A New Subgenus and Fifteen New Species of Echeandia (Anthericaceae) from Mexico and the United States Author(s):

Robert William Cruden Reviewed work(s): Source: Novon, Vol. 9, No. 3 (Autumn, 1999), pp. 325-338 Published by: Missouri Botanical Garden Press Stable URL: http://www.jstor.org/stable/3391730 . Accessed: 26/09/2012 15:13
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A New Subgenus and FifteenNew Species of Echeandia (Anthericaceae) fromMexico and the United States
Robert William Cruden Department of Biological Sciences, Universityof Iowa, Iowa City,Iowa 52242, U.S.A.

ABSTRACT.Twosubgenera in Echeandia (Anther- have a cormthatproducesnew storagerootsand OtherAnthericaceae are rhiicaceae), i.e., subg.Echeandia and subg.Mscavea, inflorescences yearly. are recognizedbased on time of flower and onlytheterminal opening, zomatous, segment produces There is no evidence color,and habitatdifferences. leaves and inflorescences. tepal shape, flower Fifteenspecies are proposed.One, E. texensis, is thata segment inflorescences in succesproduces based on fourcollectionsmade near Brownsville, sive years.Second, mostgenerain Anthericaceae The s. str.have smooth Texas, and is surelyrare,possiblyextirpated. and/or e.g., papillatefilaments, offivespecies clarifies thedistribution- Chlorophytum recognition s.1. (Kativu & Nordal, 1993) and al rangeofE. reflexa, a common species ofeastern Hagenbachia (Cruden, 1987), whereas most Mexico.Sevenspecies are endemic to Guerrero and Echeandia (63/78)have scaled filaments and those adjacentareas ofMichoacin,Mexico,Morelos, and/ withsmooth filaments occurin derivedgroups. or Oaxaca. A keyto the species ofsubgenus MscaIn addition,anthersadapted forpollination by species of subgenus vibratory bees occur in both lineages and are vea and white-flowered Echeandia is provided. s. str.In the white-flowunique in Anthericaceae ered lineage the anthersin 21 of 25 species are a cone,and in theother 4 species Echeandia, as describedby Ortega(1800: 135, connateand form the for buzz individual anthers are adapted pollitab. 18), includedNew World connate species with lineage, anthers and scaled filaments. These traits effective- nation(see below). In the yellow-flowered Echeandia and Old World Anthericum, 19 of 53 species have anthercones and in 4 adly separated anthers are adapted whichhad freeanthers and smooth or papillatefil- ditionalspecies the individual for buzz The filaments of all thesespepollination. aments.Withfew exceptions, New Worldspecies within a basipetally dorsally openwithfreeanthers were includedin Anthericum for cies are inserted which holds the anther on ing pocket, effectively thenext180 years.However, that time, during speand probably provides cies with smooth filamentswere included in thesame axis as thefilament needed to supporta bee stability Echeandia (see Weatherby, 1910), and specieswith the structural scaled filaments were described in Anthericum while it vibratesan antheror anthercone. It is constitute a parallel (Baker,1876; Greenman, 1898). Thus,forthepast possible thatconnateanthers In other Echeandia and mostother Anand apomorphy. centuryor more, New World Anthericum anthers are the versatile, thericaceae, free, usually Echeandia wereseparatedsimply on the natureof In most Chloroand dehisce laterally. theiranthers,and the unique traitsthey shared dorsifixed, more the filaments are inserted however, phytum, wereignored. Because mostNewWorld despecies the scribedin Anthericum and Echeandia share traits or less basally in a deep pit, whichrestricts ofthe anthers. thatare uniquein Anthericaceae, a corm movement including relative toEcheandia'sputative ancestor, Finally, and scaled filaments, they are now included in of primitive and Echeandia (see Cruden,1987, 1994), which,with bothlineages combinea mixture that 78 describedspecies, is the second largestgenus derivedtraits.Based on otherAnthericaceae, had whiteflowers, whichprobably opened in Anthericaceae (see Chase et al., 1996; Dahlgren ancestor in the et al., 1985: 186). Excluded from morning, early broadlyelliptical Echeandia are relatively and versatile anthers that filaments, 20 white-flowered South American tepals,smooth approximately are apoThus, yellowflowers filaments and freeanthers that dehisced laterally. species withsmooth in one lineage,as are narrowly an undescribed constitute morphic elliptical probably genus. in the other. flowers Within Echeandiathere are twodistinct lineages, innertepals and late opening witha and each includes species withfree anthersand Thus, the available data are consistent others with connate anthers. The twolineagesshare common ancestor diverginginto two lineages, at least two synapomorphies. belowas subgenera. First,all Echeandia whichare formally recognized

NovoN 9: 325-338. 1999.

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Echeanof species in subgenus Table 1. Distribution The twolineagesshareseveralsynapomorphies and of elevation. A as a function dia Mscavea and subgenus are separated of a number Further, apomorphies. by if mostof its thetwolineagestendto occurin different climatic species was assigned to a given category The range rangewas includedin thatcategory. zones (see below). I include the two lineages in elevational below from oftwospecies in Echeandia extended slightly Echeandia as subgenerabecause the differences 800 m to 2400 m. as beinghalfin one Theywerecounted thatseparatethemdo notappear to be as greatas and halfin the other category. category those separatingothergenera in Anthericaceae. a moredetailedexamination ofthe relaHowever, 1500above Below 800800 m 1500 m 2400 m 2400 m tionshipsamong genera in Anthericaceaeusing othertechniquesmight lead to a different conclu8 30 9 6 Subg. Echeandia sion. 0 12 5 8 Subg. Mscavea SubgenusMscavea is proposedto include species in the white-flowered lineage, whose flowers or in the afternoon, and open late in the morning have narrowinner tepals. This subgenushonors long capsules, whereasjust 2 species in subgenus MarionS. Cave, eminent and student Mscaveahave oblong capsules,and 17 have subgloembryologist ofthe Liliaceae s. lat.,who stimulated oblongcapsules. myinterest bose to broadly in MexicanLiliaceae in generaland Echeandia in Relativeto species in subgenusMscavea,those elin subgenusEcheandia tendto occurat higher particular. evations (Table 1; = 19.84, p<0.001). Mostspe,Echeandia (>70%) occur comEcheandia subg. Mscavea Cruden, subg. nov. cies in subgenus above 1500 m, and nineoccur TYPE: Echeandia mcvaughiiCruden,Contr. pletelyor primarily Univ. MichiganHerb. 16: 129. 1987. [Illus- above 2400 m,whereas80% ofthespecies in subbelow orprimarily genusMscaveaoccurcompletely in McVaugh, trated 1989: 187.] 1500 m, and only fourreach 2400 m. It is not AbEcheandia Echeandia interioribus an- obviousthata disproportionate subg. tepalis of species number raro albisvelcremis floribus flavis, guste ellipticis aper- in Echeanto Mscavea relative subgenus subgenus ientibus antevelpost differt. meridiem = 4.93, p<0.05). In dia occur below 800 m The twosubgenera in timeofflower differ subgenusMscavea occur open- essence, mostspecies in (, withsubtropical to warm-temperand color,capsule shape, in drierhabitats ing,tepal shape, width, and altitudinal distribution. The flowers of species ate climates,whereas most species in subgenus with mesic habitats in Echeandia subg.Echeandia open shortly before Echeandia occur in relatively orafter climates. to cold-temperate sunup.All but1 of53 species havebroadly warmis notequivof connateanthers The occurrence ellipticalinnertepals. In the 40 species forwhich I have reasonabledata, the minimum widthofthe alent in the twosubgenera. Only 19 of 53 species cominnertepal in 27 species was 4.5 mm,and theme- in subgenus Echeandia have connate anthers, dian widthexceeded 4.5 mmin 36 species. Most pared to 21 of 25 species in subgenusMscavea. ofthe species (44/53)are yellow-flowered, and six Because mostspecies in subgenusMscavea have thatspecies one might include white-flowered anticipate populations.Nine species connateanthers, In contrast, are white-flowered. in subgenus Msca- with connate anthersoccur at lower elevations. betweenoccurthereis no relationship or the However, open late in the morning vea, the flowers 24 of25 species have narrowly afternoon, elliptical rence of connateanthersand elevationin either = 1.18, p>0.5) or subinnertepals,and, in 22 of 23 species forwhichI subgenusEcheandia have reasonabledata, the maximum widthof the genusMscavea = (, 0.09, p>0.975). (, the innertepal was 4.5 mm.Mostof the species have species withconnateanthers, Amongthose withrespectto the insertion white flowers flow- two subgeneradiffer (22/25),twohave cream-colored dorsaland hidden whichis either and one, possibly ers,one has orangeflowers, two, ofthe filaments, the anthersacs and or between within a species include orange-or yellow-flowered pocket populations. essentiallybasal. In most species of subgenus the betweenthe Mscavea (12/20 examined),the distancefrom significantly Capsule shape differs twosubgenera (f = 13.91, p>0.001; analysiswith loweredge of the pocketto the base of the anther than in 18 of 19 species in subgenus a contingency table and Chi-square test).Forthose is shorter Echeandia for whichthere are Echeandia examined (0.1-0.6 mmvs. 0.6-2.0 mm; species in subgenus = 13.137, p<0.001). There is no correlation at leasttwice data,26 have oblongcapsules (length , thelower and distancefrom anther the width)and 16 have subgloboseto broadly ob- between length

