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Home , Ceratiola, Chionanthus pygmaeus, Conradina, Dicerandra, Dicerandra christmanii, Dicerandra frutescens

198 SYSTEMATIC BOTANY [Volume 14 vealed that the taxon in question was firstcol- Suffruticosechamaephyte to ca. 0.5 m tall,the lected east of Sebring in 1948 by a local floriferousand vegetative shoots stiffand as- collector,Ray Garrett.Garrett sent several spec- cending froma ramosewoody base; odor minty- imens to ErdmanWest at the Universityof Flor- cineole. Taproot branched with extensive ida, which were later identifiedas D. frutescens spreading fibrous secondary roots. Stem gla- by Daniel B. Ward (herbariumannotations, 1976; brous except for sparse unicellular-pubescence Ward 1979), and Richard P. Wunderlin (1984). at nodes. Leaves more or less sessile; apices These specimens were also reviewed by Huck rounded; bases cuneate; marginsentire; adaxial (1987), in her monographicstudy of this south- and abaxial surfacesglandular-pitted. Leaves of eastern U.S. , and placed with D. frutes- determinateflowering shoots not subtending cens.Wunderlin's (1984) attemptto relocatethese cymesnarrowly ovate to narrowlyoblong, (4)5- populations of "D. frutescens"was unsuccessful. 11(14) mm long, (0.8)1-2.5 mm wide, similarto Other recentstudents of ,L. H. Shin- but smaller than leaves of overwintering,veg- ners and R. Kral, apparently did not see Gar- etativeshoots; leaves subtendingcymes 2-8 mm rett's material because these specimens were long, 0.5-1.8 mm wide; leaves of axillary fas- neither annotated by them nor cited in their cicles 1-7 mm long, similar to bracts of cyme respective publications (Kral 1982; Shinners but slightlylonger. Internodesof upper (cyme- 1962). Withone of these specimens,Garrett had bearing) portion of determinate flowering included a note (now attachedto a sheetat FLAS) shoots 6-16 mm long, those of lower (vegeta- indicating his opinion that the plant repre- tive) portion of shoots 3-20 mm long, those of sented an undescribed in a new genus overwinteringvegetative shoots usually 3-10 closely related to Dicerandra-a conclusion not mm. Inflorescencea verticillaster,each axillary unexpected,for no suffrutescentDicerandra had cyme of 1 to 3 flowerssubsessile or with a pe- been described at thattime (see Shinners 1962). duncle to 1 mm long; pedicels 2.5-6 mm long. Recent studies of the populations near Se- Calyx 13-ribbed,bilobed, the lower lobe biden- bring have confirmedtheir distinctnessfrom tate and the upper lobe with two strongly Dicerandrafrutescens, the recognitionof the tax- winged ridges meeting at summit,6.5-10 mm on at the specific level, and the phylogenetic long, ca. 2 mm wide (at midpoint), bordered placement of the new species. Following the with an indistinctwhite band, unicellular cil- detailed field work of S. Christmanin discov- iate-margined,glandular-pitted, and with an ering populations and pointing out important internalannular band of flat,distally appressed, taxonomic characters,D. christmaniiis here de- 1 mm long multicellular (uniseriate) hairs in- scribed,illustrated, and compared with related serted 1-1.5 mm below the mouth; awns ca. 1 taxa. mm long, triangular.Corolla infundibular,ge- niculate to ca. 900; tube 7-10 mm long; limb (fromgeniculum to distal edge of upper lobe) Dicerandra christmaniiHuck & Judd,sp. nov. 5-10 mm long, the superior lobe a recurving, (figs. 1-3).-TYPE: United States, , cleftstandard, ca. 5 mm wide and set back from Highlands Co., 5.8 mi fromDesoto Ave. in the outer margin of the inferiortripartite lobe, Sebring to Central Blvd. E of Sebring, off the sinusbetween the superiorand inferiorlobes C-17 on north-southridge, scrub (fig. cut to ca. 9 mm, the midpetal of inferiorlobe 4, pop. 4), 10 Sep 1987, R. Huck, R. Wun- recurved; inner surface pubescent with multi- derlin,B. Hansen & K. DeLaney 4825 (holo- cellular conical hairs,the interiorof tube below type: FLAS; isotypes: A, BHO, DUKE, F, geniculum with longer multicellularhairs; out- FLAS, FSU, FTG, GA, K, MO, MSC, NCU, er surfacepubescent with stipitatehairs; corolla NY, SMU, TEX, UC, US, USF). buds yellow, at maturitypale cream (eventually Species haec a Dicerandra frutescerYsShin- fading to white) with vivid purple-red mark- ners antherisflavis (vs. antheris lavandulis vel ings, often trellised-patterned,on upper lobe interdumalbis), corolla cremea(vs. corolla alba), and irregularspots on lower lobe. 4, foliis (4)5-11(14) mm longis, cineolis aromati- didynamous,exerted slightly beyond the lower cis, connectivo antherae eglandulosis vel spar- corolla lobe, tiltingupward upon anthesis,then sim glandulosis, et calcaribus antherae brevio- at maturitystraight and declinate along lower ribus, i.e., 0.34-0.6 mm longis differt. lobe; filamentswhite, inserted at the same level 1989] HUCK ET AL.: DICERANDRA 199

D

A ~~~~~~~~~~3mm

t ~~~~'~ X '1~ SS '

1cm

1mm tS Giv_ _ _ 2mm2 m

FIG.1. Dicerandrachristmanii. A. Flowering branch. B. Leaf. C, D. Flower: frontaland lateralviews. E. Longitudinalsection of corolla showing upper lip (=standard) and attachmentof stamens(corolla splitthrough middle of lower lip). F, G. Anthers. H. Apex of style. I. Ovary. J.Nutlets and persistentcalyx.

withinthe corolla; connectivewidened, basally Additional specimensexamined. UNITED STATES. covered with few, small, reddish and yellow Florida: Highlands Co., T34S, R29E, Sect. 36, 2.8 mi glands or such glands lacking; anthersacs bril- SE on Airport Rd. (=Rt. 17) fromjct with Rt. 17A, liantyellow, sometimes tinged basally with red; then E on Moon Ranch Rd. 1 mi, then N on Snyder spurs brilliantyellow, 0.34-0.6 mm long, del- Rd. 0.6 mi, then E on RR trackca. 0.4-0.5 mi (pop. 1), toid,glabrous, directed upward into the corolla. Alcorn172 (FLAS, GH); along RR tracksca. 3.5 mi E white and sticky.Pistil white, 18-26 mm of Sebring (pop. 1), Christman1759 (FLAS); ca. 2 mi E of jct. of Rt. 27 and Rt. 98 on Rt. 98, then ca. 0.5 mi long, with a slender hirtellous style. Fruit a N, near RR tracks (pop. 5), Christman1840 (FLAS); of 4 schizocarp nutlets,the nutletsovoid, brown, T35S, R29E, Sect. 12, E of Central Blvd., 0.3 mi S of smooth; pericarp of compact cellulosic (slime) C-623, ca. 5.5 mi SE of Sebring (pop. 4), DeLaney 1399 cells, which, when wet, become mucilaginous. (USF); T34S, R29E, Sect. 35, scrub along W side of Flowering fromJuly to November, but primar- Snyder Rd., ca. 'A mi S of RR tracksat end of road, 3 ily during Septemberand October. mi E of Sebring (pop. 2), DeLaney 1400 (USF); T34S, 200 SYSTEMATICBOTANY [Volume 14

FIG.2. Flowers of shrubbyspecies of Dicerandra. A. D. christmanii.B. D. frutescens. C. D. immaculata. D. D. cornutissimna.

