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Crassulaceae

Chapter·April2007

DOI:10.1007/978-3-540-32219-1_12

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Crassulaceae 93

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S. ser. Pubescens [G]S. ser. Caerulea lGlS. ser. Monanthoidea [G]Aichryson

Monanthes

Aeonium

S. magel/ense [G]S. dasyphyllum [G]S. tydium lGlRosularia

S. sedoides [G]S. hispanicum [G]S. commixtum lGlPrometheum

S. gracle [G]Sedella

Dudleya

S. div. spp. [S] (Asia)

S. div. spp. [S] (Europe)

S. sexangu/are [S]S. fannosum [S]S. ser. Macaronesica [S]Villadia

S. oaxacanum lSlS. obcordafum [S]Lenophyllum

S. fernafum [S]S. hemsleyanum [S]S. fuffuraceum lSlS. sect. Pachysedum [S]Graptopetalum

Thompsonella

Echeveria

Pachyphytum

Adromischus

Kalanchoe

Tylecodon

Cotyledon

Crassu/a

Macaronesia

Europe/Mediterranean/Near EasV(CentralAsia)

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taxon distribution ::"spp.

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roots); Ieaves usually terete-subulate, * conspic-uously spurred at base; inflorescences t elongatethyrsoids, often spike- or raceme-like, with6-70(-150) cincinnoid partial inflorescences with1-5(-8) flowers; sepals (almost) free, (always?)spurred at base; petals connate at base, lobesspreading to reflexed or erect, whitish to pink orreddish; stylodia t recurved; fruits erect. n = 9-17,20-22,33 and higher. About 21 species in southernUSA (south-western Texas), Mexico, Guatemala(Baja Yerapaz), Peru, at (600-)1,500-4,000 m.

Divided into seven informal species groups(Thiede in Eggli 2003). Appears to be closest toSedum sect. Fruticisedum (IJhl and Moran, l.c.).Täxa formerly classified in sect. Altamiranoa arenow placed in Sedum (Moran 1996; Thiede and't Hart 1999).

24. Lenophyllum RoseLenophyllum Rose, Smithsonian Misc. Collect. 47:159(190a); Moran, Haseltonia 2: l-19 (1994), rev.

Herbs, roots fibrous or thickened; leaves decussatein few basal pairs, elliptic, roundish or rhombic;inflorescences thyrsoids with several cincinni,narrow thyrsoids of compact cincinni or reducedto racemes or spikes above or throughout; sepalserect or ascending, nearly equal, * as long as opencorolla; petals in upper half spreading to recurved,(ob)lanceolate, yellow(ish); stamens slightlyexserted; nectary scales subquadrate; carpelserect, narrog tapering into slender stylodia; fruitserect. n -- 22,32,33,44. Seven species, from USA(southern Texas) and north-eastern Mexico.

Genera 25-28

Stems at least basally woody, but many taxa withsessile rosettes; leaves usually thick and stronglysucculent, in t distinct rosettes; inflorescences lat-eral; petals at least basally connate. x - 30-34 withsecondary reductions. Centred in Mexico.

According to molecular data (Mort et al.2001), these genera form a distinct Americanclade (--'Echeveria group'), together with Sedumsect. Pachysedum. The latter shares the abovecharacters, except for its choripetalous flowers.

25, GraptopetalumRoseGraptopetalum Rose, Contr. U.S. Natl Herb. 13'296 (1911);

Acevedo et al., Brittonia 56:185- 194 (2004), morph. phylog.;

Acevedo et al. , Amer. | . Bot. 91:1099-1104 (2004), mol.

phylog.

Byrnesia Rose ( 1922).

Tacitus Moran (1974).