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edge of the pocket to the base of the antherin in six species ofsubgenus Mscap>0.05). Finally, vea butjust one of subgenusEcheandia, insertion ofthefilaments is basal or essentially so. the anthers of Amongspecies withfreeanthers, all fourspecies in subgenusMscavea dehisce apically comparedto 4 of 33 species in subgenus Echeandia. In threespecies of subgenusMscavea, the line of dehiscenceis fused1/?2/3 the lengthof the anther, and dehiscenceis through a relatively large teardrop-shaped opening.In one species of species ofsubgenus subgenus Mscaveaand thefour Echeandia, the line of dehiscence is completely a much smaller open, and dehiscence is through apical opening.
KEY TO MEXICAN SUBGENUS MSCAVEA AND SUBGENUS ECHEANDIA

either subgenus Echeandia (r = 0.289, n = 19, p>0.05) or subgenus Mscavea (r = 0.237, n = 20,

WHITE-FLOWERED

In any couplet,a clause precededby "if" takes precethat dence overother clauses. Leafshapefollows proposed AssociationCommittee (1962). The by the Systematics namesof species describedin thispaperare notfollowed name. The subgenusof each species is by the author's given:(E) = subg.Echeandia,(M) = subg.Mscavea.See also keysin McVaugh (1989) and Cruden(1994). 1. Anthers free .................. 2 ... 3 1. Anthers a cone ....... connate, forming 4 Filaments smooth............ 2(1). 2. 7 Filaments scaled ............ . smooth ................. 20 Filaments 3(1). 3. Filaments scaled .................. 26 AnthersFree, Filaments Smooth or glabrous .. 5 scattered enations 4(2). Scapes with or scabrous,at least in lower 4. Scapes hirsute ........... 6 half,frequently throughout notversatile, line of dehis5(4). Anthers cence partially fused,bases flared; tepals narrowly elliptic, 6-8 mm long(below1200 m,Jalisco) .... Anthers dehiscencelaterversatile, elal, bases straight; tepals broadly liptic,8-13 mm long (above 1600 to m, Durangoand San Luis Potosf Mexico) ................. E. durangensis (Greenman)Cruden (E) Basal leaves narrowly linear,veinspapillate 6(4). rebelow, marginsciliate, if tips strongly curved(1800-1900 m, Oaxaca, Puebla) E. parva Cruden(E) .................. ovate to narrowly 6. Basal leaves narrowly elveins smooth enbelow,margins liptic,flat, tireor nearlyso, tips straight above (mostly 2000 m, San Luis Potosi to Veracruzand Puebla W to Michoacin) ............ E. nana (Baker)Cruden(E) .............. AnthersFree, Filaments Scaled 7(2). Eastern Mexico (Tamaulipas,San Luis Potosi, Veracruz,Tabasco, Campeche) below 8 500 m ........................ 5.
...... E. pihuamensis Cruden (M)

at highereleElsewherein Mexico and/or 9 vations ................... ..... Flowerswhite(Tamaulipas) . 8(7). E. tamaulipensis (E) ............. Flowers yellow-orange 8. (Campeche) E. campechiana Cruden(M) ........ Basal leaves 2-6 mmwide, 5-15 cm long, 9(7). long ciliate, veins on lowersurface margin bearinglong hairs; storage papillate and/or thecorm(Oaxaca) ... roots4-10 cm from Cruden(E) E. confertiflora ............... 9. If mostbasal leaves wideror longer, margin betheveinssmooth to ciliate,and/or entire low; if storagerootsenlarged1-2 cm from 10 thecorm ...................... 10(9). Anthers dehisce apically; anther but maybe open walls notreflexed along line of dehiscence and the at apex; mostmorethan wall flared 11 3 mmlong ................ 10. Anthers dehisce laterally; anther or twisted if anthers walls reflexed, mostless than3 versatile; obviously mm long (sometimes longerin E. 14 (E)) (M) and E. durangensis gentryi whitewhen 11(10). Cauline leaves 7-11, becoming dry; storagerootsclose to the corm;basal leaves morethan20 mmwide (Jalisco) .. Cruden(M) .E. robusta ................. 11. Cauline leaves 1-6, if more,then storage thecorm;ifbasal leaves roots 4-10 cm from 12 less than20 mmwide ............. 12(11). Tepals 7-8 mmlong; line of dehisfused(Oaxaca,Puebcence partially E. mirandaeCruden(M) la) ......... 12. Tepals 11-21 mmlong; line of de13 hiscencecompletely open ...... 13(12). Ovaries 3-4 mmlong (Puebla) ....... Cruden(E) E. michoacensis (Poellnitz) ....... 13. Ovaries 2-3 mmlong (Chiapas) ...... E. matudaeCruden(E) ................. weaksometimes 14(10). Scapes pubescent, .15 ly so or onlynearthe base ...... .16 14. Scapes glabrous............. corm; 15(14). Storagerootsenlarged1--4 cm from ofbasal leaves ciliatetolongciliate; margins the base at least toward scape scabrescent, (Chihuahuaand NayaritSE to Michoacin) Cruden(E) E. scabrella(Bentham) .......... 15. corm; Storagerootsenlarged3-8 cm from marginsof basal leaves denticulate;shoot half hairsin lower minute scattered with (OaE. sp. (E) xaca) ...................... basal 9-15 mmlong;most 16(14). Filaments leaves 1-6 mmwide; if stylescaled Mixico, Morelos) ... (Guerrero, Cruden(E) E. tenuis (Weatherby) .... less than8 mmlong;or if Filaments 16. mostbasal leaves morethan6 mm ..17 wide ................... linear 17(16). Basal leaves 1-7 mmwide, narrowly moststorage to linear,margins denticulate; rootsenlarged3-8 cm from corm;capsules < twice to broadly oblong(length subglobose the width)(Oaxaca, Puebla) ......... & Galeotti (E) gramineaMartins .........E. 7.

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17.

If most basal leaves 7 or more mm wide; rootsenlarged1-3 cm from corm;if storage capsules oblong (lengthat least twice the 18 width) ......................... 18(17). Basal leaves 1-6 mmwide,narrowly in a pockinserted linear;filaments ovaries1.8-2.5 mm et,notversatile; Sinaloa, Dulong (Jalisco,Nayarit, .E. gentryi Cruden(E) rango) ....... 18. If mostbasal leaves 7 mmor more
wide; ovaries 2.5-4 mm long . . . . 19

cauline leaves 2-6 (Jalisco, Nayarit) 25.

.......

linearto narrowly ovate;an19(18). Leaves broadly thersversatile(above 1600 m, mts.of Durangoand San Luis PotosiS to Mexicoand 19.

Hidalgo) .. E. durangensis(Greenman) Cruden (E)

oblong,6-10 mm Capsules broadly long; basal leaves 7-13, if most more than 15 mm wide; cauline leaves (4-)5-10 (Sonora, Chihuahua, to Guerrero) ........... (Presl)Cruden(M) ....E. ramosissima AnthersConnate, Filaments Scaled 26(3). Eastern Mexico (Tamaulipas,San Luis Potosi, Veracruz,Campeche, Yucatan) below .27 cream-colored 1600 m; ifflowers ....... in Mexicoor above 1800 m . . . 29 Elsewhere 26. (Campeche, 27(26). Flowers cream-colored QuitanaRoo, Yucatan,Oaxaca) ... 28 white .................. 27. Flowers ofthe 7-9 mmlong,twicethelength 28(27). Anthers ciliateto longciliate leaf margins filaments; (Tamaulipas,San Luis Potosf)....... 28. thanthe 4-7 mmlong,barely Anthers longer leaf margins filaments; denticulate to ciliate E. albflora (Schlechtendal & (Veracruz)..... Chamisso)Martins (M) & Galeotti cones 1.2-3.5 mm 29(26). Apexes ofanther cones2.5-4.0 mmwide wide;anther and 4-5(-7) mmlong; basal leaves 2-7 mm wide, veins on lowersurface noticeablypapillate (Oaxaca) 29.
......... E. conzattii Cruden (E) ................ E. macrophylla Rose (M) ............ E. luteola Cruden (M)

E. parvicapsulata Cruden (M)

Leaves linear, not falcate;anthers frequently versatiledue to reflexed walls. (Guanther

anajuato) ...........

AnthersConnate, Filaments Smooth 20(3). Basal leaves abruptlyexpanded near the middleintoa broadblade, appearing petio20.
late (Guerrero) .......

E. sanmiguelensis (E)

abruptly Width of basal leaves notchanging ...............................21

E. pseudopetiolata (M)

beyond first cauline 21(20). Scapes scabrous ofleaves ifbasal margins leafand/or 22 ciliateto longciliate .......... 21. Scapes glabrousor rarelyscabrescentnearthe base ........... 25 22(21). Scapes usuallymorethan1 m high,cauline leaves 6-15; basal leaves 15-40 mm wide to short entire cilnarrower), margin (rarely Moreiate; tepals 9-13 mmlong (Guerrero, ..E. mexiae(M) los) ................... 22. 1 m high;ifcauline leaves 0Scapes rarely basal leaves less than10 mmwide, 4, ifmost theirmargins ciliate to long ciliate, and/or 23 tepals less than9 mmlong ........... 23(22). Basal leaves 2-11 mm wide, most less than 10 mmwide, flat, margin entire to denticulate;tepals 8.510.5 mm long; anthers(1.8-)2-3 times the lengthof the filaments; scapes 15-40 cm high (below 500 m, Jalisco,Sinaloa) .......... E. sinaloensis Cruden(M) ......... If most 23. ofthebasal leaves more than 10 mm wide, falcate, their tips or themargin cilrecurved, strongly iate to longciliate;iftepalsless than 8 mmlong;anthers 1-1.6(-2) times of the filaments; if scape the length morethan50 cm high......... .24 24(23). Basal leaves 6-12 mm wide, verynarrowly linearto very falelliptic, narrowly frequently withstrongly cate and/or recurved tips,freundulate, quently margins denselyciliate to denticulate or short longciliate(rarely ciliate) ColimaSE to Oaxaca, Chiapas,and (Jalisco, 24.
Guerreroto Veracruz) ...E.

