thin, merely finely ridged calyx-tube and, again, the opposite branches easily distinguish the Angulares from the Campestres. This subsection occupies the largest area of any within the genus.

These three species and the other species reported as annuals in the paper have most often in the past been treated as biennials. Field observations and limited experience with some of the species in the greenhouse has led me to believe that they are all probably annuals but this point has not been definitely established.

KEY TO THE SPECIES OF SUBSECTION ANGULARES

A, Lower portion of the stem strongly 4-angled, quadrate, the corners with conspicuous membranously-winged margins.

B. Corolla pink to roseate (very rarely white); pedicels elongate, 1 em. or more in length; lower cauline leaves usually broadly ovate to ovate- lanceolate, usually less than 3 times as long as broad; calyx-lobes usually 1.5 mm. or more in width (rarely as narrow as 1 mm.).

3. S. angularis.

B. Corolla white; pedicels short, less than 5 mm. long ; lower cauline leaves usually oblong, generally 4 or more times as long as broad; calyx-lobes less than 0.5 mm. wide.............2.......... 4. S. quadrangula.

A. Lower portion of the stem terete and not bearing thin membranous wings. 5. S. brachiata.

3. Sabatia angularis (L.) Pursh Chironia angularis L., Sp. Pl. 1: 190. 1753. C. angularis var. « latifolia Michx., Fl. Bor.-Am. 1: 147. 1803. Sabbatia angularis (L.) Pursh, Fl. Am. Sept. 1: 137. 1814. S. angularis

1955] Wilbur,— The North American Genus Sabatia 19

var. albiflora Raf., Med. Fl. 2: 77. 1830. nom. nud. S. angularis, var. elatior Raf., l.c., nom nud. S. angularis, var. latifolia Raf., l.c., nom nud. S. angularis, var pauciflora Raf., l. c., nom nud. S. angularis, f. albiflora Raf. ex House, Bull. N. Y. State Mus. 254: 566. 1924. S. angularis f. cleistantha Fern., RHopora 42: 474. 1940.

Erect annual (15-)30-50(-90) em. tall, often robust and usually with the numerous branches presenting in plants developed under favorable conditions a bushily compact aspect. Stem strongly tetragonal, hollow, the corners conspicuously wing-angled by thin membranous extensions 0.1-0.5 mm. wide. Stem green in younger portion becoming yellowish straw-colored below, (1-)2-3(5) mm. in diameter. Branches typically opposite, although ultimate-branching often alternate, usually by marked abortion of opposing bud, or generally alternate in depauperate plants, often numerous, rarely extending to base, but more typically restricted to upper third or half of stem, rather closely ascendent forming an angle of about 20-45 degrees with main stem. Branches composed of 1-10 nodes in well-developed plants and forming a convexly-corymbose or pyramidal compact crown. Roots several, fibrous, 4-10(-20) em. long, (0.2-)1-3(-4) mm. in diameter, wide-spreading and usually shallowly situated. Leaves thin membranous, spreading to ascendent, drying thin, 5-7-nerved or fewer in those leaves transitional to bracts, (1—-)2-3.5 (—4.5) em long, (0.8—)1-2.5(—3.5) em. wide, venation prominently elevated beneath. Basal rosette rarely present at anthesis, composed of spreading, shortly-spatulate, broadly oblong to obovate, obtuse leaves up to about 4 em. long and often 2-3 em. wide. Lowermost cauline leaves usually absent at time of flowering, the lowest usually broadly ovate, always strongly clasping, obtuse, typieally nearly as broad as long, the middle narrower, more typically ovate-lanceolate, obtuse or more commonly acute, and gradually reduced and modified above to the 1-nerved linear bracts. Internodes generally about 1.5-3 times as long as the leaves. Inflorescence of corymbosely- or pyramidally-arranged cymules, either or both of the lateral branches sometimes suppressed. Pedicel 5-sided, slightly wing-angled, (1—)2-3.5(—4.5) em. long, erect or strongly ascendent. Calyx-tube thin, shallow, 2-5 times exceeded by the corolla-tube, usually about 1.5 mm. long (1-2 mm.), with venation somewhat elevated. Calyx-lobes thin, narrowly linear, linear-oblong, to somewhat foliaceous, usually about 1 em. long (0.4-1.5 em.) and (1-)1.5-2.5(-3.5) mm. wide, ascendent in bud, wide-spreading at anthesis, usually exceeded by the corclla by 5-12 mm. Corolla-tube 4-7 mm. long, 2-4 mm. in diameter, cylindrical, pale greenish-white within and without. Corolla-lobes oblong, spatulate, or elliptic, obtuse, wide-spreading, (0.6—)1—1.8(-2.2) em. long, (2-)4-7(-9) mm. wide, greenish in bud turning white prior to full development and typically pink or roseate at anthesis or rarely white, coloration paler beneath, with an irregular, somewhat pointed, yellow or greenish-yellow area at base of lobe and in throat and usually bordered by a dark-red line. Filaments 2.5-4.5 mm. long, greenish- yellow to pale-yellow; anthers slender, bright yellow, 3-5 mm. long. Stig-

20 Rhodora [Vor. 57

matic lobes very slender, 3-6 mm. long. Style 4-6 mm. long, usually some- what less than length of stigmatie branches. Capsule cylindrical at ma- turity 5-9 mm. high, 4-6 mm. wide. Type Locarrrv: “In Virginia." Type: Kalm (in the Linnean Herbarium), but not seen. DISTRIBUTION: Margins of woods, fields, and prairies from southern New York south to northern Florida and west to Illinois and Texas. Occasionally introduced in Massachusetts and Connectieut. Map 3.

This species is by far the most familiar representative of the genus because of its wide range, the greatest of any in the genus, and also because of the frequency with which it is en- countered throughout most of that range. "The characteristic features are the conspicuous tetragonal stem, the corners of which are strikingly membranously wing-margined; the typically opposite pattern of branching, the ovate-clasping leaves, the roseate corolla (although albinos are encountered rather rarely), and the shallow calyx-tube, the venation of which, although often slightly elevated, is not developed into a heavy costa. None of these characters alone is enough to distinguish the species but the combination of them in one plant is unique. In the western part of the range of S. angularis, it has been confused surprisingly often with S. campestris. This latter species may be readily distinguished from the former by its typically alternate branching-pattern and especially by the turbinate and comparatively long calyx-tube with strongly developed costae. In the southeastern region, it has been confused at times with S. brachiata, but is at once separable from that species which possesses an almost terete stem and oblong, non-clasping leaves.

The species is remarkably uniform throughout its range. None of the trivial variants designated previously is here deemed worthy of recognition nor have any others been discovered. Although I have not seen even a photograph of the type of Chironia angularis B angustifolia, if it is still extant, I am ac- cepting for the time being the past interpretation that this entity is what is now called S. brachiata. Coordinate with this variety, Michaux published C. angularis x latifolia which is perhaps the typical element of S. angularis. The species remained unencumbered by the designation of minor varieties until Rafinesque. This author, after discussing S. angularis, stated that "it has some varieties: 1. Albiflora, 2. Latifolia,

1955] Wilbur,— The North American Genus Sabatia 21

S. MACROPHYLLA € VAR. MACROPHYLLA

Q VAR. RECURVANS

S.DIFFORMIS

S. ANGULARIS S.QUADRANGULA

Mars1-4. Map 1, upper left; map 2, upper right; map 3, lower left; map 4, lower right.

3. Pauciflora, 4. Elatior." None of these propositions is dis- cussed or mentioned by him further in any manner and hence are merely nomina nuda. Therefore, they are not available for transferral to any other rank. House, in providing formal nomenclature to designate the albino form, erred in transferring a name which “has no status under the Rules." For those who find it useful or satisfying to name such striking minor variations the epithet for the albino might best be written as Sabatia angularis forma albiflora House. Fernald described

22 Rhodora [Vor. 57

a teratological specimen as S. angularis forma cleistantha. It has not been seen again in any of the many hundreds of sheets of this species that I have examined. The branches are stunted and malformed bearing sessile or nearly sessile flowers which are aborted and modified in size and form as to be scarcely recognizable as flowers. Little is to be gained by formal designa- tion of such rare chance monstrosities.

REPRESENTATIVE SPECIMENS:—CONNECTICUT: Locality not stated, Wright (yu). New York: Richmond Co., near New Dorp, Britton, 9 Aug. 1879 (vv). New JERSEY: Cape May Co., Cold Spring, Pennell 2128 (us). PENNSYLVANIA! Berks Co., 0.5 mi. wnw. of Gibralter, Berkheimer 2926 (GH, PENN). DELA- WARE: New Castle Co., near Wilmington, Canby, Aug. 1878 (MO). MARY- LAND: Harford Co., se. of Flintville, Adams & Hopkins 923 (cu, PENN). DISTRICT OF COLUMBIA: Chain Bridge, Earlanson & Martin 1210 (NA). VIRGINIA: Southampton Co., se. of Ivor, Fernald & Long 10784 (GH, Mo). NORTH CAROLINA: Buncombe Co., near Biltmore, Biltmore Herb. 3303a (F, GH, MO, NY, US); Durham Co., about 4 mi. ne. of Durham, Wilbur 2898 (micu); Halifax Co., Roanoke Rapids, Godfrey 5178 (DUKE, GH, US). SOUTH CAROLINA: Williamsburg Co., 2 mi. ne. of Lane, Godfrey & Tryon 399 (DUKE, F, GH, MO, NY, PENN, TENN). GEORGIA: Bartow Co., about 4.5 mi. s. 16° east of Alla- toona Dam, Duncan 8522 (FLAS, GA, GH, MO, TENN, US); Catoosa Co., 10 mi. w. of Ringgold, Cronquist 5610 (GA, GĦ, Us). FLORIDA: Jackson Co., Sneads, Knight, 29 June 1943 (FLAs). MICHIGAN: Kalamazoo Co., Hermann 9042 (wrcH). OHIO: Coshocton Co., near Coshocton, Moldenke 13289 (OKLA, PENN, SMU). INDIANA: Morgan Co., about 2 mi. w. of Centerton, Friesner 17008 (GH, NY, OKLA); Steuben Co., near Lake Gage, Deam, 11 Aug. 1903 (GH, Us, wis, YU). ILLINOIS: Marion Co., Salem, Bebb 1860 (r, an, vv). WEST VIRGINIA: Cabell Co., overlooking Roland Park, Gilbert 778 (DUKE, F, GA, GH, MICH, MO, MT, NA, NY, OKL, PENN, RUT, TENN, Us, WIS); Putnam Co., near Hurricane, Gilbert et al., so. APP. BOT. CLUB DISTRIB. 146 (FLAS, MO, NCS, NY, OKL, PENN, TAES, TENN, WVA). KENTUCKY: Calloway Co., se. of Aurora, Smith & Hodgdon 4050 (au, Ny, us); McCreary Co., Cumberland Falls, McFarland & James, 2ND CENT. FL. KENT. 35 (DUKE, GA, MO, NY, PENN, TENN, WIS, WVA, US). TENNESSEE: Cheatham Co., Pegram, Svenson 10338 (an, TENN). ALABAMA: Lowndes Co., 23 mi. se. of Selma, Webster & Wilbur 3511 (MICH). MISSISSIPPI: Amite Co., 5 mi. sw. of Gloster, Webster & Wilbur 3277 (mica); Perry Co., 14 mi. w. of Beaumont, Webster & Wilbur 3405 (micn). Missouri: Howell Co., 4.5 mi. nw. of Willow Springs, Steyermark 23461 (F, MO). ARKANSAS: Pope Co., Nogo, Merrill 545 (TEX, UARK). LOUISIANA! Grant Parish, 15 mi. s. of Winnfield, Webster & Wilbur 3259( MICH). KANSAS: Cherokee Co., Hitchcock, Aug. 1896 (ksc). OKLAHOMA: Pushmataha Co., Antlers, Palmer 8315 (wo, wr, NY, Us). TEXAS: Smith Co., Swan, Reverchon 3120 (MO, SMU).

4. Sabatia quadrangula sp. nov.‘

Sabbatia cymosa in the sense of G. Don, Gen. Hist. 4: 207. 1838, as to

4Sabatia quadrangula sp. nov. Herba annua rosulata, caulibus quadrangularibus, ramis oppositis, pedicellis brevibus, 1-2(-4) mm. longis, corolla alba, 5-partita. Specimen ty picum legit R. Wilbur (n. 2899) prope Durham, in Carolina Septentrionali, et in Herb. Universitatis Michiganensis conservatum,

1955] Wilbur,— The North American Genus Sabatia 23

Don’s description but excluding reference to Chironia cymosa Lam., a synonym of Sabatia differmis (L.) Druce.

Sabbatia paniculata in the sense of authors after 1860 but not that of Michaux, the author of the basionym, nor of Pursh who made the combination. The earlier authors were describing the perennial now known as S. difformis (L.) Druce.

Sabatia brachiata f. candida Fern., RHopora 39: 443. 1937

Erect annual (15-)25-45(-75) cm. high, usually with but one stem

arising from each rosette, but occasionally with two to several. Roots few to more typically numerous, slender, fibrous, usually widely spreading and shallowly situated, about 4-8 em. long and less than 1 mm. in diam- eter. Branches usually restricted to upper one-quarter or one-third of stem, occasionally arising from nodes of the upper half of the stem or rarely from even the lcwermcst nodes, usually rather strongly as- cendent, forming an angle of 15-30 degrees, or occasionally up to 60 degrees, typically opposite along the stem and principal branches, the ultimate ramifications more commcnly alternate, the branches forming a flat-topped or eonvexly corymbcse loose to compact crown. Stem strongly quadrate, conspicucusly so below, 1-2(-3) mm. in diameter, strikingly although finely membranous wing-angled, wings thin, very low, less than 0.5 mm. in height in the lower portion of the stem and reduced above to the merest trace. Basal rosette present or absent at anthesis, even when present often not well-developed or conspicuous, when well-developed of spreading obovate to spatulate, obtuse to oc- casionally acute leaves tapering to an almost petiolate base, about 2-4 em. long and 1.8 em. wide. Cauline leaves membranous, ascendent, 3- nerved, or only l-nerved in smaller leaves, somewhat revolute, (0.8-) 1.6-3.2 (-5.8) em. long, (3-)4-8(-18) mm. wide, obtuse to acute, typically apiculate, usually narrowly to broadly oblong or lanceolate, usually about 3-5 times as long as broad but ranging from about 2-8 times as long as broad, typically at least the lowermost with a strongly clasping base. Internodes often equaling cr slightly exceeding the leaf, com- monly about 1-2 times as long, but ranging from 0.5-5 times as long as the leaves. Inflorescence of corymbosely to pyramidally arranged cymules, either one, both or none of whose lateral branches may be suppressed, flowers unusually closely associated in the cymule as the pedicels are always short, the flowers often sessile or nearly so, pedicels usually not more than 1-2 mm. in length, always less than 4 mm. long. Calyx-tube thin, 5 ridge-angled, about one-half as long as the corolla- tube, or about 2-3 mm. high, broadly turbinate, 1.5-3(-3.5) mm. long. Calyx-lobes thin, narrowly linear, (2-)4-8(-11) mm. long, usually about 0.5 mm. or less in width, but very rarely lobes as broad as 1.5 mm., erect in bud, and apparently ascendent at anthesis arching upward between the corolla-lobes, usually exceeded by the corolla-lobes by about 2-5 mm. Corolla-tube (2.5-)4-5(-7) mm. long, white but appearing greenish as the ovary is visible through the thin translucent wall. Corolla-lobes

24 Rhodora [Vor. 57

usually oblong, or spatulate, or somewhat elliptie, usually obtuse but not uncommonly acute, (4.5-)6-12(-15) mm. long, (1.5-)3-4(-60) mm. wide, spreading, pure white, occasionally with or more commonly without the basal yellow patch, often turning saffron-yellow upon drying, es- pecially along the reticulate veins. Filaments slender, white, 2-4 mm. long; anthers pale yellow, 1.5-3 mm. long. Stigmatie lobes greenish, lanceolate to oblong, 2-4 times the length of white style, 0.5-1.5(-2) mm. long. Capsule cylindrical, 4.5-7 mm. long, 3-4 mm. in diameter. TYPE LOCALITY: About four miles northeast of Durham, Durham County, North Carolina, rype: Wilbur 2899 (MICH). DISTRIBUTION: Fields, pine- lands and drier savannas from southeastern Virginia south to central Florida. Map 4.

This species has been known for almost one hundred years as Sabatia paniculata. This name, based upon Chironia panicu- lata Michx., has long been a source of confusion and error and has been applied by various authors to at least three different species. For almost the past century its application has been rather consistently restricted to the species which is here called S. quadrangula.

Michaux's original description in full is as follows:

paniculata. C. firmiter erecta: foliis lanceolato-linearibus; panicula multi- flora, brachiata, subfastigata; calyce subulato, corolla semibreviore.

Obs. Caulis lineis 4 prominulis quasiquadrangulus: folia inferiora interdum vali-lanceolata; suprema subulato-setacea. Co- rollae laciniae oblongae.

Hab. in Georgia et Carolina.

To be sure, there is very little descriptive information here that would be sufficient to allow anyone to be certain or even relatively confident as to which entity Michaux was attempting to characterize. Many of the phrases such as “C. firmiter erecta . . . panicula multiflora, brachiata, subfastigata: calyce subulato, corollae semibreviore . . . Corollae laciniae oblongae" might be applied with equal propriety to several other species. Of the five species included by Michaux, the flower-color of the four others is described as being rose; that of C. paniculata is not given. "Those characters of the leaf that were mentioned are rather indicative of S. difformis (“. . . foliis lanceolato- linearibus . . . folia inferiora. interdum ovali-lanceolata; su- prema subulato-setacea . . ."). The features of the stem described in the original diagnosis, especially “caulis lineis 4 prominulis quasi quadrangulus .’ are also a much more

1955] Wilbur,— The North American Genus Sabatia 29

apt description of the stem of S. difformis with its four elevated lines or ridges in the upper portion of the stem than it is of the very pronounced and strikingly 4-sided stem of S. quadrangula.

It was not surprising, therefore, to find that Michaux’s col- lection, as shown by a photograph in the Gray Herbarium of the type, was a specimen of S. difformis. This species is a perennial with a large rhizome; the stem below is terete or with four lines or ridges superimposed upon it, and above the stem becomes somewhat quadrangular; the leaves are ovate-lanceolate, lanceolate, linear to setaceous in a gradually modified sequence from the base to the inflorescence; the corolla-lobes are often 5 mm. or more longer than the longest lobes observed in S. quadrangula. The specimen in Michaux’s herbarium, as is witnessed by the very adequate photograph, is excellent and most of these features may be observed or interpreted from it. The underground portion of the plant, to be sure, is lacking. It is obvious that the lowermost part of the specimen (which can be seen even from the photograph to be terete or nearly so and approximately twice the diameter of the largest stem of the species that has most recently passed as S. paniculata) has been broken from a very substantial underground structure. Therefore Chironia paniculata is, both on the basis of its original diagnosis and upon the photograph of the specimen in Michaux's own collection in Paris, but a later synonym for Swertia difformis L. (1753), Chironia lanceolata Walt. (1788) and Chironia cymosa Lam. (1791).