R29E, Sect. 35, scrub 0.45 mi S of RR tracks,W side Rd.,then W on AirportRd. 0.9 mi, then SW on Central of Snyder Rd. (pop. 2), DeLaney 1440 (USF); T35S, Blvd,ca. 0.4 mi at "FlamingoVillas", near Sebring R29E, Sect. 12, scrub 0.6 mi S of C-623 on Central (pop. 4), Judd5570 (A, F, FLAS,FTG, MO, NCU, NY, Blvd. (pop. 4), DeLaney 1452 (USF); T34S, R29E, Sect. SMU, US, USF); T35S, R29E,Sect. 1, near Sebring, 35, scrub 0.3 mi S of RR tracks,W side of Snyder Rd. fromjct of Moon RanchRd. and AirportRd., then E (pop. 2), DeLaney 1453 (USF); T34S, R29E, Sect. 36, on Moon RanchRd. 1.4 mi, scrubat S side of road scrub along RR tracks,0.6 mi E of Snyder Rd. (pop. (pop. 3),Judd 5573 (FLAS,NCU, SMU,USF); (pop. 1), 1), DeLaney 1460 (USF); along Seaboard RR track ca. Skean 2130 (F, FLAS,FSU, GA, GH, MO, NCU, NY, 3.5 mi E of Sebring (pop. 2?), 9 Aug 1948, Garretts.n. US, USF); (pop. 2), Skean 2134 (FLAS,MO); (pop. 4), (FLAS); along Seaboard RR trackca. 4 mi E of Sebring Skean2168 (FLAS,NY); (pop. 3), Skean 2170 (FLAS). (pop. 1?), 30 Aug 1948, Garretts.n. (FLAS); along Sea- board RR track E of Sebring (pop. 1 or 2?), 25 Sep 1948, Garretts.n. (FLAS); (pop. 2), Hansen et al. 11400 Distribution and Ecology. Dicerandra (USF); T35S, R29E, Sect. 12, SW of C-623, 5.7 mi SE christmaniiis known fromonly fivepopulations of Sebring (pop. 4), Hansen et al. 11407 (USF); (pop. located to the south and southeast of Sebring 1), Judd5548 (A, DUKE, F, FLAS, FSU, MO, NY, RSA, in Highlands Co., Florida (fig. 4). All popula- Se- US, USF); T34S, R29E, Sect. 35 (NE corner), E of tions occur in openings within areas of sclero- 2.8 mi SE on AirportRd. (C-623) fromjct with bring, phyllous oak scrub on well to excessively Rt. 17A, then E on Moon Ranch Rd. 1 mi, then N on drained deep yellow sand (Astatulaand Tavares SnyderRd. 0.5 mi, near end of road just S of RR tracks (pop. 2), Judd5549 (F, FLAS, FSU, GA, MO, NCU, NY, ) with a depth to seasonal high water table US, USF); T35S, R29E, Sect. 12, fromjct of US. 27 and of 1-2 m (or more). See Appendix I for list of US. 98, E on US. 98 3.2 mi, then N on rd. to Airport associated species. 1989] HUCK ET AL.: DICERANDRA 201

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FDr 4 S f:f 202 SYSTEMATIC BOTANY [Volume 14

throughout the geographical range of both species. All fiveknown populations of D. christ- maniiwere studied in the field,along with eight of the nine extantpopulations of D. frutescens. In addition, herbariummaterial was examined and measured (see Huck 1987, and specimens cited herein, and Appendix II). These analyses 27X 1 indicate that D. christmaniiand D. frutescensare A distinguished by several morphological and chemical characters,as discussed below. Flower Color. Field study revealed that D. christmaniicould be distinguished fromD. fru- 0: tescensby the color of its anthers and corollas. Dicerandrachristmanii consistently has brilliant yellow anthers while those of D. frutescensare deep to pale purple or rarelywhite (fig.2). The corollas of D. christmaniiare cream colored with BO&,) purple-red spots, and remain so nearly throughoutthe firstday of anthesis (fig.2). To- ward the end of the firstday (ca. 1700 hr) the color of some corollas begins to fade to white (with purple-redspots). Old flowerspersisting 10 on the plant on the morningof the second day are clearlywhite with purple-redspots. In con- trast,the corollas of D. frutescens,which upon

028 opening are also cream with purple-red spots, 028? WE,) .\R ~014H 13 fade to white (with purple-redspots), in about 1 to 3.5 hr afteropening (fig.2). Thus, by noon all (or nearly all) of the flowersof D. frutescens are white with purple-redspots, while those of FIG. 4. Distributionof and D. D. christmaniiremain cream-coloredwith pur- frutescens(Highlands Co., Florida). Localities 1-5 ple-red spots into the late afternoon.Color of (stippled) = extant populations of D. christmanii;lo- both corolla anthers calities 6-14 (diagonal lines) = extantpopulations of and is distorted upon D. frutescens;solid star = extirpatedpopulation of D. drying;thus, these charactersare best observed frutescens;bold lines = major highways; circled num- in living . bers = highway numbers; thin lines = lakes; A = Foliar Aroma. Field study of D. frutescens Sebring; B = Lake Placid; dotted line = outline of and D. christmaniialso revealed that the two Lake Wales Ridge (25 m contour-lineon E side and species exhibit distinctlydifferent fragrances 30 m contour-lineon W side). when leaves are crushed;the odor ofeach species was uniformwithin and between populations. The crushed foliage of D. frutescensis similarto that of peppermintwhile that of D. christmanii TAXONOMICRELATIONSHIPS is reminiscentof eucalyptus oil. This charac- Dicerandra christmaniiis phenetically most teristicwas thus investigated in more detail similarto D. frutescens(sect. Dicerandra), a species through a survey of leaf essential oils of the occurringjust to the south of the range of D. four suffrutescentspecies of Dicerandra,which christmanii(fig. 4). The two taxa occur in similar are all Florida endemics. ,but are separated by a distance of ca. The essential oils of these species (table 1) 10.5 km (6.5 mi). Notably,a slight break in the were analyzed by gas chromatography/mass ridge occurs between the species at Josephine spectrometry(GC/MS). Fresh leaves (ca. 0.5 g) Creek, as indicated by the 25 m elevation line were placed in 1-ml glass vials and extracted on figure4. An analysis of the variationin both overnight in hexane. The resulting solutions D. christmaniiand D. frutescenswas made from were analyzed using a Hewlett-Packard5995-C 1989] HUCK ET AL.: DICERANDRA 203

TABLE 1. Percentcomposition of Dicerandraleaf essential oils, based on GC/MS analyses of hexane extracts of leaves. a Identificationstentative.

christmannzi frutescens cornutissima immaculata l-nonen-3-ola 1.2 3.8 6.1 monoterpene 1 3.3 monoterpene 2 11.5 linalyl acetate 4.8 1,8-cineole 54.3 limonene 1.1 1.0 unidentified(m.w. 154) 11.8 rose oxide isomera 7.1 rose oxide isomera 52.1 menthone 5.4 5.8 isomenthone 3.3 8.0 alpha-terpineol 17.3 isopulegone 14.7 11.9 pulegone 24.0 73.3 64.6 sesquiterpenes 5.8

