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Home , Chamaerops, Sabal, Sabal minor, Sabal palmetto

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l e95l RAMPAND THIEN: 77

Principes,39(2),1995, pp. 77-83 A TaxonomicHistory and Reexaminationof Sabalminor in the MississippiValley

Peur F. Rllrp'eNo LuonenpB. THrrN Departrnentof Cell and MolecularBiol.ogy, Tulane Uniuersity, New Orleans,LA 70118

". Sabal displaysa high degreeof variability, par- wasdescribed as having a trunk that . . scarcely ticularly in characters commonly used to delimit emergesfrom the ground. . . ,'o (petioles?) ". species(Bailey 1944, Uhl and Dransfield1987, that lack spines or teeth and are . . folded in Zona 1989). In particular, the presenceor absence a fan." The commonname for this palm was given "latanier". of a trunk (height) along with spines, and as The description fits bladefolding causedtaxonomic confusionfor early and certainly not repens which, although explorers and biologistsin . Although essentiallywithout an upright trunk and occurring S. minor (5. louisiana) has receivedgreat atten- in southeasternLouisiana, possessesteeth on the tion for its stem variability, other North American petioles. The second specieswas not named and palmettos also exhibit great trunk variability received only a short statement concerning the including S. etonia, S. palrnetto and. Serenoa leaves,which were describedas divided and folded, ". repens(Hilmon 1969, Bailey 1944,BrownL973, . . somewhat like an old-fashioned cravat or Zona and.Judd 1986). This paper reviews the collar." No mention was made of the presenceor early observationsand taxonomic history of Sabal absenceof a trunk. The description of the leaves "trunked louisiana (the palm") and utilizes pro- fits the manner in which leaves break in some tein electrophoresisto characterizeits affinity with populationsof Sabal minor (i.e., Frenier Beach, S. minor (N. J. Jacquin)Persoon. Fig. lC) in which the palms do possesstrunks. A decade later, William Darby published an "A Historyol S. minor (5. louisiana) account of his exploration of Louisiana titled geographicaldescription of the state of Louisiana of the earliest European accounts New One of . . . being an accompanimentto the map of Lou- (1691) World palmswas by Le Clercq who pub- isiana." Appearing in 1816 (secondedition in lished an account of the explorations of the Mis- o'the l8l7), this book givesa descriptionof the habitat sissippiby La Salle.He stated, whole country of a palm he named Charnaeropslnuisiana.Darby is covered with palms." In light of the present stated ". . . there is a specific difference between distribution of palms, the species observed was the palmetto l:5o6ot palmetto] S. minor. The earliest taxonomic probably treat- . . . and that of Louisiana. The Chamaerops ser- ment of the palm was apparently by Adanson rulata of Muhlenberg l:Serenoa repensl is cer- (1763) who established the Sabal presum- tainly not the same as the palmetto of Louisiana; ably in referenceIo S. minor without commenting the latter bears a much greater resemblanceto on the name'sderivation (Guernsent 1804). cabbagetree, though much more humble in ele- The earliest referenceto a speciesdistinct from vation, than to the saw-leavedpalmetto of Geor- minor in the MississippiValley is attributed Sabal gia." This statement may suggestthat some form Robin (1807). He describedtwo species.The to of trunk was present in the palms he observed. which he placed in the genus Charnaerops, first, Obviously he was familiar with the speciesoccur- ring on the east coast, so it is surprising that no comparison of this palm ,was made with Sabal I Present address:The General Biology Program, 301 Hes- minor. The habitat he described for this species ler Biology Building, The University of Temessee, Knoxvi.lle, "where Tennessee37996. is between the land sinks too low for the 78 PRINCIPES [Vor. 39

"dwarf l. Variability in the stature oI Sabal minor. A. A individual from a population near Blountstown, . B. Typical acaulescent individuals in a population near New Orleans, Louisiana. C. A caulescent individual with leaves folded in a manner fitting the description made by Robin (1807). D. Caulescent individuals growing along the Mississippi River below Venice, La. The exposed trunk of the center individual is approximately 2.4 meters in height. I