If apexes of anther cones less than 1 mm wide, or anthercones less than 2 mm wide; if basal leaves mostmore than smooth belowand/or 10 mm wide; if storagerootsen.30 corm....... larged1-3 cm from 31 30(29). Mostbasal leaves less than5 mmwide ... Mostbasal leaves morethan5 mmwide .. 34 30. 3 cm or more rootsenlarged 31(30). Storage from corm;anthers1-1.5 timesthe .32 ofthefilaments length ......... 31. Storagerootsenlarged0.5-2.0 cm 1.8-2.2 times from corm;if anthers .33 ofthe anthers ....... the length elliptic,5-7 mmwide,11.532(31). Tepals broadly 15 mm long; filaments (4-)5-7 mm long (Guerrero, Mdxico,Morelos,Oaxaca) ... 32. elliptic, 1.5-3 mm wide, Tepals narrowly 2.5-4 mmlong 7.5-12.5 mmlong;filaments
.... .......... . ................... .... E. taxcana (E)

33(31). Anthers1-1.6 times the lengthof the filaments; capsules 7-11.5 mm long (above 800 m, Nayarit) ... 33.
.........

E. parviflora Baker (M)

of Anthers 1.8-2.2 timesthe length the filaments;capsules 5-7 mm Sinalong. (below 500 m, Nayarit,
loa)........... .E.

E. flexuosa Greenman (E)

Basal leaves (8-)13-28 mmwide,flat, narrowlyovate to narrowly elliptical,margins to minutely entire denticulate (below800 m, E. atoyacana (M) Guerrero, Mexico)......... 25(21). Capsules globose,2.5-4 mm long; basal leaves 4-8, 6-15 mm wide;

parvifloraBaker (M)

the base, at least toward 34(30). Scapes pubescent, .35 so ........... and sometimes sparsely 39 34. Scapes glabrous .................. longciliate,hairsusu35(34). Leaf margins ally 0.2-0.5 mm long; scapes usubashairslongand flat; ally hirsute, al leaves (10-)15-40 mm wide

nayaritensis (M)

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Mexico,E Michoacin) (Guerrero, 35.

...............

Leaf margins to ciliate,rarely entire scabreslong ciliate; scapes mostly cent; if mostbasal leaves 2-8 mm wide .................... 36 ciliate to long ciliate, some36(35). Leaf margins timesdensely so; basal leaves 2-8 mmwide, falcate; anthers3-7 mm long, frequently most 1.2-1.8 times the lengthof the filaments;tepals 7.5-12 mmlong;basal leaves 1-8 mmwide ....... .E. parvifora Baker(M) 36. Leaf marginsentireto shortciliate, rarely basal leaves more than10 mm ciliate;ifmost 7.5-12 mmlongand/or 1.8wide; ifanthers 2.2 times thelength ofthefilaments; iftepals morethan12 mmlong .............. 37 37(36). Storagerootsenlarged4-8 cm from corm;if lowerleaf surface minutely papillate between the veins; capsules 7-9 mmlong(Guerrero) . . (M) ...................E. hintonJii 37. Storagerootsenlarged1-2 cm from smooth becorm;lowerleaf surface tweenthe veins; if capsules shorter or longer.................. 38 cones 6-11 mmlong,usuallywidest 38(37). Anther in the middle;if anthers1.8-2.2 timesthe lengthof the filaments; capsules 4-7 mm long; basal leaves 5-10, frequently falcate, if mostmorethan 15 mmwide and/or less than 25 cm long (Jalisco,Nayarit) . 38. Anther cones 5.5-7 mm long, 1-1.4 times thelength ofthefilaments, widest tousually wardthebase; capsules9-12 mmlong;basal leaves 3-5, narrowly 6-12 elliptic,straight, mmwideand more than30 cm long(Oaxaca) E. oaxacana Cruden(M) .............. rootsenlarged 3-8 cm 39(34). Moststorage from corm;anthercones widesttowardthe base, taperedgradually to the apex, usually 1-1.7 times the ofthefilaments .40 length ......... 39. Moststorage rootsenlarged1-3 cm from cones widestin corm;ifanther middle and/or 1.8-3 times the length of thefilaments ......... 41 cones 4-8 mm 40(39). Tepals 9-13 mmlong;anther ofbaslong;caulineleaves 1-5; ifunderside al leaves minutely papillate betweenveins (Guerrero) ...............E. hintonii (M) cones 8-12 40. Tepals 13-20 mmlong; anther mmlong;cauline leaves 4-10; basal leaves smoothbelow (Guerrero, Morelos) ..... E. elegans(M) ...................... cones 41(39). Tepals 9-11 mmlong;anther 5.5-8 mmlong,usuallywidestnear the base and graduallytaperedto the apex; basal leaves narrowly linear,30-45 cm long,5-14 mmwide, not falcate (Michoacin) ....... 41. cones Tepals 10-19 mmlong;anther 6-12 mm long, if cones widestin middle;if mostbasal leaves falcate,
...... E. coalcomanensis Cruden (M) ............E. mcvaughii Cruden (M)

E. hirticaulis (M)

less than 20 cm long,and/or more than15 mmwide ............. 42 linear, 42(41). Basal leaves 4-8 mmwide,narrowly ovaries1-2 mmlong(below600 m, straight; E. nayaritensis Nayarit, Sinaloa) ........ (M) 42. Basal leaves 10-30 mmwide,mostnarrowly obovateor narrowly falelliptic,frequently cate; ovaries2-3 mmlong (above 1200 m,
Jalisco, Nayarit) . . . . E. mcvaughii Cruden (M)

Fifteen below:eight in subspecies are proposed Mscavea. genusEcheandia and seven in subgenus The species in subgenus Echeandiawith connate antherscomprisethe taxonomically mostdifficult groupin the genusand includeE. reflexa (Cavanilles) Rose, the typespecies ofthe genus.Because thisspecies is quite variableand has a broadgeographicalrange,and giventhe lack of recentsystematic it is notsurprising thatspecimens analysis, with connate anthers from eastern and central Mexwereincluded, indiscrimico, as wellas elsewhere, in E. reflexa. I propose E. pseudoreflexa and inately, E. texensis to include such material from Chiapas (Cruden, 1994) and Texas (see Johnston, 1990), reE. palmeriis proposed to includespecspectively; imensfrom Durangoand Chihuahua.OnlyE. pseudoreflexa appearsto be closelyrelatedto E. reflexa oftwoadditional (see below).The description species shouldclarify thedistribution ofEcheandia in easternand centralMexico. Echeandiafalcata is to includeplantsfrom and GuQueritaro proposed broadanther short, cones, anajuatowithrelatively and E. taxcana is proposedto included narrowleaved plants withstrongly taperedanthercones from and Mexico. Oaxaca, Guerrero, Morelos, Withthe recognition ofthesespecies,E. reflexa is a morecoherent, albeitvariable, found entity primarilybetween200 and 3000 m on the eastern slopes of the Sierra Madre Orientaland adjacent from cenpartsofthe SierraVolcinicaTransversal tralNuevoLe6n and southern to southTamaulipas ern Puebla, adjacentOaxaca, and Veracruz. Populationsfrom low elevations are usuallyrobust, with and long, broad highlybranchedinflorescences, leaves. At mid elevations, the plants are straight less robust with fewer branches and narrower, shorter the leaves leaves; and at high elevations, are usually narrow and frequently falcate.Speciin Puebla and adjacent mensfrom elevations higher Veracruz and Oaxaca have falcate,undulatebasal leaves and resemblesmall specimensof E. paniculata Rose. FlowersofE. reflexa have a long (69 mm), anther slender, cone,which strongly tapered lobed apex. usuallyhas a small,minutely

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,*
O * *
U

E. pseudoreflexa E. texensis E. palmeri

E. falcata

AA E. taxcana *
.+ .*

E. attenuata
E. sanmiguelensis

a.V

E. tamaulipensis

SO

-0

Figure 1.

Distributional ranges of the eight new species in subgenus Echeandia are illustrated.