Herbs (sect. Graptopetalum) to subshrubs (sect.Byrnesia); Ieaves obovate to (broadly) spathulate,usually mucronate in sect. Graptopetalum; usually+ glaucous-pruinose; inflorescences thyrsoidswith few to many cincinnoid partial inflores-cences, or pleiochasia with few cincinnoid partialinflorescences; flowers (4)5(- 10)-merous, stamensdiplostemonous, rarely haplostemonous; sepalsappressed, free to base and + equal in size;petals slightly connate at base, spreading stellately,basicallywhitish or yellowish (to greenish), usuallywith reddish to brown cross-band markings orblotches becoming denser towards tips, rarelyuniformly coloured; stamens first erect, afteranther dehiscence spreading and the antesepa-lous stamens recurved between the petals, afteranthesis becoming erect again; carpels shortlyconnate at base; stylodia normally abruptly offset;fruits ascending to erect; seeds usually reticulate.n = 30-32, 34, 35, 62, 64, 66, 68, +93, +170,+175, 192, +204, +208, +244, +270. Eighteenspecies, USA (central and southern Arizona),Mexico (widespread from Sonora and Chihuahuato Oaxaca); rocky places, to 2,400 m.

Divided into sect. Byrnesia and sect. Grap-topetalum (incl. Tacitus). According to moleculardata, Graptopetalum and its sections are not mono-phyletic, and Tacitus, Cremnophila (= Sedum)and selected species of Sedum and Echeveria areinterspersed among its species (Acevedo et al. l.c.).

26. Thompsonella Britton & Rose

Thompsonel/a Britton & Rose, Contr. U.S. Natl Herb.12.391(1909); Moran, Cact. Succ. I. (U.S.) 64:37-44 (1992),

synopsis.

Glabrous herbs or subshrubs; leaves in rosettes,flattish, semi-amplexicaul, often glaucous, mar-gins straight or undulate; inflorescences erectnarrow thyrsoids or spicate above or as a whole,with 10-70, 1-12-flowered cincinni; flowers(almost) sessile; sepals free, (sub)equal, clavate;petals shortly connate at base, imbricate in bud,spreading from middle, outer face pale, innerface * dark purplish-red; nectary scales minute;carpels shortly stipitate, shortly connate at base;stylodia slender; fruits erect; seeds reticulatewith irregular longitudinal rows. n = 26,52. Sixspecies, central and southern Mexico; usually onlimestone.

1 1 0 |. Thiede and U. Eggli

Genera 27 + 28

Petals upright and connate for most of their length;sepals often strongly unequal in size; petals thick-fleshy; anthers (light) yellow; fruits divergent.

27, EcheveriaDC.

Echeveria DC., Prodr. Syst. Regni Veg. 3:401 (1828);Walther, Echeveria (1972), rev.Oliverella Rose ( 1 903).Urbinia Britton & Rose (1903).

Oliveranthus Rose ( 1905).

Glabrous to hirsute herbs to subshrubs; stemnone or tall, branching or not; leaves rarelyscattered along the stems, usually (ob)lanceolate

and mucronate, often glaucous or highly coloured;inflorescences, racemose, paniculate, or rarelyspicate thyrsoids, or cymose with one to severalcincinni; pedicels usually with one to severalminute bracteoles; sepals reflexed to appressed butusually somewhat expanding, almost completelyfree, equal to strongly unequal; corolla cylindri-cal to pentagonal to urceolate; petals imbricate(valvate in Ser. Valvatae), white through yellowand orange to red, rarely green(ish), inner surfaceusually with nectar-cavity at base; stamens 10,5 attached at top of nectar-cavities, 5 at top ofcorolla tube between petals; carpels connate atbase, erect at anthesis; stylodia slender; fruitswidely divergent follicles; seeds reticulate orsmooth. n = L2-34, polyploid from 28-25A. About139 species, centred in (southern) Mexico, alsosouthern USA (Texas) and Central and SouthAmerica (Venezuela, Colombia, Ecuador, Peru,Bolivia, northern Argentina).

Divided into 17 series (Kimnach in Eggli 2003).

28. PachyphytumLink, Klotzsch & Otto

Pachyphytum Link, Klotzsch & Otto, Allg. Gartenzeitung9:9-10 (1841) .

