P1. Syst. Evol. 201:243-261 (1996) Plant

Systematics a n d

Evolution © Springer-Verlag 1996 Printed in Austria

Systematics, typification, distribution, and reproductive biology of Pinckneya bracteata (Rubiaceae)

PIERO G. DELPRETE

Received July 25, 1995

Key words: Rubiaceae, Condamineeae, Rondeletieae, Pinckneya, Pogonopus. - Taxono- my, typification, BARTRAM, MICHAUX, Flora boreali-americana. - Flora of the Unites States, South Carolina, Georgia, Florida; conservation biology; ornamental shrubs; cultivated species.

Abstract: Complete taxonomic history, lectotypification, and putative generic relation- ship of Pinckneya are presented. The much debated authorship of Flora boreali-americana and the validity of W. BARTRAM'S binomials relative to this taxon are discussed. A single species, P. bracteata, is recognized. It occurs in the southeastern USA (South Carolina, Georgia, and northern Florida). Circumscription of the genus, complete synonymy, and a description of the species are presented. Distribution map, phenological diagram, notes on phenology, and medicinal and economical uses are included. A short discussion on the conservation biology of this diminishing species is included.

Taxonomical history and typification

The genus Pinckneya was first published in Flora Boreali-Americana (FB-A), a work officially attributed to ANDRE MICHAUX, but published posthumously (by his son FRANCOIS ANDR~ MICHAU×) one year after his death in 1803 (A. MICHAUX died in Madagascar in November 1802). Many botanists (PERSOON 1805: 197; W. J. HOOK- ER 1825; MARTIUS 1840: 88; GRAY 1841, 1882; PLANCHON & LINDEN 1854; SARGENT 1889; URBAN 1902: 85--86; LEROY 1957: 6, 203; etc.) have written on the validity of the authorship of FB-A, stating that F. A. MICHAU× attributed the publication to his father, but the work was almost entirely written by Louis CLAUDE MARIE RICH- ARD. WILLIAM JACKSON HOOKER (1825) stated with clarity his opinion with respect to this subject: "... that work [FB-A] ... was compiled from the materials that he [A. MICHAU×] collected during his travels in North America, is his Flora Borealis Americana, sistens Characteres Plantarum quas in America Septentrionali colleg- it et detexit. This appeared in 1803, in two volumes octavo, with fifty-one neat plates in outlines. The anonymous editor, and indeed he may justly be considered the author, was the eminent CLAUDE LOUIS RICHARD, late professor of botany at the School of Medicine in Paris, and unquestionably one of the most profound bota- nists that Europe has ever known. Regarding L. C. M. RICHARD'S authorship of FB-

244 R G. DELPRETE:

A, MARTIUS (1840: 88) wrote: "autor [L. C. M. RICHARD] quoque est, saltem pro maxima parte, Florae boreali-americanae ab MICHAUX 1803 editae." GRAY (1841) stated: "MICHAUX himself, though an excellent and industrious collector and observer, was by no means qualified for authorship; and it is to L. C. RICHARD that the sagacious observations, and the elegant terse and highly characteristic specif- ic phrases of this work are entirely due." Of the same opinion was SARGENT (1889), who referred to FB-A as that treatment which "RICHARD [L. C. M.] drew up large- ly from the plants collected by MICHAUX in North America". About L. C. M. RICH- ARD, LEROY (1957) wrote (free translation from French): "It is to him that the description of the American plants collected by ANDRE and FRANqOIs-ANDRE MICHAUX should be attributed. In conclusion, it is he that edited the posthumous work [supposedly] of ANDRE MICHAUX." Additional supporting comments on the authorship of FB-A and on MICHAUX'S and RICHARD herbaria may be found in SAYRE (1976). The most salient comment on FB-A can be found in a letter dated 12 Octo- ber, 1801 that ANDR~ MICHAUX sent from Mauritius (Mascarene Islands) [but FRANqOIS ANDRE was in America from August 1801 to Spring 1803! (SAYRE 1976)]. As a postscript, in the bottom of that letter, A. MICHAUX wrote (in SAVAGE • SAVAGE 1986: 178): "My compliments to REDOUT~ [artist and illustrator of FB-A]. At what stage is the printing of the two works: Histoire des Chines and the Flora Boreali- Americana? If some information is needed, I can send it. How has RICHARD con- ducted himself in our behalf? I can send him some shells; I have not sent them because I wished to hear from him first."

Curiously, the same problem of neglected authorship happened also to L. C. M. RICHARD'S son, ACHILLE RICHARD. The M6moire sur les Rubiac6es was presented by A. RICHARD on 7 July 1829 in front of the Acad6mie Royale des Sciences at Paris, and sent to publication the same year. Contemporarily, A. RICHARD sent one copy of the manuscript of his M6moire to DE CANDOLLE, who published it within his Pro- dromus (DE CANDOLLE 1830), without citing the original author, in September 1830 (three months before the publication of the Mdmoire). The authority of A. RICHARD'S new taxa was thereafter variably assigned to RICHARD or to DE CANDOLLE. To acknowledge the proper authority of such an honorable botanist, as recognized by STEARN (1957), A. RICHARD'S new rubiaceous taxa should be cited as: A. RICH. in DC. Prodr. 4:- (Sep 1830).

Regarding the typification of Pinckneya, one herbarium sheet has been found in the Richard Herbarium housed in the Museum of Natural History at Paris (PC, Richard Herbarium). Attached to this specimen (Fig. 1) is the complete original description in L. C. M. RICHARD'S handwriting (Fig. 2) of Pinckneya pubens, which has been partially transcribed in the same format, style and words in the descrip- tion to be found in FB-A. To confirm the previous statement, A. RICHARD (1830) in his M6moire attributed the authorship of this genus to his father as: "Pinckneya RICH. in Mica. F1. Am. B." At least three other botanists also attributed the author- ship of Pinckneya to L. C. M. RICHARD: J. D. HOOKER (1873) as "Pinckneya, RICH. in MICH. F1. Bor. Am."; BAILLON (1881: 257--503) simply as "Pinckneya L.-C. RICH.", adding "Michx. F1. B or. Am." in the footnotes; and SCHUMANN (1891) sim- ply as "Pinckneya RICH." Other authors gave authority of this genus solely to A. MICHAUX aS follows: DE CANDOLLE (1830: 341--622) as "Pinckneya [hort. var.] MICHX. fl. bor. am."; and STANDLEY (1918) aS "Pinckneya MICHX. F1. Bor. Am.".

Pinckneya bracteata 245

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Taking into consideration L. C. M. RIClqARD'S original description attached to the specimen of the Richard Herbarium, the authority chosen by his contemporary colleagues, and the arguments supported by W. J. HOOKER (1825), MARTIUS (1840), GRAY (1841, 1882), URBAN (1902), SARGENT (1889), and SCHUMANN (1891), I con- clude that the proper authority for this genus should be: "Pinckneya L. C. M. RICH- ARD in MICttAUX, F1. Am,-Bor."