Anthers 5-7.5 mmlong,equal to orslightconnate, the cones strongly taly longerthanthe filaments; to shallowlyand pered, apex barely emarginate lobed 0.5-0.8(-1.2) mmwide.Ovaries2.5broadly 4 mmlong.Capsules oblongto narrowly 9oblong, 14.5 mmlong,4.5-5.5 mmwide.In flower AugustDecember.Chromosome numbern = Radicibus a cormo 1-3 cm,foliis basalibus September, penariis 16 (Cruden 1936). linearibus vel 7-12 mm latis anguste anguste ellipticis denticulatis ad breviciliatis, temarginibus scapis glabris, filamentis antheris connatis 5-7.5 Withthe exception squamosis, palisflavis, ofmaterial collectednearLa mm conis antherarum ad apicem valdecontractis.Cidnegade Le6n, this species is knownonlyfrom longis, 1100 and 1800 m (Fig. 1). Chiapasbetween rootsenlarged1-3 cm from corm.Basal central Storage dominated bysomecombination leaves 5-8, narrowly linear to narrowly elliptical, It occursin forests to short of pine,oak, and Liquidambar. 34-64 cm long,7-12 mmwide,denticulate Materialof E. pseudoreflexa was includedin E. ciliate. Scapes glabrous,0.6-1.5 m high, 2-6in Flora Mesoamericana reflexa (Cruden, 1994) beaxis with 12-20 branched,major flower-bearing nodes. Cauline leaves (0-)2-6, lowest17-24(-32) cause of its robustshootsand leaves and strongly cone. It is distinguished from most cm long,usuallyexceedingthenextnode. Flowers taperedanther of E. reflexa nutant. broad- populations by storagerootsthatare yellow, Tepals 11-16 mmlong,inner basal ly elliptic5 mmwide,outerelliptic2.5 mmwide enlarged 1-3 cm fromthe corm,narrower scaled, narrowly leaves, and the bases ofthe previous (one flower examined).Filaments year'sleaves dorsalin a pocket. forming a fibrous collar.The storage rootsofE. reclavate,5-7 mmlong,insertion

1. Echeandia pseudoreflexa Cruden,sp. nov. TYPE: Mexico. Chiapas: road to Venustiano ca. 3 km SW ofSanta Rosalia, hillCarranza, side with Convolvulus, Calliandra, Lantana, Dahlia, 1200 m, 3 Sep. 1971, Cruden1936 UC; isotypes, MEXU, MO). (holotype,

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cauline leaves thatonlyoccasionally overlap, the corm, fewer flexa are usuallyenlarged3-6 cm from and the previous year'sleaf bases remainmoreor and anthersthat are usually equal to, or longer is easilydisEcheandiatexensis thefilaments. The long,lower caulineleafdistinguish- than, less intact. fromE. reflexaby the short,usually ofE. formosa tinguished es thisspecies from manyspecimens lobed apex. cone witha broadly broad anther Cruden. (Weatherby) Paratypes.MEXICO.Chiapas:Ruta190, 58 kmN vicinity County, Paratypes.U.S.A.Texas: Cameron La Trinitaria, 1350m,19 Sep.1988, Brownsville, El Jocote, 7 mi.from 4082 (TEX); 10 m, 25 Oct. 1945, Runyon
& Martinez 25767 (MO); Mpio. Cintalapa,near Stevens & Almeda48171 (CAS); 1230 m, 1 Dec. 1980, Breedlove

1080de Le6n, near La Ci6nega border, Oaxaca-Chiapas

4036 (TEX); Lomadel Cayo,29 Nov. Brownsville, Runyon & Tharp38b-1 (TEX). 1951, Johnston

3 kmNW Pueblo Mpio.PuebloNuevoSolistahuacin, 28 Aug.1970,Zwill293 3. Echeandia palmneri Cruden, sp. nov. TYPE: 5800ft., Nuevo Solistahuacin, 7500ft., barrio Matsab, Tuk, paraje (DS); Mpio. Tenejapa, Mexico. Durango:SantiagoPapasquiaro,Apr. 12552 (DS); Mpio.Teopisca, 30 Sep. 1965,Breedlove & Aug. 1896, Palmer410 (holotype, US; iso5900 ft.,19 Aug. 1966, of Teopisca, limits southern F, MO, GH, NY). types, 15049 (DS, F). Breedlove
2. Echeandia texensis Cruden, sp. nov. TYPE: Radicibus penariisa cormo2-4 cm, foliisbasalibus denticulatis 1-7.5 mmlatismarginibus linearibus anguste clavatis filamentis ad ciliatis,scapis glabris, tepalisflavis, 3.5-7 mmlongisad apicem squamosis,conis antherarum infirme contractis.

U.S.A. Texas: GreenIsland,23-29 June1922, TEX; isotype, US). Tharp1129 (holotype,

fo2-4 cm,scapisglabris, a cormo Radicibus penariis Storageroots1.5-3 cm long, enlarged2-4 cm 10-20mm latis 4-9 linearibus ad ellipticis liisbasalibus filamentis corm.Basal leaves 5-9, narrowly linear, denticulatis, squa- from (15-) tepalisflavis, marginibus anthera- 24-40 cm long, (1.5-)3-7.5 mm wide, margins conis connatis 4-5.5 mm antheris mosis, longis, filamentis brevioribus. 2-3 mm latisplerumque rum the midrib denticulateto ciliate, veins forming sometimes (35-)58-74 glabrous, Scapes papillate. Basal 2-4 corm. cm from roots enlarged Storage leaves 4-9, 33-60 cm long,10-20 mmwide,linear cm high,bearing0-2 branches.Cauline leaves 1mi- 2, lowest 3.5-7.6(-13) cm long, rarelyreaching to narrowly weaklyfalcate, elliptic,sometimes nutant. Tepals probably to denticulate. denticulate Scapes glabrous, nextnode. Flowersyellow, nutely Cauline leaves 4-5, 13-19 mm long, innerbroadlyelliptic,6-8 mm 56-105 cm high,unbranched. exceeding wide, outer elliptic, 3-4 mm wide. Filaments (4.3-)10-20 cm long,lowestfrequently clavate, 4-6 mm long, insertion nutant. the nextnode. Flowersyellow, Tepals 12- scaled, narrowly Anthers in a dorsal connate, (3.5-)4-7 mm pocket. 6-8.5 mmwide, 19 mmlong,inner broadly elliptic, outerelliptic,2.5-3.5 mmwide. Filaments scaled, long, the cones widest at base, taperedto apex, dorsalin 1.5-3.5 mm wide, apex (0.7-)1.1-2.1 mm wide, clavate,5-9 mmlong,insertion narrowly Ovaries 2.5coarselytoothed. a pocket. Anthersconnate,4-5.5 mm long, the broadlysometimes mmlong, mm 5.5 (9-)11-15 oblong, long. Capsules thanthe filacones 2-3 mmwide,usuallyshorter In flower mm wide. 4.5-6 August-September. mm 1-1.5 to ments, tapered, apex weakly strongly lobed. Ovaries2-3.5 mm wide,deeplyand broadly The broad,weaklytaperedanther cone, narrow long. Capsules oblong,9-13 mm long, 4-6 mm and storagerootsenlarged2 or morecm leaves, wide. In flower June,October-November. from the cormare similarto those of E. conzattii, This species is based on fourcollectionsmade E. flexuosa,and E. falcata. The veins on the unare usuTexas (Fig. 1), priorto 1952. dersideofthe basal leaves ofE. conzattii near Brownsville, In flowers. white it has and general, limited distributional a rather These suggest range, ally papillate, and cauline more have E. and E. this collections falcata and thelack ofrecent flexuosa suggests speifnotalready basal leaves than E. palmeri;the basal leaves of cies is rareand probably endangered, and thoseof it is a species manyE. flexuosaplantsare narrower The limited data suggest extirpated. falcate. are of clay dunes and Ilanos. E. falcata notEdwardPalmer, honors The specific wereincludedin E. reepithet SpecimensofE. texensis Mexican of collector ed plants. broad basal have because leaves, long they flexa cauline leaves, and oblongcapsules. overlapping Paratypes. MEXICO. Chihuahua: hills nearChihuaThe short, broad,and deeply lobed anthercones MO,NY). So1109 (MEXU, hua,6 Sep. 1886,Pringle E. pal- nora: Y6cora, 3 Sep. 1970, Pennington are similar to thoseofE. conzattii Cruden, 103 (TEX); old E. flexuosaGreenman, and E. falcata road to Maycoba,0.5 mi. E Arroyo meriCruden, Y6cora, 1550 m, 7 2478 (MO). basal leaves, Sep. 1995, Fishbein Cruden,all of whichhave narrower

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4. Echeandia falcata Cruden, sp. nov. TYPE: Mexico.Guanajuato: Ruta 49, nearKm 32, ca. 7 kmN San Miguelde Allende,grassy hillside withOpuntia, Ipomoea,1880 m,4 Aug. 1968, Cruden1459 (holotype, UC; isotypes, ENCB, F, GH, K, MEXU, MO, NY, US). Radicibus a cormo 23-5 cm, basalibus penariis foliis latislinearibus ad anguste ovatis 12(-14)mm plerumque falcatis ciliatis, marginibus scapisglabris vel ad basim filamentis clavatis flavis, pubescentibus, tepalis infirme antheris connatis 4-6.5 mmlatis,conisansquamosis, therarum ad apicem infirme contractis.

18 Sep. 1957,Paray & Rose9703 (US); Tequisquiapan, delRfo, 28 Aug.1905, near SanJuan 2500 (ENCB); Rose, & Rose9554 (NY,US); 15 kmfrom Painter Higuerillas 25 June toward 1972, Bernal, 20o54'N,99o54'W,2100m, 8100 (TEX). & Johnston Wendt, Chiang 5. Echeandia taxacana Cruden,sp. nov.TYPE: Mexico. Mexico: Ruta 55, 6.5 km NW GuerreroStateLine, 1485 m, 1 Aug. 1970, Cruden 1721 (holotype,UC; isotypes, F, GH, K, MEXU, MO, US).