l 99sl RAMP AND THIEN: SABAL 79

Arundo. . . (and). . . wheretheinundationexceeds resented by the Chamaerops palrnetto) that the in depth 15 to 20 inches." Trees associatedwith Palmetto attains a growth as gorgeouseven as in the palm were also given. These include Quercus the lower Mississippi." The palm which he observed phell.os Linnaeus, Q. rubra Linnaeus, Liquid- would have been either Sabal texana or S. mex- anb a r st y r ac if. u a Linnaeus and Ce h i s c r a ss ifo li a icana. The confusion stemsfrom his reference to "gorgeous" (C. laeaigata Willd.), speciescommonly found in a palmetto in the MississippiValley. "gorgeous" bottomland hardwood forests. If is interpreted to mean as having a During the sameyear, Rafinesque(1817) pub- large stature, as Small (L926) interpreted this lished a flora of Louisiana based on the earlier statement,then would be the most work of Robin, althoughhe never visited the region obvious comparison. However, S. palmetto doq himself (Fitzpatrick l9I I). Rafinesque assigned not occur within 300 miles of the area (the western binomials to Robin's palms. He called the first edge of its range is in the Apalachicola Valley). Sabal adansonii (:5. rninor) and the secondoa His reference may have been to stands of cau- new species,S.? adiantinurn Raf. The description lescentpalms (Sobal minor) now only represented of the latter speciesappears to be an embellish- by one known stand below Venice on the Missis- ment of Robin's description.Rafinesque stated that sippi River (Fig. lD). the new specieswas acaulescentalthough Robin The next known referenceto tnrnked pahnettos made no mention of the 's stature. Ewan in Louisiana is by Featherman in 1870. In a (1967) suggestedthat S.?adiantinurn is Serenoa discussionof the vegetation of Grand Isle, he stated "tree repens despite the specific reference Rafinesque that palmettos are seen here (behind the made to the unarmed petioles. In the same pub- beach) and there near the beach." lication, Rafinesquetook issue with Darby's spe- In 1926, J. K. Small wasin a positionto address cies Charnaerops louisiana and considered this the problem of trunked palmettos in Louisiana. to be the same as Sabal adansonii. It is unfor- He observeda population of trunked palms south tunate that Darby failed to give fuller description of Point aux Herbes (on the southeasternend of o{ the palm other than stating that it differed from Lake Pontchartrain) which were initially taken for 'omuch Serenoa repens and was more humble in Sabal palmetto. However, on closer examination elevationo' than Sabal pahnetto. It was not pos- he discovered the palm to resemble S. minor. sible for Rafinesquenor is it possiblenow to assign Small named this palm Sabal deeringiana. Darby's palm without conjecture. Included in his description of the species were Over the next 40 years there appearedseveral characters that he consideredsignificant in distin- referencesto palmsfound in the MississippiValley. guishing it from S. rninor; it is a tree to 4 meters "The In Genera of North American . . . ," with a trunk of I to 2 meters high with petioles Nuttall (1818) placed.Sabal adansonii in Loui- longer than the leaf blades,the leaf midrib extends ". siana stating that it is in . . troublesomeabun- high into the flat blade and the leaves are fila- dance around New Orleans;but lessfrequent than mentous. He also indicated that the petals are other speciesin Georgiaand Carolina." The other slightly broader and longer, the stamensare lon- speciesrefened to were S. palmetto and.Serenoa ger, and that the is more depressedthan in repens. Darby (1818) maintained the use of Cha- S. minor. The type specimenis depositedat The "The rna,erops louisiana in Emigrant's Guide, New York Botanical Garden. etc." Flint (1828) used the species name Cha- In 1929 Small made further comparisonsof S. maerops latanier, a name which must have been deeringiana with other species of Sabal. The derived from Robin's or Darby's work. Martius habitat of this palm was said to be bottomless (1838) describedthe range of Sabal adansonii gumbo, and he stated that in Louisiana, S. deer- to include the MississippiValley as well as ingiana was much more restricted and less com- and Carolina. mon than S. tninor. In this publicdtion, Small Shortly after Martius (1838) another statement provided photographsof two individualsof S. deer- was put forward which added to the somewhat ingiana at Frenier Beach on Lake Pontchartrain. confused status of the palms of the Mississippi In Small's discussion of this new species he Valley. Arthur Schott (1857), in a report on the referred to Schott's (1B57) referenceto a palm "gorgeous" -Mexican border and the lower 80 miles of of growth in the MississippiValley but "It the Rio Grande, stated is also in the lower was apparently unawareofthe earlier observations portion of this belt (where the palm tribe is rep- of trunked palmettosby Robin (1807) and Darby PRINCIPES lVoL. 39