The dedication of the name Pinckneya is clearly stated by F. A. MICHAUX (the son) in his North American Sylva (F. A. MICHAUX, 1841-42): "In testimony of his gratitude and respect, he [his father, A. MICHAUX] consecrated it dedicated to CHARLES COTESWORTH PINCKNEY, an enlightened patron of the arts and sciences, from whom my father and myself, during our residence in South Carolina, received multiplied proofs of benevolence and esteem." For some obscure reasons L. C. M. RICHARD remained anonymous, and the two MICHAUX accredited his name in none of their works. ANDR~ MICHAUX spent the early years of his life as gardener and, without receiving a formal education, his botanical knowledge was not very pro- found. This is affirmed by SARGENT (1889): "MIcHAUX'S cultivation and literary ability, judged by his journal, were not great; and his reputation as an author is due to the fact that his name was printed upon the title page of [FB-A]."

To understand his botanical preparation and the extent of his travels, I have read MICHAUX'S entire diary that was reproduced by SARGENT (1889) in the original language (French). In his diary A. Mici~AUX reported that from 30 April to 2 May 1791 he collected (his son had already returned to France) on the St. Mary (Ste Marie) River but without reporting any reference to Pinckneya. Nevertheless, in his diary, two weeks later, he recorded from the surroundings of Savannah (Geor- gia, Chatam County): "Le 18 [May 1791] herboris6 dans le camps a une grande distance et reconnu un arbrisseau qui se rapporte au genre Mussa[e]nda" [The 18 (May 1791) (I) collected in the fields at a large distance (from Savannah) and rec- ognized a shrub related to the genus Mussaenda (as Mussanda)], certainly refer- ring to Pinckneya. Additional information (but slightly contrasting) can also be found in the notes of F. A. MicttAUX (1841-42): "My father found it [Pinckneya] for the first time, in 1791, on the banks of the St. Mary [River]. He carried the seeds and young plants to Charleston, and planted them in a garden which he pos- sessed near that city. Though entrusted to ungrateful soil, they succeeded so welt that in 1807 1 found several of them twenty-five feet high and seven or eight inches in diameter; which proves that the vegetation of this tree does not require a very warm climate nor a very substantial oil."

With respect to the typification of Pinckneya pubens, it seems to be an obvious decision to select the specimen housed at the RICHARD Herbarium at Paris (PC, Richard Herbarium), bearing L. C. M. RICHARD'S description (and probably col- lected by MICHAUX) as the lectotype.

Three other specimens dubiously related to F. A. MICHAUX have been found in various institutions. In the general herbarium of Paris there is a specimen that bears (with DESFONTAINES' handwriting) the following label: "Pinckneia [hort. var.] pubens MICH. - Amer. Spt. - M~CHAUX" which attached a small printed label saying "isotype". At the Field Museum (F) has been found a fragment [one young fruit and three calycophylls] labelled as "Ex Herbario Musei Parisiensis" that bears the notation (unknown handwriting): "Pinckneya pubens Mrcnx., M~cnx", as if col-

248 R G. DELPRETE:

lected by MICHAUX, but its origin is rather dubious. Finally at the Geneva Herbari- um (G) I located one specimen that bears the following label: "Pinckneia, MICH. - plus. bot. regardant cette plante comme un Cinchona; mais c 'est un genre distinct [most botanists see this plant as a Cinchona, but it is a distinct genus]. - Amer. Sept. - MICHAUX fil. ded. [handwriting of doubtful author]". I believe that if the handwriting was that of F. A. MICHAUX (the son), he would have not made the mis- take of misspelling Pinckneya as Pinckneia, since he personally met the person to whom the genus is dedicated, and to whom he later referred (F. A. MICHAUX 1841--42). In conclusion, none of the three specimens housed at P, F, and G can be chosen with certainty as types.

To fully appreciate the complex taxonomic history of this monotypic genus we have now to go back in time. Pinckneya was originally discovered by JOHN BAR- TRAM, on 1 October, 1765, accompanied by his young son WILLIAM BARTRAM. The travel diary of the BARTRAMS was published by the son in 1971, and in three pas- sages he cited the encounters with this species. In the introduction (p. xviii) of his Travels, W. BARTRAM (1791) wrote the following passage: ".. . Others [shrubs] astonish us by their pleasing figure and disposal of their vestiture, as if designed to embellish and please the observer, as ... Bartramia bracteata ...". But W. BARTRAM'S (1791) best description of this species can be found when he recalled his very first encounter with it (in HARPER 1958: 16), when he and his father arrived "on the waters of Cat-head creek, a branch of the Alatamaha [River], near Fort Barrington" (McIntosh County, Georgia): "... The other [shrub] was equally dis- tinguished for beauty and singularity; it grows twelve or fifteen feet high, the branches ascendant and opposite, and terminate with large panicles of pale blue tubular flowers, speckled on the inside with crimson; but what is singular, these panicles are ornamented with a number of ovate large bracteae [calycophylls], as white, and like fine paper, their tops and verges stained with a rose-red, which, at a little distance, has the appearance of clusters of roses, at the extremities of the limbs: the flowers are of the C1. Pentandria monogynia; the leaves are nearly ovate, pointed and petioled, standing opposite to one another on the branches."

Later in his diary W. BARTRAM (1791, in HARPER 1958: 468) narrated his third encounter with this species, but this time proposing a different generic name: "The other new, singular and beautiful shrub*, now here in full bloom, I never saw grow but at two other places in all my travels, and there very sparingly, except in East Florida, in the neighborhood of the sea-coast ... *I gave the name of Bignonia bracteata extempore."

HARPER (1942) published a brief article in defense of the legitimacy of Bartra- mia (Bignonia) bracteata, which is validly published in having a Latin binomial and a complete beautiful description. The same viewpoint was taken by MERRILL (1945), stating that: "there is no doubt that his description and the note on p. 468 [and p. 16] belong with the binomials from the context." The argument supported by RICKETT (1944) against the validity of W. BARTRAM'S names as published in his Travels is, in my opinion, not sufficient to exclude the validity of Bartramia (Big- nonia) bracteata.