1.5-5.5cm, 1 cm longis a cormo Radicibus penariis the corm, foliis 1-4 mm latismarStoragerootsenlarged3-5 cm from linearibus basalibus peranguste velad basim denticulatis ad ciliatis, scapis glabris Basal leaves (5-)7-11(-14), ginibus usually long-tapered. clavatis filamentis flavis squatepalis velalbis, linearto narrowly ovate,weaklyto strongly falcate, scabridis, antheraconis mosis, connatis 5-7.5 mm longis, antheris 10-52 cm occasionallystraight, long, 2-12(-14) rum ad apicem valdecontractis. mmwide,margin short ciliate;veins on lowersurface smooth, rarelypapillate. Scapes glabrousor ca. 1.5-5.5 cmfrom roots corm, Storage enlarged weaklyscabrous,15-85 cm high,bearing0-1(-2) 1 cm long.Basal leaves 5-14, very linear, narrowly branches.Cauline leaves 2-4, the lowest1-7.5(- (10-)15-55 cm long, 1-3(-4) mm wide, margin nutant. 10) cm long.Flowersyellow, Tepals 13-17 denticulate to ciliate. Scape glabrousor scabresmm long,innerbroadlyelliptic,6-8.5 mmwide, centtoward 0-1 thebase, 35-85 cm high,bearing outerelliptic,2.5-5 mm wide. Filamentsscaled, branches.Cauline leaves 1-3(-5), lowest1.6-5.5 clavateto narrowly clavate,4-6 mmlong,insertion cm long. Flowersyellowor white,nutant. Tepals dorsal in a pocket. Anthersconnate,4-6.5 mm 11.5-15 mmlong,innerbroadlyelliptic,5-7 mm long,the cones weaklytapered,(1.5-)1.8-2.5 mm wide, outernarrowly elliptic,2-3 mmwide. Filawide, (0.8-)1-2(-2.8) mm wide at apex. Ovaries mentsscaled, clavate,(4-)5-7 mmlong,insertion 2.5-4 mmlong.Capsules broadly to oblong, dorsalin a pocket.Anthers oblong connate, (5-)6-7.5 mm (5-)10-13(-16) mm long, 5.5-6.5 mm wide. In long,the cones usuallylongerthanthe filaments, flower Chromosome strongly (lateJune-)August-September. tapered,(1.7-)2-2.5 mmwide,apex shaln = 16 (Cruden1456, 1459, 1803). number lowlylobed, 0.5-0.9 mm wide. Ovaries 2-4 mm 7.5-9 mmlong,4oblong, This tetraploid relative ofE. flexuosais endemic long.Capsules broadly Chromosome mm In 4.5 wide. flower July-October. to Guanajuatoand Quer6taro (Fig. 1). Two specin = 8 (Cruden1721, 1953). mens from Guanajuato have long, non-falcate number leaves, but the rest have shorter, usually narrow and strongly falcateleaves. The largeflowers with weakly tapered,usually broad anthercones are similarto those of E. flexuosafrom westernMichoacin and easternJalisco,and distinguish this bothE. reflexa and E. mexicanaCruspecies from den ofadjacentHidalgo.The falcateleaves,clavate and somewhatlarger capsules distinfilaments, E. flexuosa. guishE. falcata from are foundin a The yellow-flowered populations varietyof habitats,includingthorn scrub, oak and pine-oakwoods in Oaxaca, Guerwoodlands, and M6xicobetween1400 and 2400 Morelos, rero, m (Fig. 1). The white-flowered plantsare essentially like the yellow-flowered ones, but may be from are The sampledpopulations moremesic habitats. southeast ofthe distributional rangeofthe yellow2000 and 2600 m. between flowered populations slenderinfloresThe numerous narrow leaves, Paratypes. MEXICO. Guanajuato: Guanajuato, San Miguel de cences, and broadlyoblongcapsules are reminis1880,Dugess.n.(GH);Ruta51, between & cent of E. flexuosa,from Allende andComonfort, 2100 m,5 July 1971,Genelle whichit is distinguished de Al898 (DUKE,MO); 18 kmS San Miguel Fleming cone and smallstoranther the strongly by tapered lende, 1790m,4 Aug.1968,Cruden 1456 (ENCB,F,K, some distancefrom roots that develop usually age de MEXU, MICH, NY,UC,WIS);ca. 8 kmN SanMiguel this speleaves distinguish 1880m,23 Aug.1970,Cruden 1806(GH,UC). the corm.The narrow Allende, de Allende, 15 kmE Guana- cies from Queritaro:E San Miguel E. reflexa, and it usuallyhas morebasal state 2125m,23 Aug.1970,Cruden leaves thanE. smithii line, juato-Quer6taro Cruden. del Rfo 1803 (UC); Hacienda Ciervo, between San Juan the cityofTaxco de honors The epithet specific andCadereyta, 20 Aug.1905, Rose, Painter & Rose 9647 end ofdiswhichsits at the northwestern 20 Aug.1905,Altamirano 1597 (US); Alcar6n, (US); Del Ciervo, roadtoward 21 Aug.1903,Rose, Painter tributional Tequisquiapan, rangeofthisspecies.

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Paratypes. MEXICO. Guerrero: TaxcoRoad, 18 July A519 (F); 19 mi. from Taxco de Alarc6n, 1938, Kenoyer 3800 ft.,3 July1941, ChuteM-108 (MICH). Morelos: 1905, Rose, Painter& Rose 8609 (US). Oaxaca: Ruta 2350 m, 15 190, Km 56, ca. 16 km ESE Tamazulapan, 1953 1773 (UC), 7 Sep. 1971, Cruden Aug. 1970, Cruden (ENCB, K, MEXU, NY, TEX, UC, WIS); Ruta 190, Km 406-407, 26.5 km SE Huajuapan de Le6n, 2270 m, 30 1444 (UC). July1968, Cruden examined. MEXICO. Oaspecimens White-flowered 16 kmS San JuanMixxaca: Mpio. San JuanMixtepec, tepec, Montede Tesoro,17o14'N, 97o51'W, 2600 m, 6 Oct. 1988, Santiago 880 (MEXU); 6.7 km W of Miahuatltnon road to Piedra Larga (Coatlines),25 Oct. 1982, & Villasenor 1716 (MEXU); Tlaxiaco,San Martinez Torres, & Calzada 4439 (TEX). 6. Echeandia
17o14'34"N, 97o43'10"W, 2000 m, 3 Aug. 1994, Panero

E. flavescens it from tile anthers (Schuldistinguish tes & Schultesf.) Cruden,a commonspecies of cenofinterior forests and conifer desert grasslands Mexico. traland northern
Sinaloa on road betweenVilla Uni6nand El Salto, 7000 & Ownbey 1931 (MICH). Sinft.,28 Sep. 1953, Ownbey 2000 m, 27 aloa: Ruta 40, K 204, 5 km W El Palmito, 1827 (NY, US). Aug. 1970, Cruden 7. Echeandia sanmiguelensis

Durango:2 mi.NE El Parafso, Paratypes.MEXICO.

Km 64 on carretera Pedro Molinos, Tlaxiaco-Putla,

TYPE: Mexico.Guanajuato:Ruta 49, Km 3031, ca. 9 km N of San Miguel de Allende, grazedfieldalong road withAcacia, Opuntia, Agave, 1950 m, 4 Aug. 1968, Cruden1460 K, MEXU, MO). UC; isotypes, (holotype,

Cruden, sp. nov.

Mexico. Sinaloa: Ruta 40, Km 204, ca. 5 km W El Palmito, seepage area with Begonia,Bi2000 m, 18 dens,Dahlia, Castilleja,Cosmos, UC; isoSep. 1971, Cruden1997 (holotype, types,ENCB, F, GH, K, MEXU, MO).

attenuata Cruden, sp. nov. TYPE:

452-3 cm,scapis a cormo Radicibus glabris penariis 7-13 falcatis ovatis basalibus 75 cmaltis, anguste foliis lon5-6 mm filamentis mm albis, latis, squamosis tepalis libris nonversatilis. gis,antheris corm,3-5 Storagerootsenlarged2-3 cm from 5cm long.Basal leaves linear,attenuate, falcate, ciliate. 8, 17-33 cm long,7-13 mmwide, margin 45-75 cm high,bearing0-2(-4) Scapes glabrous, lower Cauline leaves 0-2, whenpresent, branches. cernuous18-67 mmlong.Flowerswhite, aristate, eldeclinate.Tepals 11-15 mmlong,inner broadly liptic, 5.5-7 mm wide, outerelliptic,3-4.5 mm wide. Filamentsscaled, linear,5-6 mm long,insertiondorsal in a pit. Anthers free,1.5-2.5 mm anther due to thereflexed walls, long,notversatile dehiscence lateral. Ovaries 2.5-3.5 mm long. belowthe stigma. Capsules obStylesbentupward long,(8.5-)10-16 mm long,3.5-6.5 mmwide. In late July-August. flower