Pursh (1814) transferred Michaux’s epithet to Sabatia (as Sabbatia) listing Swertia difformis as a synonym, stating by way of explanation that this species “certainly is the long lost Swertia difformis, as the specimens in the Herbarium of Sir Joseph Banks, sufficiently prove." Pursh also formally named as varieties the broad- and narrow-leaved forms of this species with C. lanceolata Walt. as a synonym of the latter. Elliott (1817) rather hopelessly confused the white-flowered Sabatias nomenclatorially and the influence of his splendid "Sketch" was rather long-lasting. The very different S. brevi- folia Raf. (S. elliottii Steud.) was very well-described, no doubt for the first time, but called S. paniculata with the following note by way of explanation: “Though the description of Michaux

26 Rhodora (Vou. 57

applies more peculiarly to the S. corymbosa [S. difformis], yet as this species was definitely included, and is the only one to which the term paniculata is correctly applicable, I have referred to him here." This freedom of interpretation on the part of Elliott caused confusion that persisted for many years. Grise- bach (1839) pointed out the confusion but apparently felt that it would be best to apply the name S. paniculata in the sense of Elliott, who had been followed by all American authors, and to adopt the name S. corymbosa for the perennial species. His list of synonymy showed a very complete understanding of the nomenclatoral propositions that had been made for this last-mentioned entity for the names of Linnaeus, Walter, La- marck, Michaux, Pursh, and finally Baldwin (or Elliott) are arrayed there. Grisebach apparently included the opposite- branched annual, here called 5. quadrangula, in the same concept for a duplicate of the specimen of Beyrich cited by him under S. corymbosa is the earliest collection of S. quadrangula known to me. Chapman (1860) was the first to define S. paniculata in the sense that it has been applied in recent times. His action in this, as in most other matters, was heavily influenced by ray as 1s shown from a letter (in the National Herbarium, dated December 1883) from Chapman to Dr. J. H. Mellichamp which is here quoted in part.

. . . I well remember the quarrel Gray and I had over it, I contending that the one we now call S. elliottii, our really only paniculate species, should bear that name. However, for the sake of uniformity as to our northern and southern Floras, I yield.

The appearance of the Synoptical Flora (1878) firmly es- tablished the usage of S. paniculata as the name of the white- flowered, opposite-branched annual and this name has been so applied since that date.

Rafinesque (Med. Fl. 2: 77. 1830.) published a description that in some ways strongly indicates this entity. This name has never been identified and I am at a loss to know what species Rafinesque had. Sabbatia nivea was described by him as having snowy-white, trichotomose flowers, a four-angled stem and oblong leaves which all would indicate S. quadrangula as well or better than any other species. However, the species was definitely stated to have been discovered by him in eastern

1655] Wilbur,—' The North American Genus Sabatia 27

Kentucky near the Cumberland River. There is no species known to me from that area which matches his description.

G. Don (1838) transferred Lamarck's C. cymosa to Sabatia applying it to what is here called S. quadrangula. This usage was not followed by any other author. The photograph of the specimen upon which Lamarck’s species was based is again what is now known as S. difformis.

Gray (1878) placed S. paniculata var. latifolia Pursh under the synonymy of what is now considered S. difformis while placing S. paniculata var. augustifolia Pursh under the species which I am calling S. quadrangula and which he called S. panicu- lata. In other words he felt that Pursh was including two species under S. paniculata and Gray also implied that Michaux had done so as well. I am aware of no evidence that has been presented that would warrant such a conclusion.

The only name that has been published previously for this species, whose identity has been for so long known to American botanists, is S. brachiata forma candida Fernald. This name must be assigned to the synonymy of S. quadrangula rather than to S. brachiata as the type specimen, Fernald & Long 6346, is not an albino of S. brachiata as claimed by Fernald, but rather is a specimen of the white-flowered species, S. quadrangula. Its identity is shown by the stems which below are strongly quadrate with conspicuously membranous-margined angles and whose pedicels are 2 mm. or less in length so that the flowers appear almost sessile. The only specimen cited by Fernald besides the type was Seymour 33 which came from Sussex Co., in south- eastern Virginia as did the type-collection of Fernald’s form. Seymour's plant, seen only at the Gray Herbarium, is S. brachiata as is indicated by the stem being terete below and some of the pedicels being over 5 mm. long and many of them over 2 mm. long. The flowers are bleached on the herbarium specimen but, as the sheet was originally determined as S. paniculata, they were very likely white. I also have seen white flowered specimens of S. brachiata in the field. However, since the designated type of Fernald's form is what is here called S. quadrangula, it wil be necessary for those who feel it useful formally to designate such minor variations to provide another name for albino plants of S. brachiata. Fernald's name could have

28 Rhodora [Vor. 57

been elevated to specifie rank for the annual species with a quadrate stem instead of providing an entirely new name. This procedure would seem undesirable as Fernald's intention in describing the form is clear; his choice of type was unfortunate. Besides this consideration, isotypes are not available for that number. Therefore, a new name has been provided and a type designated of which there are many duplicates.

REPRESENTATIVE SPECIMENS:—VIRGINIA: Chesterfield County, near Chesterfield Courthouse, Fernald & Long 6342 (an); Greensville Co., nw. of Taylor's Millpond, Fernald & Long 10785 (GH, PENN); Hanover Co., 1 mi. s. of Ashland, Ward, 8 Aug. 1885 (vs); Henrico Co., west of Elko Station, Fernald & Long 8818 (au); James City Co., south of Hotwater, Fernald & Long 8817 (DUKE, GH); Pittsylvania Co., Falls Creek, Heller 1105 (wv, us); Prince George Co., w. of Prince George Courthouse, Fernald & Long 8816 (am); Sussex Co., ne. of Homeville, Fernald & Long 6341 (an, MO, NY, PENN, US). NORTH CAROLINA: Anson Co., 6 mi. w. of Wadesboro, Smith, 30 July 1884 (au, us); Columbus Co., Nakina, Schallert, 30 June 1928 (oxra); Dare Co., Old Battle Ground, Bartley & Pontius 489 (Ny); Davidson Co., High Rock, Schallert 8660 (puKE); Durham Co., about 4 mi. n.e. of Durham, Wilbur 2899 (micu); Forsyth Co., without exact locality, Schallert, 18 Aug. 1940 (MO, UARK, wis); Granville Co., Oxford, Godfrey 5464 (DUKE, GH); Johnston Co., 0.5 mi. n. of Cox's Mill, Foz 4942 (micn); Orange Co., Hillsboro, Gray & Carey, July 1841 (au); Randolph Co., Asheboro, Hood 2399 (rLAs); Rowan Co., vicinity of Salisbury, Heller 130 (F, Mo, NY, PENN); Wake Co., just w. of Morrisville, Godfrey 49518 (DUKE, FLAS, GA, GH, MICH, NCS); Wayne Co., Goldsboro, Burlingame, July 1887 (BRU). SOUTH CAROLINA: Aiken Co., Vancluse, Eggert, 6 Aug. 1898 (Mo); Anderson Co., Long Branch Church, Davis, 29 July 1919 (mo, TEX, US); Beaufort Co., Bluffton, Mellichamp, 1884 (r, vs); Berkeley Co., 8 mi. s. of Monks Corner, Wiegand & Manning 2557 (mixed sheets) (cv, aH); Georgetown Co., 12 mi. n. of Georgetown, Godfrey & Tryon 26 (DUKE, F, GH, MO, NY, US); Jasper Co., Ridgeland, Mohr, 1893 (mo); Lexington Co., 8 mi. s. of Columbia, Godfrey & Tryon 1229 (mixed sheet) (xv); Williamsburg Co., Lanes Station, Burlingame, 20 July 1894 (BRU). GkoRGIA: Emmanuel Co., near Graymont, Harper 990 (au, wv, vs); Gwinnet Co., between the Alcovy River and No Business Creek, Small, 14 July 1893 (F, GH, MO, NY, US); Johnson Co., 2.5 mi. w. of Wrightsville, Pyron & McVaugh 3083 (GA); Mitchell Co., few miles e. of Pelham, Thorne 5078 (cu); Newton Co., 4 mi. n. of Coving- ton, Pyron & McVaugh 3036 (G^, NA); Sumter Co., without exact locality, Harper, July 1897 (Ny); Telfair Co., McRae, Biltmore Herb. 4511 (us); Wash- ington Co., 3 mi. n. of Harrison, Pyron & McVaugh 3104 (NA, NY). FLORIDA: Bay Co., Panama City, Knight, 6 July 1943 (Frias); Calhoun Co., without exact locality, Chapman (Mo); Dixie Co., near east limits of Jena, Beaman 280 (mic); Franklin Co., Apalachicola, Chapman (au); Gulf Co., north of Port Saint Joe, Small, DeWinkeler & Mosier, 11 July 1924 (DUKE, FLAS, GH, MICH, MO, NCU, NY, PENN, TENN, WVA); Levy Co., Rosewood, Garber, June 1876 (BRU, F, NY); Hamilton Co., near Jennings, Bright 3902 (wis); Taylor Co., between Perry and the Gulf of Mexico, Small, Small & DeWinkeler 11452 (Ny, vs); Wakulla Co., prope St. Marks, Kugel May 1843 (mo, us); Walton Co., Freeport Mohr, 18 June 1880 (vs).

1955] Wilbur,—The North American Genus Sabatia 29

5. Sabatia brachiata Ell., Sk. Bot. S. C. & Ga. 1: 284. 1817. Chironia angularis var. B angustifolia Michx., Fl. Bor. Am. 1: 147. 1803. Sabbatia concinna Wood, Class-Book ed. 2. 451. 1847. S. angustifolia (Michx.) Britton, Mem. Torr. Club 5: 259. 1889.

Erect annual (10-)25-50(-67) em. tall, usually but one stem arising from each rosette, more rarely with 2 or 3. Branching usually restricted to the upper one-third or half of stem but in more robust plants with branches sometimes developing from the lowermost nodes, usually rather strongly ascendent forming an angle of 15-40 degrees with the main stem, rarely spreading to as much as 70 degrees, typically opposite, al- though rarely alternate along main stem or principal branches, or ultimate branches somewhat more commonly alternate, forming convexly corym- bose or somewhat pyramidal loose to compact crown. Stem 1-2(-4) mm. in diameter, terete, conspicucusly so below and smooth, not wing- angled, although sometimes noticeably lined or finely ridged. The stem above and upon the branches becoming subquadrate or even quadrate and bearing very slight wings at the angles. Roots few to numerous, slender, fibrous, spreading or descendent, about 4-8(-10) em. long, 0.5-1.5 mm. in diameter. Basal rosette typically present at time of flowering, usually conspicuous, typically composed of numerous overlapping, spread- ing, usually broadly spatulate, rarely elliptie, obtuse to seldom acute leaves tapering strongly to an almost petiolate base, (8-)15-30(—45) mm. long, (6-)10-14(-18) mm. wide. Cauline leaves membranous, ascendent, 3-nerved or rarely only I-nerved in smaller leaves, (1—)1.5-3 (—4.5) em. long, (3-)4-10(-16) mm. wide, often obtuse, especially below, or acute, commonly slightly callose-apieulate, oblong, with more or less parallel margins, or more rarely elliptie or somewhat lanceolate, tapering somewhat to base, at least usually not strongly or broadly clasping. Internodes usually about 1.5-3 times longer than the leaves, but ranging from but about three-fourths to about 4-times as long as the leaves. Inflorescence of corymbosely to pyramidally arranged cymules either 1, both, or none of whose lateral branches may be surpressed. Pedicels 5-sided, noticeably thin-ribbed, (1-)2-8(-13) mm. long. Calyx-tube thin, noticeably but finely 5-ribbed, usually about half as long as the corolla-tube, (1—-)1.5-3(-4) mm. long, turbinate to campanulate. Calyx- lobes thin, narrowly linear, (4-)7-10(-15) mm. long, 0.3-1.0 mm. wide, as- cendent in bud, wide-spreading at anthesis, usually exceeded by the corolla lobes by 3-8 mm. Corolla-tube (3-)4-5(-6) mm. long, 1.5-3 mm. in di- ameter, cylindrical, greenish within and withcut. Corolla-lobes oblong, or narrowly to broadly spatulate, usually obtuse, or occasionally tapering to an acute apex, (5-)7-14(-20) mm. long, (2-)3-6(-8) mm. broad, wide- spreading at anthesis, pale-pink to darker roseate, rarely white, with low triangular greenish-yellow area at base of lobe, usually bordered by reddish line. Filaments slender, pale yellow to nearly colorless, (1.5-)2-3(-4) mm. long; anthers bright yellow, usually about (2-)3(-4) mm. long. Stigmatic lobes slender, (2-)3-5(-7) mm. long; style 2-4 mm. long, usually less than the stigmatic branches in length. Capsule cylin-

30 Rhodora [Vor. 57

drical, 5-8 mm. high, 3-4.5 mm. in diameter. TYPE LOCALITY: “Grows in the middle and upper country of Carolina. Near Columbia," Lexing- ton Co., South Carolina. type: Herbemont s. n. (CHARL!). DISTRIBU- TION: Fields, dry and open oak and pine woods ranging from south- eastern Virginia south into Georgia and westward to southern Missouri and Louisiana. Map 5.

This well-marked species has been rarely confused with any other, except occasionally with specimens of S. angularis or S. quadrangula (which for almost a century has been called S. paniculata). Both of these last-mentioned species are, however, readily distinguished from S. brachiata by their strongly tetra- gonal stems whose angles are strikingly membranous winged. S. brachiata is strongly contrasted to those two species by pos- sessing, especially below, a smooth, terete stem. Above and on the branches the stem is somewhat finely ridged and angled but not conspicuously so. The corolla of S. brachiata is typically pink or roseate and very robust specimens, especially in areas where 5. angularis is more commonly to be expected, have been rather often mistaken for that wide-spread species. Albinos of S. brachiata are rarely encountered, but when they are found or when the corollas become entirely bleached as they often do in drying, these plants are often mistaken for what has previously been called S. paniculata (= S. quadrangula). The type of S. brachiata forma candida (Fernald & Long 6346), is not S. brachiata, which normally has a rose-colored corolla, but S. quadrangula (S. paniculata in the sense of recent authors) which always has a white corolla. Fernald’s misidentification of the type-sheet is easily demonstrated: the lower portion of the stem of the type is very distinctly quadrate with winged margins which is a distinctive feature of S. quadrangula. The only other specimen cited with the original description of this "form" was Seymour 33 which is definitely a specimen of S. brachiata. A new name will be required by those who feel it essential to designate formally such minor variations.

Chironia angularis B augustifolia, of Michaux, has been as- signed to the synonymy of this species at least since the ap- pearance of the Synoptical Flora, where it was so listed. Not having seen an authentic specimen from Michaux's herbarium, it is only tentatively that I place the name in the synonymy of S. brachiata. The original characterization of the variety was

1955] Wilbur,—The North American Genus Sabatia 3l

very brief ("8 augustifolia: folis quasi lanceolatis; supremis etiam linearibus") and certainly states nothing that would exclude forms of S. angularis from consideration or indicates clearly that the entity described was S. brachiata. Both this variety of Michaux's and his latifolia, which has been considered the sole element of Michaux's concept that is S. angularis as now interpreted, were arranged under the generalized diagnosis of the species. The stem in that account of the species is stated to be “marginato-quadrangulo” and this would seem to exclude the entity now known as S. brachiata. Perhaps Gray saw an authentic specimen at Paris and hence may be correct in his treatment; in any event the question is yet to be settled.

S. concinna Wood is assigned to the synonymy of this species upon the basis of the original description which led Gray as early as 1856 to the same conclusion. It is perhaps of some significance that Wood himself later listed the name as a syn- onym of S. brachiata. The oXginal description follows:

3. S. concinna. Wood (Nov. sp.) Elegant Star Flower. St. slender, sub- quadrangular, internodes 2—4-times longer than the leaves; branches oppo- site, sub-erect; lvs. linear and lance-linear, lower ones ovate, all acutish, sessile, panicle oblong; cal. segments linear, twice longer than the tube, twice shorter than the corolla; cor. 5-parted, segments oblong-obovate, obtuse, light purple.—Dry grassy prairies, Ia.! abundant. Stem a foot high, few or many-flowered. Leaves 9-12" by 1-3", flowers 15" diam., of a delicate blush purple, the star in the center yellow, bordered with green. Jl. Aug.

Merrill (RHopora 50: 127. 1948.) concluded that “it is the same as S. campestris Nutt., not S. brachiata Ell. as the latter species does not occur in Iowa; Wood's type was from Iowa." However, Wood described the branches as opposite and the calyx-segments as linear and only half as long as the corolla and these features would exclude from consideration anything but an unusual specimen of S. campestris. If'the plant were S. campestris, it would hardly seem possible that the very peculiar and prominent costae of the calyx-tube could be overlooked in even the most cursory examination. I have seen no specimen of S. campestris from Iowa. Merrill placed considerable weight of evidence upon the locality of the plant but his interpretation of the abbreviation “Ia.” is certainly an error. In 1846 between

32 Rhodora [Vor. 57

the first and second editions of his Class-Book, Wood, enlarging the area to be covered by his flora, made a short collecting trip to Indiana. S. concinna was presumably seen upon that trip. The explanation of geographical abbreviations appearing in

the second edition informs one that “the names of . . . states . are often abbreviated, and always in the same manner as in other works; thus . . . Ia. or Ind., Indiana, &c." In this

same edition (1847, p. 3) the area of the flora was said to be "essentially the States lying north of the Ohio River and Mary- land." One also learns in Wood's ‘American Botanist and Florist" that the “geographical limits of the present flora are the same as those adopted in the Class-Book: viz, all the States of the American Union lying east of the Mississippi River." Although the area covered by Wood's publications was expanded in later years, they were never intended to cover the region west of the Mississippi River except indirectly. Wood meant Indiana by the abbreviation “Ia.” Fernald (Rhodora 47: 404— 405. 1945) concluded as much from indirect evidence.

S. brachiata is not represented from Indiana in any of the herbaria that I have studied and neither it nor S. campestris were included by Deam in his Flora of that state. In fact, Deam (1940, p. 1080) after considering the evidence and after finding that that species was not represented in either the Gray Herbarium or in that of the New York Botanical Garden defi- nitely excluded it from the known flora of the state. Fernald (1950) lists the range of the species as including southern Indiana. S. brachiata is not represented from Indiana, Illinois, or Kentucky in any of the collections that I have studied. The nearest known stations are southeastern Missouri and in Tennessee.

The description of the stem as being ''subquadrangular" is also a bit puzzling if the name really is synonymous with S. brachiata, for that species is very conspicuously marked by a decidedly terete stem in at least the lowermost portion. The upper part of the stem and the branches do become rather strongly angled and, if S. concinna is the same as S. brachiata, this explanation probably accounts for that at first puzzling statement in the description. The disposition of Wood’s name, in the absence of authentic material, will remain tentative, but the evidence available indicates that it was S. brachiata.