RAFINESQUE (1820) merely listed the genus [as Pinckneya Mx.], but without attaching it to any specific epithet. Several years later RAFINESQUE published a short article (in the series "The School of Flora"), both in the Saturday Evening Post

Pinckneya bracteata 249

(RAF~NESQUE 1827a) and in Casket (RAFINESQUE 1827b), where he legalized the com- bination Pinckneya bracteata (W. BARTRAM) RAF., with a complete description of the plant, along with an account of its medicinal and ornamental uses. The previ- ous publication (RAFINESQUE 1827a) clearly states on its front page that this was published on 19 May, 1827, but the second (RAFINESQUE 1827b) simply states May 1827. Because of the impossibility to ascertain the exact date of the second publi- cation, I conclude that RAFINESQUE'S valid combination was first published in the Saturday Evening Post (RAFINESQUE 1827a). In the same article RAFINESQUE (1827a, b) also cited (under P. bracteata) the combination Mussaenda bracteata, which he reported as one of the many names previously proposed for this species, but with- out citing any authority (DE Juss~Eu?). The same inconsistent author, three years later (RAFINESQUE 1830: 57, fig. 72), took up the name P. pubens MICttx., without even reporting P. bracteata as synonymous! To add more confusion MERRmL, while previously recognizing BARTRAM'S name as valid (MERRmL 1945), in his Index Rafinesquianus (MERRILL 1949) inexplicably failed to recognize RAFINESQUE'S P. bracteata, reducing it to synonymy under P. pubens!

At the end of the analysis of such a complex and confounding taxonomic histo- ry, I conclude that Bartramia bracteata W. BARTRAM is validly published, and that the combination P. bracteata (W. BARTRAM) RAF. is the correct one to be used for this species.

HARPER (1942), purposely or otherwise, overlooked RAFINESQUE'S (1827a, b) combination, proposing his own combination, stating: "Since BARTRAM'S specific name bracteata has 12 years' priority, it must replace MICHAUX'S pubens. Accord- ingly, the Georgia bark is to be known hereafter as Pinckneya bracteata (BARTRAM) F. HARPER, n. comb." Since RAFINESQUE'S (1827a, b) combination has priority, HARPER'S combination is superfluous.

In selecting the lectotype of Bartramia (Bignonia) bracteata, EWAN'S (1968) reproduction of the 'Botanical and zoological drawings' by WILLIAM BARTRAM, 1756--1788, has been of cardinal importance. EWAN (1968: 152) transcribed the personal diary and copied the drawings of WILLIAM BAaTRAM, and specifically about Pinckneya he wrote: "This very elegant flowering Tree or Shrub, I [W. BAR- TRAM] discovered in the maritime parts of Georgia & E[as]t Florida, above 20 years ago, when attending my Father John Bartram on Botanical researches; but that botanical excursion being in Autumn, We saw only the dry seed vessels, But about 15 years ago when on discoveries in the employ of Doctor Fothergill I revisited the same place, in the Spring Season, when I had the pleasure and satisfaction of seeing it in perfection, in full flower together with Franklinia [=Gordon& alata- maha!] which then flourish'd in sight of it." After a detailed description of this species WILLIAM BARTRAM concluded: "I know not to what Genus it belongs to; In its fructification is seems somewhat allied to the Yellow Jasmine But I believe it to be entirely new. I should be glad to know Mr. Walter's determination (I have sent a specimen of the flowers, a small part of a panicle). If it should prove to be a New Genus, I have a Request that it may be called BARTRAMIA (bracteata) in memory of my Father John Bartram deceas'd, The American Botanist & Natural- ist Whose labours, Travels, collections & Communications to the Curious in Europe hath contributed perhaps as much as that of any Man of his time toward increasing the Stores of Botanical knowledge with respect to North American production".

250 R G. DELPRETE:

The drawing which accompanied the above account is missing, but the drawing of W. BARTRAM for Dr. FOTHERGILL is reproduced in EWAN (1968: plate 8).

At the Natural History Museum of London (BM) has been found one herbari- um specimen (of which I have seen photocopies) that was written (handwriting of DRYANDER) on the verso "In Maritimis Georgia et Florida Orientalis. WM. BAR- TRAM". This specimen is one of those collected by W. BARTRAM and belongs to a series entitled "Georgia Plants for Dr. FOTHERGILL". I have selected this specimen as the lectotype of Bartramia (Bignonia) bracteata W. BARTRAM. Duplicates of BARTRAM'S collections were sought - but not found - at LINN, OXF, BM-SLOANE, P, P-Juss~Eu, P-LAMARCK, and P-DURAND, because their collections have been par- tially exchanged with these institutions (STAFLEU t~ COWAN 1976--1988).

POIRET (1804) described Cinchona caroliniana (= P. bracteata) based on mate- rial collected by L. A. G. Bosc in South Carolina. Bosc was a French consul in Carolina resident in Charleston. As gardener and amateur-naturalist, Bosc was also in charge of the MICHAUX garden in Charleston, and responsible for the ship- ment of M~CHAUX'S living plants and seeds to France (PoIRET 1808). In addition, Bosc went sporadically on botanical expeditions, one of them in South Carolina (1789-1800). Regarding the typification of C. caroliniana, one specimen has been found in the Paris Herbarium (P, ex Poiret Herbarium), which I examined, bearing the label "Cinchona Caroliniana -Bosc - Caroline". It seems to me an obvious decision to select this specimen as the lectotype of Cinchona caroliniana.

Proposed generic relationship: a brief history WILLIAM BARTRAM ( 179 l) in describing Bartramia (Bignonia) bracteata recognized that it belonged to the class "Pentandria monogynia," but he was obviously con- fused even to which family he should assign this new genus. ANDR£ MICI-IAUX in his diary, before deciding its name, simply described it as "a shrub related to Mus- sa[e]nda." POIRET (1804) placed it under Cinchona (Cinchoneae). In F-BA, L. C. M. RICHARD (in MICHAUX 1803) treated it as a genus related to Cinchona. DE JussIEU (1820) apparently related it to the genus Mussaenda (Mussaendeae) (probably because of its calycophylls), a genus less closely related than Cinchona.

In addition to a brief description (which differs from that of his father in FB-A), A. RICHARD (1830), observed that Pinckneya had been - erroneously - reunited to Mussaenda (Mussaendae) by D~ JUSS~EU (1820), but differed from it in having long filaments attached at the base of the corolla tube. ACHILLE RICHARD (1830) was the first to observe the resemblance of many of its characters to those of the closely related Macrocnemum corymbosum R. & R [= Condaminea corymbosa (R. & R) Dc., Condamineeae]. DE CANDOLLE (1830) placed Pinckneya (as Pinkneya) in the tribe Cinchoneae (as Cinchonaceae), subtribe Cinchoninae (as Cinchoneae), between Bouvardia and Calycophyllum, while placing Condaminea and Macro- cnemum (which included the not yet founded Pogonopus) in the tribe Hedyoti- deae, subtribe Rondeletiinae (as Rondeletieae).