1-2 cm,3-4 cm longis, a cormo Radicibus penariis attenuatis basalibus 1-5(-7)mm anguste linearibus foliis velad basim scabris 30-90 cmaltis, latis, scapisglabris 2-6 longiattenuatis, flavis caulibus ellipticis tepalis foliis inmarvaldereflexis, filamentis unumquidque squamosis nonver4-6.5 mmlongis, antheris libris supioinsertis dehiscens lateralibus. satilis 31-2 cm from thecorm, roots Storage enlarged 4 cm long.Basal leaves narrowly attenuate, linear, dentic23-53 cm long,1-5(-7) mmwide,margin ulate to serrulate. Scapes glabrousor scabrescent towardthe base, 30-90 cm high, bearing 0-4 branches.Cauline leaves (2-)3-5(-6), long attenceruate, lowerto 17.5 cm long. Flowersyellow, nuous-declinate. reflexed, (6.5-)8Tepals strongly 13 mmlong,innerelliptic,3.5-5 mmwide, outer narrowlyelliptic, 2-2.5 mm wide. Filaments dorsalin scaled, linear,4-6.5 mmlong,insertion 2.5-3.25 mmlong,notvera pocket.Anthers free, satile,dehiscencelateral.Ovaries 1.5-3 mmlong. Capsule (onlyone seen) 6.5 mmlong,5 mmwide. In flowerlate August-September. Chromosome

collections from This species is known onlythree just one or two populations possiblyrepresenting near San Miguelde Allende(Fig. 1). The twoCrua single large popuden collectionsmay be from lation. and declinateflowers, The falcateleaves, white, the corm to close roots suggesta enlarged storage withE. scabrella (Bentham)Cruden, relationship number n = 8 (Cruden 1827). which has smaller flowers, subglobosecapsules, and scabrescentscapes. The few aristatecauline Echeandia attenuata is known only fromseepage broaderlanceolateleaves, and 2000 and 2200 m alongthe Duran- leaves, if present, areas between E. tamaulipenitfrom differentiate the border It is longer capsules 1). by (Fig. distinguished go-Sinaloa thisspedistinguish leaves, both basal and cauline, sis Cruden.The whiteflowers long-attenuate which occurs in similar flowers cernuous-declinate with smallish, relatively cies fromE. flavescens, fewindividualsof the latter reflexed and strongly narrow tepals,and filaments habitats.In addition, the an- species have storageroots enlargedclose to the in a pocket,whichforces thatare inserted with 5-8 basal leaves that thers to extend alongthesame axis as thefilaments. corm in combination The narrow, reflexed strongly tepals and nonversa- tendto be falcate.

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Paratypes. MEXICO. Guanajuato: Ruta49, Km 3132, ca. 8 km N of San Miguel de Allende, 1950 m, 23 Aug. 1970, Cruden 1805 (ENCB, F, GH, MICH, NY, UC, 26 July 1950, SpivUS); San Miguelde Allende,6500 ft., ey 177 (UC).

9. Echeandia atoyacana Cruden, sp. nov.TYPE: with Mexico.Guerrero: Matalea, outcrop rocky Commelina, Cnidosculus, Ruta 95, Km 103104, ca. 31 Km S bridgeover Rio Papagayo, 300 m, 4 Aug. 1970, Cruden1734 (holotype, F, GH, K, MEXU, MO). UC; isotypes, 8. Echeandia tamaulipensis Cruden, sp. nov. 6foliis basalibus 3-8 cm, a cormo Radicibus penariis TYPE: Mexico. Tamaulipas: Papolote de la ad denticumm ovatis 12 marginibus integris latis 8-28 3 mi. SSW ofheadquarters, Loreto Mirandena, latis, scapisscabris ad 1 m altis, foliis caulibus 2-4, teRanch, 24?20'N, 98?W,16 Sep. 1960, John- palisalbis, flexus intro linearibus filamentis super glabris ston& Crutchfield 5550 (holotype, connatis. antheris MEXU; iso- ovarium, LL, MICH, TEX, UC). types, Storageroots3-8 cm from corm,1-2 cm long. elto narrowly ovate leaves Basal 6-12, narrowly Radicibuspenariisa cormo(1-)3-6 cm clavatis,foliis 4-12 mm velsca- liptic,(12-)23-52 cm long, (8-)13-28 mm wide, basalibus linearibus latis, glabris scapis or serrulate. entireto denticulate bris,tepalisalbis,filamentis pauci squamosis4.5-6.5 mm margins Scapes antheris librisnonversatilis dehiscenslateralibus. scabrous, 33-97 cm high, bearing (0-)1-4(-9) longis, branches.Cauline leaves 2-4, reduced to bracts. corm, Flowerswhite,nutant, Storagerootsenlarged(1-)3-6 cm from openingin earlyafternoon. 3-9 cm long.Basal leaves 3-7, linear, longtapered, elliptic,6.5-9 mm long to 1 mm Tepals narrowly 20-65 cm long,denticulate to shortciliate,4-12 wide. Filaments smooth, linear,bentor pinchedin mmwide.Scape glabrous orscabrescent, 40-60 cm basal or 2.5-3.3 mm long,insertion above ovary, Cauline leaves 1high,bearing0-2(-3) branches. 3-5 mmlong,thecones so. Anthers connate, nearly cer4, reducedto bracts.Flowerswhite, probably tapered.Ovaries 1-2 mm long. Capsules strongly nuous-declinate. Tepals 10-15 mm long, inner mmlong.In flower 4-6.5 July-September. globose, 4.5-5 mmwide,outer el- Chromosome broadly elliptic, narrowly n = 16 (Cruden1732, 1733, number 1.5-3.5 mmwide.Filaments weaklipticto elliptic, 1734). usuly scaled, linear,4.5-6.5 mmlong,insertion GuerThis tetraploid onlyfrom species is known ally dorsalin a deep pit or pocket,if in a shallow 250 and 650 m thereflexed anther walls holdtheanther on reroand adjacentM~xico between pocket, of small flowers, short, same radiusas thefilaments. Anthers 1.7-2.5 (Fig. 2). The combination free, tapered anther cones, small capsules, mmlong,notversatile, dehiscencelateral.Ovaries strongly scapes is unique 2.5-3.5 mmlong. Capsules broadlyoblongto ob- broadbasal leaves,and pubescent Mscavea. long,8-11 mmlong,5-5.5 mmwide.In flower Sep- in subgenus ofAtoythetown The specific tember-October. recognizes epithet ac de Alvarezand the Rio Atoyacin whosevalley This white-flowered species is known onlyfrom I first collectedthisspecies. the coastal plain of centraland southern TamauliGuerrero: Paratypes.MEXICO. ca. 9.5 kmN Atoybe confusedwitheitherE. ac, roadtoSan Vincente pas (Fig. 1). It might 300 m,3 de Gallo, andPuerto or E. chandleri & Thompson) Aug.1970,Cruden (Greenman flavescens 1732 (ENCB,K, UC); ca. 6 kmN 1733 (MEXU, 260 m,3 Aug.1970,Cruden MO, M. C. Johnston, which both have yellowflowers. Atoyac, 25-300m,10 Atoyac, The former occurs at muchhigherelevationsand NY,UC,US,WIS);Dist.Galeana, Hinton et al. 14526 (ARIZ, GH, LL (2), MO, has versatile has somewhat Aug. 1939, whilethelatter anthers, NY, US); 23 kmN Ruta200 (coastroad)on roadto and strongly scaled filaments. The Ciudad Altamirano, largerflowers 500 m, 26 Sep. 1983, Anderson Dist.Temascaltepec, Mexicancollections ofE. chandleri are from higher 12816 (MICH).Mexico: Palmar, al. 6321 (GH, K, NY, Hinton et 21 1934, [650 July m], elevations or from northern Tamau(900-1000 m) US). lipas. 10. Echeandia elegans Cruden,sp. nov.TYPE: with oak woodland, Mexico.Guerrero: Cosmos, Commelina, Calochortus, Sessilanthera, Agave, 4540 (MICH,TEX); Sierra de Tamaulipas, ca. 40 km NNW Aldama,ca. 23o14'N, 98?10'W,above JuanTomis, ca. 15 km SW Xochipala,road to Puertode E Las Yucas, 13 Oct. 1957, Dressler 2384 (GH, MO); 15 Gallo, 1950 m, 21 Sep. 1973, Cruden2096 27 Sep. 1959, mi.from on theMante Tampico highway, UC; isotypes,ENCB, F, GH, K, (holotype, Graham& Johnston 4087 (MEXU, MICH, TEX). MEXU, MO, NY, US). The following species are includedin subgenus 5foliis basalibus 2-6 cm, a cormo Radicibus penariis Mscavea. latis 7-18 mm obovatis ad anguste linearibus 8 anguste
& Graham 7 mi. N of Altamira, 27 Oct. 1959, Johnston

MEXICO. S Lomas delReal, Tamaulipas: Paratypes.