(1816). Had he known of these,he surely would trunked individuals with a stem height of approx- have noted the similarity of these descriptions. imately I - I .5 meters (total height near 4-5 The similarity can be seen by comparing Small's meters).The tallest individualshave tnrnks greater photographs of S. deeringiana with Robin's than three meters (total plant height approxi- descriptionof the manner in which the leavessplit mately six meters, Fig. 1D). The largest individual and break over (Fig. lC). known, tentatively identified as S. rninor, has a Bomhard (1935) reviewed these accounts of l9-foot trunk (5.8 meters) with a total height of trunked palmettosand consideredDarby's specific 27 feet is located in Brazoria County, Texas (Lan- epithet to have priority, thus creating the binomial don Lockett, personal communication). Sabal louisiana .(Darby) Bomhard. Bomhard (1937,1940, 1943) providedphotographs and a Gel ElectrophoresisMethods map of population sites. She extended the range of the speciesto easternTexas acrossto the west- Plant material was collected from six taxa rep- ern portion of the Florida panhandleand north to resenting three species in two genera. Serenoa southern . She stated that S. louisiana repens (Bartram) Small was collected from a pop- is stemlessunder someconditions while producing ulation near Orlando, Florida and'Sabal palmetto an abovegroundtrunk under others. Unfavorable (Walt.) Loddigesex J. A. & J. H. Schulteswas conditionsthat contribute to retarded trunk devel- collected near Crystal River, Florida. Four pop- opment include exposureto brackish water, direct ulationsof Sabal minor wereexamined. An acau- sunlight, and the deposition or removal of soil lescent population samplednear Savannah,Geor- "dwarf around the plant's base (Bomhard 1943). Bom- gia (SVGA), a population of individuals hard suggestedthat the largest individuals she near Blountstown, Florida (BTFL)' a caulescent observedwere in excessof 200 years old. population at Frenier Beach, Louisiana (FBLA), In his first monograph of the genus, Bailey one of the sites of Sabal louisiana describedby (1934) recognizedS. louisiana; however in his Bailey (1944) and Bomhard (1943), and a cau- second monggraph (1944), he combined Sabal lescent population of Sabal m.inor locared below minor and Sabal louisiana. He consideredthe VeniceoLouisiana actually growing in the Missis- latter to be the final, emergent stageof the former sippi River (MRLA). This population previously "no stating that clear definite . . . differencesare had been found to have the greatestmean genetic recorded betweenthe acaulescentand caulescent distance(Nei's index, I972) {rom I3 populations phases . . ." and that if these are two species, distributed from Georgiato Texas (Ramp, 1989). characters other than stature need to be found. Leaf portions were cut into small pieces then Since this tlme, Sabal lttuisiana has been.consid- ground with mortar and pestlena7 .5 pH grinding ered a synonym of S. minor (except by Bomhard buffer composedof 0.I M Tris, 1.0 mM EDTA, 1950 and Vines 1960 who maintainedit as a I0 mM KCl, l0 mM MgCl, 14 mM 2-mercapto- distinct species).In a recent monograph of the ethanol, and57o polyvinylpyrrolidone- 0 (Gottlieb genus, Zona did not separate S. louisiana ftom l98lb). This wasabsorbed onto 3 x l5 mm paper S. minor (Zona 1990). wicks (Beckman) and placed tnto I2'97o star.ch Acaulescent populations of S. minor occur gels. throughout the species'range while trunked palms A lithium hydroxide buffer system (Soltis et al. occur mostly in the western portion of the range 1983) wasused to examineseven enzyme systems; (specifically in LA and TX, Figs. lC and lD). alcohol dehydrogenase(ADH), leucine aminopep- Typical, acaulescentpopulations are composedof tidase (LAP), glutamate-oxaloacetatetransami- individuals with leaves reaching 1.5 to 2 meters nase (GOT), peroxidase (PER), superoxide dis- in height (Fig. lB). A populationof unusually small mutase (SOD), phosphoglucosisomerase(PGI) and individualswas observed on the ApalachicolaRiver malic enzyme (ME). Staining proceduresfollowed flood plain near Blountstown' Florida. Here, the Soltiset al. (f 983). largest reproductively mature individuals scarcely The unbiasedestimate of mean heterozygosity reached 40 cm in height (Fig lA). A similar pop- per locus (H) for each taxa and the unbiased "dwarf ulation of individuals was described by genetic distances were calculated (Nei 1978). Bailey (1944) from Angelian Countyo Texas. Although sample sizes are small for Sabal pal- : In contrast, it is not uncommon in the western metto (n : 5) and Serenoa'rePetus, (n l0) half of the species' range to find populations of empirical studies have found that samplesof 8- I 99sl RAMPAND THIEN:SABAL 8l