KLOTZSCH (1853) founded Pogonopus, separating it from Macrocnemum, but without discussing its generic relationships. HOOKER (1873) established the tribe Condamineeae which he divided into three subtribes; in the subtribe Pinckneyinae (as Pinkneae), where he included Pinckneya and Pogonopus. BAILLON (1881)

Pinckneya bracteata 251

treated Pogonopus and Eupinckneya (= Pinckneya) as the single genus Pinc- kneya, which he placed between Condaminea, Rustia, and Rondeletia as part of Portlandieae (or Portlandia Series). SCHUMANN (1891) maintained Pinckneya next to Pogonopus with the Condamineeae, but without recognizing any subtribal divi- sion. VERDCOURT (1958) also did not recognize the validity of the three subtribes, and even included the Condamineeae under the tribe Rondeletieae (where he placed Pinckneya).

KIRKBRIDE (1985) described the monotypic genus Kerianthera, placing it within the Condamineeae, close to Pinckneya and Pogonopus because of its valvate corolla, capsule with many horizontal ovules, and presence of calycophylls. Keri- anthera differs from the other two genera in having 4 calyx lobes, 7-8 corolla lobes, very short filaments attached at the top of corolla (at the base of the corolla lobes), anthers with ca. 300 locelli, septicidal capsules, seeds with irregular fringed wing, and pentaporate pollen with smooth exine (vs. tricolporate with reticulate exine).

ROBBRECHT (1988) simply listed Pinckneya among the genera of the Condami- neeae but without recognizing any tribal subdivisions. BREMER & JANSEN (199 l), in their phylogenetic analysis based on cpDNA, mapped Pogonopus and Pinckneya next to each other, on the same clade with Calycophyllum (Calycophylleae) and Mussaenda (Isertieae).

I regard the genera Pinckneya and Pogonopus as sister genera, and morpholog- ically similar in having terminal frondose inflorescences, tubular flowers, valvate- reduplicate corolla aestivation with contact zone, long slender filaments attached near the base of the corolla tube, minute oblong anthers exserted at anthesis, one or few calyx lobes expanded into calycophylls, many-seeded capsules dehiscing loculicidally, and bark containing quinine-related alkaloids; Pinckneya differs from Pogonopus in having creamy-white corollas (vs. pink-red), entirely glabrous within (vs. a ring of hairs at base within), narrowly-triangular corolla lobes coiled outward at anthesis (vs. triangular-ovate and gradually expanded), much bigger capsules, and seeds with a broad orbicular wing (vs. minute non-winged seeds).

Taxonomy Pinckneya RICH. in MICHX., F1. bor.-am. 1: 103-105, t.13. 1803.

Type: Pinckneya pubens RICH. in MICHX. [= P. bracteata (W. BARTRAM) RAF.]. Selected references. BAILLON, H., Hist. P1.7: 334, 472. 1881. - BARTON, W. R

C., F1. N. Amer.: pl. 7. 1820. - BRITTON, N. L., N. Amer. Trees: 841-842. 1908. - Brown, C. L., KIRKMAN, L. K., Trees of Georgia and adjacent states: 247-248, pl. 78. 1990: - CANDOLLE, A. R DE, Prodr. 4: 366. Sept. 1830. - HOOKER, J. D., in BENT- HAM, G., HOOKER J. D., Gen. P1. 2: 47. 1873. - MICHAUX, A., F1. bor.-amer. 1: 103-105, t. 13. 1803. - MICHAUX, F. A., N. Amer. sylv. 3: 180-181, pl. 49. 1841-42. - RICHARD, A., M4m. Faro. Rubiace6s 197. Dec. 1830. - SARGENT, C. S., Silva 5: 107-109. pl 227-228. 1893. - SARGENT, C. S., Man. Trees: 875-877, pl. 771. 1922. - SCHUMANN, K., in EN~LER, A., PRANTL, K., Nat. Pflanzenfam. 4(4): 21. f6, M-O. 1891. - STANDLEY, R C., Rubiaceae in N. Amer. F1.32: 16. 1918. - TOR- REY, J., GRAY, A., F1. N. Amer. 2(2): 36-37. 1842.

Synonymy. = Pinknea PERS., horth, vat., Syn. P1.1: 197. 1805. =- Pinckneya

252 R G. DELPRETE:

RAF., horth, var., Ann. G6n. Sci. Phys. 6: 81. 1820. ----- Eupinckneya BAILL., Hist. P1. 7: 472. 1881. Not Pinckneya BAILL. (incl. Pogonopus), Hist. P1.7: 334, 472. 1881. Not Pinckneya ALLEMAO & SALDANHA (incl. Simira), Conf. Est. Bot. Dos Veg. Sec. 3: 7. 1865. = Mussaenda RAF., Sat. Eve. Post 6(303): [4]. fl., 19 May 1827. (reimpr. Casket 1827: 194. f 17. May 1827). = Bartramia W. BARTRAM, Travels xviii, 16. 1791. Not Bartramia L. (= Triumfetta, Tiliaceae). Not Bartramia SAHSm (= Pen- stemon, Scrophulariaceae). Not Bartramia HEDW. (Bryophyta). = Bignonia W. BARTRAM, Travels 468. 1791. Not Bignonia L. (Bignoniaceae).

Description. Shrubs to trees; commonly much branched shrubs. St ipules interpetiolar, free at base, small, narrowly-triangular and acuminate, caducous; stipules in apical buds with basal colleters that secrete a sticky yellow resin (exu- date). Leaves petiolate, blades ovate to elliptic, foliaceous; domatia absent (sometimes a tuft of pilose hairs). I n f lo re scences terminal and frondose, openly corymbose-paniculate, general shape shallowly ovoid, each lateral branch sub- tended by leaf-like bracts and terminating into a small cyme. F lowers protan- drous; hypanthium narrowly-obconical. Calyx reduced but with long-linear lobes, deciduous; calyx lobes 5, in some flowers 1-2(-3) calyx lobes expanded into foliose expansions (calycophylls). Ca lycophy l l s petiolate with blades ovate to elliptic. Corol la tubular and entirely glabrous inside, fleshy when fresh, cream- white; corolla tube cylindrical; corolla lobes 5, narrowly-triangular, coiled out- ward at anthesis; aestivation valvate-reduplicate with contact zone; glabrous and without basal callous zone inside. S tamens 5, exserted well above the corolla, fil- aments attached near the base of the corolla tube; anthers elliptic-oblong, dorsi- fixed at medial zone, dehiscing by longitudinal slit; filaments thin and glabrous, without a tuft of hairs at base. Style exserted well above the corolla; style branches ovate. Ovary two-celled, placentation axile, turbinate, glabrous, with many ovules in each locule horizontally inserted. Immature fruits green and semi- carnose. Mature capsules subglobose and bilobed, shallowly-convex at apex, with many white lenticels; dehiscing loculicidally, disk-septicidal dehiscence present in some old capsules. Seeds many and 2-ranked, horizontal, with a broad thin orbicular wing compressed. Po l len tricolporate, with reticulate exine.