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* Cd. Hid go

Ai

MEXICO
SToluca

E. atoyacana
O E. elegans E. I O hirticaulis E. hintonii E. mexiae

SO O
* Temascaltepec

D.F.
Cuernavaca

MICHOACAN

gMORELOS

"

E. pseudopetiolata

A5

P
*Chilpancingo

Olc

aU
A
GUERRERO

Mscavea.The namesofthe statesare capitalizedand Figure2. Distributional rangesofsix new species in subgenus towns and cities are indicated is notillusby a small solid circle.The distributional rangeofEcheandia nayaritensis trated.

ad breviciliatis, marginibus intergris scapisglabris glau- 2). The available material includestwotypes.The caulibus albis13-20mm cis,foliis 4-10,tepalis longis, specimens from Temiscohave 5-9 branches, anther filamentis clavatis antheris connatis 8-11.5 squamosis, cones that are and have weakly tapered, may large mm longis. from centralGuerrero storageroots.The material Storageroots2-6 cm fromcorm, 1.5-2.5 cm has 0-3 branches,strongly taperedanthercones, linearto narrowly and quite slenderstorageroots.Vegetatively, long.Basal leaves 5-8, narrowly this 20-70 cm long,7-14(-18) mmwide,mar- species is difficult obovate, to distinguish from E. hintonii. gins entireto shortciliate. Scape glabrous,glau- Relativeto thelatter species,E. eleganshas longer 0-9 branches.Cau- tepals and anthers, cous, 0.7-1.5 m high,bearing an anther cone thatis usually line leaves 4-10, lowest 9-30 cm long, others twicethelength ofthefilaments, and smooth lower reducedupward. Flowers in leaf surfaces. white, nutant, opening late morning. Tepals narrowly elliptic,13-20 mm Guerrero: 14 mi.from Mex 95 Paratypes.MEXICO. long,inner3.5-5 mmwide,outer1.5-3 mmwide. on roadto Chichihualco, 26 Oct. 1970, Graham 1226 Filaments scaled, clavate,3.5-7.5 mmlong,inser- (MICH).Morelos:Ruta95 (libre) in Temisco, ca. 9 km 22 Sep. 1974, Cruden tion dorsal in a pocket.Anthers 2173 (GH, K, connate,8-11.5 S Cuernavaca, MICH,UC). mmlong,1.5-2 timesthe length of the filaments, MEXU, the cones usually strongly tapered,rarelyweakly 11. Echeandia hintoniiCruden, sp. nov.TYPE: so. Ovaries2.5-4 mmlong. Capsules broadly obMexico. Guerrero: on steep, damp bank with long, 8-9.5 mm long, 5-5.5 mm wide. In flower Adiantum, Oxalis,Cuphea,in pineBlechnum, Chromosome numbern = 8 September-October. oak woods,Ruta 95, nearAgua de Obispo,ca. 2096, 2173). (Cruden 39 km S Chilpancingo, 780 m, 2 Aug. 1970, This is a species ofoak woodlands between 1900 Cruden1729 (holotype, UC; isotypes, ENCB, and 2300 m in Guerrero and adjacentMorelos F, GH, K, MEXU, MO, NY, US, WIS). (Fig.

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4-8 cm, basalibus 3- 12. Radicibus a cormo penariis foliis ad anguste obovatis 5-20 mm linearibus 5(-8) anguste caulibus latis,scapisglabris 1-4(-5), vel scabris, foliis albis 10-13 mm filamentis clavatis tepalis longis, squaconis 4.5-8 mm antheris connatis antheramosis, longis, rum ad apicem valdecontractis. corm,1-2 cm long. Storageroots4-8 cm from Basal leaves 3-5(-8), narrowly linear to narrowly obovate,in manyminutely papillate betweenthe veins on lowersurface, 13-56 cm long,5-20 mm entire to denticulate. wide,margin Scapes glabrous or scabrescent,37-111 cm high, bearing 0-4 branches.Cauline leaves 1-4(-5), the lower8.5in late 28 cm long.Flowerswhite, nutant, opening morning. Tepals narrowly elliptic,10-13 mmlong, inner3.5-4.5 mmwide,outer1.5-2 mmwide. Filaments scaled, clavate, sometimesnarrowly so, 3.5-5 mmlong,insertion basal or dorsalin a pocket. Anthersconnate,4.5-8 mm long, the cones ofthe filstrongly tapered,1-1.7 timesthe length aments.Ovaries 2-4 mm long. Capsules broadly 7-9 mmlong,4-5 mmwide. In flower late oblong, n = 8 (Cruden Chromosome number June-October. 1723, 1729, 2102).

Echeandia hirticaulis Cruden, sp. nov. TYPE: Mexico.Mexico:Ruta 130, ca. 2.5 km oak woods withTigridia, NE Temascaltepec, 1750 m, 11 Cuphea,Eryngium, Sphenostigma, UC; isoSep. 1971, Cruden1967 (holotype, MEXU, ENCB, K, GH, MO). types,

2-4 cm,foliis basalibus a cormo Radicibus penariis latismarginibus ovatis longicilia(8-)15-42mm anguste in dimidio hirsutis tis,scapisplerumque inferiore, foliis antheris filamentis caulibus albis, 3-4, tepalis squamosis, connatis.

Storageroots2-4 cm from corm,1-2 cm long. Basal leaves 4-8, narrowly ovate,(17-)25-55 cm longciliateand long,(8-)15-42 mmwide,margins usually undulate. Scapes usually hirsuteon the lowerhalf,hairs long and flat,0.2-0.5 mm long, with just a fewhairsabovethelowest rarely cauline whichin leaf,0.6-1.3 m high,(3-)5-9-branched, branches. bear secondary largeindividuals Cauline reduced leaves 3-4, lowest12-54 cm long,others to bracts, nutant, white, openlongciliate.Flowers elliptic,12Tepals narrowly ing in late morning. 17 mmlong,1-2 mmwide. Filaments scaled, cladorsal in a vate, (3-)3.5-5.5 mm long, insertion pocket. Anthersconnate,(6-)7.5-10.5 mm long, Mscaveawith narrow Among species in subgenus thecones strongly usuallytwicethelength tapered, leaves and scaled filaments, thisspecies is distin- of the filaments. Ovaries 1-3 mm long. Capsules well-removed roots. guishedby its slender, storage broadlyoblong,6-8 mm long, 4-5 mm wide. In All collections are from oak and pine-oak woodsin flower number October.Chromosome August-mid central and north-central Guerrero between750 n = 8 (Cruden1967, 1970). and 2350 m (Fig. 2). Specimenswithpubescent between1000 and 2000 m This species is found scapes are a littlelargerand have fewercauline leaves and tinypapillae on the lowersurfacesof in openingsin oak and pine-oakwoodsin a small thebasal and caulineleaves. Thereis no geograph- regionin western Mexico,easternMichoacin,and Guerrero ic pattern to plants withglabrousand pubescent northern (Fig. 2). The closest relativeof Rose, whichis of Agua de Obispo the thisspecies maybe E. macrophylla scapes, and in the vicinity 1000 and 1500 m in San Luis Potosf between leaves ofspecimens withbothtypesofscapes were found species in theSierra papillate; thus,thereis littlereason to recognize and Tamaulipas.Otherrobust smoothfilahave Transversal straight, Volcinica the variation formally. that flowers and/or cauline numerous leaves, ments, B. This species honors Hinton whose colGeorge The specific epithetdeto open in the afternoon. lectionsfrom Guerrero have added considerably scribesthe hirsute scapes. our understanding ofthe region's Echeandia. Dist. Paratypes. MEXICO. Guerrero: Manch6n, Dist.Mon- Mina, MEXICO.Guerrero:Pasi6n, Paratypes. etal. 9596 (GH,K, LL,NY, 27 Sep. 1936,Hinton et al. 10763 US). Mexico: between tes de Oca, 1000 m,5 Oct. 1937,Hinton Dist. and Las Anonas, Ziticuaro (GH,LL, NY,US); Ruta95, ca. 17 kmS Chilpancingo, et al. 13146 1600 m,23 Aug.1938,Hinton Ziticuaro, 1723 (ENCB,GH,K, (GH,K, LL, NY,US); Ixtapan, ca. 1250m,2 Aug.1970,Cruden 1000 Dist. Temascaltepec, 230 (TEX); m,5 Aug.1932,Hinton Aug.1965,Irving UC);61 mi.NE Acapulco, 1263 (GH,K, MICH,NY,UC, onroadtoPuerto ca. 29 kmSW ofXochipala, de Gallo, US); Carboneras, 2030 m,27 Sep. Dist.Temascaltepec, 2102 (MEXU, 2225 m,22 Sep. 1973,Cruden UC); 65.3 1932,Hinton 1850 1852 (K); 4.5 mi.NE Temascaltepec, oftheAcapulco-M6xico kmfrom the"desviaci6n" & Laskowski 3934 (ENCB, high- m, 11 Oct. 1966,Anderson Romo 321 (MEXU); de MICH);5 km 2350m,21 Oct.1986, 1900m,3 Sep.1965, Agua way, S Temascaltepec, Roe, & Kenton Roe,Mori & Rzedowski 1100m,10 Oct.1986, Romo 1707 (WIS);roadtoTingambato, Palomino, Obispo, 12 July 308 (MEXU); 239 (GH),23 E San Nicholas, 2169 1600 m,20 Sep. 1974,Cruden Taxco, 1932,Abbott ManAbbott 160(GH);N Taxco, 25 June Clark (UC).Michoacain: 20 Aug.1953, 7 mi.E Hidalgo, 1935, 1936, July 531089(GH); Ruta15, nearKm116, 7238(NY). ning& Manning

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1970 2000 m,11 Sep. 1971, Cruden ca. 9.5 kmS Tuxpan, (ENCB, GH, K, UC).

ofCuerGH, K, MO, NY, U, UC, US). Morelos: vicinity & Lemmon navaca, 1905, Lemmon s.n. (UC).