GC/MS systemequipped with a 30-meterDB-1 terial,a phenetic investigationof several mor- capillary column and a capillary direct inter- phological characterswas conducted based on face. Oven temperaturewas programmedfrom measurementsfrom collections made by W. S. 35-100?C at 4?C/minuteand from100-210?C at Judd and J. D. Skean fromall known popula- 12?C/minute.Mass spectra of each compound tions of both D. christmaniiand D. frutescens, were searched against the NBS mass spectral except populations 12 and 14 (fig. 4). To eval- library;tentative identifications of compounds uate the phenetic separation between the two are based on these matchesand retentiontimes. species evident using only characters observ- The results(table 1) are based on an analysis able on herbariumsheets, a Principal Compo- of three populations of D. frutescens,two of D. nentsAnalysis of 70 specimensfrom both species cornutissimaHuck, two separate gatherings (in was conducted using the CLUSTAN Computer May and September)from the same population program, version 3.2 (Wishart 1987); see dis- of D. christmanii,and a single population of D. cussion of this ordination technique in Wiley immaculataLakela (table 2). Values for those (1981) and Sneath and Sokal (1973). Values for species with two or three gatheringsare aver- eight continuously varying and meristicchar- aged because they were essentially identical. acters,mainly of vegetative morphology(table The essential oil composition of Dicerandra 3), were scored foreach specimen and the taxa christmaniiis distinctive and shares no major ordinated along the firsttwo principal com- compounds with any other shrubbyspecies of Dicerandra(table 1); its oil is dominated by 1,8- cineole and alpha-terpineol. The oil of D. fru- TABLE 2. Localitiesof Dicerandraspecies used in tescensis also unique, containinglarge amounts gas chromatographicanalysis. of several compounds tentativelyidentified as isomersof rose oxide. The oils of D. cornutissima D. christmanii:Highlands Co., T34S, R29E,Sect. 12 and D. immaculataare nearly identical,and pos- (pop. 4, wild),sampled twice (May and September, sess a strongminty odor due to the presence of 1987). pulegone and menthoneisomers. The chemical D. cornutissima:Marion Co., T17S, R21E, Sect. 2 (wild); data thus give strongsupport to the conclusion T17S,R21E, Sect. 11 (nursery). that D. christmaniiand D. frutescensare specifi- D. frutescens:Highlands Co., T36S, R29E, Sect. 25 (pop. cally distinct. 6,wild); T36S, R30E, Sect. 20/32 (pop. 7, wild); T36S, R29E,Sect. 24 (nursery). Vegetative Morphology. Because the char- D. immaculata:St. Lucie Co., T34S, R40E, Sect. 8 (nurs- acters of corolla and color and foliage ery). odor are not usually available in herbariumma- 204 SYSTEMATIC BOTANY [Volume 14

TABLE 3. Morphologicalcharacters used in PCA analysisof variationwithin and betweenDicerandra -o frutescensand D. christmanii.

0 0 0 1. Lengthof leaves on lowerportion of determinate 1 0 o? 0 reproductivebranches; i.e., those leaves not subtend- 0 0 X@0~ .@0 000. ~O O 08a, o00 o ing cymes. 2. Widthof leaves on lowerportion of I @0 00 -1 @$ 0 determinatereproductive branches. 3. Length of * 0 00 0 0 leaves subtendingcymes on determinatereproduc- 0 0 0 tivebranches. 4. Widthof leaves subtending cymes on determinatereproductive branches. 5. Lengthof -2* ? leaves of fasciculateaxillary shoots on determinate reproductivebranches. 6. Internodelength on low- er portion of determinatereproductive branch- -3 -2 -1 0 1 2 3 4 5 ? es. 7. Numberof flowersper cyme. 8. Lengthof PC. 1 53% anther-horns. FIG. 5. Plotof thefirst two principal components resultingfrom phenetic analysis of Dicerandrafrutes- censand D. christmanii.Open circles = D. frutescens; closedcircles = D. christmanii;OTU's of bothspecies the total range of leaf lengths,(4)5-11(14) mm identifiedon basis of antherand corollacolor and long in D. christmaniivs. (7.5)9-23(30) mm long odorof crushedleaves. See discussionin text. in D. frutescens,there is no overlap in the average leaflength per plant on the lower portion of the floweringshoots of the two species (based on ponents (fig.5). Identificationof the OTU's used 70 specimens, measuring three leaves per in this analysis was based on antherand corolla shoot-one froma node just below the lower- color and odor of crushed foliage. The resulting mostnode producing a cyme,one at or near the diagram (describing66.5% of the variationcon- base of portion of the shoot produced in the tained in the original data set) indicates that spring, and one in between). In addition, the most individuals of D. christmaniiand D. frutes- length of the leaves subtending cens are separable phenetically on the basis of (2-8 mm in D. christmaniivs. 3-15 mm in D. the morphological charactersincluded in the frutescens)and antherspur length (0.34-0.6 mm analysis. Charactersheavily loaded on the first in D. christmaniivs. 0.45-0.8 mm in D. frutescens) principal component,which best separates the provide some informationuseful in species de- species, are: 1) length of leaves on lower, veg- termination.Dicerandra christmanii also tends to etative portion of determinate flowering have shorter internodes and fewer-flowered shoots, 2) length of leaves on upper portion cymes than D. frutescens,but there is too much of floweringshoots, i.e., those leaves bearing overlap in these featuresfor them to be of di- cymes in their axils, 3) length of fasciculate agnostic value. leaves in axils of leaves on lower portion of Anther Morphology. Anther spurs of Dic- determinateflowering shoots, and, to a lesser erandraare highly variable among species and extent, 4) internode length on flowering functionas triggermechanisms in the dispersal shoots, and 5) anther spur length. Thus, the of pollen by insects. Huck (1987), examining a best characterto use in distinguishingherbar- single specimen of each species of Dicerandra, ium material of D. christmaniifrom that of D. comparedanther micromorphology using scan- frutescensis the length of the leaves. The flow- ning electronmicroscopy (SEM), and construct- ering shoots of D. christmaniihave leaves that ed a preliminarykey to species based on char- consistentlyhave shorteraverage lengths than acters of the anther spurs. To present similar those of D. frutescens;i.e., leaves of the lower data forD. christmanii,nine anthers fromSkean portion of determinateflowering shoots, those 2168were fixedin FAA forthree days, and taken notsubtending axillary cymes. The average leaf throughan EtOH dehydrationseries of two suc- lengthper plant in D. christmaniiis 5.3-10.3 mm, cessive 15-minutewashes each at 70%,90%, 95%, while thatof D. frutescensis 11.2-25.7mm (based and threesuccessive 15-minutewashes at 100%. on our sample). Although there is overlap in The antherswere then critical-point-driedafter 1989] HUCK ET AL.: DICERANDRA 205 four15-minute washes of liquid CO2 in a Balzers dermal cells, with one species, D. odoratissima CPD-010 critical-pointdrier, attached to stubs Harper (ibid., fig. 21D), having the cells at the with redimounts,plated with gold in an Eiko apex of the spur with striationsterminating in IB-2 ion coater forsix min at 8 mA, and exam- a rosettepattern. Some spur cells of D. christ- ined and photographed using a Hitachi S-450 maniialso appear similarin thisrespect (fig. 6D), scanning electron microscope. while othercells lack these radiatingstriations. The anthers of D. christmanii(fig. 6A-D) are It should be emphasized thatnearly all of these most similar to those of D. frutescens.Figure 6A SEM observationsare based on a single anther shows an entire anther with two pollen sacs, per species; thus more study is needed to de- spurs,and a raised region that is oftennot this termineinfraspecific variation. pronounced,on the connective.Figure 6B shows . Although investigations of a single open pollen sac and a spur that is be- pollination biology are preliminary,they sug- ginning to wither.When compared with figure gest that of the fourperennial species of Dicer- 18D (D. frutescens)in Huck (1987), it appears andra,D. immaculataand D. cornutissimaare - thatthe spurs of D. christmaniiare slightlysmaller pollinated. Both these species have hairyanther than those of D. frutescens.This differencealso spurs (Huck 1987). In contrast,D. frutescensand was indicated above, with some overlap, when D. christmaniihave more or less smooth anther spur length of both species was measured for spurs and are visited regularlyby bee-flies(Ex- the PCA analysis. oprosopafasciata), the latter species almost ex- Perhaps the greatestdifference between the clusively so. Bee-fliesprobe the corollas appar- anthers of D. christmaniiand D. frutescensis an ently in search of present on the hairs apparent reduction in the number and size of of the tube below the geniculum. Dicerandra anther glands in the former.In the close-up of frutescensis visited by a varietyof insects(Huck the connective and the bases of the pollen sacs 1987). Halictid (Augochlorellagratiosa) have (fig.6C), note the single, small (ca. 53 ,umlong) been observed visiting the anthers of this gland located near the base of the left pollen species, possibly attractedby the pollen, oils, sac. All species of Dicerandraillustrated by Huck and the glistening glands on the anther con- (1987) have larger,more numerous glands scat- nective (Huck 1987). Connective glands, as in- tered on the bases of the pollen sacs and con- dicated above, are few or wanting on D. christ- nective. In most anthers of D. christmaniiex- manii, and no such visits to anthers were amined in thisstudy (e.g., fig.6A, B) such glands observed. In addition, other bees (Apis mellifera were absent, but in some anthers there were and Bombusimpatiens) were seen on D. frutescens one or two such glands. In Huck (1987, figs. (Huck 1987), but were not observed to visit the 18D, 19F) the anthersof D. frutescenshave more flowers of D. christmanii.Cineole is the domi- numerousand larger(ca. 71-85 ,umlong) glands. nant essential oil in the leaves of this species, Thus, D. christmaniiappears not only different but it is not known if this chemical plays an fromD. frutescens,but unique among all other importantrole in attraction. membersof the genus in having this reduction Summary. Dicerandrachristmanii is clearly in gland size and number. specificallydistinct from D. frutescens.It is dis- The anther spurs of D. christmanii(fig. 6D), tinguishedby: 1) yellow (vs. purple to white) like D. frutescens(see Huck 1987,figs. 18D, 21B), anthers, 2) persistentcream-colored corolla (vs. have apices that terminaterather abruptly. Al- corolla quickly fading to a stark white), 3) though the epidermal cells are heavily striated, foliagewith distinctsmell of cineole (vs. foliage the spurs themselves are relatively smooth, with mintysmell), 4) shorterleaves, and 5) lacking the knoblike projections of epidermal anther connectives with fewer and smaller cells, i.e., "hairs," found in D. densifloraBenth. glands. These differencesare maintained in ex DC., D. immaculata,and sometimes also D. plants grown in a common garden at Bok Tower cornutissima(ibid., figs. 18C-D, 20, 21B, C). Un- (S. Christman,pers. comm.). Additionally, the like D. frutescens(ibid., fig.21B), the epidermal habit of D. christmaniiis more compact-spread- cells at the tip of the spur in D. christmaniiappear ing than that of D. frutescens(fig. 3), and this to be relatively straight,rather than slightly differencemay be reflectedin the pheneticanal- twisted together.Huck (1987) also differentiat- ysis where internode length, i.e., 4-20 mm in ed patternsof striationson the individual epi- D. christmaniiand 5-37 mm in D. frutescens,was 206 SYSTEMATIC BOTANY [Volume 14