1. Pinckneya bracteata (W. BARTRAM) RAy., Sat. Eve. Post 6(303): [4]. f 1. 19 May 1827. (Reimpr. Casket 1827: 194. f 17. May 1827.)

Basionym: Bartramia (Bignonia) bracteata W. BARTRAM. Synonymy. = Mussaenda bracteata (W. BARTRAM) RAF., Sat. Eve. Post 6(303):

[4]. fl., 19 May 1827. (Reimpr. Casket 1827: 194. f 17. May 1827.) =- Pinckneya bracteata (W. BARTRAM) ]7. HARPER, Bartonia 21: 7. 1942. ----- Bartramia bracteata W. BARTRAM, Travels xviii, 16. 1791 (and personal notes reproduced in EWAN 1968). --= Bignonia bracteata W. BARTRAM, TRAVELS 468. 1791. Not Bignonia brac- teata CHAM., in Linnaea 7: 692. 1832. [= Adenocalymna bracteatum]. Type: "In maritimis Georgia et Florida Orientalis," ca. 1770, W. BARTRAM s.n. (Lecto type here se lec ted , BM [photocopies examined], Bartram Herbarium).

Pinckneya pubens RICH. in M~cHx., F1. bor.-amer. 1: 103-105. Table 13. 1803. =- Pinknea pubescens PERs., horth, vat., Syn. P1.1: 197. 1805. Type: Georgia, "ad ripas fluvii Santae Marie," 1791, MICHAUX s.n., ( lec to type here se lec ted , PC!, Richard Herbarium).

Cinchona caroliniana POXR., In Lain. Encycl. 6: 40. 1804. Type: South

Pinckneya bracteata 253

Carolina, 1798-1800, Bosc. s.n. (Lecto type here se lec ted , PC!, Poiret Herbar- ium).

Description. Shrubs to t ree le ts 3-6 m tall (exceptionally trees 8 m tall and 20 cm at dbh); much branched and with spreading crown, main trunk terete; bark pale-brown. Young branchlets sparsely to densely pubescent, terete; older branches sparsely short pubescent, becoming glabrous, with cream-white lenticels, puncti- form to linear, up to 3.0 mm long. St ipules narrowly-triangular (or narrowly ovate); sparsely pilose (rarely glabrous) and minutely ciliate at margin, with elon- gated colleters (up to 0.5 mm long) at base inside; 5-10 mm long, 3-5 mm wide at base, caducous; the winter stipules thickened and scale-like, persistent through the winter until early spring. Leaves petiolate, petioles 5-30 mm long, 1-3 mm thick, adaxially concave to flattened, sparsely pubescent; blades (6-)8-19(-27) cm long, (3-)4-8(-13) cm wide, ovate to narrowly elliptic, cuneate to decurrent at base, acute and often short-acuminate at apex, acumen 5-10 mm long; dark green above and pale green below, stiff-foliaceous to thinly-coriaceaous; drying olive- green, stiff-chartaceous; sparsely-pilose above, sparsely to densely pilose below. Primary and secondary veins sparsely to densely pilose, secondary veins 5-8(-10) each side, arcuate-ascending and anastomosing 2-4 mm from the margin; tertiary veins starting subparallel and subreticulate in the middle. Domatia absent (rarely a tuft of pilose hairs). I n f lo re scences laxly paniculate with opposite decussate branches, each lateral branch subtended by leaf-like bracts and terminating in a small cyme; 4-12 cm long, 4-15 cm wide; basal branches 1.5-5.0 cm long, later- al branches 2-3 pairs. Rachis decussately compressed; rachis, branches, and branchlets densely pilose; flowers on distal branches in lax cymules of 6-25 flow- ers. Bracts subtending lateral branches usually leaf-like, petiolate, decreasing in size distally; bracteoles subtending the flowers 4-14 mm long, 0.5-1.5 mm wide, linear (often trifid), pilose. F lowers pedicellate, pedicel 1-5 mm long, densely adpressed to erect pubescent; hypanthium narrowly obconical, 5-7 mm long, 2-3 mm wide, adpressed to erect pubescent. Calyx reduced but with long-linear lobes; calyx tube 0.5-2.0 mm long, 2.5-3.5 mm wide, sparsely pilose; calyx lobes 5, linear, (6-)10-20 mm long, 0.5-4.0 mm long, pale-green to pinkish, caducous; in some flowers 1-2(-3) calyx lobes expanded into ca lycophy l l s . Calycophylls petioles 0.5-1.5 cm long, blades 3.5-7.5 cm long, (1.5-)2.5-5.0 cm wide, narrow- ly to broadly ovate to elliptic, base acute to obtuse, apex acute (with the very tip rounded), palmately veined, glabrous to sparsely pilose; primary and secondary veins sparsely pilose; cream-white, pink-white, pale-lavender, to pale-lilac. C oroll a tubular with short reversed lobes, 3-5 cm long, aestivation valvate-redu- plicate with contact zone, thick and fleshy when fresh, cream-white, yellow, greenish-yellow, pinkish, often with red or brown streaks inside; young flower buds minutely-pilose throughout (hairs up to 0.5 mm long); corolla tube cylindri- cal and gradually expanding, 2.0-3.5 mm long, 2.5-4.5 mm wide at base and 4-7 mm wide at orifice; densely sericeous outside, glabrous inside; corolla lobes 5, coiled outward at anthesis, 7-15 mm long, 2.5-4.5 mm wide at base, oblong- linear and acute at apex, densely sericeous outside, curled short-pubescent inside. S tamens 5, subequal, filaments attached 2-4 mm from the base of the tube, anthers narrowly-elliptic, 2-3 mm long, 0.5-0.6 mm wide, dorsifixed at medial zone, base rounded, apex acute, pale-purple when fresh and drying black; fila-

254 R G. DELPRETE:

ments 3.2-5.5 cm long, glabrous throughout, white. S ty le exserted, 3.8-6.1 cm long, glabrous, green; style branches broadly ovate, 0.5-0.8 mm long, 0.5 mm wide, reversed; stigmatic surface microscopically (40×) papillose. C a p s u l e s subglobose and bilobed, 1.0-2.5 cm long, 1.5-2.5 cm wide, with many punctiform white lenticels, dark brown, glabrescent; disk shallowly convex, with a white tubercle at the point of attachment of the style, glabrous, black and without lenti- cels, exceeding the calyx. Seeds (from STANDLEY 1918) pale brown, the body 2-3 mm long, surrounded by a thin orbicular wing 1-7 mm wide. C h r o m o s o m e number unknown.