1955] Wilbur,— The North American Genus Sabatia 33

REPRESENTATIVE SPECIMENS:—VIRGINIA: Princess Anne Co., Little Neck, Fernald & Long 4134 (GH, PENN); Prince George Co., just s. of Disputanta, Fernald & Long 8815 (GH); Sussex Co., northeast of Homeville, Fernald & Long 6344 (GH, NY, PENN, US). NORTH CAROLINA: Anson Co., 4 mi. n. of Ansonville, Boyce 1127 (Ncs); Beaufort Co., 7 mi. s. of Washington, Godfrey 4400 (GH, US); Brunswick Co., 3 mi. n. of Bolivia, Wilbur 2888 (micu); Bladen Co., without exact locality, Biltmore Herb. 4511a (GH, MO, NCU, NY, PENN, US); Carteret Co., 3 mi. se. of Newport, Godfrey 48192 (Ncs); Cumberland Co., 7.5 mi. n. of Fayetteville, Godfrey & For 49446 (DUKE, Ncs); Harnett Co., 6 mi. e. of Cameron, For 2546 (ncs); Montgomery Co., 1 mi. n. of Ether, Wiegand & Manning 2543 (cu); Moore Co., 2 mi. s. of West End, Fox & Whitford 3903 (mica, Ncs); New Hanover Co., Wilmington, McCarthy, Aug. 1885 (vs); Onslow Co., at Richlands, Godfrey 4475 (an, vs); Sampson Co., Roseboro, Godfrey 4532 (DUKE, GH, US); Scotland Co., 12 mi. n. of Laurinburg, Godfrey 5046 (DUKE, GH, US); Wake Co., Raleigh, Godfrey 4918 (GH, NCU, NY); Wilkes Co., up Pores Knob, Radford & Stewart 1743 (NCU). SOUTH CAROLINA: Chesterfield Co., near Cheraw, Ward, 25 June 1895 (Ny, us); Darlington Co., Society Hill, Canby, July 1878 (r, ny); Horry Co., without exact locality, Adams 30 (PENN); Lexington Co., 8 mi. s. of Columbia, Godfrey & Tryon 1229 (GH, NY [a mixed sheet], vs). GEoRGIA: Bartow Co., 1.25 mi. e. of Emerson, Duncan 8548 (GA, MO); DeKalb Co., northern slope of Stone Mountain, Small, 17 July 1893 (r, xv); Douglas Co., 10 mi. s. of Douglasville, Duncan 3648 (micH); Pickens Co., near Jasper, Biltmore Herb. 4511a (us); Richmond Co., Augusta, Cuthbert, Aug. 1876 (RUT); Taylor Co., without exact locality, Pyron, 3 Aug. 1930 (DUKE). TENNESSEE: Coffee Co., near Manchester, Clebsch & Clebsch 4770 (TENN); Franklin Co., between Tullahoma and Estill Springs, Svenson 9211 (au); Grundy Co., east of Altamont, Svenson 7137 (aH, TENN); McNairy Co., without exact locality, Bain 431 (an, Ny); Van Buren Co., Falls Creek State Park, Shanks, Clebsch & Sharp 3431 (MICH, TENN); White Co., sw. of Clifty, Shanks, Clebsch & Sharp 2916 (TENN). ALABAMA: Mobile Co., about 8 mi. sw. of Mobile, Webster & Wilbur 3479 (mica); Wash- inton Co., Fruitdale, collector not stated, July 1904 (Mo). wississiPPI: Coving- ton Co., 1.5 mi. se. of Mt. Olive, Webster & Wilbur 3308 (icu); Forest Co., about 14 mi. s. of Hattiesburg, Webster & Wilbur 3397 (micu); Harrison Co., 3 mi. s. of Saucier, Webster & Wilbur 3442 (mic); Jackson Co., Ocean Springs, Pollard 1083 (mo, NY, us); Jones Co., Laurel, Tracy 3356 (Ny); Pearl River Co., Poplarville, Tracy 1687 (Gu, vs); Pike Co., Holmesville, Wheeler (mo); Stone Co., 8 mi. s. of Wiggins, Webster & Wilbur 3440 (mica); Wayne Co., 6 mi. nw. of Ala.-Miss. state line, Sargent, 18 June 1950 (OKL, NCS). MISSOURI: Butler Co., 12 mi. n. of Poplar Bluff Steyermark 11593, (MO). ARKANSAS: Faulkner Co., Conway, Haas 1629 (us); Drew Co., Ladelle, Demaree 22321 (MO, OKLA); Lonoke Co., Grand Prairie, Demaree 22333 (MO, NY, OKL, OKLA); Prairie Co., DeValls Bluff, Demaree 22176 (mo); Pulaski Co., near Little Rock, Carpenter, June 1938 (UARK). LOUISIANA: Calcasieu Parish, Lake Charles, Daves, Aug. 1888 (F); Grant Parish, 4 mi. s. of Pollock, Webster & Wilbur 3257 (MICH); Orleans Parish, New Orleans, Drummond 224 (an, x); Rapides Parish, 12 mi. se. of Hineston, Webster & Wilbur 3275 (micu); St. Tammany Parish, vicinity of Covington, Arséne 11741 (vs); Tangipahoa Parish, 6 mi. e. and 1 mi. n. of Hammond, Nease, 1945 (okL); Vernon Parish, 2 mi. w. of Leander, Webster & Wilbur 3228 (wicH); Washington Parish, Bogalusa, Cocks, 5 June 1917 (xo).

(To be continued)

34 Rhodora (Vor. 57

ILEX GLABRA AND A NEW STATION FOR KALMIA LATIFOLIA IN New HawrsnuiRE.— The Inkberry, Ilex glabra (L.) Gray, has never been validly reported to occur naturally in New Hamp- shire. However, it has long been known to be present in Maine on Isle au Haut as well as in Nova Scotia.

The frequency of stations for it in Essex County south of the Merrimack River, suggests that one might expect to find it in similar habitats along the coast in Rockingham County, New Hampshire.

In October 1950 some fragments of Inkberry were mailed to me from Seabrook accompanied by a brief diseussion of the plants' whereabouts, the inference being that the station was within the boundaries of New Hampshire.

Not until 1954, was it convenient to arrange a visit to the Ilex-colony. But in March of that year, though temporarily viewed with some quite undeserved suspicion as a vandal, I was permitted to see the somewhat straggling and recently molested colony, and was even encouraged to bring away a small living plant. The herbarium specimen which unfortu- nately is meagre, though quite adequate for verification, is deposited in the University of New Hampshire Herbarium. The colony definitely is in New Hampshire, but rather less than a half mile from Massachusetts.

It will be interesting to watch the progress of this colony. Recent growth of shading Pinus Strobus in the immediate vicinity suggests vegetational changes perhaps detrimental to the plant’s growth. Possibly in earlier times the colony was more vigorous and extensive and what we have now is a mere remnant of a much larger stand. Out of respect for the wishes of the owners in the matter, more precise information concerning the plant's whereabouts will not now be published.

Kalmia latifolia L. is one of numerous species which have mostly disjunet ranges in New Hampshire. Reaching the northeastern limit of its range in south-central and southeastern Maine, Mountain-laurel becomes increasingly more frequent as one approaches its more or less continuous area beginning in New Hampshire, west of the Merrimack River and including those upland townships which are adjacent to Massachusetts or which lie not many miles to the north. Unlike many other

1955] Hodgdon,— Ilex glabra 35

species of southern distribution, Kalmia latifolia does not extend northward appreciably in the Connecticut Valley. Fernald? summarizing botanical studies made in western Cheshire County during three summers, reported it from only one station. He described it as “Forming an extensive thicket by the Ashuelot River, Gilsum," while H. G. Jesup? mentioned only the well known Squam Lake Station. Kalmia latifolia is a normal constituent of the acid upland forests of southwestern New Hampshire becoming only occasional in the different type of forest of the Connecticut Valley. To the east of the Merrimack River, Mountain-laurel again is very localized in the dominant White Pine Region of southeastern New Hampshire. Specimens are represented in herbaria from favorable sites in southeastern New Hampshire where locally the plant is luxuriant, in the townships of Pelham, Windham, Sandown and Barrington. Doubtless it occurs elsewhere in this region but to most persons living there, Kalmia latifolia is either totally unknown as a native or is exceedingly rare.

Farther northward in New Hampshire as in Maine the plant becomes less frequent. Reports from creditable observers have reached me of colonies in New Durham and Effingham. These I have not visited, but an herbarium specimen from Chase Hill in Albany, confirms the existence of a vigorous colony there with which I am familiar.

In central New Hampshire, there is the isolated colony near Squam Lake in Holderness, evident from the highway and familiar to all. This I had always believed to be the most outlying stand in this part of New Hampshire. But in mid- July 1954, in company with Mr. Radcliffe Pike, I visited Peaked Hill Pond, in Thornton, Grafton County, to investigate a report of “Rhododendron” in the vicinity. A half-acre of ''Rhodo- dendron" near the southern shore turned out to be two or more acres of Mountain-laurel. The colony occurs in deciduous woods, with a scattered admixture of large hemlocks and is somewhat away from the water on the southern and southeastern sides of the pond. "Phe growth of laurel is noticeably dwarf, most of the plants not exceeding 3 or 4 feet in height, in marked

1 RHODORA, 3: 235, September 1901. . . ? A catalogue of the Flowering Plants and Higher Cryptogams found within about thirty miles of Hanover, New Hampshire, 26 (1891).

36 Rhodora [Vor. 57

contrast to some of the colonies in southeastern New Hampshire which contain plants averaging much higher than a man’s height. Here at nearly 1200 feet altitude in an area exposed to winds from the north the necessity for protection by deep snow in severe winter weather may account in some way for the low stature of the plants. Peaked Hill Pond is rather less than 30 miles northeast of Hanover and not more than 15 miles, as the crow flies, from the area of Kalmia latifolia at Squam Lake. Specimens from Thornton are deposited in the Herbaria of the University of New Hampshire and the New England Botanical Club.—A. R. HopGDON, UNIVERSITY OF NEW HAMPSHIRE.

A NEw VARIETY OF SOLIDAGO ULMIFOLIA.—Solidago ulmifolia Muhl., var. palmeri Cronq. var. nov. Caulibus pilis patentibus obtectis.

TYPE: Palmer 24111, shaded sandstone slopes, north side, near top of Magazine Mt., Logan Co., Arkansas, October 14, 1923; deposited at the New York Botanical Garden. ADDITIONAL SPECIMENS: Arkansas: Garland Co., Palmer 29092, 29194; Demaree 11001, 20498, 20512, 21836. Logan Co.: Palmer 24180; Demaree 8072. Perry Co.,: Demaree 20145. Pope Co.: Demaree 19876. Yell Co.: Demaree 15942. Alabama: County undetermined: Buckley s.n., October, 1838, without further data; Biltmore Herb. 15008, Avondale.

The var. ulmifolia, with the stems essentially glabrous below the inflorescence, is largely replaced in the Ouachita region of Arkansas by the var. palmeri. The existence of two specimens of var. palmeri from Alabama in the herbarium of the New York Botanical Garden indicates that the hairy-stemmed phase of the species is not purely local, however, and the limits of its range remain to be determined.

Several of the specimens (including the type) here cited as S. ulmifolia var. palmeri were annotated by K. K. Mackenzie with an apparently unpublished binomial using the epithet palmeri in conjunction with the genus Solidago. "The treatment of the new variety was inadvertently omitted by me from an earlier paper dealing with eastern American goldenrods, with the result that the name as used on page 428 of volume 3 of Gleason’s Illustrated Flora is technically a nomen nudum.— ARTHUR CRONQUIST, THE NEW YORK BOTANICAL GARDEN.

Volume 56, no. 672, including pages 261-286, was issued 10 January 1956,

Dodora

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club. by REED CLARK ROLLINS, Editor-in-Chiet

ALBERT FREDERICK HILL ~. STUART KIMBALL HARRIS l

RALPH CARLETON BEAN Associate Editors RICHARD ALDEN HOWARD (

CARROLL EMORY WOOD, JR.

Vol. 57 February, 1955 No. 674 CONTENTS:

Studies in the Hippocastanaceae, I. Variation within the Mature Fruit of Aesculus. James W. Hardin..................... 37

A Revision of the North American Genus Sabatia (Gentianaceae). Robert L. Wilbur (continued from page 33)................. 43

Epifagus virginiana. Samuel C. Palmer........... cee cece cece 71

Calycera balsamitaefolia in the United States. Julian A. Steyer- mark. and Floyd A. Swink... reaa ua aiena raah 72

The New England Botanical Club, Jne.

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RHODORA.—A monthly journal of botany, devoted primarily to the flora of the Gray's Manual Range and regions floristically related. Price, $4.00 per year, net, postpaid, in funds payable at par in United States currency in Boston; single copies (if available) of not more than 24 pages an ! wich 1 plate, 40 cents, numbers of more than 24 pages or with more than 1 plate mostly at higher prices (see 3rd cover-page). Back volumes can besupplied at $4.00. Some single numbers from these volumes can be supplied only at advanced prices (see 3rd cover-page). Somewhat reduced rates for complete sets can be obtained on application to Dr. Hill. Notes and short scien- tific papers, relating directly or indirectly to the plants of North America, will be considered for publication to the extent that the limited space of the journal permits. Illustrations can be used only if the cost of engraver's blocks is met through the author or his institution. Forms may be closed five weeks in advance of publica- tion. Authors (of more than two pages of print) will receive 15 copies of the issue in which their contributions appear, if they re- quest them when returning proof. Extracted reprints, if ordered in advance, will be furnished at cost.

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Rbodora

JOURNAL OF

THE NEW ENGLAND BOTANICAL CLUB

Vol. 57 February, 1955 No. 674

STUDIES IN THE HIPPOCASTANACEAE, I. VARIATION WITHIN THE MATURE FRUIT OF AESCULUS

JAMES W. HARDIN!

A wIDE variation in fruit size and shape is readily noticeable from observations of an Aesculus tree or shrub in late summer (Fig. 1). Upon closer examination it becomes apparent that this variation is due in part to the number of maturing seeds in the ovary.

The ovary is typically 3-locular and 3-carpellate with two ovules in each locule. These ovules are oriented differently, the lower descending with a dorsal raphe, and the upper hori- zontal or more often ascending with a ventral raphe. Each fruit, therefore, is potentially six-seeded.

The literature is not clear in regard to the number of seeds usually found in the mature fruit. Gray (1849) stated that the locules are one seeded by the abortion of one ovule in each, and by suppression the fruit is often 1- to 2-celled and 1- to 2- seeded. Pax (1895) said that the mature fruit is mostly 1- seldom 2-locular with one seed. Rendle (1925) also stated that only one seed per fruit develops due to the crushing of two out of three cells by the considerable growth of the one ovule. Chapman (1897) on the other hand described the fruit as being l- to 3-seeded. The more recent literature presents the same information: Fernald (1950)—3-seeded or usually by abortion 2- or I-loeular and 2- or 1-seeded; Lawrence (1951)— usually 1-loeular and I-seeded; Gleason (1952)—seeds by

! My sincere thanks go to Professor Rogers McVaugh, University of Michigan, for his helpful suggestions during the preparation of the manuscript.

38 Rhodora [Vor. 57

abortion only one and subglobose, or 2 and semiglobose, or rarely 3 with flattened sides. Summarizing this information then, one might expect to find usually one, or possibly up to three, seeds per fruit.

Since there seems to be as much variation in the literature as in the fruit, and more than three seeds per capsule are often found, it was thought necessary to make some actual counts in conjunetion with a much broader study of the entire genus.

Collections of fruit from six species were made during the summer of 1954 and studied from the point of view of the num- ber of mature seeds, the position of these seeds within the capsule and the number of locules present. "The result of this analysis is presented here.

Each fruit was examined and the number of mature seeds and their position recorded. In Table 1, "top" and “bottom” refer to the ascending and descending ovules respectively. The fifteen position-combinations were determined from the abortive ovules more easily than from the mature ones since the former more nearly retain their original position after the great en- largement of the seeds. More combinations could be recognized if, e.g. with 3 and 4 seeds, it were determined whether one or both ovules mature within the same locule. This information was kept only for 2-seeded combinations—both seeds in the same or different locules. Also given in Table 1 is the number of fruits found with a particular combination and the percentage based on the total fruits examined for each species.

From three trees of A. hippocastanum L. growing on the University of Michigan Campus and from two trees along a highway in Franklin County, Ohio, a total of 107 fruits was collected and examined. Table 1 indicates that the 1-seeded fruit was the most common type and that the ratio of “top” to *bottom" was essentially 1:1. In the 2-seeded type the most frequent combination was the 1 — 1 diff., meaning one ascending and one descending seed but in different locules.

An analysis was made of 153 fruits of A. glabra Willd. which were collected from a small population in Washtenaw County, Michigan. The l1-seeded capsule was again found to be the most common (Table 1) and maturation of ovules occurred in essentially the same numbers for ascending and descending.

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40 Rhodora [Vor. 57

Aside from percentages which differ between A. hippocastanum and A. glabra, the most obvious difference is the occurrence in A. glabra of fruit with four, five and six mature seeds. Only one fruit each of the 5- and 6-seeded type was found, but it is significant that this condition does occur, even though seemingly rare for the species as well as the entire genus.

Fifty five capsules of A. octandra Marsh., from a population in Haywood County, North Carolina, were studied. Over half were of the 1l-seeded type, although 2-, 3- and 4-seeded forms were found in decreasing frequencies.

A total of 145 capsules of A. sylvatica Bartr. were examined from two populations, one from Clarke County, Georgia and the other introduced from Georgia and now growing at the University of Michigan Botanical Garden. Again there was a high percentage of l-seeded fruits, but also relatively high percentages of 2- and 3-seeded forms were found.

A very small and probably clonal population of A. parviflora Walt. was visited in Early County, Georgia during June 1954. Mature fruits were not available, but a population sample of partially mature fruits was collected. Analysis of these showed only the one-seeded type. This lack of variation is of interest since the population is probably one clone which occuples an area of nearly 400 square feet and includes approxi- mately 75 trees.

Sixteen capsules were kindly sent from a small population of what is tentatively considered A. discolor Pursh growing near Old Sutherland Springs, Wilson County, Texas. It was unusual to find no 1-seeded fruits among this collection. Over half were of the 2-seeded type and the rest 3- and 4-seeded.

Considering all six of these species together, percentages are given (Table 1) for the genus based upon the random samples totaling 504 capsules. The emphasis which previous authors have placed upon the 1-seeded type is not justified on the basis of these figures which show that this form is present in barely more than one half of the total. Over one quarter of all fruit was of the 2-seeded form and just under one eighth had three seeds. "The most significant deviation from the literature is the nearly 4% of the total which contained four to six seeds.

Correction should be made of the error found in the literature regarding the interpretation of the number of locules present

1955] Hardin,—Studies in the Hippocastanaceae 41

in the mature fruit. Regardless of the number of seeds maturing, and the crushing of the locules by the excessive growth of the seeds, each of the locules remains at least partially distinct with its abortive ovules in their original position. The typical fruit then, is always 3-locular regardless of whether 1-seeded or 2- to 6-seeded.

Exceptions to this 3-locular condition were found in two of the species. Four-carpellate, 4-locular and in maturity 4-valved fruits of A. hippocastanum were found among the collections (Fig. 2). Also 2-locular and 2-valved fruits of A. sylvatica were found (Fig. 3). Of the 107 capsules of A. hippocastanum examined, thirteen of them, or 12%, were 4-locular. The majority of these were of the l-seeded type, only a few having 2 seeds. Each of the four locules was distinct at maturity and each contained the expected two ovules. Only two capsules of A. sylvatica out of the 145 examined were 2-locular. One of these was l-seeded and the other 2-seeded. The expected total of four ovules was present in both cases. The anomalous condition of 2 or 4 locules is rare, or at least infrequent in A. hippocastanum. Just what evolutionary significance these conditions have, if any, is not yet known.

At the outset of this study, the question arose as to whether only the ascending or descending ovules matured, or if both, in what proportions. The answer is shown in Table 1. A 1:1 ratio was found in the 1-seeded forms. In the 2-seeded and 3-seeded capsules only slightly more of the ascending ovules matured. These ratios show that the position of the maturing ovule in respect to ascending or descending is entirely random, and abortion is not due to a morphological degeneration of a portion of the ovary or placenta.

The cause of abortion in the majority of ovules is not known. According to Gray (1849) and Rendle (1925) the pressure from the maturing ovules is the important factor. If this were true, one would expect to find some degree of enlargement in all six ovules. Fig. 4 illustrated one of the few cases seen where pressure was effective in retarding the growth of ovules. In this case the lower two ovules did enlarge greatly until presumably stopped by pressure from the top. The smaller two were not true abortives since they expanded to full size after the capsule was opened. In practically every other case, the abortive

42 Rhodora [Vor. 57

ovules were 1-4 mm. in diameter—the size being nearly constant within a single capsule. Even when one mature seed was found in a locule, the second and abortive ovule in that locule showed no signs of being crowded. The “pressure” theory then, does not seem to be the answer in the majority of cases.

Incomplete fertilization could possibly be the determining factor. If this were true, then the unfertilized ovules would not ordinarily enlarge, which is found to be the case in these capsules. This incomplete fertilization could arise because of an incomplete pollination, or more probably caused by some morphological, physiological or genetic sterility factor. The answer to this question involves a major research problem in itself.