Subshrubs; stems first erect, with age usually de-cumbent to pendent, not or few-branched; leavesobovate, spathulate, elliptic-oblong or lanceo-late, usually very thick, usually conspicuouslyglaucous-farinose; inflorescences almost alwayssimple cincinni, first drooping,later * erect; floralbracts 5-9 mm in sect. Diotostemon or usually13-30 mm in sect. Pachypltytum; sepals erect,appressed, almost equal (sect. Diotostemon) or (of-

ten strongly) unequal (sect. Pachyphytum); petalserect (sect. Diotostemon) or spreading to divaricate

(sect. Pachyphytum), * oblong to oblanceolate,white to pink, rarely orange to red(dish), innerface in upper part often with red blotch in sect.P a chy phy t um, Iaterally ne ar b as e with app endageswhich form two free, 1-2 mm large scales beneathfilaments; antetepalous filaments connate withcorolla, antesepalous ones (almost) free; nectaryscales oblong, * yellowish(-white); carpels erectat anthesis, f free; stylodia inconspicuously offsetto abruptly narrowing; fruits usually divergentfollicles; seeds fairly smooth. n - 3l-33,62,64,66,96,99, +I24, +128, +160, +186. Fif teen species,in eastern central Mexico, (600-) I,200-2,500 m.

Divided into sections Diotostemon and Pachy-phytum (Thiede in Eggli 2003). The morphology ofthe petal scales was studied by Leinfellner (195a);similar scales also occur in som e Echeveria spe-cies. Pachyphytum maybe nested within Echeveriaand closest to its sect. Urceolatae (Thiede in Eggli2003).

II. SuspA,r,r. KaTRNcnoIDEAE A. Berger (1930).

Shoots * woody*; tissues (always?) with crystalsand*; petals connate to strongiy developed corollatubex; anthers with terminal, f spherical connec-tive appendage*; seeds with few (4-6) costae* inside view, coronate* . x - 9*.Genera 29-32.Note that Kalanchoideae are morphologicallyhighly derived, although cladistically they are thesecond clade branching off from the remainder ofthe family.

29. AdromischusLem.

Adromischus Lem., lard. Fleur. misc. 2:58-59 (1852);

Pilbeam et al., Adromischus (I998), synopsis.

Shrublets to * 20 cm; stems fleshy-woody; leavesflat to almost terete, glabrous or glandular-hairy,often with thick wax bloom; inflorescences erectspike-like thyrsoids or spikes without terminalflower, to 55 cm, with few to numerous, 1-5-flowered dichasia; flowers usually erect, rarelypendulous (A. phillippsiae (Marloth) Poelln.);corolla usually long and narrow; petals white topink to red, rarely bright orange, Iobes at sinusesjoined by thin membrane; filaments slightlyexserted or included, papillate where connatewith corolla tube; carpels elongate; fruits follicles,(always?) dehiscing completely along ventral

Crassulaceae

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suture. n = 9. About 28 species, Namibia, South

Africa (especially Succulent and Little Karoo).

30. Kalanchoe Adans. Fig. 28

Kalanchoe Adans., Fam. Pl. 2:248 (1763); Hamet, Bull.

Herb. Boiss. I I ,7:870-900 (1907) & 8:17-48 (1908); Hamet

& Lapostolle, Genre Kalanchoe au |ardin Botanique "Les

Cödres" (I96a); Raadts, Willdenowia 8: 101- 157 (1977 ), rev.

E. Afr.; Fernandes, Bol. Soc. Brot. II, 53:325-442 (1980)'

African taxa; Boiteau & Allorge-Boiteau, Kalanchoe de

Madagascar (1995); Gehrig et al., Pl. Sci. 160:827-835

(2001), mol. phylog.

Bry ophyllun Salisbury ( I 805).

Kitchingia Baker ( I 881 ).

Shrublets to shrubs, rarely rosulate or small trees,or biennial to annual; leaves usually decussate,rarely alternate, verticillate or subrosulate, * flat,rarelyterete, sometimes t incised or 3- to 5-foliate,margins usually crenate, serrate or dentate, partlywith bulbils (usually in sect. Bryophyllum), rarelyentire; inflorescences rarely axillary, corymboseor paniculate thyrsoids, partial inflorescencesdichasial, rarely inflorescences few- to 1-flowered;flowers 4-merous, * erect (usually in sect. Kalan-choe) or pendent (usually in sects. Kitchingia andBryophyllum); sepals free, connate or forminglong, sometimes t inflated tube (usually in sect.Bryophyllum); petals usually brightly coloured,lobes shorter than corolla tube, erect, spreading orreflexed; filaments exserted or included, connateto corolla tube at base (sect. Bryopltyllum) or at orabove middle (sects. Kalanchoe and Kitchingia);carpels free to somewhat connate at base, erect orsomewhat spreading (sect. Kitchingia); fruits erectfo l l ic les. n = usual ly 17, a lso 18,20,34,35,36,51,85. About 144 species, mainly Madagascar, easternand southern Africa, to tropical Africa, Arabia andtropical and Southeast Asia; some taxa (especiallyK. pinnata (Lam.) Pers.) are neophytic invadersthroughout the tropics.