Representative herbarium specimens examined. South Carolina. Beaufort : Pritchardville, 26 May 1960 (ft.), AHLES C~Z HAESLOOP 53498 (BR, FLAS, LL, NY, UC, TEX, US, USCH); Bluffton, MELLICHAMP s.n" 1883 ( (A, F, GH, U-S), 1884 (A, US), t886 (A, F, GH, LL, NY, US). Charleston: Michaux Garden Site (cultivated), N of Charleston Airport, 13 Apr. 1946 (old fr.), RAVENELS c~; al. 3574 (CLEMS). Darlington: Kalmia Gar- dens of Coker College (cultivated), 5 Aug. 1975 (st.), SAWYER 2624 (USCH). Jasper: Switzerland, just N of S-110, 29 Sept. 1983 (st.), AULBACH-SMITH 2889 (USCH).

Georgia. Appling: 1 km from Hwy GA-15, 16.8 km S of Baxley, 16 May 1970 (ft.), KISER s.n. (MO); 11.2 km N of Baxley, 19 Aug. 1967 (ft.), BLAKE & JONES 15114 (GH). Atkinson: 16 km S of Pearson, 3 May 1974 (ft.), RODGERS & al. 74100 (FLAS, MO). Bacon: 9.3 km W of Alma, 19 May 1953 (ft.), DUNCAN & HARDIN 16255 (GA). Ben Hill: 19.2 km NE of Fitzgerald, 22 May 1953 (ft.), WILBUR 3350 (FSU, GA); 2.7 km SE of Bowen's Mill, 9 Oct. 1952 (fr.), DUNCAN ~Z HARDIN 14350 (GA). Berrien: Hwy 82, 3 km W of Alapaha, 300 m S of hwy, 25 May 1995 (ft.), DELPRETE ~; SHARIF 6473 (FLAS, FSU, GA, GH, MO, NY, TEX, US). Brantley: 12 km E of Hortense, just E of Fendig, 1 June 1949 (ft.), DUNCAN 9620 (GA, MO). Brooks: 800 m NE of Barney, 24 May 1953 (ft.), WILBUR 3424 (FSU); 8.8 km NNW of Quitman, 1 July 1965 (fr.), FAIRCLOTH 1992 (GA). Bryan: Conochee River, 800 m N of Hwy 144, 16 km W of Richmond Hill, 25 May 1977 (fl.), PATTI 166 (CLEMS). Bulloch: rd to Statesboro, 1 km from Upper Lott's Creek Church, 10 June 1961 (ft.), BOOLE 1109 (FSU, GA, GH); Altamaha River, near Bloys, 10 June 1901 (ft.), HARPER 856 (A, [7, GH, MO, NY, US). Burke: Sugar Creek on Trinity- Cedar Park rd, ca. 12.8 kin SE of McRae, 20 May 1981 (ft.), AULBACH-SMITH 1469 (USCH). Calhoun: 6.4 km NE of Morgan, 24 May 1947 (ft.), TttORNE 4142 (GA, NY, US). Camden: 6.9 k m N of White Oak, 8 June 1931 (ft.), McKAY s.n. (GA). Candler: 3.2 km W of Metter, 25 Aug. 1940 (fr.), McVAuGH 5317 (A, UC). Charlton: W side of Right of Way on Hwy 1,800 m N of Okefenokee Swamp Park entrance, 27 May 1961 (ft.), CYPERT 410 (GA). Chatham: Savannah, date and collector unknown (NY). Clinch: Hwy 441 (Hwy 89), 18 km S of Homerville, 27 May 1995 (ft.), DELPRETE • SHARIF 6477 (FLAS, FSU, GA, GH, MO, NY, TEX, US). Coffee: W of Douglas, 28 Aug. 1941 (fr.), DUNCAN 3982 (GA). Colquitt: Okalpico Creek, 18 km N of Moultrie, 12 July 1966 (old ft.), FAIRCLOTH 3429 (MO). Cook: Hutchinson Mill Creek, 2.3 km NE of Cecil, 2 July 1965 (ft.-ft.), FAIRCLOTH 2050 (GA). Decatur: Willacoochee Creek, 4.8 km NE of Face- ville, 27 July 1946 (fr.), DUNCAN 6709 (GA). Echols: 11.7 km NE of Stateville, 30 June 1953 (st.), DUNCAN & QUARTERMAN 16707 (GA). Effingham: 400 m from Chatham county line, 1958 (ft.), MELLINGER s.n. (GA). Emanuel: ca. 17.6 km W of Metier, 16 May 1953 (fl.), WILBUR 3303 (FSU, TEX); SE of Summertown, 20 Nov. 1964 (st.), HENRY 7295 (GA). Evans: Road 209, 2.5 km W of Jct with Road 129, 26 May 1995 (ft.), DELPRETE & SHARIF 6474 (FLAS, FSU, GA, GH, MO, NY, TEX[2], US), Road 209, 2.3 km E of Jct with Road 129, 26 May 1995 (fl), DELPRETE & SHARIF 6475 (FLAS, FSU, GA, GH, MO, NY, TEX[2], US). Grady: Birdsong Nature Center, 8 km SW of Beachton (1 km N of Florida State line), 23 July 1993 (st.), ANDERSON 14502 (FSU). Irwin: 8 km N of Irwinville, 23 May 1953 (ft.), WILBUR 3352 (FSU). Jeff Davis: 5 km W of Snipesville, S of Hwy 107, 28

Pinckneya bracteata 255

Fig. 3. Inflorescence, flowers, and habit of Pinckne.ya bracteata, a inflorescence with flo- wers and calycophylls; b top view of the corolla, showing red streaks, and exerted anthers; c side view of one flower showing calyx lobes, coiled corolla lobes, and exserted anthers; d general habit