rx~~~~~~~~~~~~~~r

FIG. 6. Anthers of Dicerandrachristmanii. A. Anther. B. Pollen sac and anther spur. C. Close-up of anther connective,note single small gland. D. Close-up of anther spur showing ornamentation. 1989] HUCK ET AL.: DICERANDRA 207 fairlyhighly loaded on the firstprincipal com- linated by a wide variety of insects. The two ponent. Dicerandrachristmanii also shows a ten- species are completely allopatric and are thus dency to have shorteranther spurs. Finally,Di- reproductivelyisolated. cerandrachristmanii appears to be pollinated A key to the species of Dicerandrarevised from mainly by a bee-flywhile D. frutescensis pol- that presented in Huck (1987) is given below.

1. Corolla infundibular;tube geniculate; superior lobe a standard; stamens and style exserted; style free; all 4 filamentsinserted at same level within corolla; anther spurs pointing upward or laterally into corolla (sect. Dicerandra). 2. Plants herbaceous; annuals. 3. Anther spurs obtuse or barely acute, with minute domes of digitate hairs; cymes epedunculate with flowers nearly sessile in compact verticils; stamens subequal in length; pollen white to pale yellow ...... D. densifloraBenth. ex DC. 3. Anther spurs acuminate, acicular, hairless; cymes pedunculate, with pedicels 3-6 mm long, and flowerswith pedicels 3-9 mm long; stamens unequal in length; pollen brightyellow ...... D. linearifolia(Elliott) Benth. 2. Plants woody; chamaephyticperennials. 4. Corolla standard lacking spots; tube smoothlybent; spurs hairy ...... D. immaculataLakela 4. Corolla standard spotted and patterned;tube abruptlygeniculate; spurs hairless or nearly so. 5. Corolla reddish-purple;style with few hairs or glabrous; anther spurs more than 1 mm long; leaves ca. 1 mm wide ...... D. cornutissimaHuck 5. Corolla white or cream-colored;style with numerous stiffconical hairs; anther spurs less than 1 mm long; leaves usually more than 1 mm wide. 6. Antherslavender or occasionally white,with glands on connectiveabundant; crushed leaves with a mintyfragrance ...... D. frutescensShinners 6. Anthersbrilliant yellow, with glands on connectiveinconspicuous or lacking; crushed leaves with fragranceof cineole (odor similar to that of medicinal eucalyptus oil) ...... D. christmaniiH uck& Judd 1. Corolla tubular, straightor slightly curved; superior lobe cucullate; stamens and style included; style clamp present;4 filamentsinserted at 2 differentlevels withincorolla; antherspurs pointing downward into the corolla (sect. Lecontea). 7. Corolla tube to 18 mm long; corolla orifice2 mm wide; hood narrow,tent-like and apiculate; leaves linear ...... D. odoratissimaHarper 7. Corolla tube more than 20 mm long; corolla orifice4 mm wide; hood wide and apiculate to truncately rounded; leaves narrowlyoblong ...... D. radfordianaHuck

and apomorphic (derived: scored as 1) states REVISED CLADISTIC ANALYSIS (tables 4, 5). A few of the charactersare quan- OF DICERANDRA titive,and the states of such charactersare nec- The phylogeneticanalysis presentedby Huck essarily somewhat arbitrarily defined. The (1987) is revised here by the addition of new guidelines outlinedby Almeida and Bisby(1984) characterinformation and the newly described were followed in delimitingbinary-state char- species, D. christmanii.This revised phyloge- actersfrom continuously varying measurement netic analysis of Dicerandrawas conducted us- data. Following Huck (1987), most characters ing the branch-and-bound algorithm (Hendy were polarized using /Calaminthaas and Penny 1982) and Wagner parsimony al- an outgroup (see Maddison et al. 1984; Stevens gorithm (with global branch-swapping and 1980, 1981; Wheeler 1981; Wiley 1981). Char- multipleparsimony options) of the PAUP (Phy- acter E was polarized using a generalized mint logenetic Analysis Using Parsimony) version outgroup;and the polarityof several characters 2.4 computer software for inferringphyloge- (I, L, P, R, U, W, X, and Y) was determinedby nies under the principle of parsimony (Swof- using the D. odoratissima-radfordianaclade as a ford 1985). functional outgroup (Watrous and Wheeler Twenty-fivecharacters were delimited and 1981), afterinitial analyses indicated that this assigned plesiomorphic (ancestral: scored as 0) clade was the sister group to the other species 208 SYSTEMATIC BOTANY [Volume 14

TABLE4. Charactersused in the cladistic analysis of Dicerandra.