Table 1. Allozymes obseraedfor three Ameri- indicate that for interspecific comparisons,a sam- can palmettos. Sabal minor populations are giu- ple size of one is sufficient to estimate genetic en Ln text. distance within 0.I in over 9O% of the cases studied and that the error is to systematically Sabal Sabal minor overestimate D (Nei 1978, Gorman and Renzi Serenoa pal- 1979). Both heterozygosityand genetic distance repens metto SVGA BTFL FBLA MRLA estimatesare much more severelyaffected by the PGI.I r.0 1.0 L0 ' number of loci sampled.Gorman and Renzi(1979) I.0 found that a sample of two individuals scored for 0.50 0.0 0.0 0.0 0.0 PGI-2a (23) gave a better estimate gf b 0.50 0.0 0.0 0.0 0.0 all loci studied c 0.0 0.07 0.20 0.89 0.50 heterozygositythan all individuals(20-4 I per spe- .d0.0 0.93 0.80 0.1l 0.50 cies) scored for a subset(up to 20) of the loci in PER-I r.0 eight speciesof Anolis. PER-2a 0.0 1.0 t.0 1.0 I.0 1.0 b I.0 0.0 0.0 0.0 0.0 0.0 LAP-la I.0 0.0 0.0 0.0 0.0 0.0 Resultsand Discussion b 0.0 I.0 1.0 0.0 I.0 1.0 LAP-2a I.0 0.0 0.0 0.0 0.0 0.0 Thirteen loci were resolved (Table I ). Mean b 0.0 1.0 1.0 t.0 I.0 I.0 heterozygosityvalues for these loci (on the diag- GOT-la 1.0 0.0 0.0 0.0 0.0 0.0 onal of Table 2) were similar for all taxa ranging b 0.0 1.0 0.0 0.0 0.0 0.0 from H : 0.081 in BTFL minor) to H : c 0.0 0.0 0.83 0.73 0.53 0.50 {Sabal d 0.0 0.0 0.17 0.27 0.47 0.50 0.000 in Sabal palmetto. Mean heterozygosities ME-la 1.0 0.0 I.0 1.0 1.0 I.0 for these palms are low compared to the average b 0.0 1.0 0.0 0.0 0.0 0.0 of H : 0.156 found in plants (Hamrick et al. SOD-l:i 1.0 0.0 0.0 0.0 0.0 0.0 1979) but are similar to those calculated for b o.0 1.0 1.0 1.0 I.0 : Washingtonia filifera, H 0.009 (McClenaghan soD-2 1.0 1.0 I-0 1.0 1.0 t.0 and Beauchamp1986). When values for these soD-3 1.0 I.0 1.0 I.0 I.0 palms are compared with other long-lived peren- SOD- a 0.0 1.0 1.0 1.0 t.0 nials (averageheterozygosity, H : 0.27), the dif- I.0 0.0 0.0 0.0 0.0 b ference is striking. It would be interesting to know 0.0 0.98 0.85 0.41 0.33 ADH-2a if low heterozygosityis characteristic of palms in b-I.0 0.02 0.r5 0.89 0.67 general or if the low heterozygosityobserved here is the result of similar evolutionary histories(e.g., populationsundergoing similar bottlenecks). 12 individuals usually give heterozygosity esti- The smallest genetic distances were between mates within l% of estimates based on much larger the four populations of Sabal rninor, ranging samples and that samples even as small as two between0.011 and 0.I32. Geneticdistances and individuals generally give estimates withn 2.57o identity are given in Table 2 and a phenogram of (Gorman and Renzi 1979). There is even a smaller genetic identity in Figure 2. The high genetic effect of sample size on genetic distance. Results identity within the populations SVGA, BTFL,

Table 2. Nei's (1978) unbiasedestimates of genetic identity (abouediagonal), genetic d'istance (below d.iagonal), and. heterozygosities(on the rnajor diagonal).