256 R G. DELPRETE:

June 1985 (fl.-fi'.), CAaTER & CARTER 4224 (FLAS, GA). Lanier: 800 m W of Stockton, 21 Sept. 1940 (fr.), DUNCAN 3025 (CA). Liberty: Altamaha River, 18-21 June 1895 (fl.), SMALL s.n. (A, F, NY). Long: Fort Stewart Military Reservation, Training Area 15, Glenn- ville NE Quad, 3 June 1992 (fl.), ZEBRYK & al. 337 (GA). Lowndes: Naylor, 22 June 1936 (fl.), CORREI+ 5543 (GA, GH[2]); Withlocoochee River, near Valdosta, 6-12 June 1895 (fl.), SMALL s.n. (A, F, NY). McIntosh: Altamaha River, near Ft. Barrigton, 13 June 1940 (ft.), EYLES 7303 (FLAS, FSU, MO[2], NY[2], TEX, UC[2]). Montgomery: NW of Uvalda, 320 m SE of GA-130, 29 May 1988 (fl.), Wood & BOUFVORD 4870 (GH, NY). Pierce: Big Satilla River, 11 Sept. 1966 (old ft.), DUNCAN 22981 (CA). Randolph: Cuth- bert, 10 May 1938 (fl.), GooLsBY s.n. (GA, USCH). Screven: Cameron, 18 May 1928 (fl.), READE 1622 (GA). Sumter: Kinchafoonee Creek, 22.4 km W of Sumter, 11 Aug. 1965 (old fr.), PARKER 83 (GA); Americus, 300 In, 29 Aug. 1900 (fr.), HARPER 538 (NY, US). Tattnall: along Hwy 301, 5.6 km N of Glennville, 30 June 1959 (fl.-fr.), DRESS & READ 7792 (BR). Telfair: 3.2 km N of Lumber City, 12 April 1940 (old ft.), DUNCAN 1937 (GA, UC). Terrell: N of Dawson on Haines rd, 30 May 1973 (ft.), FITZGERALD 677 (GA). Thomas: 4.8 km E of Ochlocknee, 23 May 1953 (fl.), WILBUR 3410 (FSU, CA) and 3412 (FSU). Tift: Hwy 82, 3.2 km E of Tifton, 25 May 1995 (fl.), DELPRETE & SHARIF 6472 (FLAS, FSU, CA, GH, MO, NY, TEX[2], US). Toombs: 4.8 km E of Vidalia on Hwy US-280, 18 May 1976 (fl.), SALOMON 2062 (MO). Turner: Pocosin between Wolf and Deep Creeks, 6.5 km ENE of Ashburn, 6 May 1967 (ft.), FA~RCLOTH 4394 (GA, MO). Ware: near Waycross, 7 June 1950 (ft.), GODFREY 50417 (FSU, GA, NY). Wayne: Hwy 301 (Hwy 23), just N of McKinnon, 26 May 1995 (ft.), DELPRETE & SnARIF 6476 (FLAS, FSU, GA, GH, MO, NY, TEX, US). Wilcox: 17.6 km SW of Abbeville, 17 May 1953 (fl.), DUNCAN & HARDIN 16111 (GA). Worth: 12.8 km E of Sylvester, 23 May 1953 (fl.), WmBUR 3394 (FSU, GA). Unknown County: Ogeechee River, August 1881 (ft.), CuRxISS 1131 (F, G, GH, K, MO, NY, UC, US[3]).

Florida. Bay: Little Bear Creek, jct Hwy 388 and Hwy 231, Youngstown, 24 April 1972 (st.), MILLER s.n. (GA). Bristol: 1 June 1941 (ft.), STEARNS 17 (GA). Calhoun: 2.9 km N of Blountstwon on rd 71, 28 May 1973 (ft.), BOUFmRD & AnLES 9525 (A, MO); Clarksville, 22 April 1949 (F1.), Hood 2014 (FLAS). Clay: 4.8 km S of Middleburg, 21 May 1940 (fl.), MERRILL s.n. (CA, MO[2]); 4.8 km W of Green Cove Springs, 15 June 1939 (fl.), MURRILL 313 (MO, US). Franklin: Apalachicola National Park, 1 May 1964 (fl.), DEMAREE 50295 (A); Rte 65, jct of entrance to Wright Lake, just S of Sumatra, 17 May 1980 (ft.), FOLSOM & MAUSETH 7762 (TEX). Gadsden: near Quincy, 5 June 1987 (fl.), CURTISS 587 (F, FLAS, FSU, G, CA, GH, K, NY[2], UC, US); Wallwood Scout Reserva- tion, between Council Fire and Chapel, 10 July 1976 (st.), ANDERSON 4315 (FSU). Gulf: 6.4 km W of Wewahitchka, 3 Sept. 1958 (fr.), GODFREY 57596 (FSU). Jackson: Marian- na, 10 June 1936 (ft.), KNIGHT s.n. (FLAS, GA). Jefferson: 8 km W of Monticello, on US-90, 10 June 1961 (ft.), SMITH & MYNT 263 (FLAS, FSU, GH). Leon: 24 km W of Tal- lahassee, 26 May 1956 (ft.), GODFREY & KURZ 54866 (FLAS, FSU, GH); Greensboro, 25 July 1956 (fl.-fr.), GODEREY 54987 (FSU); Apalachicola National Forest, forest rd 304, 16 May 1990 (ft.), GODFREY 83858 (A, NY). Liberty: 11.2 km N of Sumatra, rte 65, 19 May 1981 (fl.), GODFREY 78686 (FSU). Marion: Orange Springs, just S of Orange Creek, rt 315, 4 Sept. 1980 (old fr.), JUDD & SIMONS s.n. (FLAS). Putnam: 11 km N of Palatka, 4 June 1940 (fl.), WATSON & al. 221 (FLAS). Wakulla: Apalachicola, 18 May 1892 (fl.), CHAPMAN 4351 a (A, MO, NY, US). Was higto n: 3 km W of rte 77 and Greenhead commu- nity, 26 May 1990 (fl.), ANDERSON 12836 (FSU). Unknown County: Rock Bluff, 24 May 1930 (ft.), BLANTON 6583 (F, GH, MO, US); Florida reed., Ocolockne River, June 1845 (ft.), RUGEL 316 (F, MO, NY, US); "Florida", CHAPMAN s.n. (CA, GH, K[2], US); "swamp", 1-2 June 1941 (fl.), STEARN 17 (US).

Pinckneya bracteata 257

M I L E S

0 I 0 0 2 0 0 3 0 0 4 0 0 I , I ~ I ~ I , I , I , I J I , I

9 0 ° 85 ° ~ / ¢ 8 0 o

Fig. 4. Distribution of Pinckneya bracteata

25o :,;~ -"

Notes on distribution, phenology, reproductive biology and uses

Distribution (Fig. 4). Swamp forests, edges of boggy areas, moist bays, bay- heads, bayswamps adjacent to sandridges, poorly drained mucky soils, springy areas, along small rivers, drainage areas in low pine woods, pitcher plant bogs, and moist pine-cypress savannas, of South Carolina, Georgia, and northern Flori- da. Pinckneya bracteata is commonly found in association with Gordonia lasian- thus (L.) ELL. [described to be growing with G. alatamaha (BARTR. ex MARSH) SARa. by W. BARTRAM 1791], Viburnum nudum L., Rhus vernix L., Myrica hetero- phylla RAF., Cyrilla sp., Nyssa biflora WALT., Sorbus sp., Cliftonia monophylla (LAM.) BRITT. ex SARG., Magnolia virginiana L., Liriodendron tulipifera L., Pinus taeda L., Persea palustris (RAF.) SARG., Quercus sp., and Acer rubrum L.