Radicibuspenariisa cormo0.5-2 cm, foliisbasalibus linearibus, tepalisalbis 12-19 mm scapis glabris, anguste mm latis brev- longis,filamentis obovatis connatis marginibus (10-)15-40 fusiformis 5 anguste squamosis,antheris 6- 6-11 mmlongis,conis antherarum 1-1.3 m altis, caulibus iciliatis, scapisscabris foliis filamentis duplo lonlinearibus antheris gioribus. 15, tepalisalbis,filamentis glabris, 3-9 cm,foliisbasalibus3Radicibuspenariisa cormo

13. Echeandia mexiae Cruden,sp. nov.TYPE: 14. Echeandia nayaritensis Cruden, sp. nov. deciduouswith Mexico.Guerrero: tropical palTYPE: Mexico. Nayarit: Mpio. Ruiz, 1-3 km mettopalm, Salvia, Dahlia, Bursera,4.5 km W El Venado,road from Ruiz to JesusMaria, SW Xochipala,1280 m,21 Sep. 1973, Cruden 60 m, 9 Aug. 1980, Breedlove& Almeda 2094 (holotype,UC, isotypes, F, GH, K, 45284 (holotype, CAS). MEXU, MO, US).

connatis.

the corm, Storagerootsenlarged3-9 cm from 1.5-3 cm long.Basal leaves 3-5, narrowly obovate, short 37-45 cm long,(10-)15-40 mmwide,margin 1-1.3 m high, ciliate. Scape scabrousthroughout, some nodeswithtwobranches (3-)5-14-branched, and some branchesbearinga secondarybranch. Cauline leaves 6-15, reduced to bracts.Flowers in late afternoon. white, nutant, opening Tepalsnarrowly elliptic,9-13 mmlong,inner2-3 mmwide, outer1-2 mmwide. Filamentssmooth, linear,2basal or nearly so. Anthers 3.5 mmlong,insertion taconnate,4.5-6.5 mm long,the cones strongly pered.Ovaries1.7-2.5 mmlong.Capsules broadly Au5-8 mmlong,4-5 mmwide. In flower oblong, n = 8 (Cruden number Chromosome gust-October. 2086, 2087, 2088, 2094, 2172).

corm,1Storagerootsenlarged0.5-2 cm from 4 cm long.Basal leaves 3-10, narrowly linear,16sometimes 38 cm long,(2-)4-8 mmwide, entire, 41-58 cm high,bearing falcate.Scapes glabrous, 0-3 branches. Cauline leaves 1-2, bract-like. 12nutant. Flowers white, elliptic, Tepals narrowly 19 mm long,inner2-3 mmwide, outer1-2 mm 3.0-4.5 mmlong, wide. Filaments scaled,fusiform, basal or nearlyso. Anthers insertion connate,(6widest tapered, )7.5-11 mmlong,thecones weakly twice in lowerhalf, in the middleor parallel-sided Ovaries 1-2 mmlong. of the filaments. the length Capsules broadlyoblong,5-7 mm long, 4.5-5.5 mmwide. In flower July-August.

This little-collected species is endemic to the and adjacent coastal lowlandsof northern Nayarit Sinaloa, whereit occurs in open areas and palm All collectionsof this species were made in savanna below 400 m. The long tepals, long annorth-central Guerrero and adjacentMorelos(Fig. thers,and storageroots enlarged and clustered 400 and 1500 m in relatively 2) between dryhab- close to the corm withE. suggesta relationship deciduousthorn scrub.Severalcolitats, including whichoccurs at higherelevaCruden, mcvaughii lectionsare from limestone-derived soils. The nuMichoactin. tions in Nayarit, Jalisco,and western merous cauline leaves and smooth filaments The scape of the latterspecies is usuallypubesE. rawiththe widespread suggesta relationship cent, and the basal leaves are usuallymorethan mosissima (Presl) Cruden of western Mexico. 10 mmwide and falcate. are usuallyglabrous, scaScapes ofthelatter rarely brescentnear the base, and the basal leaves are 26.8 mi.S ofSinaloa Nayarit: Paratypes.MEXICO. 24 July stateline,200 ft., 1975, Dunn,LeDoux& Wallace usuallynarrower. ca. 149 kmN Tepic, Ynes Mexia,whocol- 21850 (LL). Sinaloa: La Jarretadera, The specific honors epithet 19 Aug. 1988, Walker (K, MEXU, MO, UC). s.n. in Mexicoand SouthAmericain lectedextensively the 1920s and 1930s. 15. Echeandia pseudopetiolata Cruden, sp. Paratypes. MEXICO. Guerrero: Caion de la Mano Ruta 95, Km nov. TYPE: Mexico. Guerrero: Negra,near Iguala, 11 Aug. 1905, Rose,Painter& Rose near El Rinc6n,ca. 44.5 km S Chil44-45, 9328 (US); Ruta 95D, Km 63-64, ca. 12 km S Morelos 600 m, 2 Aug. 1970, Cruden1731 pancingo, 2086 (ENCB, stateline, 1070 m, 20 Sep. 1973, Cruden GH, K, TEX, UC, WIS); Ruta 51, ca. 22 km W Iguala, UC). (holotype,
915 m, 20 Sep. 1973, Cruden2087 (ENCB, F, GH, K, UC); ca. 35 kmW Iguala, 1500 m,21 Sep. 1973, Cruden 2088 (UC); ca. 36 kmW ofIguala, 1000 m,30 Sep. 1982, Iltis,Benz & Burd28634 (IA, WIS); Ruta 55, Km 118119, ca. 30 km N Taxco, 1150 m, 21 Sep. 1974, Cruden 2172 (MEXU, MO, NY, UC); Temisco,top of barranca 380-400m,30 Oct.1937,Mexia 8711 (ARIZ,F, Limo, Radicibus penariisa cormo1-3.5 cm, foliisbasalibus 4-6, 43-56 cm longisdimidioinferiore angustelinearis ovatis 3-5 mmlatisdimidio expansis magnopere superiore foad ellipticis(21-)26-35(-45) mmlatis,scapis glabris, linearibus pauliis caulibus 2-4, tepalisalbis, filamentis connatis. antheris cisquamosis,

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Storageroots 1-3.5 cm fromcorm, 1-1.5 cm CAS, DS, DUKE, ENCB, F, GH, IA, K, LL, MEXU, long. Basal leaves 4-6, oblanceolate,43-56 cm MICH, MO, NY, PAUH, TEX, UC, US, WIS. linear,3-5 mmwide,exlong,lowerhalfnarrowly Cited panded in the upper half into a broad, ovate to Literature ellipticalblade, (21-)26-35(-45) mm wide, mar- Baker,J. G. 1876. Revisionof the generaand species of and Eriospermeae. Anthericeae J. Linn. Soc., Bot. 15: ciliateto ciliate.Scape glabrous, 88-106 ginsshort 253-363. cm high, Cauline leaves 2-4, the up to 7 branches. Chase, M. W., P. J. Rudall & J. G. Conran.1996. New lowest 12.5-36 cm long, resemblingthe basal lilies: of asparagoid and a newfamily circumscriptions the "petiole." Flowerswhite, leaves, but without Kew Bull. includedin Anthericaceae. Generaformerly nutant, openingin the afternoon. Tepals narrowly 51: 667-680. elliptic,6-9 mmlong, 1-1.5 mmwide. Filaments Cruden,R. W. 1987. Hagenbachia,a misplacedgenusof Liliaceae. NordicJ. Bot. 7: 255-260. New World wrinkled and/or weaklyscaled, linear,sometimes . 1994. Echeandia. Pp. 27-30 in: G. Davidse, M. dorsalin a pocket. 3-4 mmlong,insertion twisted, Flora Mesoamericana, Sousa & A. O. Chater(editors), Anthers 4-5.5 mmlong,thecones strong- Vol. Alismataceaea Cyperaceae.Universidad Naconnate, 6, Bode Mexico,Mexico,D.F.; Missouri cionalAut6noma ly tapered. Ovaries 1-2 mm long. Capsules not St. Louis:The Natural Museum, History seen. In flowerlate July-August. Chromosome tanicalGarden, London. n = 8 (Cruden1731). number & P. F. Yeo. 1985. The R. M. T., H. T. Clifford in This species is known from a fewcollections lin. Heidelberg, Germany. south-central Guerrero between600 and 1100 m MexJ.M. 1898. Diagnosesofnewand critical Greenman, Daedalus. Proc. Amer.Acad. Arts ican Phanerogams. from alongthehighway Acapulcoto Iguala (Fig. 2). 33: 471-489. It occurs in pine-oakand oak woods,as well as Johnston. M. C. 1990. The VascularPlants of Texas, 2d disturbed roadsides.The specific describes epithet Ed. PublishedPrivately, Austin, Texas. the distinctive basal leaves. pseudopetiolate ofAfrican S. & I. Nordal.1993. Newcombinations Kativu,
Paratypes. MEXICO. Guerrero: Agua de Obispo, 1100 m, 12 Aug. 1962, Kruse 796 (ENCB); 1000 m, 8 Aug. 1972, Boege 2441 (MEXU); Hwy.to Acapulco,Km 3000 ft.,20 Aug. 1948, 338, 3 km beyondAcahuizotla, Moore& Wood 4675 (F, GH, MICH, UC, US). Dahlgren, BerFamilies of the Monocotyledons. Springer-Verlag,

Nordic (Anthericaceae). species in genusChlorophytum J. Bot. 13: 59-65. Vol. 15. BroR. 1989. Flora Novo-Galiciana, McVaugh, meliaceae to Dioscoreaceae. Univ. MichiganHerbarium,Ann Arbor. horaut rariorum C. G. 1800. Novarum, plantarum Ortega, decades,... Matriti, ti reg.botan.matrit. discriptionum I thank Diana Horton, FredIbarriana. Ex Typographia Acknowledgments. TerforDescriptive AssociationCommittee erickH. Utech,and Wendy B. Zomlefer for helpful Systematics of simple symmetrical 1962. Terminology minology. and constructive comments on themanuscript. Salplane shapes. Taxon 11: 145-156, 245-247. material ofEcheandia Weatherby, ly and TimWalker provided ofthegeC. A. 1910. A preliminary synopsis herbaria Curatorsof the following nus Echeandia. Daedalus. Proc. Amer.Acad. Arts45: nayaritensis. 387-394. made materialavailable for study:ARIZ, BRIT,