The phylogenetic significance of seed number in the genus Aesculus is not completely understood at this point. If one extreme in number (l-seeded or 6-seeded) is more primitive than the other in terms of evolution of the family, genus or species, the interpretation may be made only after a more complete study of the related taxa within the Sapindales.

DEPARTMENT OF BOTANY, UNIVERSITY OF MICHIGAN,

LITERATURE CITED

CHAPMAN, A. W. 1897. Flora of the Southern United States. Ed. 3. Amer. Book Co., N. Y.

FrERNALD, M. L. 1950. Gray’s Manual of Botany. Ed. 8. Amer. Book Co., N. Y.

Gurason, H. A. 1952. The New Britton and Brown Illustrated Flora of the northeastern United States and adjacent Canada. Vol. II. Lan- caster Press, Pa.

Gray, Asa. 1849. The genera of the plants of the United States. Vol. II. George Putnam, N. Y.

Lawrence, G. H. M. 1951. Taxonomy of Vascular Plants. Macmillan Co., N. Y.

Pax, F. 1895. Hippocastanaceae. In Engler and Prantl, Natürl. Pflanz- enfam. III (5): 273-276.

RENDLE, A. B. 1925. The Classification of Flowering Plants. Vol. II. Cambridge.

1955] Wilbur,—The North American Genus Sabatia 43

A REVISION OF THE NORTH AMERICAN GENUS SABATIA (GENTIANACEAE)

Rosert L. WILBUR (Continued from page 33)

C. Subsection campresTREs Blake, Ruopora 17: 56. 1915.

Neurola Raf., New Fl. 4: 92. 1838.

Annuals, branches typically alternate, at least above. Basal rosette never present. Leaves either thin and membranous or thick and suc- culent, margins not scarious. Calyx-tube conspicuously ribbed or costate by the prominent development of the fused lateral traces and bearing a thin membranous wing. TYPE SPECIES: Sabatia campestris Nutt.

This subsection contains but two species which are obviously closely related to one another and which are strongly and unique- ly distinguished from the other groups by the prominently developed calycine costae. In addition, the typically alternate branching pattern sets this small species-group apart from subsection Angulares, to which it is probably most closely related. Also, the Angulares possess a very well-developed basal rosette at some stage in their life-cycle although it is not always per- sistent at time of flowering; the Campestres are never rosulate. An additional indication, besides the aspect of the plants, the texture of the leaves, and perhaps the habit, of the relationship of this group to the opposite-branched Angulares is the pronounced quadrate stems in the species of both subsections (at least above) which are margined along the angles by a thin, hyaline membrane.

The distribution of the Campestres is almost entirely west of the Mississippi River where the subsection occupies an ex- tensive area. S. campestris accounts for the bulk of the sub- sectional range. Sabatia arenicola is very obviously related to S. campestris and is believed to have been derived from that species or from its precursor. S. arenicola is at present known only from the Quaternary littoral deposits along the beaches from Louisiana to southern Tamaulipas.

KEY TO THE SPECIES OF SUBSECTION CAMPESTRES

A. Leaves and calycine lobes thin and membranous, neither succulent nor heavily cutinized; leaves broadest near the base, usually strongly clasping the stem, the midvein at least prominently elevated beneath; calyx-lobes usually 2-6 times longer than the calyx-tube, generally 4-8 times longer than broad; corolla-lobes typically equaling or exceeding the calyx-lobes,

44 Rhodora [Vor. 57

usually longer than 1 em. and wider than 7 mm.; stigmatic lobes over 5 mm. long; anthers longer than 2 mm.; widespread in prairies and fields from Texas to Mississippi and north to Illinois and Kansas. 6. S. campestris. A. Leaves and calycine lobes thick and succulent, heavily cutinized; leaves broadest above the base, tapering somewhat to the sessile, nonclasping base, venation obscure and flush with the surface; calyx-lobes usually less than twice the length of the calyx-tube, generally less than 4 times as long as broad; corolla-lobes usually equaled or exceeded by the calyx-lobes, the corolla-lobes usually less than 1 em. long and narrower than 7 mm.; stigmatic lobes less than 5 mm. long; anthers shorter than 2 mm.; restricted to the strand and dunes from Louisiana southward into Mexico. 7. S. arenicola.

6. Sabatia campestris Nutt., Trans. Am. Phil. Soc. n. s. 5: 197. 1835. S. nervosa Raf., New Fl. 4: 92. 1838. Neurola arkanzica Raf., l. c., an alternate name. Sabbatia formosa Buckl., Proc. Acad. Phila. 1862: 7. 1863. S. campestris f. albiflora Moore, Proc. Ark. Acad. Sci. 1: 26. 1941.

Erect annual (6—15-30(-40) em. high; stems 1-3 mm. wide, strongly quadrate throughout, the corners winged by a thin hyaline membrane, 0.1-0.2 mm. high. Branches usually few or absent from the lower half of the stem, but when present sometimes opposite, typically alternate throughout, usually widely divergent, often forming angles of 60-90 degrees with stem, giving plant a loose, strikingly divaricate or even geniculate aspect or, more rarely, forming a fuller, more compact, corym- biform crown, or in unfavorable situations sometimes unbranched. Root system of few to several fibrous roots or of a slender, wiry tap-root usually 5-10 em. long bearing several slender laterals. Leaves thin, membranous, neither succulent nor heavily cutinized, primary venation apparent even in living material and at least the midrib noticeably elevated on the lower surface, conspicuously 3-5-nerved in dried speci- mens. Lowermost leaves rarely conspicuously different in either texture or shape from those at the middle or upper nodes, except usually some- what smaller in size, rarely persistent at anthesis; or very rarely quite different in being broadly elliptie to oblong, tapering to the base and up to 8 em. long, above narrowly lanceolate to broadly so, or ovate- lanceolate, rarely somewhat oblong or even ovate, typically broadest at or very near the base, generally broadly clasping, usually those at the lower nodes obtuse while the upper are acute, the tip callose-mucronu- late, (0.8-)1.5-3.0(-4.0) em. long, (0.3-)0.8-1.5(-2.0) em. wide, usually about (1-)1.5-2(-3) times as long as broad. Internodes (1-)2-5(-10) em. long, usually about 1.5-4 times the length of the leaves. Inflorescence usually of loosely-arranged, divergent, reduced, 1-2-flowered cymules forming a more or less open corymbose cluster. Peduncles (or pedicels) wiry, rigid, pentagonal, (1-)2-5(-10) em. long, bearing fine hyaline membranous wings at the angles. Calyx-tube broadly campanulate, usually about 1-1.5 times as long as broad, (3-)4-6(-8) mm. long; strongly pentagonal with 5 thin, membranous costae or ribs elevated about 1 mm. with fine hyaline margins. The intercostal tube very thin, membranous,

1955] Wilbur,—' The North American Genus Sabatia 45

translucent, appearing green due to ovary within. Calyx-lobes thin, membranous, not heavily cutinized, strongly spreading at anthesis, (1.5-)2-4(-6) times longer than the tube, acute, apiculate, linear, (0.4—) 1.0-2.2(-3.2) em. long, usually 4-8 times longer than broad, strongly resembling the leaves in texture. Corolla-tube usually 1-2 mm. longer than the calyx-tube, colorless except for the extension of the yellow "star" patches in the uppermost portion. Corolla-lobes typically equaling or exceeding the calyx-lobes, occasionally shorter, ascendent to rather strongly spreading, oblong, elliptical, or most commonly broadly obovate or spatulate, acute to obtuse, (1.0-)1.3-2.0(-2.3) em. long, (0.4-)0.6-1.3 (—1.5) em. wide, rose to pale pink or rarely white with a 3-4 mm. by 1-1.5 mm. greenish-yellow patch at the base of the lobe, bordered by a white area and sometimes by a more densely roseate region. Anthers bright yellow, narrowly linear, 2.5-5(-6) mm. long; filaments white to pale yellow, slender, 3-6 mm. long. Style slender, white to pale yellowish, 3-4 mm. long; stigmatie branches greenish at first, turning yellow at maturity, 5-8 mm. long. Capsules 5-9 mm. long, either equaling and included within the calyx-tube or exserted about 2-4 mm. TYPE LOCALITY: “in the open prairies of Arkansas and Red River." Type: Nuttall's collection, presumably in the Herbarium of the Philadelphia Academy of Science, but not seen. DISTRIBUTION: Fields and prairies from Illinois to Kansas south throughout most of the eastern half of Texas and east- ward to Mississippi. Very rarely introduced farther east. Map 6.

This species is easily recognized by its prominently ribbed calyx-tube together with its thin, membranous leaves.

Nuttall's type, or authentic material known to have been in his possession when the original description was made, has not been seen by me but his diagnosis is so ample as to leave no doubt as to the plant he had at hand. As yet but four species of the genus are known from Arkansas and of these Nuttall's excellent description fits but one. The original collection is stated to have been made “in the open prairies of Arkansas and Red River."

If one were to accept the date of publication that appeared with the works of Nuttall and Rafinesque describing this species, a different conclusion than that adopted here would be reached as to which name has priority. Nuttall’s “Collections towards a Flora of the Territory of Arkansas" appeared in Vol. 5 of the “Transactions of the American Philosophical Society." It has therefore been dated as 1837, the year that that volume was apparently completed. Foster (RHopomRA 46: 1506-157. 1944.) has pointed out that that date is certainly later than the publication and distribution of Nuttall's contribution.

46 Rhodora [VoL. 57

W. J. Hooker was familiar with Nuttall's account and cited it and descriptions provided in it on several occasions. The original diagnosis of S. campestris was quoted by Hooker (Comp. Bot. Mag. 1: 171. 1836.) and its place of publication cited to the page. Since Sprague (Kew Bull. 1933: 362-364.) has provided very precise dates for the publication of the various parts of Hooker's "Companion to the Botanical Magazine," it is certain that the whole of Nuttall's Flora, and not just the first twenty-one pages, as implied by Foster, was published and had reached England some time prior to the publication of Vol. 1, No. 6, of the Companion, which appeared January 1, 1836. The four parts or volumes of Rafinesque's New Flora were issued, not in 1836 as stated on the title page, but at various dates between very late in 1836 and late in 1838. This has been demonstrated rather conclusively by Barnhart (Torreya 7: 177-181. 1907.). The names of Sabatia appeared in the part which appeared in 1838.

The Rafinesquian names here considered as synonymous with S. campestris were all published in one paragraph and were intended for the same entity. His account is quoted in full as it seems the surest way to prove the identity of the plant de- scribed by him and also to explain his peculiar nomenclature.

975. Sabbatia ? nervosa Raf. ramose pauciflore, branches anceps, leaves ovate lanceolate trinerve acute, peduncles axil. and terminal, calix base campanulate 5 gone, segments linear elongate nervose, corolla as long as calix, segments broad obovate, trinerve at base—anonymous sp. of Nuttal, which deserves perhaps to be a genus Neurola Raf. by calix 5 gone corolla nerved, stamens rolled up but not twisted, style declinate and with linear stigmas as in Sabbatia. From Arkansas, 8 inches high, leaves small remote semi-uncial, flowers large over uncial incarnate ? Neurola arkanzica will be a better name.

This description, which alone is rather convincing, coupled with the information that the specimen was one of Nuttall’s collections from Arkansas leaves no doubt as to the identity of the plant. Nuttall found but two species on that journey, S. campestris and the well-known S. angularis. Rafinesque (Aut. Bot. p. 55. 1840.) was still uncertain as to the proper rank to bestow upon this species for he referred to it as '*Sab- balia nervosa or Neurola arkanzica . . . fine subg. perhaps a Genus . . ."

The final proposition of specific rank made for this species was that of Buckley. The specimen upon which this was

1955] Wilbur,— The North American Genus Sabatia 47

based has not been seen by me but again the description is such that determination is certain. "The original publication reports it to have been collected in Llano Co., Texas, from whence numerous specimens of S. campestris have been examined. 'That is the only species of the genus known from Central Texas. Buckley's proposal was promptly relegated to synonymy by Gray (Proc. Acad. Nat. Sci. Phil. 1862: 166. 1863.) whose regret as to the “hundred worse than useless synonyms" published with S. formosa, especially after the caustic criticism strongly reminiscent of an even later Fisher Professor of Natural History, was doubtless shared by Buckley.

Merrill (Rnopona 50: 127. 1948.) equates S. concinna Wood with S. campestris rather than with S. brachiata to which Gray long before had reduced it and to which Wood himself later considered it synonymous. The evidence that it is anything but S. brachiata is not convincing and the name is discussed more fully under that species.

REPRESENTATIVE SPECIMENS:—ILLINOIS: DuPage Co., Hinsdale, Smith 497 (F). MISSISSIPPI: Covington Co., about 13 mi. nw. of Hattiesburg, Webster & Wilbur 3359 (micH). Mrissounr: Christian Co., 2 mi. s. of Saunderds, Steyer- mark 23232 (F, MO, NY, TENN, WIS); Jasper Co., near Asbury, Palmer 34661 (MO, NY). ARKANSAS: Benton Co., 9 mi. ne. of Springdale, Moore 350339 TYPE of S. campestris f. albiflora Moore (UARK); Bradley Co., Warren, Demaree 19268 (MO, NY, OKLA). LOUISIANA: Calcasieu Parish, vicinity of Lake Charles, Allison 279 (Ny, vs); Winn Parish, 10 mi. w. of Winnfield, Webster & Wilbur 3261 (MIcH). KANSAS: Bourbon Co., 4 mi. w. of Fort Scott, Horr E165 (DUKE, FLAS, NCS, OKL, OKLA, PENN, SMU, US); Cherokee Co., Baxter Springs, Oyster 5445 (F, MO, NY); Montgomery Co., 5 mi. ne. of Caney, Rydberg & Imler 408 (KSC, MO, NY, US). OKLAHOMA: Cleveland Co., Norman, Demaree 13112 (Gu, MO, NY, OKL, US); Comanche Co., near Cache, Stevens 1328 (GH, MO, NY, OKL, OKLA, US); Creek Co., 7 mi. n. of Sapulpa, Ownbey 1606 (GH, MICH, MO, NY, OKL, PENN, TEX, US); Johnston Co., near Mannsville, Griffith 3474 (GH, Mo, NY). TEXAS: Brazoria Co., Columbia, Bush 76 (an, Mo, NY, US); Calhoun Co., Port Lavaca, Tharp, 22 May 1930 (DUKE, KSC, MO, MT, OKLA, TAES, TEX, WVA); Frio Co., near Moore, Palmer 33868 (Gu, Mo, NY, vs); Harris Co., Houston, Hall 508 (BRU, F, GH, NY); Llano Co., Enchanted Rock, Tharp, 11 June 1930 (GH, MO, NY, SMU, TAES); Parker Co., Weatherford, Tracy 8045 (cU, F, GH, MO, NY, TEX, US); Rockwall Co., ca. 30 mi. e. of Dallas, Webster & Wilbur 2960 (MrcH); Travis Co., Austin, Tharp, 16 May 1939 (MICH, MO, NA, NY, SMU, TEX).

7. Sabatia arenicola Greenm., Proc. Am. Acad. 34: 569. 1899. S. carnosa Small, Fl. SE. U. S. 927. 1903.

Erect annual (3-)10-20(-32 or up to 45 in greenhouse-grown plants) em. high; stems rigid, 1-3 mm. in diameter, succulent when young but becoming less so with maturity, more or less terete in the lowermost internodes but soon becoming strongly quadrate above and the corners

48 Rhodora [Vor. 57

winged by a fine, hyaline membrane about 0.1-0.3 mm. high that is barely discernable in dried material. Branches most commonly arising from near the base and then often opposite but becoming alternate above, or in more loosely developed plants absent from the lower half of the plant and alternate throughout, strongly ascendent or at first widely divergent (up to 60 degrees) and, in plants grown under the normal highly exposed environmental conditions, typically presenting an almost globose form or a compact corymbiform to convex crown. Root-system most commonly a strongly developed tap-root or almost as commonly with several almost equally developed fibrous roots; these 4-20 cm. long and bearing few to several slender laterals. Leaves thick, widely spreading, strikingly succulent, shiny, heavily cutinized, venation almost completely obscured in living material, not at all elevated on either surface, appearing 3-5 nerved in dried specimens, the lowermost narrowly spatulate to somewhat elliptie, strongly tapering to the almost petiolate base, above becoming elliptical to ovate-lanceolate or rarely somewhat oblong, usually broadest one-fourth to one-third the length from the base or even near the middle, and somewhat tapering to the sessile, non-clasping base, (0.7-)1.2-1.8(-2.7) em. long, (2.0-)5.0-8.0 (-13.0) mm. wide, usually about 1.5-4 times as long as broad, obtuse to acute with a minute, non-chlorophyllous callosity forming a mucronu- late tip. Internodes (0.3-)1.0-2.0(-8.0) em. long, usually 0.5-2 times the length of the leaves but in more loosely developed plants not uncom- monly up to 4 or more times as long as the leaves. Inflorescence of corymbosely to pyramidally arranged cymules compactly clustered or borne loosely in plants grown under less rigorously exposed conditions. Cymules 1-2-flowered. Pedicels often somewhat succulent, rigid, pentagonal, bearing 5 fine, hyaline, membranous wings, 0.5-3(-7) em. long. Calyx-tube broadly campanulate to urceolate, usually about twice as long as broad, (0.3—)1.0-2.0(-8.0) mm. long, becoming broadly turbinate in fruit, strongly pentagonal due to thick, somewhat succulent, costae or ribs about 0.5-0.7 mm. high, the internal portion of which is chlorophyllous, the outer finely hyaline margined. Intercostal tube very thinly membranous, colorless but translucent, thus appearing green due to the ovary within. Calyx-lobes thick, succulent, heavily cutinized, erect in bud but becoming strongly spreading at anthesis, (0.5—)1—1.5(-3) times longer than the tube, (0.3-)0.5-1.2(-2.0) em. long, oblong to most commonly linear-lanceolate, usually about 3 times as long as broad, acute, minutely hyalinely callose-tipped, becoming prominently nerved when dried, in general very similar to leaves in texture. Corolla-tube exceeding calyx-tube as well as the ovary by about 1-2 mm., lower portion colorless, the upper colored by extension of the yellow patches at lobal bases. Corolla-lobes more or less equaling or somewhat shorter than the calyx-lohes, or rarely longer, ascendent to strongly spreading, broadly spatulate, obovate, or oblong, obtuse, (0.5-)0.7-1.0(-1.3) cm. long, 3-7.5 mm. wide, deeply roseate or pure white, both forms with a 2-3 mm. long, 1.5 mm. wide basal, oblong, yellow patch, which in roseate forms is often bordered by deeper shade of red. Anthers bright-yellow, oblong,

1955] Wilbur,—The North American Genus Sabatia 40

1.5-2 mm. long; filaments white to pale-yellow, slender, 2-4 mm. long. Style slender, colorless to pale greenish-yellow, 2-3.5 mm. long, stigmatic branches green to greenish-yellow, 2.5-4 mm. long. Capsule 5-9 mm. long, equaling and included within the calyx-tube or exserted by 1-3 mm. TYPE LOCALITY: “on damp sands of seacoast near Tampico, State of Tamaulipas, Mexico." Type: Pringle 6808 (Gu!). DISTRIBUTION: Beaches and dunes from Louisiana southward to at least the southern part of the State of Tamaulipas, Mexico. Map 7.

The pronounced succulence of this species is perhaps its most striking characteristic and that by which it is most readily distinguished from the closely related but perfectly distinct S. campestris.

Many of the differences noted between the two species of subsection Campestres are such that one might expect them to be merely response of a highly plastic organism to an environ- mental extreme. Succulence and diminution of the various organs and structures and the assumption of an almost globose, compact form are features that logically would be expected among the littoral members of a species of wide environmental variation. Sabatia arenicola, found along the strand and dunes from Louisiana to at least Tampico in Mexico, possesses such characteristics and accordingly suspicion is raised as to its status.