Divided into three sections: the widespreadsect. Kalanchoe, and the Malagasy sects. Kitchin-gia and Bryophyllum.

31. Tylecodon Toelken

Tylecodon Toelken, Bothalia 12'378 (197S); van Jaarsveld &

Koutnik, Tylecodon & Cotyledon (2004), rev.

Shrublets or dwarf geophytes to pachycaul dwarftrees to 2.5 m; stems succulent, rarely woody,usually with flaking bark; leaves usually crowdedat stem tips, soft-herbaceous, with elongate

epidermal cells with sinuate anticlinal walls,often (always?) with bladder-cells idioblasts' usu-ally completely drought-deciduous; inflorescencesthyrsoids with one to several dichasia; petals white,greenish, yellowish or mauve, rarely reddish; fila-ments usually exserted, hairy where connate withcorolla tube; fruits follicles, dehiscing apicallyonly; seeds with irregular costae. Forty-six species,winter-rainfall regions of Namibia and SouthAfrica, mainly Succulent Karoo. Growing seasonautumn to early summer; flowering t in summer.

Sister to Cotyledon, according to moleculardata (Mort et al. 2001). The two genera have incommon basally hairy fi.laments.

32. CotyledonL.

Cotyledon L., Sp. Pl.:429 (1753); van Jaarsveld & Koutnik,

Tylecodon & Cotyledon (2004), rev.; Mort et al., Amer. |. Bot.

92:1170-1176 (2005), mol. phylog.

Procumbent to erect shrublets to shrubs, rarely

climbers; stems usually becoming woody; leaves

decussate, flat or terete, rarely lobed or orbicular,

glabrous or (glandular) hairy; inflorescences thyr-

soids with several dichasia, ending in monochasia

with one to many pendent flowers; corolla tube

hairy or glabrous; dried calyxlcorolla complex cir-

cumscissile along basal groove; filaments exserted,hairy where connate with corolla tube; carpels ta-

pering into erect stylodia;nectary scales * cuplike.

n = g.Eleven species, southern and eastern tropicalAfrica, south-western Arabian Peninsula.

According to molecular data (Levsen et al., l.c.),

the variable Cotyledon orbiculataL. is polyphyletic.

The circumscissile calyx/corolla complex present

in all Cotyledon is found also in at least some TyIe-

codon and Kalanchoe spp. (Moran 2000), and may

represent a synapomorphy for these three genera.

I I I . SusrA,M. CnRssur.oIDEAE Burnett (1835).

Leaves decussate*, rarely ternate or whorled; flow-

ers haplostemonous*; anthers slightly introrsex,

nucellus tenuinucellate*; fruits opening * com-

pletely along whole suture, but releasing seeds

through apical pore*; seeds sinuate-unipapil late*.Genera 33 + 34.Note that Crassuloideae are morphologically

highly derived, although cladistically they are the

first clade branching off from the remainder of the

family.

t12 J. Thiede and U. Eggli

Fig.31. Crassulaceae. Crassula columnaris. A Floweringplant. B Flowet opened out. C Young plant seen from above.(Berger 1930)

33. Crqssu laL. F ig .31

Crassula L., Sp. Pl;282 (I753); Toelken, Contr. Bolus Herb.

8 (1977), rev. southern African taxa; Tölken, f. Adelaide

Bot. Gard. 3:57-90 (1981), rev. Austral. taxa; Blwater &

Wickens, Kew Bull. 39:699-728 (1984), rev. New World

taxa; Mort et. al., IOS Bull. 12:35-36 (2004), mol. phylog.(abstract).

Ti l laeaL. (17s3).

Rochea DC. (1802) .