Code Ancestralstate Derived state A. Shrubby herbaceous B. Leaves leptophyllous nanophyllous C. 1-3 flowers/cyme often3-7/cyme D. Perianth rounded, closed infundibular E. Corolla tube ? straight geniculate F. Upper corolla lobe cucullate standard G. Style clamp absent present H. Stamens hidden under upper lobe declinate along lower lobe, flaring I. Spur direction similar variable J. Stamens straight inverted K. Anthersnot spurred spurred L. Spurs smooth ornamentationconic or cylindric M. Corolla white, pink, to purple consistentlywhite N. Nectar guides present lacking 0. Nectar guides present only on lower lip absent on lower lip, but usually presenton upper lip P. Anther gland not sugary sugary Q. Flower odor minty cinnamon-like R. Leaf odor minty cineole odor S. Pedicel '3 mm long >3 mm long T. Pollination by bees or flies crawling insects U. Leaves usually >11 mm long <11 mm long V. Plants on quartz soil calcareous W. Antherswhite to purple yellow (to reddish-brown) X. Anthersnot reddish-brown reddish-brown Y. Pollen white to pale yellow ? brightyellow

of Dicerandra.The cladograms were rooted us- C, and M (tables 4, 5). The variation seen in ing a hypotheticalancestor with all characters habit (A) is especially noteworthy.Dicerandra scored as plesiomorphic. christmanii,D. frutescens,D. immaculata,and D. The PAUP analyses resulted in the genera- cornutissimawere scored with a "O" because they tion of three of 35 steps and consistency are woody, as are the outgroup genera (table 4). indices of 0.714 (or 0.643 when autapomorphies However, these fourspecies show some signif- and uniform characters are eliminated). The icant differencesfrom Conradina and Calamintha preferredtree, agreeing in most respects with in their growth architecture,possibly indicat- the published by Huck (1987), is shown in ing that their woody habit may have evolved figure7. All threegenerated cladograms are very independently from that of the woody out- similar, differingonly in the placement of D. groups. In the fourwoody species of Dicerandra, densifloraand D. linearifolia.In the preferredtree the floweringshoots are determinate,and die (fig. 7) these species are basal members of the back to near the base of the plant at the end of clade leading to the shrubby Florida endemic the growing season (see description),while in species. In the other trees, D. densifloralinks Conradinaand Calaminthathe floweringshoots with D. cornutissimaand D. immaculataon the are indeterminate(and do not die back). The basis of ornamentedanther spurs (L), while D. distinctivehabit of the woody species of Dicer- linearifolialinks with D. christmaniiand D. fru- andra is like that of the herbaceous species of tescenson the basis of long pedicels (S) and the genus in thatboth produce morphologically either a white corolla (M-but reversed in D. similar floweringshoots that die at the end of linearifolia)or yellow anthers (W-but reversed the growing season, and this observationlends in D. frutescens).These two treesare considered support to the hypothesis that these species implausible due to their high level of homo- evolved fromherbaceous ancestorssimilar to D. plasy in several morphologicalfeatures; e.g., A, linearifoliaand D. densiflora;i.e., the tree topol- 1989] HUCK ET AL.: DICERANDRA 209

TABLE 5. Charactervalues for species used in cladistic analysis. See table 4 for characters.

Character

Species A B C D E F G H I J K L M N O P Q R S T U V W X Y D. radfordiana 1 1 1 0 0 0 1 0 0 1 1 0 0 0 0 0 1 0 0 1 0 0 0 0 0 D. cornutissima 0 0 0 1 1 1 0 1 1 0 1 1 0 0 1 0 0 0 1 0 0 0 0 0 0 D. immaculata 0 0 0 1 1 1 0 1 1 0 1 1 0 1 1 1 0 0 0 0 0 0 0 0 0 D. frutescens 0 0 0 1 1 1 0 1 0 0 1 0 1 0 1 0 0 0 1 1 0 0 0 0 0 D. odoratissima 1 0 1 0 0 0 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 D. densiflora 1 1 1 1 1 1 0 1 0 0 1 1 0 0 1 0 0 0 0 1 0 1 0 0 0 D. linearifolia 1 0 0 1 1 1 0 1 0 0 1 0 0 0 1 0 0 0 1 0 0 0 1 0 1 D. 1. var. robustiorHuck 1 1 1 1 1 1 0 1 0 0 1 0 0 0 1 0 0 0 1 0 0 0 1 1 1 D. christmanii 0 0 0 1 1 1 0 1 0 0 1 0 1 0 1 0 0 1 1 0 1 0 1 0 0