The few specimens of S. campestris observed in the field in almost similar situations, although stunted or dwarfed, failed to show the peculiarities characteristic of the littoral species. Furthermore, herbarium specimens of plants that were un- doubtedly S. campestris, as indicated by their morphological characters, were collected, according to their accompanying labels, from what was approximately the environment of S. areni- cola. ln addition, numerous plants of S. arenicola grown in the greenhouse from seeds collected at various stations along the Texas coast retained the pronounced succulent nature so charac- teristic of the littoral species. The greenhouse-grown plants were larger, more elongate, and much more diffuse than the majority of specimens of S. arenicola seen by me either in the field or in the herbarium but are approached to some extent by those growing in nature in the more sheltered areas. The principal distinguish- ing characters mentioned in the key remain constant and the plants of the two species grown side by side are readily separ- able.

50 Rhodora [Vor. 57

S. BRACHIATA

S. CAMPANUL ATA

Maps 5-8. Map 5, upper left; map 6, upper right; map 7, lower left; map 8, lower right.

Two specimens were annotated by Small as S. carnosa. These are the only representatives of the littoral species that were available to him at New York prior to 1903 when his Flora was published. None of the specimens on either of the sheets is as much as 3 dm. tall, the maximum height mentioned in the original description. However, it is likely that these were the only examples seen by him before publication; they were certainly the only two annotated by him as that species and one of them should be chosen as the type of the name. The only collection of the two that matches the statement “‘often branched at the base" is that of A. Schott taken on the beach at Brazos

1955] Wilbur,— The North American Genus Sabatia 51

Santiago near the mouth of the Rio Grande on the expedition led by Major Emory to survey the Mexican boundary. The other specimen annotated as S. carnosa by Small is one of Drummond’s but no other information, other than that it was from Texas, is presented on the accompanying label. However, the great similarity in aspect and in state of preservation strongly suggests that it is a duplicate of these specimens seen at both the Gray Herbarium and Kew which bear Drummond’s collection number ‘59’ and which came from the “Rio Brassos" (Rio Brazos) Texas. The New York specimen may well be taken as the lectotype of S. carnosa. These specimens, the basis of Small’s S. carnosa, can in no way be distinguished from Green- man's S. arenicola which has four years priority.

This species and S. campestris are usually readily separable if the specimens are adequate. A few collections have been seen, however, that indicate that hybridization may be taking place between them in the region where they come into contact. Careful observation in the field should do much toward settling this suspicion.

REPRESENTATIVE SPECIMENS:—LOUISIANA: Jefferson Parish, Grand Isle, Cangemi & Andrus, 13 June 1931 (Ny); Terrebonne Parish, Timbalier Island, Tharp, 29 July 1929 (mo, TEX). tTExas: Aransas Co., n. of Rockport, White- house 18232 (smu); Brazoria Co., Rio Brazos, Drummond 59 (GH, K, NY); Calhoun Co., beach near Port O'Connor, Webster & Wilbur 3134 (wich); Cameron Co., beach at Brazos Santiago, Schott 27 (F, Ny), about 3 mi. w. of Boca Chica, Lundell 8635 (Gn, MICH, NY, SMU); Galveston Co., beach on Gal- veston Island, Webster & Wilbur 3153 (micu); Harris Co., Morgan’s Point, Palmer 11966 (Mo, Us); Kennedy Co., El Toro Island, Tharp 49218 (TEX); Kleberg Co., Kingsville, Sinclair, 1940 (Tex); Matagorda Co., beach beyond Sargent, Webster & Wilbur 3152 (micH); Nueces Co., north end of Padre

Island, Webster & Wilbur 3092 (micH); San Patricio Co., 10 mi. s. of Aransas Pass, Whitehouse 18213 (swv).

Mexico: TAMAULIPAS: Washington Beach, Runyon 461 (aH, Ny, us); San Jose, LaSueur 349 (TEX); vicinity of LaBarra, 8 km. e. of Tampico, Palmer 297 (F, GH, US); Tampico, Pringle 6808 (BRU, F, GH, NY, UC, US).

D. Subsection CAMPANULATAE Blake, RHopona 17: 56. 1915.

Annuals or perennials with either slender tap-root, several fibrous roots, or an erect, branched, perennating caudex. Branches typically alternate. Corolla 5-parted, rose, pink, or white. Calyx smooth or finely ribbed, not wing-costate. TYPE SPECIES: Sabatia campanulata (L.) Torr.

The species forming this subsection are not of uniform habit of growth as three of the species are annuals, while one, the type, is a perennial

52 Rhodora [Vor. 57

with a subterranean system unique in the genus. This is the only species- group showing diversity in life-form. The branching pattern in this group is typically alternate, but this character is highly variable, as it is throughout most of the genus. This subsection is not set off by one strong characteristic or even by a series of lesser ones. Segregation of the annuals, leaving S. campanulata to form a monotypic subsection characterized by its shortly branched, erect, caespitose caudices, would result in two subsections, each certainly more homogeneous than the present one. For the moment at least, I am hesitant about erecting additional subgeneric units as their utility is defeated when they become so numerous as to be composed of only one or two species. The genus is already, for one so small, burdened with a large number of subgeneric units,

KEY TO THE SPECIES OF SUBSECTION CAMPANULATAE

A. Caespitose perennial from branched, erect caudex; calyx-lobes usually more or less equaling the corolla in length......... 8. S. campanulata.

A. Annuals with either a tap-root or several fibrous roots; calyx-lobes usually less than three-fourths the length of the corolla.

B. Calyx lobes linear to almost filiform, usually more than 8 mm. long; corolla usually roseate, rarely white; style 2 mm. or more in length; branches usually not numerous, mostly restricted to the upper half of the stem; nodes along the main axis not numerous, usually less than 15; root-system typically of several, almost equal, fibrous roots.

C. Leaves drying thin and smooth, the uppermost equaling or exceeding the stem in diameter; corolla-lobes usually less than 1.5 cm. long; calyx-tube usually turbinate or narrowly campanulate; known from Mexico, Cuba, the Bahamas, Florida west to Louisiana and north to southern Massachusetts. ........ 0.00000 eevee 9. S. stellaris.

C. Leaves drying thick and very noticeably rugose; above typically less than the stem in diameter; corolla-lobes usually longer than 1.5 cm.; calyx-tube usually broadly campanulate; known only from Cuba and peninsular Florida.................. 10. S. grandiflora.

B. Calyx-lobes setaceous or subulate, less than 8 mm. long; corolla white; style 1 mm. long or less; branches usually numerous and typically arising from throughout the length of the stem; nodes very numerous and typically more than 20 along the main axis; root-system of one prominently developed, wiry tap-root with several slender laterals.

11. S. brevifolia.

8. Sabatia campanulata (L.) Torr., Fl. N. & Mid. U.S. 217. 1824. Chir- onia campanulata L., Sp. Pl. 1: 190. 1753. C. gracilis Michx., Fl. Bor. Am. 1: 146. 1803. C. campanulata 8 gracilis (Michx.) Pers., Syn. Pl. 1: 282. 1805. Sabbatia gracilis (Michx.) Salisb., Parad. Lond. t. 32. 1806. S. campanulata f. albina Fern., Ruopora 18: 151. 1916. 5$. Tracyi Gandoger, Bull. Soc. Bot. Fr. 65: 61. 1918. S. campanulata var. gracilis (Michx.) Fern., RHopora 39: 444. 1937.

Caespitose perennial from an erect, short, much-branched, underground caudex 1-4 em. long. Stems few to numerous, erect, (15-)30-60(-90) cm. high, 1-3 mm. in diameter, terete but often strongly ridge-angled

1955] Wilbur,— The North American Genus Sabatia 53

from fine elevated internodal lines, becoming hollow below. Branches typically alternate throughout, or occasionally those at lower nodes opposite, rarely opposite even above; usually restricted to the upper half of the stem but occasionally arising from throughout the length, strongly divergent forming an angle with the stem of 50-80 degrees or, especially in the northern portion of its range, strongly ascendent forming angle of about 20-40 degrees; few to numerous, slender, bearing but few nodes (more common in the northern portion of the range) or several. Root- system of few to many, slender, but apparently succulent, roots descending from the caudex, about 5-10 em. long and 1 mm. thick bearing few to several very slender, fibrous laterals. Leaves ascendent to very com- monly (especially the upper) strongly spreading, midvein noticeably elevated beneath, venation otherwise obscure, margins somewhat thick- ened and slightly revolute, drying thick, somewhat chartaceous, smooth or slightly rugose, obtuse to acute, broadly sessile or somewhat tapering in broader leaves to the base, the lowermost rarely ovate; the lower cauline (1-)1.5-3(-4) em. long, (1-)2-7(-12) mm. wide, usually 5-12 times as long as broad, narrowly lanceolate, oblong, or linear gradually reduced above to very narrowly linear or even filiform; those of the branches narrowly linear to filiform. Internodes usually 1-2 times the length of the leaves. Flowers apparently solitary, borne on alternate, or rarely opposite, ascendent to widely divergent, slender branches bearing 1 to several nodes, or in more obvious cymules which are often reduced. Pedicels (2-)4-7(-9) em. long, slender, wiry, slightly angled. Calyx-tube shallowly turbinate or more rarely somewhat campanulate, smooth or nearly so, not conspicuously nerved, scarious, usually about as broad as long, (1—)1.5-2.5(-3) mm. long. Calyx-lobes linear to very narrowly so or very commonly acicular or setaceous, erect to more com- monly widely spreading, (6-)7-17(-23) mm. long, typically less than 0.5 mm. wide but occasionally as wide as 1 or even 2 mm., usually 5-15 times as long as the calyx-tube, generally nearly equaling the corolla in length, or but several millimeters shorter, rarely exceeding it. Corolla- tube cylindrical, about 2-3 times as long as the calyx-tube or exceeding it about 2-3 mm., colorless to white or pale greenish-yellow, (2-)3-5(-6) mm. long. Corolla-lobes wide-spreading, oblanceolate, oblong, or elliptic, acute to obtuse, (0.6-)0.9-1.8(-2.4) em. long, (3-)4-7(-9) mm. wide, usually about 2-4 times as long as wide, rose, pink, or rarely white with an usually unlobed yellow area at base of lobe 2-3 mm. long, often bordered by a dull red irregular line. Anthers slender, bright yellow, (2-)3-4(-5) mm. long; filaments pale yellow, slender, 2-4 mm. long, usually shorter than the anthers. Style 2-5 mm. long, usually shorter than its branches, green to greenish-yellow; stigmatic lobes 3-7 mm. long, pale-green turning yellowish (with pollen?) at maturity. Ovary pale-green, slightly exserted from the corolla-tube. Capsule cylindrical, 5-7 mm. long, 2.5-4 mm. in diameter, usually 1.5-2 times as long as wide. TYPE LOCALITY: “in Canada." This plant is not known from farther north than Massachusetts. type: Kalm (Photograph of the

54 Rhodora (Vou. 57

Type in the Linnaean Herbarium seen in the collection of the Gray Her- barium). DISTRIBUTION: Peaty bogs and savannas along the Coastal Plain from Massachusetts south to northern Florida and westward to Louisiana and Arkansas; also found in the southern Appalachians and locally in Kentucky and Illinois. Map 8.

Fernald recognized two populations within this species. The supposed differences as defined by him are presented below.

campanulata gracilis Primary cauline leaves oblong- Lower cauline leaves linear, the linear to lanceolate. upper ones very narrowly so. Pedicels naked or only slightly Pedicels mostly leafy-bracted. bracted. The linear calyx-segments (except in The linear-acicular calyx-segments in small secondary flowers) 1-2 6-14 mm. long. em. long. The corolla-segments 1-1.7 cm. long. The corolla-segments 6-14 mm. long. Branches erect or strongly ascendent. Branches more divergent. Upland and Piedmont region, Ga., On the Coastal Plain from Florida and Ala., n. to Va., and Ind., to Louisiana and se. Va. and on or near Coastal Plain to se. Mass.

Many of the specimens studied could be assigned to one or the other group with a fair degree of assurance by applying these criteria. Yet there are a number of puzzling collections which aroused a strong suspicion as to one's ability to determine "correctly" a large series of specimens if one were unaware of their geographic origin. For example, Deam (1940) considered the material from southwestern Indiana to be a representative of the typically Coastal Plain population while Fernald (1950) mentioned Indiana only in the range of the campanulata-group.

Fernald (Ruopona 34: 27. 1932.) originally had considered the plants found along the southern Coastal Plain to be “a rather well defined species" although admitting that “. . . it may be better to treat it as a southern Coastal Plain variety . . ." After encountering specimens in southeastern Virginia that possessed some of the supposed distinguishing characteristics of both varieties, Fernald (Rhodora 39: 444. 1937.) concluded that “. . . S. gracilis should be treated as a geographical vari- ety." Those intermediate specimens combined the presumably characteristic narrow leaves and calyx-segments of the southern Coastal Plain population with the longer lobes of both calyx and corolla of the more northern collections. Since numerous

1955] Wilbur,— The North American Genus Sabatia 55 large-flowered specimens had been observed throughout the supposed range of the gracilis-population, the several hundred available measurements were tallied for comparison and are presented below. The specimens from southeastern Virginia which Fernald considered atypical as they showed features of both populations were not included.

LENGTH OF CALYx-LonBEs LENGTH OF CoROLLA-LOBES No. of lobes No. of lobes No. of lobes No. of lobes mm. gracilis-range | campanulata-range gracilis-range | campanulata-range

6 6 0 1 0

7 23 5 4 0

8 30 6 4 1

9 42 18 16 7 10 40 22 34 16 11 50 31 44 38 12 60 42 63 38 13 45 36 52 43 14 45 20 36 38 15 27 19 32 19 16 19 Lr 17 7 17 10 19 15 0

18 TA 9 10 0

19 1 5 4 2 20 4 2 2 0 21 2 1 0 1 22 0 1 0 0 23 0 1 e 0 24 0 0 0 1

These hundreds of measurements indicate that, although the smallest flowers were found on the specimens from the gracilis-range and the largest from the range of the more northerly population, no reliance can be placed upon the length of either the calyx- or corolla-segments as a distinguishing characteristic between the two supposed varieties. These measurements show that the range of flower-size of the plants from the Coastal Plain is considerably greater than that known to Fernald and for diagnostie purposes the range of the length of the lobes of either floral series is so broadly overlapping as to be unworkable.

Nor am I able to detect any consistent differences in the shape of the calyx-lobes between those specimens of the Coastal Plain south of Virginia and those from the Appalachian region and the Costal Plain north of Virginia. Numerous specimens from Florida and the other states of the supposed range of gracilis have linear calyx-lobes presumably characteristic of the northern variety.

56 Rhodora [Vor. 57

By ''pedicels" Fernald apparently meant lateral branches bearing flowers. "These branches in the Massachusetts material, with which Fernald had field experience, are typically short and rather strongly ascendent and their one or two nodes bear bracts that are foliose in comparison to the narrowly linear to even filiform bracts of the typical southern coastal material. A tendency for opposite branching in the upper part of the stem is also more pronounced, especially in more robust specimens, and then the unreduced cymose pattern of inflorescence is some- what more apparent. These rather striking characteristics found with considerable constancy in the few New England stations are far less apparent in the specimens seen from the more southern part of the range of the supposed typical variety. Numerous specimens from the mountainous regions of Georgia, Alabama, the Carolinas, Tennessee and Kentucky, as well as Deam’s Indiana station, possess elongate, widely divergent branches with several to numerous nodes bearing narrowly linear to even filiform bracts. The extremes of leaf-type from the two ranges are often most striking and clear-cut but many specimens within the range attributed to one of the varieties would be assigned to the other upon the application of this criterion. These considerations have convinced me that it would be difficult to recognize varieties in this species in spite of some rather strong tendencies which are somewhat geograph- ically segregated and whose extremes are of rather striking difference in aspect.

Gandoger based the description of S. tracyi upon the specimen in his herbarium of Tracy 6468 collected at Biloxi, Mississippi. I have not seen his type but four specimens of the same collection number are located at the Gray Herbarium (2 sheets), New York Botanical Garden, and the National Herbarium. They are presumably isotypes. Gandoger described his specimen as an annual and this would relate or equate S. tracy? with S. stellaris, the only annual of subsection Campanulatae known to the state. The presumed isotypes, however, are perennials with narrowly oblong-linear to linear leaves gradually reduced above and on the branches to filiform bracts; their branches, which are alter- nate, are elongate, bear numerous nodes, and are very strongly divergent. There is still a trace of rose-pink color on the corolla-

1955] Wilbur,— The North American Genus Sabatia 57

lobes of at least three of the sheets seen although even on those sheets most of the flowers appear yellowish-white. Gandoger’s species was described as having white flowers which together with the more slender leaves and mucronate sepals supposedly distinguished it from S. gracilis. S. tracyi is synonymous with S. campanulata, or, if one prefers to recognize what now seems to me to be a variation too weak to maintain, to S. campanulata var. gracilis (Michx.) Fern. It may be matched by a majority of the specimens collected along the Gulf and Southern Atlantic Coastal Plain.

REPRESENTATIVE SPECIMENS:—MASSACHUSETTS: Barnstable Co., Barn- stable, Fernald & Svenson, PL. EX. GRAYANAE 479 (in many herbaria); Nan- tucket Co., Almanac Pond, Williams, 30 July 1911 (aH); Plymouth Co., Pem- broke, Foster, 10 Sept. 1884 (aH). NEW YORK: Nassau Co., East Rockaway, Bicknell 7089 (Ny); Suffolk Co., Oakdale, Ferguson 7775 (NY). NEW JERSEY: Ocean Co., e. of Silverton, Fogg 4908 (GH, NY, PENN, TENN). PENNSYLVANIA: Bucks Co., Tullytown, MacElwee, 28 July 1894 (aH, NY). DELAWARE: Kent Co., Felton, Canby, July 1878 (us). MARYLAND: County unknown, Delaware Beach, Hood 2370 (FLAS). DISTRICT OF COLUMBIA: Virginia Highlands, Griggs, 17 July 1923 (us). VrRGINIA: Greensville Co., ne. of Gaskins, Fernald & Long 13421 (GH, MO, TENN, US); Sussex Co., about 4 mi. nw. of Homesville, Fernald & Long 6351 (GH, NY, PENN). NORTH CAROLINA: Columbus Co., 2 mi. w. of Freeman, Wilbur 2882 (micu); Henderson Co., 1 mi. s. of East Flat Rock, Oosting 1790 (puKE); Northampton Co., near Margarettsville, Heller 1160 (CU, F, GH, KSC, MO, MT, NY, PENN, US). SOUTH CAROLINA: Anderson Co., Belton, Smith, 23 July 1881 (cH, vs); Georgetown Co., 12 mi. n. of George- town, Godfrey & T'ryon 66 (DUKE, F, GH, MICH, NY, PENN, TENN, US). GEORGIA: Bartow Co., 4.5 mi. s. 16° east of Allatoona Dam, Duncan 8521 (FLAS, GA, MO, SMU, TENN, US); Thomas Co., about 1 mi. n. of Coolidge, Duncan 8466 (FLAS, GA, MO, TENN). FLORIDA: Bradford Co., near Lawtey, Murrill 528 (DUKE, MO, US); Holmes Co., Bonifay, Curtiss 6481 (GH, MO, NY, SMU, US); Washington Co., 8 mi. s. of Chipley, Webster & Wilbur 3615 (MICH). INDIANA: Daviess Co., about 4 mi. n. of Washington, Deam 52742 (F, GH, MO, NY); Jasper Co., about 3 mi. se. of Wheatfield, EK, 30 July 1940 (GĦ). KENTUCKY: Fayette Co., Lexington, Short, 1831 (Ny); Whitley Co., n. Jellico, Braun, 21 July 1933 (GH, NY). TENNESSEE: Coffee Co., n. of Manchester, Svenson 9219 (DUKE, GH, MO, NA, NY, WIS, US); Franklin Co., near Tullahoma, Sharp, Clebsch & Fairchild 9938 (DUKE, GA, NCU, OKLA, SMU, TENN, TEX, US). ALA- BAMA: Conecuh Co., about 9 mi. w. of Evergreen, Webster & Wilbur 3514 (mich); DeKalb Co., Lookout Mountain, Biltmore Herb. 4510g (us); Mobile Co., about 8 mi. sw. of Mobile, Webster & Wilbur 3480 (MICH). MISSISSIPPI: Covington Co., 1.5 mi. se. of Mt. Olive, Webster & Wilbur 3336 (wicm); Forest Co., 18 mi. nw. of Beaumont, Webster & Wilbur 3424 (micu); Jackson Co., Ocean Springs, Pollard 1010 (cv, F, GH, MO, NY, US). ARKANSAS: Lonoke Co., Grand Prairie, Demaree 22329 (wo); Pulaski Co., Little Rock, Harvey, July 1882 (UARK). LoUuiSIANA: Calcasieu Parish, Lake Charles, Cocks, Sept. 1906 (No); Tangipahoa Parish, 3 mi. e. of Robert, Correll 9307 (DUKE, Mo, TEX).