Perennial or rarely annual herbs to (sub)shrubs,rarely tuberous geophytes; glabrous, papillateor hairy; leaves decussate or rarely in whorlsof 4, partly with bladder-cell idioblasts and leafbases connate within a pair (usually in subg.Crassula); inflorescences thyrsoids with 1 tomany dichasia, sometimes partial inflorescencesglomerate, monochasia or reduced to solitaryflowers; corolla urn-shaped to tubular or stellate;flowers (2-)5(-12)-merous, usually small; sepalsshortly connate at base; petals shortly connate

at base, partly with apex papillate on outer faceand with distinct appendage (usually in subg.Crassula) often * whitish; filaments shortly adnateto petals at base and alternate with these; carpelsusually free; fruits rarely nutlike and indehiscent.n = 8,7 and polyploids. About 195 species; mainlysouthern Africa, a few species in sub-SaharanAfrica and south-western Arabia, some ephemeralherbs ('TiIIaea') distributed worldwide, and theoniy genus of the family in Australia.

Divided into the paraphyletic subg. Disporo-carpa with nine sections (hydathodes of type I,rarely type II; n = 8, rarely 7) and subg. Crassulawith eleven sections (hydathodes of type IIl' n - 7with two satellites; Friedrich 1.973; Toelken 1977l.c.; Martin and von Willert 2000). The ephemeralherbs of sects. Helophytum and Glomeratae, oftensegregated as genu s Tillaea, are nested within Cras-sula, according to molecular data ('t Hart unpubl.data).

34. Hypagophytum A. Berger

Hypagophytum A. Berger in Engler & Prantl, Nat. Pflanzen-fam., ed. 2, l8a:467-468 (1930); Gilbert, Opera Bot.12r:47-50 (1993).

Tuberous geophytes; stems one to few, drought-deciduous; leaves ternate, sessile or with petiole-like base, somewhat spurred at base, flat;inflorescences usually with 3 monochasia belowterminal flower; flowers 10-l2-merous, stellate;sepals free; petals white or with faint pink tinge;carpels free, laterally compressed, constricted intotwo segments, upper part spiny-papillate, withlong filiform stylodia; fruits 2-seeded, breakingtransversely at the constriction, upper seeddispersed within the upper part of the carpel,lower seed released separately. Only 1 species,H. abyssinicum (Hochst. ex A. Rich.) A. Berger innorth-western Ethiopian highlands.

Characterised by a unique combination of spe-cialised features, which all occur in Crassula (tu-bers with annual shoots, whorled leaves, hydath-odes along leaf margins, haplostemonous and poly-merous flowers, and the peculiar fruits). The seedsurface structure was given as costate by Gilbert(1989 and l.c.) and Knapp (1994), which prompted't Hart (1995: 169) to place the genus in his 'Se-

doideae'. However, according to Knapp (1997), theseed surface structure in fact corresponds to thesinuate-papillate (Crassula-) type, clearly favour-ing the placement in Crassuloideae. Hypagophy-tum may be nested within Crassula and closest to

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its sect. Petrogeton,whichshares the tuberous habitand exhibits in some species leaves with short peti-ole and in whorls of 4, monochasial inflorescenceswith stellate and polymerous flowers, and long fili-form stylodia. The same peculiar fruit type is foundin sect. Glomeratae p.p.(cf. Stopp 1957).

Selected Bibliography

Akiyama, S., Ohba, H., Wu, S.-K. 2001. A new variety ofSinocrassula paoshingensis (S.H. Fu) H. Ohba et al.(Crassulaceae). I . |ap. Bot. 76:222-226.

Alm, T. 2004. Ethnobotany of R hodiola rosea (Crassulaceae)in Norway. Sida 21, l:321-344.

Bahadur, 8., Ramaswamy, N., Srikanth, R. 1986. Studies onthe floral biology and nectar secretion in some Kalan-choe species (Crassulaceae). In: Kapil, R.P. (ed.) Pol-lination biology -an analysis. New Delhi: Inter-IndiaPublication s, pp. 251-259.

Baskin, J.M., Baskin, C.C. I97 2. Germination characteristicsof Diamorpha cymosa seeds and an ecological interpre-tation. Oecologia (Berlin) 10:17 -28.