Sabal minor Serenoa Sabal rePens palnetto SVGA FBLA MRLA

Serenoa repens (0.043) o.372 0.3?6 0.388 0.386 0.381 Sabal palmetto 0.989 (0.000) 0.578 0.6I9 0.725 0.684 Sabal minor (SYGA,) 0.978 0.548 (0.038) 0.989 0.9r0 0.876 Sabal minor (BTFL) o.947 0.480 0.0II (0.08r) 0.924 o.924 Sabal m.inor (FBLA) 0.952 o.322 0.094 0.079 (0.076) 0.980 Sabal minor (MRLA) 0.965 0.380 0.I32 o.o79 0.020 (0.07s) PRINCIPES lVoL. 39

FBLA Ltrenerunr CIrBo

MBLA ADANSoN,M. 1763. Familles des Plantes. Vincent, Paris. Sabal BAILEY, L. H. 1934. American palmettoes. Gentes Herba- rm 3: 275-339. BTFL 1944. Revisionof the American palmettoes.Gentes Herbarum 6:365-459. SVGA BoMHARD,M. L. 1935. Sabal louisiana, the correct name Ior the polymorphic palmetto of louisiana. J. Wash. Acad.

Sabal palmeta of Sci. 25: 35-44. 1937. What pahns grow in louisiana. Louisiana Conservation Review, Autumn, pp. 37-42. Serenoa rebe6 1940. What palms grow in Louisiana. Louisiana Conservation Review, Autm, pp. 43-47. 1943. Distribution and character oI Sabal louis- .60 .50 iona. J. Wash. Acad. Sci. 33: 170-I82. 1950. Palm trees in the .U.S. Depart- 2. Phenogram of Nei's ( I 978) unbiasedgenetic identities for ment of Agriculture, Agriculture Information Bulletin No. Serenoa repens, Sabal palmetto and three populations of 22. Sabal minor. Bnovn, K. E. 1973. Ecologicallife history and geographical distribution of the cabbage palm, Sabal Palnetto. Ph.D. Thesis, State University at Raleigh. Cnawrono, D. J. 1983. Phylogenetic and systematic infer- studies. In: O. Tanksley MBLA" and FBLA indicated that Sabal louisiana ences from electrophoretic S. and T. J. Orton (eds.). Isozymes in plant genetics and is probably not genetically isolated from S. minor breeding, Part A. Elsevier, Amsterdam. and supports Bailey's (1944) conclusion that S. Deney, W. 1816. A geographical description of the state Louisiana should not be separatedfrom S. rninor. of Louisiana, the southern part of the state of Mississippi, Studies of other specieshave found that genetic and territory of . . . Together with a map, from actual survey and observation . . . of the state of Loui- identity betweenintraspecific populationsare usu- siana, and adjacent countries. JamesOlmstead, publisher, ally quite high, nearly always 0.90 or above (Got- New York. theb 1977, l98la, Crawford1983). 1818. The ernigrant's guide to the western and Between congenericspecies, most studieshave southwestern states and territories: comprising a geo- identity are much lower than graphical and statistical description of the states . . . found that genetic Accompanied by a map of the United States. Kirk and within species(Gottlieb l98la, Crawford l9B3). Mercein, New York. For 13 speciespairs in 8 genera,Gottlieb (L977) EvAN, J. 1967. ClassicaBotanica America. Vol. 5: C. S. found a mean genetic identity of I : 0.67. Iden- Rafinesque; Florula Ludoviciana; or, A Flora of the State tities betweenthe three populationsof Sabal minor of Louisiana. Hafner Publ. Co., New York. (I : FEATHERMAN,A. I 870. Report of botanical survey of south- and S. pahnetto certainly fall in this range ern and central a Louisiana made during the year 1870. 0.662). New Orleans. Between Serenoa repens and the two species FITzIATRIcK, T. J. lglf. Rafinesque.A sketch of his life of. Sabal sampled, the mean genetic identity is with bibliography. The historical Dept. of lowa, Des Moines. for comparisonsbetween genera 241 pages. 0.379. However, FLINT, T. 1828. A condensedgeography and history of the and higher taxonomic levels,the utility of allozyme Western States, or the Mississippi Valley. E. H. Flint, data is increasingly limited. This is due to under- Cincinnati. lying assumptionsin data interpretation. To inter- C,onuen, G. C. et.lo J. Rrnzr, JR. 1979. Genetic distance and heterozygosity estimates in electrophoretic studies: pret two different electromorphs from diferent Effects of sample size. Copeia 1979(2): 242-249. genera as representing genetic divergence equal C,orTrtrn, L. D. 1977. Electrophoretic evidence and plant to two different electromorphswithin a genusdis- systematics.Ann. M. Bot. Gard. 64: 16l-180. torts the data, forming closer genetic affinities l98la. Electrophoretic evidence and plant popu- betweentaxa than may exist. Also, the assumption lations. Prog. Phytochem. 7: l-46. I98Ib. Gene numbers in species of Asteraceae that electromorphs with identical mobilities rep- that have different chtomosome numbers. Proc. Nat. Acad. resent identical proteins when derived from widely Sci. USA 78:3726-3729. separated taxa is undesirable (Crawford 1983). GuenNsnnt, L. B. 1804. Observationssur le Sabal d'Adan- The results of these taxonomic comparisonsindi- son. Bullentin Societ6 Philomatique de Pais 2: 2O6. B. LTNHARTero B. MrrroN. 1979. techniquesto Heumcr. J. L.. Y. J. cate that the use of electrophoretic Relationships between life history characteristics and elec- detect genetic differencesshould be a valuabletool trophoretically detectable genetic variation in plants. Amu. in nnraveling the systematicsof Sa.bal. Rev. Ecol. Syst. l0: 173-2OO. =