58 Rhodora [Vor. 57

9. Sabatia stellaris Pursh, Fl. Am. Sept. 1: 137. 1814. Chironia amoena Raf., Med. Repos. II. 5: 359. 1808. not Salisb., Prodr. 137. 1796. C. stellata Muhl., Cat. Pl. Am. Sept. ed. 2, 23. 1818. C. stel- laris [Pursh] Eaton, Man. ed. 2, 204. 1818. Sabbatia maritima Raf., Med. Fl. 2: 77. 1830. S. amoena (Raf.) G. Don, Gen. Hist. 4: 207. 1838. S. stellaris 8 pumila A. Gray ex Griseb., Prodr. 9: 49. 1845. Eustoma maculata Benth., Pl. Hartw. 292. 1848. Sabbatia gracilis 8 stellaris [Pursh] Wood, Am. Bot. & Flor. 267. 1870, without basionym. S. nana Featherman, Rep. Bot. Surv. S. & Cent. La. 71. 1871. 5$. maculata (Benth.) Benth. ex. A. Gray, Proc. Am. Acad. 22: 438. 1887. S. Palmeri A. Gray, Proc. Am. Acad. 22: 438. 1887. S. stellaris f. albiflora Britton, Bull. Torrey Club 17: 125. 1890. S. simulata Britton, Bull. N. Y. Bot. Gard. 3: 448. 1905. S. Purpusii Brandegee, Univ. Calif. Pub. Bot. 4: 275. 1912. Sabatia amoena f. albiflora (Britton) Fern., HnHopon4 34: 26. 1932. Sabbatia campanulata var. amoena (Raf.) Monachino, Torreya 41: 99. 1941.

Erect annual (2-)15-50(-80) em. high; stem 1-4 mm. in diameter, terete to strongly angled due to several somewhat irregularly disposed fine lines or ridges extending between nodes, pith parenchymatous in young living specimens but becoming hollow in older or dried specimens. Branches almost always alternate, very rarely one or so nodes with op- posite branches, usually restricted to upper half or two-thirds of the stem but sometimes arising throughout the length and even from the base, typically strongly divergent, forming an angle of about 30-70(-85) de- grees with the stem, few to numerous, slender, wiry, bearing few to numerous nodes. Hoot-system of several to numerous, usually slender, but occasionally thickened, fibrous roots 2-10 em. long, 1-3 mm. in diameter or occasionally with one principal root and several laterals. Leaves ascendent, above often closely appressed to stem, succulent, thick, rubbery in texture, midvein conspicuous and elevated beneath, venation otherwise obscure; after drying leaves commonly darken, usually thinly membranous or occasionally the lowermost slightly rugose, the midvein alone usually prominent. Lower leaves broadly to narrowly elliptic, or even linear, rarely spatulate or obovate, acute to rarely obtuse, often apiculate, usually tapering to both ends, typically about 5-10 times as long as broad, (0.5-)1.5-3(-6) em. long, (2-)3-8(-15) mm. wide. Upper leaves more narrowly elliptie to linear, the uppermost sometimes very narrowly so to even almost filiform, usually 7-15(-20) times as long as wide, 1-4 em. long, (0.5-)1.5-3(-5) mm. wide, typically exceeding the diameter of the stem or at least equaling it. Internodes usually 1-2 times the length of the leaves. Flowers appearing solitary but usually arranged in very loose and reduced eymules and these sometimes aggre- gated in loose clusters. — Pedicels slender, straight, ascendent to divergent, (1-)4-10(-15) em. long, usually about 0.5 mm. in diameter. Calvx-tube turbinate, or occasionally even narrowly campanulate, usually rather gradually narrowing to the base, smooth or but very faintly nerved, thin, somewhat scarious, usually 1-2 times as long as broad, (1.5-)2-4(-6) mm.

1955] Wilbur,— The North American Genus Sabatia 59

long. Calyx-lobes narrowly linear to almost filiform, ascendent or more typically wide-spreading, (4-)6-15(-22) mm. long, typically less than 0.5 mm. but rarely as wide as 1 mm., usually 3-6 times as long as calyx-tube, in general considerably shorter than the corolla, rarely equaling or ex- ceeding it, apparently slightly hyaline-scarious margined. Corolla-tube about 2-3 times as long as wide, usually exceeding calyx-tube by 1-3 mm., translucent but appearing greenish due to ovary, (3-)4-6(-8) mm. long. Corolla-lobes strongly spreading, elliptic, oblong, spatulate, or obovate, obtuse to somewhat acute, (0.5-)1—-1.5(-2) em. long, (2-)4-8(-10) mm. wide, rose, pink, or more rarely white, with a usually irregularly 3-lobed yellow area at the base often bordered by a distinct, bright or dark red line which in turn is sometimes bordered by a white area of variable width. Anthers linear, yellow, (2-)3-4(-5) mm. long; filaments pale yellow to greenish, (1.5-)2-3(-4) mm. long. Style 2-4 mm. long, usually about half the length of its branches; stigmatie lobes 3-8 mm. long. Capsule from almost globose to cylindrical, (4-)6-8(-14) mm. long, (3-)4-5(-6) mm. wide, usually 1.6-2 times as long as broad. TYPE LOCALITY: “in salt marshes: New York, New Jersey &c." DISTRIBUTION: Salt marshes and sandy (usually littoral) places from southern Massa- chusetts south along the coast and throughout much of Florida and west into Louisiana. Also known from the Bahamas, western Cuba and the central plateau of Mexico. Map 9.

Sabatia stellaris is the most variable species of the Campanulatae but within this wide range of diversity there appears to be no sharply differentiated population that might be more naturally treated as a subspecies or a variety. Rather, there is an almost imperceptible change and as a consequence specimens from the northern limits of the range are often strikingly different from those seen at the extreme southern portion of its distribution. Typically, plants collected in southern Florida will be taller, more profusely branched, often from near the base, and these branches will be much more elongate, wiry, and slender. The leaves, for the most part, will be narrowly linear and only the uppermost with indications of a more elliptical or spatulate nature. The northern material will be on the average smaller, less branched, and then the branching usually restricted to the upper portion of the stem; the branches usually short and bearing few nodes. Also the leaves in most cases will be linear only in the very upper portion of the stem and otherwise will often be broadly elliptical. The sum of these differences results in plants of very dissimilar appearance. However, examination of an ample series taken from throughout the range will convince

60 Rhodora [Vor. 57

one, I believe, that these differences are not sufficient for the recognition of subspecific categories. Rather these differences should in part be attributed to the great diversity in environment encountered through the range of more than twenty degrees of latitude and especially the difference in the length of the growing season between the Massachusetts coast and the shores of the Caribbean.

After initially attempting (1903) to distinguish between 5$. campanulata and 5. stellaris, Small abandoned the effort and thereafter applied the former name to specimens of both species in the Southeast. The confusion between the two species is understandable due to the often misleading distinguish- ing criteria published in most works previous to the detection of the strikingly obvious differences by Bicknell (Bull. Torrey Club 42: 30-32. 1915.) and especially by Fernald (RHODORA 18: 145-147. 1916.). "Thereafter there should have been little diffculty in distinguishing the two. However, most of the southern material of S. stellaris has passed as S. campanulata or the larger-flowered specimens as S. grandiflora due to the influence of Small’s work. Sabatia campanulata is not known in peninsular Florida or from the West Indies.

Gleason's treatment (1952) illustrates the confusion that still exists in this species-group. He ascribes a salt or brackish marsh habitat to both S. campanulata and S. stellaris and includes the West Indies in the range of the former species rather than that of the latter. Furthermore he states that S. stellaris “and S. campanulata constitute the local extremes of a variable population which may represent only one species or as many as four. Even in our range numerous intermediate forms occur and their assignment to a species depends on the general prominence of characters of which the proportion of the petals is probably the most reliable. In the southeastern part of our range and in the southeastern states both species pass into forms with lvs. all linear (S. gracilis (Michx.) Salisb.)." "The true picture is by no means as confusing or as complex as Gleason suggests. Sabatia stellaris and S. campanulata are separable by strong morphological features and the two species occupy mutually exclusive habitats.

It should be pointed out that while S. stellaris is rather closely confined to the littoral region in the northern part of its range,

1955] Wilbur,— The North American Genus Sabatia 61

specimens from inland peninsular Florida are fairly numerous. In spite of the information available in the literature, S. stellaris, at least in the southern part of its range, 1s not restricted to a salt-marsh or littoral environment and in fact is at least more commonly collected in inland sandy areas.

Britton (Bull. N. Y. Bot. Gard. 3: 448. 1905.) and later Britton & Millspaugh (1920) recognized two species as occurring in the Bahamas, S. campanulata and the supposedly endemic S. simulata. This latter species was to be distinguished from what has been called S. campanulata by 1ts more slender habit and smaller white flowers. All the material seen from the Bahamas has proven to be annual and consequently the larger- flowered plant with the rose-pink corollas is definitely not S. campanulata; it is S. stellaris. There is no break in flower-size between the pink and white flowered specimens although the latter are on the average smaller. Several of the larger-flowered specimens (2.5 cm. or more in diameter) had been designated by Britton as S. simulata apparently on the basis of the flower color. The stigma of S. simulata was described as ''spatulate" in contrast to the "oblong-linear" stigma of “S. campanulata.” I was unable to discover any consistent difference between the stigmas of the specimens. There was the usual variation dependent upon age. S. simulata is, I believe, merely a white- flowered form of S. stellaris and is conspecific with the pink- flowered specimens of the Bahamas. These color-phases are indistinguishable from much of the material from southern Florida and, for the present at least, all had best be treated as but another trend in the variation apparent in this species throughout its extensive range.

One name deserving brief discussion 1s S. nana of Featherman, which is not listed in “Index Kewensis." The description is here quoted in full:

Sabbatia Nana.—Stem simple, low, somewhat angled. Leaves small, sessile, the lower spatulate lanceolate, the upper linear lanceolate. Lobes of the corolla five to six, one third longer than the narrow linear calyx lobes. Root perpendicular, slender. Flowers rose-colored. Stem three to four inches high. Blooms in August.

Habitat—Marshy soil of Grande Isle.

The diagnosis is certainly brief but the shape of the leaves, the ‘narrow linear” calyx-lobes two-thirds as long as the corolla, the habit and the habitat all together make it quite certain

62 Rhodora [VoL. 57

that S. stellaris was the plant being described. This Louisianan record is at or near the westernmost extremity of the species’ present known range in the southeastern states.

Type-specimens of S. maculata, S. palmeri, and S. purpusii have all been examined and compared both with the few addi- tional Mexican collections of the genus that have been made and specimens of the widespread species of eastern North Amer- ica. Two of the species seemed very different from each other and at first I thought that I could easily distinguish them both from any others in the genus. Sabatia palmeri, on the other hand, very quiekly seemed to be indistinguishable from 5. maculata. After examining more material, especially from Florida, it became obvious that any of the Mexican specimens could be matched or very nearly matched by a considerable number of plants of 5. stellaris. I am certain that I could not distinguish a large unlabeled series of specimens made in Florida and another large series made in Mexico. Therefore, all three names are here treated as synonyms of S. stellaris, a wide- ranging, variable species.

The blackened patches at the base of the corolla-lobes of the type of S. maculata are very striking and the epithet chosen by Bentham points out this feature. However, the dark black color ean almost certainly be attributed to discoloration upon drying and has been noted in many specimens found as far north as New Jersey. The type itself seems indistinguishable from many specimens from Florida of which Cuthbert 1525 (FLAS) from Manatee Co. is a good example. The type of S. palmeri is very similar in appearance to that of S. maculata.

S. purpusit is indistinguishable from many of the wiry Floridian specimens. For example, Pennell’s collection (18048) from the state of San Luis Potosí seems remarkably similar to the collection of Hood 3565 (FLAS) among others from western peninsular Florida. In the Mexican specimens a tendency was noted for the calyx-tube to be more cylindrical with sides more parallel rather than strongly tapering and whose length is typically in the upper range of that of the specimens from the eastern United States. Also, the few capsules seen in the Mexican specimens of the purpusii-aspect appeared to be more slender and longer than those of most of the specimens seen from the principal area of distribution of the species.

1955] Wilbur,—The North American Genus Sabatia 63

The disjunct distribution of this species is somewhat perplexing. Fernald (RHopora 18: 146. 1916.) stated that the range of S. stellaris was from “Massachusetts to eastern Mexico" and at that time he considered the plants called by some S. purpusiz as part of the wide ranging species as is shown by his annotation labels. In the most recent edition of Gray’s Manual the western limit of the range was stated to be Louisiana. The apparent absence of this species from the Texas coast and its eventual reappearance in eastern Mexico far south of the border and, most surprisingly of all, in the central plateau of that country is not easily explained. It was with considerable reluctance that attempts to distinguish these species were abandoned but there seemed to be no alternative. The absence of the plant from Texas is in particular inexplicable as there seem to be enormous stretches of territory offering an environment apparent- ly similar to that in which it is found so abundantly in the Atlantic and Gulf Coastal Plain.

REPRESENTATIVE SPECIMENS:—MASSACHUSETTS: Bristol Co., Dartmouth, Hervey (au); Dukes Co., Tisbury, Cushman, July 1911 (wis), RHODE ISLAND: Washington Co., Westerly, Weatherby 3311 (NCS). CONNECTICUT: Fairfield Co., Stratford, Eames, 16 Aug. 1895 (NCU, NY, US). NEW YORK: Kings Co., Barren Island, Svenson 8242 (GA, GH, MO, PENN, TENN); Richmond Co., vicinity of New Dorp, Small, 2 Aug. 1890 (F, GH, MO, NA, NY, US, YU); Suffolk Co., mouth of Wading River, Muenscher & Curtis 6420 (cU, GH, MT, NY, Us). NEW JERSEY: Cape May Co., Cold Spring, Pennell 2179 (an, Mo, vs); Salem Co., about 4 mi. s. of Canton, Fogg 6054 (GH, PENN, TENN). DELAWARE: Sussex Co., 2 mi. s. of Rehoboth, Larsen 989 (GA, PENN, US). MARYLAND: Caroline Co., between Choptank River and Bethlehem, Killip 7278 (an, vs). VIRGINIA: Princess Anne Co., along Back Bay, Fernald & Long 11113 (DUKE, GH, MO, NY, US); Surry Co., Hog Island, Fernald & Long 12774 (GA, GH, MO, US). NORTH CAROLINA: Brunswick Co., near Southport, Biltmore Herb. 8304a (GH, Mo, NCU, NY, Us); New Hanover Co., Fort Fisher, Godfrey 4726 (DUKE, GH, NCU, NY, US). SOUTH CAROLINA: Georgetown Co., Pawleys Island, Godfrey & Tryon 306 (DUKE, F, GH, MO, NY, PENN, TENN, US); Horry Co., about 2 mi. sw. of Ocean Drive, Wilbur 2901 (micH). GEORGIA: Chatham Co., Tybee Island, Harper 924 (an, Mo, NY, US); Glynn Co., Sea Island, Cronquist 5356 (FLAS, GA, GH, NO, SMU, US). FLORIDA: Brevard Co., Titusville, Nash 2308 (F, FLAS, GH, K, MICH, MO, US); Lee Co., vicinity of Fort Myer, Standley 12 (F, GH, MO, NY, US); Levy Co., 1 mi. se. of Chiefland, Webster & Wilbur 3639 (micu); Monroe Co., Big Pine Key, Small 8148 (DUKE, GH, NCU, NY, PENN, WVA); Orange Co., about 2 mi. e. of Fort Christmas, Moldenke 210 (DUKE, MO, NY, PENN, US); Sarasota Co., Longboat Key, Tracy 6808 (F, MO, NY, US). ALABAMA: Mobile Co., saline marshes, Mohr, Aug. 1892 (NY). MISSISSIPPI: Harrison Co., Cat Island, Tracy, Earle & Seymour 101 (cu, an, MO, NCU, WIS). LOUISIANA: Plaquemines Parish, Breton Island, Tracy & Lloyd 343 (cv, F, GH, MO, NY, US). BAHAMA ISLANDS: Abaco Is., Brace 1793 (xv); Andros Is., Brace 7116 (rF, NY, vs); Cat Is., Britton & M illspaugh 5780

64 Rhodora (Vor. 57

(r, NY); East Bimini, Howard 10274 (smu, us); Eleuthera Is., Britton 6427 (F, GH, NY, US); Great Bahama Is., Brace 3518 (F, Ny, us); New Providence Is., Britton 58 [ryeE of S. simulata Britt.] (Ny); Curtiss 200 (F, GH, MO, NY, US). CUBA: Without more exact locality but very likely from Pinar del Rio, Wright 412 [sheets sometimes mixed with S. grandiflora] (BRU, GH, NY, US). This is the same number but from a different collecting trip as the type of S. gracilis var. cubensis Griseb. (= S. calycina). MExico: Coahuila: Cuatro Cienagas, Marsh 2068 (F). | Michoacán: 60-70 mi. e. of Lake Pátzcuaro in bog 4 mi. w. of Hidalgo, Hitchcock & Stanford 7196 (GH, vc). Jalisco: Lagos de Moreno, Hartweg 1615 [rype of Eustoma maculata Benth.] (x); Rio Blanco, Palmer 668 [ryrE of S. palmeri Gray] (GH). Querétaro: Querétaro, Arséen 10281 (vs). San Luis Potosí: Media Luna near Rio Verde, Palmer 80 (F, GH, MO, NY, UC, US) and Palmer 80.5 (GH, Mo, Ny, US); Minas de San Rafael, Purpus 5845 [TYPE of S. purpusit Brandegee at uc] (F, GH, MO, NY, UC); Hacienda de Angostura, Pringle 3810 (BRU, F, GH, MICH, MO, NY, UC, US); Las Tablas, Pennell 18048 (us). Tamaulipas: Jaumave, Rozynski 377 (F).

10. Sabatia grandiflora (A. Gray) Small, Fl. SE. U. S. 928. 1903. Sabbatia gracilis var. grandiflora A. Gray, Syn. Fl. N. Am. 2: 115. 1878. Sabatia campanulata var. grandiflora (A. Gray) Blake, Rnopona 17: 52. 1915. Sabbatia Alainii Marie-Victorin, Contr. Inst. Bot. Univ. Montreal No. 63: 73. 1948.