ogy indicated in figure 7. It should be noted, maniiand D. frutescensare likely sister species, however, that breeding data (Huck 1987) sup- as indicated in figure7. portsthe non-preferredtrees in suggestingthat All cladograms also indicate a close relation- woodiness is primitivein Dicerandra,and it is ship between D. immaculataand D. cornutissima; clear that additional phylogenetic study is re- these species are linked by charactersof anther quired,especially an investigationutilizing data spur orientation and ornamentation (I, L). In on foliar essential oils of all species. contrast,the Wagner Groundplan Divergence Dicerandrachristmanii is hypothesized to be diagram in Huck (1987) indicates that D. cor- monophyleticin that it shows the autapomor- nutissimais cladisticallycloser to D. frutescenson phies of presence of cineole in the leaves (R) the basis of 1-2 floweredcymes, with the char- and leaves usually less than 11 mm long (U). actersof antherspur orientationand ornamen- This species also shows the apomorphic char- tation evolving in parallel. The linking of D. acter,yellow anthers (W), a traitthat may have cornutissimawith D. immaculata,instead of with evolved independently in D. linearifoliaif the D. frutescens,is more parsimonious-even using preferred tree is accepted. In addition, this Huck's (1987) data. However, the present anal- species is distinctive in lacking conspicuous ysis reverses the polarityof cyme flowernum- glands on the anther connective. These glands ber (C) because few-floweredcymes are found are found on all other species of the genus. in the outgroup genera. Thus, the presence of Dicerandrafrutescens is also likelymonophyletic; few-floweredcymes in both D. frutescensand D. in the cladisticanalysis the only autapomorphy christmaniiis considered a symplesiomorphyand indicatedis thatof pollination by various crawl- ing insects (T; see Huck 1987), which shows RAD ODO DEN LIN ROB FRU CHR COR IMM homoplasy (fig.7). However, this species is also B ~~~~~B R SN unique in having isomers of rose oxide as the majorcomponent of the leaf essentialoils. These in the three other L L compounds are lacking G shrubby species of Dicerandra,but this infor- Q mation was not included in the cladistic anal- ysis because chemical analyses of the herba- ceous species of the genus have not been EF conducted, and because this species has a pep- H permintsmell that is not easily distinguished K fromthat of D. cornutissimaor D. immaculata.All FIG. 7. 35-stepcladogram indicating relationships three cladograms indicate that D. christmaniiis withinDicerandra generated by PAUP (see discussion more closely related to D. frutescensthan either in text);synapomorphies indicated on cladogram,re- D. cornutissimaor D. immaculatafor both share versalsindicated by asterisk(see table4 forcharac- the derived character of having consistently ters);taxon names abbreviated by first three letters of cream to white corollas (M). Dicerandrachrist- specificor varietalepithet (see table5). 210 SYSTEMATIC BOTANY [Volume 14 cannot be used to indicate phylogenetic rela- -sclerophyllousoak scrubvegetation of the tionship. ridge, particularlyin the southern part, is rich In other respects,the preferredPAUP clado- in endemic and near endemic (>90% of range gramis similarto thatpresented by Huck (1987), in Florida) plant and animal taxa, many often exceptthat the Florida endemic species D. christ- in disturbed openings; examples include: As- manii,D. frutescens,D. immaculata,and D. cor- clepiascurtissii A. Gray,Asimina obovata (Willd.) nutissimaare here hypothesizedto forma mono- Nash, Bonamiagrandiflora (A. Gray) Heller, Cal- phyletic group on the basis of a reversal to a aminthaashei (Weatherby)Shinners, Caryaflor- shrubby/chamaephytichabit (A). This group is idana Sarg., Chapmanniafloridana Torrey & A. paraphyleticin Huck's (1987) analysis as well Gray, Chionanthuspygmaeus Small, Clitoriafra- as in the two non-preferredbut equally parsi- gransSmall, Eriogonumlongifolium Nutt. var. gna- monious trees (one showing two reversals to a phalifoliumGandoger, Garberiaheterophylla (Bar- shrubbyhabit, and the second with three dis- tram)Merr. & Harper, Ilex opaca L. var. arenicola tinct evolutionary origins of the herbaceous (Ashe) Ashe, Lupinusaridorum McFarlin ex Beck- condition, with the shrubby habit of the out- ner, Nolina brittonianaNash, Osmanthusmega- group taxa and the above listed Dicerandra carpus(Small) Small ex Little,Palafoxia feayi A. species considered a symplesiomorphic simi- Gray,Persea humilis Nash, Polygalalewtonii Small, larity).As discussed above, a treehypothesizing Polygonellamyriophylla (Small) Horton, two independent evolutionary events leading geniculataHarper, Quercus inopinaAshe, Sabal to a chamaephytichabit with determinate(and etoniaSwingle ex Nash, Sisyrinchiumxerophyllum annual) floweringshoots seems implausible; as Greene,Stylisma abdita Myint, amplexifolia is one that considers this habit, which is likely (Nutt.) Nutt., and W. carteriSmall. Of the 272 not even homologous to the woody condition known Florida endemics and near endemics (D. present in the outgroup, to be plesiomorphic. Hardin, pers. comm.) approximately13% occur Kral (1982) considered the shrubby species of in the southern Lake Wales Ridge region. In Dicerandrato be of comparativelyrecent deri- addition to Dicerandrachristmanii, seven species vation fromD. linearifolia-likeancestors; our re- are restrictedto Polk and Highlands counties sults (fig. 7) lend support to this conclusion. at the southernmostpart of the ridge:Dicerandra The monophyly of Dicerandrasects. Diceran- frutescensShinners, Eryngiumcuneifolium Small, dra and LeconteaHuck is confirmed,as well as Hypericumcumulicola (Small) Horton, Liatrisoh- that of the genus itself,in all generated clado- lingerae(S. F. Blake) B. L. Robinson, Paronychia grams. The presence of anther spurs (K) is a chartaceaFern., Polygonella basiramia (Small) Ne- strongsynapomorphy of the genus. In addition, son & Bates,and Ziziphuscelata Judd & Hall. The all species of Dicerandrapossess determinate latter was known from only two herbarium floweringshoots thatdie at the end of the grow- specimens until recentlyrediscovered by K. R. ing season-likely an additional synapomor- Delaney. Associates of Dicerandrachristmanii in- phy. The evolutionaryrelationships of the ge- clude 27 Florida endemics or near endemics nus, and the species within it, are discussed in representingnearly one-thirdof the species re- more detail by Huck (1987). corded at the type locality (Appendix I). Although endemism in Florida has been ad- dressed by several workers (e.g., Hardin 1974; ENDEMISM WITHIN THE Harper 1949a; 1949b, 1950; James 1961; Judd LAKE WALES RIDGE 1983;Judd and Hall 1984; Nash 1895; Neill 1957; The Lake Wales Ridge is the most southerly Ward 1979; Woodson 1947; Zona and Judd1986), and easterlyof a series of localized areas of high it is stillpoorly understood. We believe the high ground that comprises the Central Highlands degree of endemism on the Lake Wales Ridge of Florida (White 1970). It is a narrow ridge, is due to a combination of both historic and only a few kilometerswide but more than 150 edaphic factors.Although most of peninsular km long, extendingsouthward fromsoutheast- Florida was inundated in the past, particularly ern Lake County to southernHighlands Coun- during the Pliocene and Pleistocene intergla- ty.It consistsof xeric,residual sand hills, beach cial times,the southern Lake Wales Ridge has ridges, and dune fields with numerous sink- apparentlybeen emergentand suitablefor plant hole lakes and basins. The predominant sand habitation since the late Miocene or the early 1989] HUCK ET AL.: DICERANDRA 211