Phenology (Fig. 5). Specimens with flower buds were collected in the last week of April. Flowering period extends from the first week of May to the last week of June with average flowering peak ranging from May 16 to June 10 (see Fig. 5). Specimens in late-blooming/early-fruiting were collected in the last two weeks of June and first week of July. Specimens with mature capsules (and with mature seeds) were collected from the second week of July to the third week of

258

" / 5 -

P.G. DELPRETE:

U~

o . u

o 0

20

15

10

Anthesis

In te rmedia te

I I F r u i t i n g

stage

E - I

Z

innlh m - ~, o T , F , - -

= N N N

May June July August September

Fig. 5. Diagram showing anthesis and fruiting stage of Pinckneya bracteata determined from label data of 150 specimens collected from the year 1800 to present

September. The old capsules tend to stay attached on old branches up to the fol- lowing year, but without containing any viable seeds.

Reproductive biology and herbivory. Anthesis of any one flower of P. brac- teata regularly last for three days. The first one-two days the anthers open and release pollen. Usually by the third day the anthers are dried-up and no longer functional, while the stigma is protruded well above the corolla and starts to be receptive (pers. obs.). The flowers of P. bracteata are intensely visited by adults of Phoebis sennae L. (Cloudless sulphur, fam. Pieridae, subfam. Coliadinae) during late mornings and early afternoons (pers. obs.). Adult individuals of P. sennae visited the flowers, evidently feeding on nectar while performing a pumping motion, and I have personally observed their proboscis and mouth parts to be charged with pollen following such activity.

I have also observed the foliage to be eaten by caterpillars of Epargyreus cla- rus CRAM. (Silver-spotted skipper, ram. Hesperiidae). In some leaves of P. brac- teata the caterpillars of E. clarus have been observed to produce a silk mat into which they will later transform into a chrysalis.

Conservation biology. Taking into account the relatively few recent collec- tions of this species, P. bracteata is certainly a threatened species in South Caroli- na. In southern Georgia and northern Florida several relatively recent collections show that this species is still relatively frequent in this area. Nevertheless, human

Pinckneya bracteata 259

pressures (due to modern agricultural and industrial development) on swampy habitats in this region likely account for its increasing rarity.

Common names. Pinckneya (US), Georgia Bark (US), Fever Tree (Georgia), Maiden Blushes (Florida), Georgia Fever Bark (US), Georgia-Rinde (Germany), Ecore de Georgie (France), Quinquina de Caroline (France, POIRET 1804). RAFI- NESQ~'E (1827a, b) reports the following names: Pinckney Bark (English), and Quinquina Pinckney (French).

Medicinal and economic uses. This species was used with considerable suc- cess during the Civil War (and in more recent times) for the treatment of malaria and intermittent fevers, apparently due to the content of quinine in its bark (hence some of the common names). In North American sylva F. A. MICHAUX (1841--42) reports: ".. . its inner bark is extremely bitter, and appears to partake of the febri- fuge virtues of Cinchona, for the inhabitants of the southern parts of Georgia employ its successfully in the intermittent fevers which, during the latter part of summer and the autumn, prevail in the United States. A handful of the bark if boiled in a quart of water till the liquid is reduced one-half, and the infusion is administer[ed] to the sick."

Pinckneya bracteata has been sporadically used as an ornamental in gardens of the southeastern United States (and rarely European botanical gardens). Because of its beauty and increasing scarcity in the natural environment, I strongly encour- age local nurseries to implement its cultivation into private and public gardens.

Pinckneya bracteata is one of the most beautiful shrubs native of the United States, with its inflorescences crowned by creamy, delicately fragrant flowers, sur- rounded by white to deep-pink calycophylls (Fig. 3 a), generally appearing like a scattered arrangement of elegant bouquets (Fig. 3 d). At a closer look, its corolla throats and lobes display numerous and random red streaks (Fig. 3 b), adorned with slender white filaments topped by pale-purple anthers. Its habitat is common- ly swamp areas and poorly drained soils; for this reason, when introduced to cul- tivation, this shrub should be planted in relatively humid areas or else watered rather frequently. Under natural conditions (as pointed out above), it is becoming an increasingly rare species. Label data on some of the old collections of P. brac- teata report this species to be a nearly dominant shrub in some small areas, a con- dition which is sadly impossible to find at the present time.

Pinckneya bracteata is usually encountered as a shrub 3-5 m tall, but upon occasions can grow into a medium sized tree. Label data on one herbarium speci- men (STEARN 17, US) noted the plant to have been collected in a Florida swamp (precise locality unknown) from a tree 7.5 m tall with a main trunk 7.5 cm dbh. Attached to the sheet was included the wood sample from one of its basal branches. Counting the annual rings of this wood sample, I estimated its age as about 30 years.

This study was based on field work in Georgia and examination of 446 herbarium sheets from the following institutions (number of sheets shown in parenthesis): A(24), B(1), BR(4), CLEMS(4), F(19), FLAS(34), FSU(51), O(12), 0A(73), 0H(34), K(5), LL(5), M(5), MO(33), NY(48), P(22), TEX(16), UC(18), US(47), USCH(11). While only representative specimens are cited in this study, the distribution map (Fig. 4) was con- structed from all of the sheets concerned. I am grateful to the directors and staff of each

260 R G. DELPRETE:

for the loan of the material, and to the many colleagues that collaborated in clarifying the complex taxonomic history and lectotypification of this taxon: to Dr ELMER ROBBRECHT (BR) for sending xeroxed copies of RAFINESQUE'S publication (Ann. G6n. Sci. Phys. 6: 81. 1820); to NANCY ELDER of the Life Science Library (TEX) for finding RAFINESQUE'S remote publications both in Casket (1827) and in the Saturday Evening Post (1827); to JAN BAR- BER and Dr SCHUYLER (PH) for searching for RAFINESQUE'S types in their institution and sup- plying additional literature on RAF1NESQUE; to DOUG GOLDMAN (TEX) and Dr C. E. JARVlS for finding W. BARTRAM'S collection of Bartramia (Bignonia) bracteata preserved at the Brit- ish Museum (BM); to Miss S. K. MARNER (OXF) for looking for BARTRAM'S types in the Sherardian and Dillenian Collection in the Oxford Herbaria (OXF); to Mister S. BARRIER (P) for looking for BARTRAM'S types in the JussiEu, LAMARCK, and DURAND Herbaria (P). - Special thanks should go to Dr C. E. JARVIS (BM) for type searches at LINN, BM-Sloane and BM general herbarium, for addition information regarding the BARTRAM'S collection of Bartramia bracteata (selected as lectotype) present at BM, for sending copies of EWAN'S reproduction of the Botanical and Zoological Drawings of WILLIAM BARTRAM, and for his skilled and friendly collaboration. I am finally grateful to BRIAN BOOM (NY), PAUL FRYXELL (TEX), and BILLIE TURNER (TEX) for reviewing the manuscript and for their helpful com- ments.

References

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P i n c k n e y a b r a c t e a t a 261

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Address of the author: PIERO G. DELPRETE, Department of Botany, University of Texas, TX 78713, USA.

Accepted October 25, 1995 by B. L. TURNER