Baskin, |.M., Baskin, C.C. 1977. Germination ecology ofSedum pulchellum Michx. (Crassulaceae). Amer. |. Bot.64:1242-1247.

Behnke, H.-D. 1991. Distr ibution and evolut ion of formsand types of sieve-element plastids in the dicotyledons.Al iso 13:167-182.

Berger, A. 1930. Crassulaceae. In: Engler, A., Prantl, K.,Die natürlichen Pflanzenfamilien, ed. 2, 18a. Leipzig:W. Engelmann, pp. 352-483.

Bland, K.P. 1995. Phytomyza rhodiolae Griffiths, 1976(Diptera: Agromyzidae), a leaf-miner in roseroot,Sedum rosea (Crassulaceae), new to Britain. Entomol.Gaz.46:267-269.

Boiteau, P., Allorge-Boiteau, L. 1995. Kalanchoe (Crassu-lacdes) de Madagascar. Systdmatique, dcophysiologieet phytochimie. Paris: Karthala,

Böttcher, W., läger, E.J. 1984. Zur Interpretation derVerbreitung der Gattung Sedum L. s.l. (Crassulaceae)und ihrer Wuchsformtypen. Wissensch. Z. Univ. Halle33:127-14I .

Bowman, R.N. 1983. Intraspecific variability of leaf cuti-cle alkanes in Sedum lanceolatum along an elevationalgradient. Biochem. Syst. Ecol. 11:195-198.

Braun, U.1987. A monograph of the Erysiphales (powderymildews). Nova Hedwigia, Beih. 89:l-700.

Braun-Blanquet, |., Sutter, R. 1982. Zur Kenntnis derCrassulaceen-Pioniergesellschaften in den Bünd-ner Alpen. |ahresber. Naturf. Gesell. Graubünden99:75-83.

Burchard, O.1929. Beiträge zur Ökologie und Biologie derKanarenpflanzen. Bibl. Bot. 98:l-262, pls. 1-78.

Byalt, V.V. 1997. Meterostachys sikokianas (Crassulaceae),a new species and genus for the flora of China (inRussian with English summary). Bot. Zhurn. (Moscow& Leningrad) 82:128-1 30.

Byalt, V.V. 1998. Orostachys paradoxa, a rare species fromthe Russian far East. Cact. Succ. I. (U.S.) 70:262-263.

Bywater, M. 1980. Observations on seeds of Crassula sect.Rosulares. Kew Bull. 35:401-402.

Bywater, M., Wickens, G.E. 1983. New world species of thegenus Crassula. Kew Bull. 39:699-728.

Caballero, A., |imdnez, M.S. 1977. Contribuciön al estudioanatömico foliar de las crassuläceas canarias. Vieraea7:I l5-132.

Calie, P.l. 1981. Systematic studies inSedum section Ternata(Crassulaceae). Britton ia 33:498-507 .

Candolle, A.P. de 1828. Mdmoire sur la famille des Crassu-lacdes. Paris: Treuttel & Würtz.

Clausen, R.T. 1959. Sedum of the Trans-Mexican VolcanicBelt: an exposition of taxonomic methods. Ithaca:Comstock.

Clausen, R.T. 1975. Sedum of North America north of theMexican Plateau. Ithaca: Cornell University Press.

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c

d

The Familiesand Generaof Vascular PlantsEdited bv K. Kubitzki

T \f Flowering Plants . EudicotsI /\ Berberidopsidales, Buxales, Crossosomatales,

Fabales p.p., Geraniales, Gunnerales, Myrtales p.p.,P r o t e al e s, S axifr agal e s, Vit aI e s, Zy go phyll aI e s,Clu s i a c e a e Alli an c e, P a s s ifl o r a c e a e Alli an c e,Dilleniaceae, Huaceae, Picramniaceae, Sabiaceae

Volume Editor:K. Kubitzkiin Collaboration with C. Bayer and P. F. Stevens

With 174 Figures

üspringer

Prt. r tessor Dr. Kraus KusrrzrrL'niversität HamburgBiozentrum Klein-Flottbek und Botanischer GartenOhnhorststraße 1822609 HamburgGermany

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