Pliocene (ca. 12 million yr B.P.). More recent scrub endemics [e.g., Hypericumcumulicola, ap- floristicstability is seen in fossil pollen records parentlymost closely related to the annual H. dating from 37,000 to 13,000 B.P. from Lake gentianoides(L.) Brittonet al. (Adams 1962); and Annie in southern Highlands County, which Polygonellaspp., apparentlyderived fromances- indicatethe presence of a dominant scrub com- torssimilar to the annual P. articulata(L.) Meiss- munity characterized by ericoides ner (Horton 1963)]. Michx.,Polygonella fimbriata (Elliott) Horton (?=P. Dicerandrafrutescens and D. christmaniiare re- robusta(Small) Neson and Bates),Polygonella cil- strictedto the southern Lake Wales Ridge and iata Meissner, Selaginella arenicola Underw., D. cornutissimato northernSumter and southern Quercus spp., and various (Watts Marion counties in the SumterUplands (White 1975). These species still occur locally in the 1970). Both regions are of late Miocene or early surroundingareas. It is probable that the Lake Pliocene origin(White 1970). On the otherhand, Wales Ridge served as a refuge for plants and D. immaculatais endemic to northernSt. Lucie animals during times of higher sea levels. and southernIndian River counties and found Equally importantfor development of ende- on the Atlantic Coastal Ridge, which consists mism is that the deep, dry, sandy soils of the of relic beach ridges of late Pleistocene (Pam- paleo-dune fields and ridges of the Lake Wales lico) origin (White 1970). During pre-Pamlico Ridge define a unique xeric environment to times, Dicerandramay have spread from the which a large number of taxa have become Central Highlands into geologically more re- adapted. cent areas. Interestingly,an entityhas just been The endemic plant species of the southern found in northernBrevard County (on the At- Lake Wales Ridge are likely of various origins lantic Coastal Ridge), which may constituteyet and ages. Many appear to be "neo-endemics" another species of Dicerandra.It resembles D. thatshare a recentcommon ancestor or are de- immaculatain many respects.This entityand its rived fromwide ranging eastern North Amer- relationshipto the other Dicerandraspecies are ican taxa (e.g., Perseahumilis Nash and P. borbonia currentlyunder investigationby R. B. Huck and (L.) Sprengel, Osmanthusmegacarpus (Small) R. P. Wunderlin. Furtherwork, particularlyes- Small ex Little and 0. americanus(L.) A. Gray, sential oil analysis of the non-woody members Chionanthuspygmaeus Small and C. virginicusL.), and cytological analysis, is needed to under- while othersappear to have a southwesternU.S. stand betterthe evolutionaryhistory of the ge- floristicaffinity (e.g., (A. Gray) nus. Heller, Carya floridanaSarg., Liatrisohlingerae, Peninsular Florida is being subjected to rapid Palafoxiafeayi A. Gray,and Ziziphuscelata); see urban and agriculturaldevelopment. The Lake also Zona and Judd (1986). Wales Ridge is no exception,where the natural Accepting the hypothesis that the communitiesare rapidly being destroyed Florida membersof Dicerandra(D. christmanii,D. by the expansion of the industry,and cornutissima,D. frutescens,and D. immaculata)are more recently,unbridled urban development. derived froma common ancestor,as suggested It is estimatedthat less than 10% of the original by the cladistic analysis, it follows that geo- Lake Wales Ridge scrubvegetation remains, with graphical isolation either through a fragmen- less than 3% of the total land of the endemic- tation of a widespread ancestral taxon or short rich southernLake Wales Ridge protectedfrom distance dispersal, possibly by water (see Huck development (Peroni and Abrahamson 1985). 1987), to "ecological islands" and subsequent For this reason, action to preserve the distinc- genetic differentiationgave rise to the four al- tive biota of the Lake Wales Ridge is essential lopatricFlorida species we see today.These pos- and additional systematicstudies of its endemic sibilitiesare comparable to what is believed to species are urgentlyneeded. Furtherstudy of have occurred in other genera having high Dicerandraand other scrub endemics should endemism in Florida (e.g., Polygonella,Polygo- provide considerable insightinto the evolution naceae, Horton 1963; Chrysopsis,Asteraceae, of the biota. Semple 1981). The perennial condition in Florida Dicerandra ACKNOWLEDGMENTS. Special thanksare due Ste- is seen as an adaptation to the xeric habitatlike ven P. Christman,field associate of theFlorida State that proposed for certain other central Florida Museum,who brought to our attention the distinctive 212 SYSTEMATIC BOTANY [Volume 14 species describedherein, mapped itsgeographic range, MADDISON, W. R., M. J.DONOGHUE, AND D. R. MAD- pointed out several of its most distinctive features, DISON. 1984. Outgroup analysis and parsimony. and provided the color photos representedin figure Syst. Zool. 33:83-103. 2. It is a pleasure to name this interestingspecies for NASH, G. V. 1895. Notes on some Florida plants. him, in acknowledgment of his contributionsto our Bull. TorreyBot. Club 22:141-161. knowledge of this species, and of scrub plants in gen- NEILL, W. T. 1957. Historical biogeography of pres- eral. We thank Steve Christmanand Daniel B. Ward ent day Florida. Bull. Florida State Mus., Biol. Sci. fortheir many comments.The authorsare gratefulto 2(7):175-220. Peter Alcorn and Bruce Hansen for their assistance PERONI, P. A. and W. G. ABRAHAMSON. 1985. Vege- in the field,and to Wendy B. Zomlefer forher illus- tation loss on the southern Lake Wales Ridge. tration(fig. 1). We thank the Florida Game and Fresh Palmetto 5:6-7. Water Fish Commission's Nongame Wildlife Pro- SEMPLE, J. C. 1981. A revision of the goldenaster gram,grant no. GFC-84-010,for support of S. Christ- genus Chrysopsis(Nutt.) Ell. nom. cons. (Com- man's scrub surveys that led to the discovery of this positae-Astereae). Rhodora 83:323-384. species. Financial supportfor the color plate was pro- SHINNERS, L. H. 1962. Synopsis of Dicerandra(Labia- vided by GFC-84-010; other publication costs were tae). Sida 1:89-91. supported by the Department of Botany, University SNEATH, P. H. A. and R. R. SOKAL. 1973. Numerical of Florida. We thank the curators of SMU and the .San Francisco: W. H. Freeman & Co. Archbold herbariafor their loans of comparativema- STEVENS, P. F. 1980. Evolutionarypolarity of char- terial,and Kent Perkins (Manager of SystematicCol- acter states.Annual Rev. Ecol. Syst. 11:333-358. lections,FLAS) forthe processing of specimen loans. . 1981. On ends and means, or how polarity criteriacan be assessed. Syst. Bot. 6:186-188. LITERATURE CITED SWOFFORD, D. L. 1985. PAUP: Phylogeneticanalysis usingparsimony, version 2.4. Urbana: Illinois Nat- ADAMS,P. 1962. Studies in the Guttiferae.II. Taxo- ural History Survey. nomic and distributionalobservations on North WARD, D. B., ED. 1979. Rare and endangeredbiota of American taxa. Rhodora 64:231-242. Florida,vol. 5, Plants. Gainesville: Univ. Presses ALMEIDA,M. T. and F. A. BiSBY. 1984. A simple meth- of Florida. od for establishing taxonomic characters from WATROUS, L. E. and Q. D. WHEELER. 1981. The out- measurementdata. Taxon 33:405-409. group comparison method of characteranalysis. HARDIN,J. W. 1974. Studies of the southeastern Syst. Zool. 30:1-11. United States flora.IV. . Sida 5:274-285. WATTS, W. A. 1975. A late Quaternary record of HARPER,R. M. 1949a. A preliminarylist of the en- vegetation fromLake Annie, south-centralFlor- demic floweringplants of Florida. Part I-Intro- ida. Geology 3:344-346. duction and historyof exploration.Quart. J.Flor- WHEELER, Q. D. 1981. The ins and outs of character ida Acad. Sci. 11:25-36. analysis: A response to Crisci and Stuessy. Syst. . 1949b. A preliminarylist of the endemic Bot. 6:297-306. floweringplants of Florida. Part II-List ofspecies. WHITE, W. H. 1970. Thegeomorphology of theFlorida Quart. J.Florida Acad. Sci. 11:39-57. peninsula.Geological Bull. 51. Tallahassee: Florida 1950. A preliminarylist of the endemic flow- Dept. of Natural Resources. ering plants of Florida. Part III. Quart. J.Florida WILEY, E. 0. 1981. Phylogenetics.New York: John Acad. Sci. 12:1-19. Wiley & Sons. HENDY,M. D. and D. PENNY. 1982. Branchand bound WISHART, D. 1987. Clustanuser manual,version 3.2. algorithms to determine minimal evolutionary St. Andrews, Scotland: Computing Laboratory, trees. Math. Biosci. 59:277-290. Univ. St. Andrews. HORTON,J. H. 1963. A taxonomicrevision of Polyg- WOODSON,R. E., JR. 1947. Notes on the "historical onella (Polygonaceae). Brittonia15:177-203. factor" in plant geography. Contr. Gray Herb. HUCK,R. B. 1987. Systematicsand evolution of Di- 165:12-25. cerandra(Labiatae). Phan. Monogr. 19:1-343. WUNDERLIN, R. P. 1984. Status report on Diceranda JAMES,C. W. 1961. Endemism in Florida. Brittonia frutescensShinners. Unpublished report pre- 13:225-244. pared forthe U.S. Fish and WildlifeService, Jack- JUDD,W. S. 1983. The taxonomicstatus of Stipulicida sonville, Florida. filiformis(Caryophyllaceae). Sida 10:33-36. ZONA, S. AND W. S. JUDD. 1986. Sabal etonia(Palmae): and D. W. HALL. 1984. A new species of Systematics,distribution, ecology, and compari- Ziziphus(Rhamnaceae) fromFlorida. Rhodora 89: sons to other Florida scrub endemics. Sida 11: 381-387. 417-727. KRAL,R. 1982. Some notes on Dicerandra(Lami- aceae). Sida 9:238-262. 1989] HUCK ET AL.: DICERANDRA 213

APPENDIXI. Associates of Dicerandrachristmanii. Endemics or near endemics (>90% of range in Florida) are indicated by an asterisk(*). Voucher specimens are deposited at FLAS and/or USF.