Erect annual (15-)40-90(-110) em. high; stem (1-)2-5(-7) mm. in diameter, terete, although in dried specimens with several fine lines run- ning between the nodes. Branches, rarely absent, almost entirely re- stricted to the upper third or half of the stem and almost invariably alternate, typically strongly divergent, forming angles of about 50-70 degrees, stiffly geniculate, bearing but few nodes. Plant appearing vir- gate from elongate, rigid internodes. Root-system of several to numerous fibrous roots 3-8 em. long, 1-3 mm. in diameter. Leaves erect, at least above closely appressed against the stem, succulent, drying thick, rigid, rugose-roughened, venation obscure in both living and dried material; the lower spatulate, elliptie, oblong-linear or even linear, obtuse to acute, (1-)2-4(-5) em. long, (2-)4—7(-10) mm. wide, typically about 5-10 times longer than wide, usually several times exceeding the typically short, 0.5-3 em. long internodes; gradually to abruptly reduced above to very narrowly linear or even filiform, the width noticeably less than the diam- eter of the stem, (1-)3-5(-12) em. long, (0.5—)1-2(-3) mm. wide, generally 20-60 times as long as wide, hyaline or callose-apiculate. Internodes above typically greatly elongate, often 2-5 times as long as the leaves. Inflorescence of (1-)2-3(-4) or rarely more flowers in reduced cymules; the flowers borne on slender, but rigid, elongate, terete pedicels (2-)4-8 (-12) em. long and about 0.5 mm. in diameter; the flowers thus appearing solitary. Calyx-tube campanulate, usually broadly so, the sides more or less parallel, abruptly contracting to base or but gradually tapering in the lower half, typically 1—1.5(-2) times as long as broad, 2-4(-6) mm. long, generally very smooth or but very finely lined, the wall thin and somewhat hyaline-scarious especially with age. Calyx-lobes erect to spreading, very narrowly linear or filiform, (0.6—)1—2(—3) em. long, usually

1955] Wilbur,—' The North American Genus Sabatia 65

S. GRANDIFLORA

S. BREVIFOLIA

Maps 9-12. Map 9, upper left; map 10, upper right; map 11, lower left; map 12, lower right. about 3-6 times as long as the tube, typically less than three-fourths as long as the corolla, rarely equaling or exceeding it, the tip callose-apiculate. Corolla-tube cylindrical, (3-)5-7(-8) mm. long, exceeding calyx-tube by about 2-4 mm., usually 1.5-2 times as long as wide, apparently colorless. Corolla-lobes strongly spreading, oblanceolate, obovate, broadly spatulate, oblong, or elliptic, usually obtuse or more rarely acute, (1.3-)1.8-2.5(—3) cm. long, (5-)7-12(-15) mm. wide, deep rose to pale-pink or white, with oblong basal vellow area 2-4 mm. long, rather irregularly lobed or toothed, and in the roseate forms usually bordered by an intense red line or area. Anthers linear, dark yellow, (3-)5-7(-8) mm. long; filaments yellow, slender, 2-4 mm. long. Style 2-5 mm. long; stigmatie lobes (4-)6-8(-9) mm. long. Capsule usually narrowly cylindrical, (6-)8-10(-15) mm. long, 4-5(-7) mm. wide, generally 2-3 times as long as broad. TYPE LOCALITY: “Coast of E. Florida." Lectotype: Palmer 430 (GH!) from ‘Indian River." DISTRIBUTION: Everglades, pinelands and sandy places in penin- sular Florida and western Cuba. Map 10.

In spite of the numerous striking characters which distinguish this species from any other, too much reliance has been placed

66 Rhodora [Vor. 57

upon floral size as the criterion for the recognition of S. grandi- flora. As a consequence one finds under this name in most herbaria a miscellaneous assemblage of large-flowered specimens belonging to subsection Campanulatae. Small never pointed out the distinctive characteristics of S. grandiflora and a con- siderable number of the specimens labeled or annotated by him are merely large-flowered S. stellaris. The identity of these incorrectly determined specimens is shown by the smooth, thin texture of the dried leaves which above, although narrowly linear, are typically as broad or broader than the diameter of the stem. Also these large-flowered specimens of 5. stellaris, which have been so often mistaken for S. grandiflora, are generally more profusely branched, especially below the middle of the stem. S. grandiflora may be readily distinguished from S. stellaris by its thick rugose-roughened or wrinkled leaves which above are almost filiform and less in width than the diameter of the stem. Furthermore, it is typically a larger and coarser plant with branching more or less restricted to the upper third or half of the stem. Another fairly reliable feature that may be utilized in recognizing S. grandiflora is that its calyx-tube is most often broadly campanulate, while that of S. stellaris is more typically narrowly turbinate. That much-relied-upon character, the length of the corolla-lobes, is at best a subsidiary one because of the considerable overlap in the size of flowers between the two species.

In spite of the inferences that might be derived from the synonymy, the relationship of S. grandiflora is closer to S. stellaris than to S. campanulata (= S. gracilis). Numerous differences might be listed to distinguish S. grandiflora from S. campanulata with which it was originally and even later considered to be a variety. However, the most obvious and perhaps most fundamental is that the latter is a caespitose perennial while S. grandiflora, like S. stellaris, is an annual. The perennial species is not known in peninsular Florida. S. stellaris has been collected throughout the state, while in Florida, S. grandiflora is almost entirely restricted to the peninsular portion. Judging from the number of mixed sheets of S. stellaris and S. grandiflora, there is considerable overlap in range of habitat between these two species if their environmental toler- ances in Florida are not exactly the same.

1955] Wilbur,—The North American Genus Sabatia 67

In the United States other than in Florida, S. grandiflora is represented only from Louisiana by a specimen attributed to Josiah Hale deposited in the Gray Herbarium. This col- lection is S. grandiflora but more recent collections from Louisi- ana seen in the herbaria under this name have always proven to be large-flowered specimens of S. stellaris. Confusion in labeling should be considered a strong possibility for this speci- men reputedly from Louisiana.

After the short original diagnosis, Gray presented the col- lections known to him in the following manner: ‘coast of E. Florida, Leavenworth, Buckley, Palmer & c.". Theirs are the only specimens available to him now preserved in either the Gray Herbarium or that of the New York Botanical Garden; who the other collectors were is unknown to me. Palmer’s collection, made in 1874, is definite and all four sheets at the three different herbaria bear the information “Indian River, East Florida" as well as the number ‘430.” These specimens are also equal to or better than the others and, in spite of being collected after a varietal status had been decided upon (with another epithet) based on the other collections, Palmer's col- lection (no. 430) at the Gray Herbarium is here designated the lectotype.

The recently proposed S. alainii from Cuba here is considered to be but a synonym of S. grandiflora; its distinctness has cer- tainly not as yet been demonstrated. The original description of this supposedly distinct Cuban entity was accompanied by the following discussion in reference to its relationship with S. grandiflora. ‘Cette espèce se rapproche du Sabbatia grandi- flora par sa grande corolle, ses curtes feuilles, l'absence d'oeil jaune, sa capsule plus petite." The meaning of this is not altogether clear. It would seem as though a word or so had been omitted in printing and that the first two characters men- tioned were meant as features shared in common by S. grandi- flora and the newly proposed species and that the second two phrases were supposed distinctions. The stated absence of the yellow eye in the throat of the corolla-tube and at the base of the lobes of S. alainii is not borne out by examination of either the type material, distributed isotypes or information (“white with light yellow eye") provided on the label of Britton et al. 7166 which was collected in the same region. The few

68 Rhodora [Vor. 57

capsules available were smaller but did not appear to have fully developed. The Cuban representatives of this group are, I believe, indistinguishable from those of Florida.

REPRESENTATIVE SPECIMENS:—FLORIDA: Alachua Co., Earlton Beach, Loucks, 5 Sept. 1927 (Frias); Brevard Co., between Cocoa and Lake Poinsett, Rhoads, 23 May 1937 (Frias); Broward Co., Ft. Lauderdale, Small & Wilson 1785 (Ny); Charlotte Co., without exact locality, Frye, 17 May 1946 (Frias); Clay Co., Keystone Heights, West, 29 June 1930 (FLAs); Collier Co., 2 mi. s. of Immokalee, Sheehan 28 Feb. 1919 (Ny); Dade Co., w. of Camp Jackson, Small & Wilson 1991 (Ny); DeSoto Co., Lacy, West, 23 Sept. 1938 (FLas); Duval Co., without exact locality, Fredholm 5 (au); Glades Co., Lakeport, Lovett 264 (DUKE); Hardee Co., Limestone, Kirk, 1 July 1942 (FLAs); Hendry Co., se. corner of county, Davis, 13 Sept. 1947 (FLAs); Hernando Co., without exact locality, Hitchcock 1283 (v, Mo); Highlands Co., near Brighton, McFarlin 10913 (GH, PENN); Hillsborough Co., Tampa, Garber, May 1876 (BRU, F, NY, us, YU); Indian River Co., near Felsmere, Small 8917 (Ny); Lake Co., Eustis, Nash 768 (cv, F, GH, MICH, MO, NY, US); Lee Co., about 8 mi. se. of Fort Myers, Standley 446 (F, GH, MO, MT, vs); Levy Co., Bronson, Watson & Murrill, 18 June 1939 (rLas); Manatee Co., near Palma Sola Bay, Cuthbert, 23 June 1916 (FLAS); Marion Co., Lake Kerr, West & Arnold, 21 July 1935 (rLas) Martin Co., Stuart, Atwood 1917 (cv); Okeechobee Co., n. of Fort Drum, West, 22 Apr. 1946 (FLAS); Orange Co., near Oakland, Curtiss 6624 (CU, GH, MO, NY, us); Osceola Co., Kissimmee, Singletary 136 (pUKE, Ncv); Palm Beach Co., along the Palm Beach Canal, Small 8250 (FLAS, NY, TENN); Pasco Co., near Zephyrhills, Hood 3483 (rr4s); Pinellas Co., near St. Petersburg, Deam 1901 (MicH); Polk Co., n. of Davenport, Hood 3546 (FLAS); Putnam Co., Welaka, DeVall, 23 June 1939 (rLas); Sarasota Co., Myakka River State Park about 18 mi. se. of Sarasota, Wilbur & Webster 2502 (wicum); Volusia Co., near Ariel, Moldenke 180a (Ny, PENN); County unknown, Indian River, Palmer 430 [LeEcTOTYPE of S. gracilis var. grandiflora] (F, GH, NY). CUBA:—Pinar Del Rio: au bord des lagunes de Santa Maria, San Luis, Marie-Victorin & Alain 369 [rve& of S. Alainii] (an, wr); Borders of Laguna Santa Maria, Britton, Britton & Gager 7166 (Ny, us). Province not stated, Wright 412 [sheets sometimes mixed with S. stellaris] (BRU, NY, US).

11. Sabatia brevifolia Raf., Atl. Journ. 1: 147. 1832. S. Elliotii [Elliottii] Steud., Nom. Bot. Ed. 2. 2: 489. 1841. S. paniculata B Elliottii [Steud.] Wood, Am. Bot. & Flor. 266. 1870, without basionym.

Erect annual (15-)30-60(-70) em. high; stem (1-)2-3(-4) mm. in diameter, terete, but usually finely ridged or lined by slight elevations extending between the nodes, pith hollow, at least below. Branches typically alternate, always so above, but in robust, profusely branched plants occasionally with few to several branches of the main stem and even near base of principal branches opposite; in large well-developed plants usually branching throughout the length of the stem, in smaller plants often restricted to the upper half of the stem, strongly ascendent to more commonly widely divergent, usually forming an angle with the stem of (20-)40-60(-80) degrees, slender, very wiry, usually bearing numerous nodes, the internodes not elongate; the aspect of the plant strikingly wiry-virgate. Root-system a pronounced slender or occasionally thick-

1955] Wilbur,—The North American Genus Sabatia 69

ened tap-root 2-8 em. long, (1-)2-3(-4.5) mm. in diameter, bearing few to several very slender laterals. Leaves ascendent and apparently ap- pressed on the main stem and the more or less vertical branches but on strongly divergent branches usually strikingly upwardly secund and often appearing faleate, midvein alone easily detected and that elevated be- neath, apex very minutely apiculate, the lowermost somewhat obovate, oblanceolate, oblong, elliptic, or linear, (0.5-)1-2(-3) em. long, (1-)2-5(-7) mm. wide, typically obtuse or sometimes acute, usually tapering to the sessile base, generally 3-6 times as long as wide; those from about the mid- dle of the stem narrowly oblong or more commonly linear, 1.5-2.5 em. long, 1-3 mm. wide, acute; the upper cauline leaves and those of the branches narrowly linear to filiform or even subulate, gradually reduced to about 3-5 mm. long and less than 0.5 mm. wide, and almost invariably less than 2 mm. wide or twice the diameter of the stem but often the same width, usually 10-25 times as long as broad. Internodes usually 1-2 times the length of the leaves, occasionally greater or sometimes, especially near the base, less. Flowers appearing solitary but arranged in reduced often 1-2-flowered usually secund eymules on the upper side of the divaricate branches. Pedicels very slender, wiry, erect (1—)2-4(-5) em. long. Calyx-tube turbinate, sides sloping to the base, smooth or more commonly noticeably finely nerved, internerval wall thin, often scarious, about 1—2 times as long as broad, (1-)1.5-2.5(-3) mm. long. Calyx-lobes setaceous or subulate, ascendent to wide spreading, (3-)4-7(-8) mm. long, tapering from about 0.5-1 mm. base, usually 2-4 times as long as the calyx-tube, usually not exceeding half the length of the corolla. Upper half of corolla- tube usually cylindrical, tapering below middle to the base, about 1.5-2 times as long as wide, usually 1-2 mm. longer than the calyx-tube or about 2 times as long, (2.5-)3.5-4.5(-5) mm. long. Corolla-lobes spread- ing, elliptic, oblong, oblanceolate, or broadly spatulate, (0.6-)0.9-1.3(-1.8) em. long, (2-)3-5(-7) mm. wide, usually obtuse but not uncommonly somewhat acute, white with a greenish-yellow to yellow patch at base of lobe. Anthers linear, yellow, (2-)3-5(-6) mm. long; filaments greenish- yellow to pale yellow, 1-2 mm. long. Style 1 mm. long or less; stigmatic lobes 3-5 mm. long. Capsule cylindrical, 3-6 mm. long, 2-4 mm. wide. TYPE LOCALITY: Florida. Type: unknown. Described by Rafinesque along with thirty other species from Florida *'seen in gardens and herbals.” DISTRIBUTION: Pinelands and savannas from southeastern South Carolina along the Coastal Plain throughout most of Florida and west at least to Alabama. Map 1l.

This species was described originally by Elliott with the acknowledged aid of Baldwin but the name applied to it was Sabbatia paniculata. This erroneous nomenclature was gen- erally employed even after it was understood to be a mistake. Chapman (1860) was persuaded by Gray to take up the sub- stitute name first proposed by Steudel nearly twenty years

70 Rhodora [Vor. 57

before. Elliott’s application of Michaux’s epithet which initi- ated the confusion is curious in view of his understanding of the subject as shown by the following quotation:

Though the description of Michaux applies more peculiarly to the S. corymbosa, yet as this species was certainly included, and is the only one to which the term paniculata is certainly applicable, I have referred to him here.

This change in concept of the species passing as S. paniculata was pointed out by Grisebach (1839) but he chose to continue the use of the name in the sense of Elliott. Steudel simply substituted the epithet elliottii (as Elliotii) citing in synonymy "S. paniculata Ell. (non Pursh)." The original spelling of the name need not be retained in a case such as this as it was clearly an unintentional orthographic error. Steudel, in listing Stephen Elhiott’s name, spelled the name, “Eliot.” It seems hardly possi- ble that anyone would carry the prerogative of modifying a name when latinizing it to a bibliographic listing; the change must have been unintentional. Wood's solution to the confusion that had enveloped the nomenclature of the white-flowered sabatias was to reduce S. elliottii to varietal status of S. panicu- lata. This indicated no more than a vague bibliographie rather than biological familiarity with the plants.

Nine years before Steudel's publication of S. elliottii, Rafin- esque in a paper describing over thirty species from Florida be- longing to various genera ''seen in gardens and herbals” published S. brevifolia, the original description of which is here quoted in full:

8. Sabbatia brevifolia Raf. Stem dichotomus filiform leaves short subulate acute, flowers terminal white, calyx shorter than corolla setaceous, segments of corolla obovate. Near to S. brachiata and stellaris.

This description, although very brief, seems very apt in pointing out the stronger features that characterize the species that has long been called S. elliottii. Even in the absence of an authentic specimen I feel no hesitation in taking up Rafinesque's name other than that caused by the regret of seeing a well- established name replaced by one long-forgotten. It would be difficult to present a better description in so few words of the plant first characterized by Elliott than Rafinesque’s attempt. Every descriptive phrase fits Elliott's plant very well.

REPRESENTATIVE SPECIMENS:—NORTH CAROLINA: without locality, Delile (Nv). Since this species is not otherwise known north of Charlestown, S. C.,

1955] Palmer,—Epifagus virginiana 71

its presence in North Carolina seems most doubtful SOUTH CAROLINA: Beaufort Co., Bluffton, Mellichamp, 1886 (r, Mo, Ny, vs); Charleston Co., Adams Run, Godfrey & Tryon 1544 (DUKE, F, GH, MICH, MO, NY, PENN, TENN, us); Hampton Co., about 4 mi. se. of Hampton, Wilbur & Webster 2826 (MrcH); Jasper Co., Ridgeland, Mohr, Nov. 1893 (Mo, NY, US). GEORGIA: Camden Co., w. of St. Marys, Small, DeW inkeler & Small 10545 (DUKE, WVA); Charlton Co., Okefenokee Swamp between Chesser Island and Camp Cornelia, Thorne & Ford 2064 (cv, Ga); Coffee Co., without exact locality, Harper 681 (NY, vs); Pierce Co., 3 mi. n. of Blackshear, Thorne & Norris 6202 (cu, GA); Ware Co., 5 mi. se. of Waycross, Wilbur & Webster 2731 (MICH). FLORIDA: Brevard Co., Merritts Island, Curtiss 2227 (F, GH, MO, Ny, us); Clay Co., about 2 mi. n. of Orange Park, Moldenke 162 (DUKE, MO, NY, PENN, US); DeSoto Co., 4 mi. w. of Arcadia, Webster 4224 (micu); Duval Co., near Jack- sonville, Curtiss 5114 (cU, F, GH, NCS, NY, US); Franklin Co., Apalachicola, Biltmore Distrib. Chapman Herb. 903b (GH, MO, NCS, NCU, NY, PENN, US); Hernando Co., between Brookside and Bayport, Jones 43 (cu, vs); Hills- borough Co., Tampa, Nash 2422 (F, GH, MICH, Mo, MT, NY, US); Indian River Co., near Felsmere, Small 8902 (Fuas, GH, NY, US); Jackson Co., 4 mi. e. of Marianna, Wiegand & Manning 2559 (cu, au); Lake Co., near Cassia, Hood, 1 Sept. 1911 (FLAs, GH); Lee Co., s. of Fort Myers, Moldenke 909 (DUKE, Mo, NY, PENN, Us); Liberty Co., near Roy, Wiegand & Manning 2558 (cv, an); Manatee Co., Bradenton, Tracy 7080 (CU, F, FLAS, GH, NY, PENN, TAES, US); Orange Co., about 2.5 mi. ne. of Apopka, Wilbur & Webster 2650 (MicH); Palm Beach Co., Earman, Randolph 26 (cv, GH); Pinellas Co., about 4 mi. w. of St. Petersburg, Deam 2840 (micu, vs); Putnam Co., 10 mi. se. of Inter- lochen, Fox 5700 (micu, Ncs); Seminole Co., Lake Monroe, Garber, Mar. 1896 (BRU, F, US, YU); Volusia Co., Beresford, Hood, 21 Sept. 1910 (FLAs, Mo). ALABAMA: Mobile Co., between Theodore and Hollander’s Island, Pennell 4512 (NY, PENN). LOUISIANA: without locality data, Featherman (mo). This species should not be included in the flora of the state without a better substantiated record. It seems likely that there is confusion in labeling in regard to this collection. (to be concluded)

EPIFAGUS VIRGINIANA.—On the northern exposure of a deciduous woods, consisting chiefly of oaks and beeches, there is at Swarthmore, Delaware County, Pa., a large colony of Epifagus which is butter-yellow in color. Nearby a number of hemlocks are growing, but this plant is not found under these, but confines itself to the deciduous trees. A year ago I counted 300 of this yellow Epzfagus without materially changing my position, and this year (’54) I counted 175 plants along a wood’s lane within the space of 100 feet. A fair estimate of the plants on this northern hillside would be more than 1000.