l99sl RAMPAND THIEN: SABAL

HILMoN, BnIsro, Jn. 1969. Autecology of saw pahnetto ... Suivis de la Flore Louisianaise.vol. 3. F. Buisson, (Serenoa repens (Bartr.) Small). Ph.D. Thesis, Duke Uni- Paris, versity. Scnorr, A. 1857. Substmce of the sketch of the geology Lr Crrncg, C. 1691. Premier etablissementde la foy dans of the lower Rio Bravo Del Norte. 1z: William H. Emory la Nouvelle . A. Auroy, Paris. (ed.). Report on the United Statesand Mexican Boundary MARTIUS,C. F. P. 1838. historia naturalis palmarum, Vol. Survey. 1(2): 28-48. 3. T. O. Weigel, Lipsiae. Sruerr, J. K. 1926. A new palm from the MississippiDelta. 'McCrnNecneu, L. R.. Jn. eno A. C. BEAUCHAMP. 1986. Torreya 26: 33-35. Low genetic differentiation among isolated populations of 1929. Palmetto-with-a-stem-Sabal deeringiana. the California fan palm (Washingtonia f.lifera). J. N.Y. Bot. Gard. 30: 273-244. 4O:315-322. Sorrrs, D. E., C. H. Haunrrn, D. C. Dennow AND G. J. .Moonr, H. E., JR. 1971. Notes on Sabal in cultivation. Gesronv. 1983. Starch gel electrophoresisof ferns: r{, Princioes15:69-73. compilation of grinding buffers, gel and electrode buffers, NEI, M. f972. Genetic distance between populations. Am. and stainingschedules. Am. Fern. I.73:9-27. Nat. I06: 283-292. Uur, N. W. ANDJ. Dnervsnrnrn. 1987. Genera Palmarum. 1978. Estirnation of average heterozygosity and A classificationof palms based on the work of Harold E. genetic distance from a small number of individuals. Moore, Jr. Allen Press, Lawrence, Kansas. Genetics89: 583-590. VItrs, R. A. 1960. Trees, and woody vines of the NUTTALL,T. 18f8. The genera of North American plants, Southwest, University of Texas Press, Austin, Texas. and a catalogue of the species, to the year 1817. Phil- ZoNe, S. 1999. A monograph of Sabal (: Cory- adelphia. phoideae).Ph.D. Dissertation, Claremont, California. RAFINESeUE,C. S. 1817. Flora ludoviciana; or; A flora of 1990. A monograph oI Sabal (Arecaceae: Cory- the state of Louisiana. Wiley and Co. New York. phoideae).Aliso l2(4): 583-666. Reprp, P. F. 1989. Natual history of Sabal minor: Demog- eNo W. S. Julo. 1986. (Palmae): raphy, population genetics and reproductive ecology. Ph.D. systematics, distribution, ecology, and comparisons to other Dissertation. Tulane University, New Orleans, LA. Florida scrib endemics.Sida ll: 417-427. RosIN, C. C. 1807. Voyagesdans l'interieur de la Louisiane

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