Amorphaherbacea Walter, Aristidapurpurescens Poiret, Asclepiascurtissii A. Gray*, A. humistrataWalter, A. tuberosaL. subsp. rolfsii(Britton) Woodson, Asiminaobovata (Willd.) Nash*, Balduina angustifolia(Pursh) B. Robinson, Befariaracemosa Vent., Berlandierasubaculis (Nutt.) Nutt.*, Bonamiagrandiflora (A. Gray) Heller*, Bulbostylisbarbata (Rottb.) Clarke, B. ciliatifolia(Elliott) Fern.,B. warei(Torrey) Clarke, Bumeliatenax (L.) Willd., Callicarpaamericana L., Caryafloridana Sarg.*, Ceratiolaericoides Michaux, Chamaecristafasciculata (Michaux) E. Greene, Chapmanniafloridana Torrey & A. Gray*, pygmaeus Small*, Clitoriafragrans Small*, Cnido- scolusstimulosus (Michaux) Engelm. & A. Gray,Commelina erecta L., Crataeguscf. lepidaBeadle, Cuthbertiaornata Small*, Cyperuslecontei Torrey, C. retrorsusChapman, Dalea feayi(Chapman) Barneby,Eriogonum longifolium Nutt. var. gnaphalifoliumGandoger*, Froelichiafloridana (Nutt.) Moq., Galactia volubilis(L.) Britton,Garberia heterophylla(Bartram) Merr.*, Gaylussacia dumosa (Andrews) Torrey& A. Gray,Helianthumum nashii Britton*, Heterothecasubaxillaris (Lam.) Britton& Rusby,Indigofera carolina Miller, Lecheadeckertii Small, L. sessilifloraRaf., Liatrisohlingerae (S. F. Blake) B. Robinson*,L. tenuifoliaNutt., Licania michauxii Prance, Linaria floridana Chapman, Lupinusdiffusus Nutt., Lyoniaferruginea (Walter) Nutt., L. fruticosa(Michaux) G. S. Torrey,L. lucida (Lam.) D. Don, Matelea publiflora(Decne.) Woodson, Nolina brittonianaNash*, Opuntiahumifusa (Raf.) Raf.,Palafoxia feayi A. Gray*, Fern.*, P. herniarioides(Michaux) Nutt.,P. patulaShinners, Persea humilis Nash*, (Chapm. ex Engelm.) Vasey ex Sarg.*, P. elliottiiEngelm., Pityopsisgraminifolia (Michaux) Nutt., Polanisiatenuifolia Torrey & A. Gray,Polygonella basiramia (Small) Nesom & Bates*,P. gracilis(Nutt.) Meissner, P. myriophylla(Small) Horton*, P. polygama(Vent.) Engelm. & A. Gray, P. robusta(Small) Neson & Bates*, Prunusgeniculata Harper*, Quercuschapmanii Sarg., Q. geminataSmall, Q. inopinaAshe*, Q. laevis Walter, Q. myrtifoliaWilld., Q. virginianaMiller, Rhuscopallina L., Rhynchelytrumrepens (Willd.) C. E. Hubbard, Rhynchosia cineriaNash*, Rhynchosporamegalocarpa A. Gray,Sabal etoniaSwingle ex Nash*, Schizachyriumsanguineum (Retz.) Alston, Schrankiamicrophylla (Dryander ex Smith) J. F. Macbr., Selaginellaarenicola Underw., Serenoarepens (Bartram)Small, Seymeriapectinata Pursh, Sisyrinchiumxerophyllum Greene*, Smilaxauriculata Walter, Stillingia sylvaticaL., Stipulicidasetacea Michaux, Stylismaabdita Myint*, Tephrosia chrysophylla Pursh, Tillandsiarecurvata (L.) L., T. setaceaSw., T. usneoides(L.) L., Tradescantiaroseolens Small, Tragiaurens L., Trichostemadichotomum L., Vacciniumdarrowii Camp, V. stamineumL., Vitisrotundifolia Michaux, Yuccafilamentosa L., and Ximeniaamericana L.

APPENDIXII. Dicerandrafrutescens specimens examined as part of this taxonomicstudy (in addition to those cited by Huck 1987).

UNITEDSTATES. FLORIDA. Highlands Co.: ca. 200 yd N of Highway Patrol Station on US hwy 27, just SW of Lake Placid Medical Center at jct of Tomaka Blvd. and US hwy 27, just N of Lake Placid (town) (pop. 6), Alcorn184 (FLAS); Lake Placid (pop. 6?), Brass 15604 (Archbold); Brass 33333 (Archbold); (pop. 6), Judd5550 (FLAS, FSU, USF); T37S, R29E, Sect. 13, S on US hwy 27 ca. 1.5 mi fromjct of US hwy 27 and Rt. 621, then W on Lake MirrorRd. 1 mi, then SE on Placid View Dr. 0.6-0.7 mi, just NW of Lake Placid (lake) and S of Lake Placid (town) (pop. 8), Judd5553 (A, F, FLAS, MO, NY, SMU, USF); T38S, R30E, Sect. 16, ca. 2 mi S on US hwy 27 fromjct of US hwy 27 and Rt. 70, on W side of rd. by abandoned gas station,S of Lake Placid (pop. 13), Judd5555 (FLAS, FTG, NCU, RSA, US, USF); T37S, R30E, Sect. 34, E on Rt. 70 1.3 mi fromjct of Rt. 70 and US hwy 27, then N on highlands Blvd. 0.9 mi, just N of East Orange St., SE of Lake Placid (pop. 11), Judd5557 (FLAS, NCU, SMU, USF); T37S, R30E, Sect. 28, 1.1 mi N on US hwy 27 fromjct of Rt. 70 and US hwy 27, on E side of rd., SE of Lake Placid (pop. 10), Judd5562 (FLAS, US, USF); ca. border of T37S, R30E, Sect. 8 & 17, fromjct of US hwy 27 and Co. Rt. 29, E on Rt. 29 0.2 mi, "Y.M.C.A. site" (pop. 9), Judd556 (FLAS, MO, USF); T36S, R30E, Sect. 29-32 border,Lake Placid (town), Highland Park Estates,0.5 mi E of jct. of Lake Clay Rd. & Central Ave. on Central Ave., vacant lot near home of B. Cavander (pop. 7), Judd5569 (A, Duke, F, FLAS, FSU, NY, SMU, US, USF); (pop. 8), Skean2130 (FLAS, NCU, USF); (pop. 11), Skean2149 (FLAS); (pop. 10), Skean 2153 (FLAS); (pop. 7), Skean 2165 (FLAS); Archbold Biol. Station, Tract 18, W545, 8 mi S of Lake Placid, (pop. 12), 7 Oct 1987, VanderKloets.n. (Archbold), Tract 30619, (pop. 12), 15 Jan 1987, VanderKloets.n. (Archbold).

Note: Rapid destructionof the vegetation of the Lake Wales Ridge is underscored by the factthat, while this paper was in press, population 9 of D. frutescenswas destroyed by urban growth. Population 1 of D. christmanii,the only population of significantsize that is not subdivided, is scheduled for conversion to a citrus grove within the year. Continued survival of D. christmaniiand other Lake Wales Ridge endemics appears doubtful.