The plant resembles the species closely except for color. If any difference is to be noted I think the yellow form tends to be more bushy and is perhaps more fleshy. However, there

72 Rhodora [Vor. 57

are numbers of single-stemmed plants scattered about. Some of these are pure yellow, but others show touches of reddish brown on the stem scales.

The usual color-formof E. virginiana does not appear within the area occupied by the main growth of the yellows, but as the yellows diminish along the edges the usual form appears. "The whole area is about an acre in extent.

'The concentration of the yellow forms on this hillside suggests a common origin for them, but does not explain the absence of the usual form. So many plants of this one color within the limited area indicated with the usual form of the species on the periphery offers à nice problem of distribution and might provide an interesting population study for the appropriate person.— SAMUEL C. PALMER, SWARTHMORE, PA,

CALYCERA BALSAMITAEFOLIA IN THE UNITED STATES.— Recently there were found in the Herbarium of the Chicago Natural History Museum two sheets of Calycera balsamitaefolia (Juss.) Rich., of the family Calyceraceae. "They were included in the J. T. Rothrock collection, when that herbarium was purchased by the Chicago Natural History Museum in 1909. Both sheets were collected on ballast at Kaighns Point (spelled Kaigns on the label) in New Jersey. This station is near Cam- den opposite Philadelphia. One sheet has the label of ‘Her- barium of Isaac Burk, Philadelphia; Pa.", the other sheet ac- companied by a label of the “Herbarium of University of Penn- sylvania." No date is indicated on either label.

It is of interest to note this collection, because the species is a native of Chile, and is not mentioned in either the eighth edition of Gray’s Manual, Gleason’s New Britton & Brown, or any of the floras of New Jersey and Pennsylvania. Probably col- lected at the station given above in the latter decades of the nineteenth century or at the beginning of the twentieth century, the species apparently has never been re-collected either at the Kaighns Point station or elsewhere. Botanists from the vicinity of Philadelphia and New York should give special attention to ballast sites similar to the one where Calycera was found.— JULIAN A. STEYERMARK AND FLOYD A, SWINK, CHICAGO NATURAL HISTORY MUSEUM AND COLLEGE OF PHARMACY, UNIVERSITY OF ILLINOIS,

Volume 57, no. 673, including pages 1-86, was issued 28 February, 1955.

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A NOTE ON THE NAME CALAMINTHA Gorpon P. DEWorr, Jn.

THE genus Satureja, L. as interpreted in the 8th edition of “Gray’s Manual of Botany" includes plants that have been included in four genera: saTUREJA (S. hortensis L.); CALAMINTHA (S. Calamintha, (L.) Scheele — C. officinalis Moench; S. glabella (Michx.) Briq. = C. glabella (Michx.) Benth.; S. arkansana (Nutt.) Briq. = C. glabella (Michx.) Benth., var. angustifolia (Torr. DeWolf [Satureja glabella (Michx.) Briq., var. angusti- folia (Torr.) Svenson]; AciNos (S. Acinos (L.) Scheele = A. arvensis (Lam.) Dandy; and cLinopopium (S. vulgare (L.) Fritsch — C. vulgare L.).

Study of material of species of this group known to be in cultivation has convinced me that these four taxa are not con- generic, and that current European practice of recognizing them as distinct genera should be followed. A fuller discussion of the taxonomic aspects of the problem has been published else- where (Baileya 2(4): 142-150. 1954 [Jan., 1955]). Here I should like to discuss a purely nomenclatural problem.

In the 52nd volume of Fedde's “Repertorium specierum novar- um regni vegetabilis," Heft 2, pp. 144—161, 1943, there appears the second of a series of articles by Erwin Janchen entitled “Zur Nomenclature der Gattungsnamen." My attention was drawn to this paper by the inclusion in the 1952 edition of the "International Code of Botanical Nomenclature," p. 131, of the name Calamintha, Lamarck as a nomen conservandum pro- positum, the respective nomen propositum rejiciendum being Clinopodium, Linnaeus. It seems to me that certain data have been overlooked in the framing of this proposal, and hence, further, and perhaps fuller, discussion is in order.

74 Rhodora [Vor. 57

In the 1943 paper, p. 156, no. 7305, Janchen refers to a paper by Janchen and Neumayer, entitled “Beiträge zur Benennung, Bewertung und Verbreitung der Farn- und Blütenpflanzen Deutschlands" in the “Oesterreichische Botanische Zeitschrift," volume 91, 1942, pp. 209-298, in particular to p. 274. At this place the identity of Calamintha, Moench (1794) [non Lamarck! 1778] and Clinopodium, Linnaeus is asserted, and their dis- tinetness from Satureja, Linnaeus is affirmed. Further, Janchen and Neumayer stated: *. . . Mit Rücksicht auf die Artenanzahl ist Calamintha gegenüber dem älteren Gattungsnamen Clinopodi- um auf der Ausnahmsliste zu schützen . . ." They also refer to a paper in the “Acta Horti Gotoburgensis," volume 13, 1939, pp. 335-380, in particular p. 349, in which essentially the same thought is expressed by Handel-Mazzetti.

Objections to this proposal arise on two points:

1. Calamintha, Lamarck (1778) is a later homonym of Calamintha, Miller (1754) Calamintha, Trew (1754) and Calamintha, Adanson (1763) (and/or Scopoli, 1772), and is synonymous with Calamintha, Miller and Calamintha, Trew.

2. Calamintha, whether of Miller, Trew, Adanson, Scopoli, Lamarck, or Moench, did not contain any species then ascribed to Clinopodium. The two taxa were not confused until 1891 when O. Kuntze reduced Calamintha, Moench emend. Bentham to Clinopodium, Linnaeus emend. O. Kuntze.

Reference to Pfeiffer's “Nomenclator Botanicus,” volume 1,

part 1, of 1873 indicates that Calamintha was first used as a post-Linnaean name by Adanson, in the “Families des Plantes" of 1763, volume 2, p. 192, for the Linnaean genus Glechoma. This same usage was maintained by Scopoli in the first volume of the second edition of the “Flora Carnioliea" (1772), at p. 423, where synonymy and a good description of the single species [Glechoma hederacea] are given. The next entry should be to the usage of Lamarek in the second volume of the first edition of the “Flore Francoise," 1778, p. 393, no. 432. Pfeiffer missed this, however, so the next actual entry is to Moench's “Methodus Plantas . . ." of 1794.

A note about Calamintha, Moench is perhaps in order. In 1794 the “Methodus Plantas Horti Botanici et Agri Marburgen- sis, a Staminum Situ Describendi" of Konrad Moench was published. In this work, at least so far as the labiates are

1955] DeWolf,—A Note on the Name Calamintha 75

concerned, very good generic and specific diagnoses are given. It was used by Bentham during the preparation of his ‘‘Labia- tarum, Genera et Species," of 1832-36, and, in particular, Bentham accepted Moench’s circumscription of Calamintha over that of Lamarck. As published by Moench, three species were accepted: C. grandiflora, (L.) Moench; C. officinalis, Moench; and C. trichotoma, Moench = C. Nepeta, (L.) Savi.

Meanwhile, Clinopodium had received a very uniform treat- ment. In 1753 Linnaeus included three species in his genus, viz., Clinopodium vulgare; C. incanum = Pycnanthemum incanum, (L. Michx., fide Bentham, "Labiatarum . . ." p. 327; and C. rugosum = Hyptis capitata, Jacquin (ex Jamaica) and H. radiata, Willdenow (ex Carolina), fide Bentham, “Labiatarum

. ." pp. 104 and 108. Of these three species, Clinopodium vulgare has been consistently retained in the genus since its publication, and its immediate relatives have been placed with it.

In 1754 Philip Miller published a three volume, octavo, abridgement of the last folio edition (? 6th, 1752) of “The Gar- deners Dictionary." In this work binomial nomenclature was not used, but the generic descriptions are well drawn!. We are, therefore, faced with the necessity of considering Miller's genera. We find that Miller defined a genus he called “Cala- mintha.” The generic circumscription is tolerably specific, and, of the six taxa listed, the first three are the same as those accepted by Moench. They are: Calamintha vulgaris, vel officinarum Germanicae, of Caspar Bauhin = C. officinalis, Moench; Cala- mintha pulegii odore, foliis latioribus, of Paul Hermann = C. Nepeta, (L.) Savi; and the Calamintha magno flore, of Caspar Bauhin = C. grandiflora, (L.) Moench.

When I discussed the taxonomic aspects of this problem in Baileya (l.c.) the name Calamintha was ascribed to Philip Miller. This occurred because of ignorance of the fact that also in 1754 Christopher Jacob Trew, in his Nuremburg edition of Elizabeth Blackwell’s “A Curious Herbal," which is usually referred to as the "Herbarium Blackwellianum," had used the name Cala- mintha. Trew validated his name by reference to the generic descriptions of pre-Linnaean authors, including Ray and Tourne-

! After Tournefort, fide G. C. Druce in Rpt. Botanical Exchange Club of the British Isles 3: 426-428, 1913.

76 Rhodora [Vor. 57

fort. Further he gave two excellent plates: no. 166, Calamintha moniana — C. officinalis, Moench and no. 167, Calamintha officinalis — C. Nepeta, (L.) Savi.

In the past there has been a good deal of discussion as to the advisability of rejecting all names published in works dated after 1753 which do not use binomial nomenclature. As late as 1935 a formal proposal? was made to the 6th International Botanical Congress that such be done, and a list of proscribed works be made. This seems, however, to have met with no success. We are left, then, with the tacit understanding that generic names, so long as validly and legitimately published after 1753, may be taken up even from works not using binomial nomenclature.

In the present case we are concerned with homonymous and/or synonymous names, published in the same year, but in unknown sequence. It behooves us, then, to take up for purposes of nomenclature, the most adequately circumscribed of the two names. Miller provides a generie description in English, and cited, with description, also in English, six species. Trew gave no generic description as such, but validated his name implicitly by reference to previously effectively published generic deserip- tions, especially those of Ray and Tournefort. Trew also gave two excellent plates of the two species which he accepted and full specific descriptions in Latin and German. Therefore, for purposes of citation I am accepting Calamintha, Trew.

Lamarck’s circumscription of Calamintha on the other hand, is not good, nor does he refer to descriptions by previous authors. The genus may be identified with certainty only by recourse to the included species. Of these, the first two, C. alpina and C. arvensis, are now placed in Acinos; C. cretica is referred to Micromeria; and only the last two, C. parviflora and C. montana are now retained, under earlier names, in Calamintha (C. Nepeta, (L.) Savi and C. officinalis, respectively, fide Bentham “Labia- tarum . . ." p. 387-388).

Until the publication of O. Kuntze (Revisio Generum Plan- tarum, volume 2, pp. 513-516) in 1891, there was little con- fusion of the two taxa which may be distinguished taxonomieally as follows:

? Wilmott, A. J.—Kew Bull. 1935 66. 3 Little, E. L.—Madrono 7: 240—242, 1944.

1955] DeWolf,—A Note on the Name Calamintha 71

Calamintha Clinopodium Calyx tubular, straight. Ver- Calyx tubular, curved. Ver- ticellasters on relatively long ticellasters sessile or nearly so.

pedicels, sub-secund.

After the genera were firmly established, with good circum- scriptions, by Moench in 1794, in the “Methodus Plantas . . .," they were generally treated as taxa of correlative rank. Ben- tham treated them as sections of Melissa, L. in the *Labiatarum

. ." of 1832-36, and as sections of Calamintha, Moench emend. Bentham in the 12th volume of de Candolle's "Prodromus Systematis Naturalis . . ." of 1848. Briquet, in IV Teil, 3 Abteilung, a, of" Die natürlichen Pflanzenfamilien" of 1897, treated them as sections of Satureja, L. Neither Bentham nor Briquet considered them synonymous, as did O. Kuntze and Janchen and Neumayer.

Two facts emerge from this discussion. In the first place, Calamintha and Clinopodium, as originally proposed, and as generally used to 1891, apparently had no species in common. 'There was, and is today, controversy over the rank of the taxa, but not over their identity or composition. In the second place, the proposal by Janchen is to conserve a poorly circumscribed later name over an adequately circumscribed Linnaean name. Further, the name proposed for conservation is both a later homonym and synonym of earlier names, facts not mentioned by the propositor.

The “International Code of Botanical Nomenclature" (1952), at Article 24, stated: “. . . These names [for conservation] are preferably such as have come into general use in the fifty years following their publication, or which have been used in monographs and important floristic works up to the year 1890

. ." Up to the year 1890 both Calamintha, Moench (non Lamarck) and Clinopodium, Linnaeus were in general use for the respective taxa concerned. "There was no confusion between them.

Finally, if Clinopodium, Linnaeus is declared a nomen rejicien- dum, those who desire to recognize the taxon which has borne that name as a genus will be faced with the necessity of pub- lishing a new name for it.

For these reasons it is recommended that the proposal for the

78 Rhodora (Vor. 57

conservation of the name Calamintha, Lamarck, and the rejec- tion of the name Clinopodium, Linnaeus not receive favorable

actlon.— BAILEY HORTORIUM, CORNELL UNIVERSITY, ITHACA.

A REVISION OF THE NORTH AMERICAN GENUS SABATIA (GENTIANACEAE)

ROBERT L. WILBUR (Continued from page 71)

E. Subsection DopECANDRAE subsect. nov.?

Subg. Plurimaria Raf., Med. Fl. 2: 76. 1830, in part, not Plurimaria Raf. (as genus), Fl. Tell. 3: 31. 1837.

Pleienta Raf., Fl. Tell. 3: 30. 1837, in part, an illegitimate name since its type, designated by Rafinesque, is the same as that of Sabatia. Sect. Pleienta (Raf.) Blake, Ruopora 17: 56. 1915, an illegitimate

name. Dodecandrae Small, Man. SE. Fl. 1049. 1933, a category of undesig- nated status.

Rhizomatose perennials with at least the secondary and very often the primary branches alternate. Strongly pronounced tendency towards plurimerous flowers ranging from 5-12(-14)-parted. Flowers typically large and pedicels usually longer than 1 em. TYPE SPECIES: Sabatia dodecandra (L.) BSP.

This subsection contains but four species which, except for one species, are restricted to the Coastal Plain of the United States. S. calycina occurs in addition on at least two islands of the West Indies.

S. calycina, I feel certain, has very little in common with the species of subsection Campanulatae with which it has been for- merly associated. Its proper relationship is more aptly shown by grouping it with the rhizomatose, pluripetalous species of the dodecandra-alliance. The strongly pronounced tendency for this species to possess flower-parts more numerous than five, the large, often foliose calyx-lobes, the often conspicuous, elongate rhizome, the broad leaves and the broadly campanulate calyx-tube are features of all or most of the dodecandra-group

5 Subsectio Dodecandrae, Perennes rhizomatibus praeditae. Rami secondarii et interdum primarii alterni. Flores saepe plurimeres, inter 5-partiti et 14-partiti, plerumque in specie una 5-6-partiti, in speciebus aliis ca. 9- partiti, typice grandes,

pedicellis quam 1 cm. longioribus. Species typica, Sabatia dodecandra (L. BSP. (Chironia dodecandra L.)

1955] ^ Wilbur,—The North American Genus Sabatia 79

and are present in none of the species with which it has formerly been allied. I do not feel that the Dodecandrae are so much more strongly differentiated than the other subsections as to warrant sectional rank which Blake accorded to them.

KEY TO THE SPECIES OF SUBSECTION DODECANDRAE

A Calyx- and corolla-lobes 5-6-parted or rarely up to 7; corolla-lobes (0.6—) 0.7-1.3(-1.5) em. long; cauline leaves spatulate, tapering into a much- narrowed, sometimes almost petiolate base............ 12. S. calycina.

A. Calyx- and corolla-lobes 7-13-parted, usually 9-12 in number, corolla- lobes (1.2—)1.6-2.4(-3.5) em. long; cauline leaves elliptic, lanceolate, or linear, the median and upper at least not strongly tapering into an almost petiolate base.

B. Upper cauline leaves less than the diameter of the stem or but little wider, very narrowly linear, contrasting strongly with the broadly spatulate, rosulate, basal leaves; leaves thick and succulent, often drying somewhat rugose; the larger roots succulent; calyx-lobes strongly subulate, often succulent and even semicircular in cross- Secuont Mc vu PME ree ee mere re 15. S. bartramit.

B. Upper cauline leaves considerably wider than the diameter of the stem, lanceolate, elliptic, or linear; basal leaves often absent, when present linear to lanceolate, and not strongly contrasting with the lower cau- line leaves; leaves drying smooth and usually thin in texture; roots fibrous, non-succulent; calyx-lobes linear, thin and flattened in cross-section.

C. Calyx-lobes hyaline-margined; plants strongly stoloniferous; primary branches very often opposite; terminal flower short-pedicellate, usually considerably exceeded by the first internode of the lateral branches arising at the same node............ 14. S. kennedyana.

C. Calyx-lobes not hyaline-margined; stolons usually lacking, if present neither numerous nor strongly developed; primary branches usually alternate; terminal flower typically long-pedicellate, usually about equaling or exceeding the first internode of lateral branch arising at the same node.............. cece eee eee eee 13. S. dodecandra.

12. Sabatia calycina (Lam.) A. Heller, Bull. Torr. Bot. Club 21: 24. 1894. Gentiana calycina Lam., Encyc. 2: 638. 1788. Chironia dicho- toma Walt., Fl. Car. 93. 1788. C. calycosa Michx., Fl. Bor.-Am. 1: 147. 1803. Sabbatia calycosa (Michx.) Pursh ex Sims, Curtis’s Bot. Mag. pl. 1600. 1813. S. gracilis var. cubensis Griseb., Mem. Am. Acad. 11: 521. 1862. S. dichotoma (Walt.) Trelease ex Branner & Coville, Ann. Rept. Geol. Surv. Ark. 1888. 4: 204. 1891. Sabbatia cubensis (Griseb.) Urb. Symb. Ant. 8: 536. 1921.

Perennial herb (although flowering the first year) (8-)15-40(-50) cm. high, with a slender to thick rhizome (1-)2-4(-10) em. long, 1-3 mm. in diameter. Stems usually solitary, or sometimes several and clustered, erect, rigid, smooth above or but very slightly ridged, hollow at least below, (1-)2-3(-4.5) mm. in diameter, branching usually restricted to the upper half or third of the stem but may arise from nearly throughout the entire length. Branches generally alternate but commonly opposite

80 Rhodora [Vor. 57

even above along the main stem, ascendent or more typically divaricate, further branching also strongly divergent, thus the plant typically pre- sents a strikingly geniculate aspect, branching usually restrieted to the second or third order, usually less than 20 em. long, bearing but few nodes. Root-system of several to numerous slender, fibrous roots arising from the rhizome or from the base of the stem. Leaves thin, drying very thinly membranous and hence venation conspicuous although only the midvein even slightly elevated beneath. Basal rosette lacking and the lower cauline leaves not conspicuously differentiated in shape from the median or upper leaves and usually the reduction in size is gradual and slight. Leaves ascending or more often strongly spreading, elliptical to broadly spatulate, mostly obtuse but still commonly acute, tapering into a con- spieuous, much-narrowed or even petiolate base, (1.5-)2.5-6(-10) em. long, (4-)10-18(-20) mm.