the scilla plumbea puzzle-present status of the genus scilla sensu lato in southern africa and...

INTRODUCTIONIn 1830 J Lindley first described Scilla plumbea

only known from a drawing made by the late MrSydenham Edwards in 1813 in Kew Gardens (Fig 1A)from a bulb that had been imported from the Cape (SouthAfrica) Attempts to find the new taxon in living collec-tions near London were without success (Lindley 1830)Kunth (1843) recognized Scilla plumbea (as ldquospeciesanomalardquo) from a collection by Dregravege (no 1997) at theDutoitskloof (South Africa) Kunth realized howeverthat Dregravegersquos plant had narrower leaves than the icono-type of Scilla plumbea Baker (1897 482) opined thatldquothe little known Scilla plumbea is probably a form ofS natalensis [= Merwilla natalensis Fig 1B] and maybe the same as [S natalensis] var sordidardquo Kunthrsquos(1843) opinion was supported by Lewis (1947) who gavean illustration (Fig 1C) and a rather detailed descriptionof a plant collected by J S Linley sn in Baviaanskloofoff Bainrsquos Kloof not far from Dutoitskloof (SouthAfrican Museum Herbarium No 56004 compare withFig 1D) He mentioned three differences between thisnew collected plant and the iconotype of Scilla plumbealdquo(i) In Lindleyrsquos type the leaf is considerably broader andappears to be flat not convolute (ii) Lindleyrsquos plant has

dull blue flowers with slightly shorter and broader peri-anth-segments which with the exception of one or twoflowers only are shown spreading from the base (iii)The style is very much shorter in Lindleyrsquos typerdquo Basedon examination of Lindleyrsquos iconotype Speta (1998a)transferred Scilla plumbea Lindl into the genus Merwillaas M plumbea (Lindl) Speta Most recent treatments ofthe South African flora have accepted Kunthrsquos andLewisrsquo view of a broad species concept for S plumbea(Jessop 1975 Reid 1993 Goldblatt amp Manning 2000Williams 2000) According to Reid (1993) and Williams(2000) the genus Scilla L is represented in southernAfrica by at least four species (S firmifolia S natalen-sis S nervosa S plumbea) Speta (1998a) not only pro-vided an extensive historical treatment of Scilla but alsopresented a new generic concept He viewed Scilla Lsstr as being restricted to the Northern Hemisphere intribe Hyacintheae Dumort whereas South African taxawere treated as belonging to tribe Massonieae BakerTaxa in these two regions therefore should be placedinto different genera Speta (1998a) placed Scilla speciesrecognized by Reid (1993) into the genera MerwillaSpeta (Fig 1A) Schizocarphus Merwe (Fig 2A) andPseudoprospero Speta (Fig 2B) He also adopted a nar-rower species concept than that of Reid (1993) accept-

75

Wetschnig amp Pfosser The Scilla plumbea puzzle52 February 2003 75ndash92

The Scilla plumbea puzzlemdashpresent status of the genus Scilla sensu lato insouthern Africa and description of Spetaea lachenaliiflora a new genus andspecies of Massonieae (Hyacinthaceae)

Wolfgang Wetschnig1 amp Martin Pfosser2

1 Institute of Botany University of Graz Holteigasse 6 A-8010 Graz Austria wolfgangwetschnig uni-grazat (author for correspondence)

2 Department of Higher Plant Systematics and Evolution Institute of Botany University of Vienna Rennweg14 A-1030 Vienna Austria martinpfossermailboxunivieacat

We have studied a taxon from South Africa that has been hidden for more than 150 years within Scilla plum-bea (Hyacinthaceae) However phylogenetic reconstruction based on trnL-F data morphological evidence(including seed characters) and karyology suggest independent status DNA data place it close to theDaubenya alliance but not close to any particular genus of Massonieae previously included within Scilla slUnique characters of seeds indicate that this taxon does not belong to the Daubenya clade Analysis of the icon-otype of Scilla plumbea Lindl on the other hand reveals morphological details suggesting membership in theScilla natalensis clade (= genus Merwilla in the recent classification of Speta) Merwilla however occupies abasal position among Massonieae and is not related to the Daubenya clade Based on all data we describe thistaxon as a new genus and species Spetaea lachenaliiflora Wetschnig amp Pfosser

KEYWORDS chromosome numbers flora of southern Africa Hyacinthaceae Massonieae plastid DNAsequences Scilla plumbea seed morphology Spetaea lachenaliiflora

Dedicated to Doz Dr Franz Speta on the occasion of his 60th birthday

Wetschnig amp Pfosser The Scilla plumbea puzzle 52 February 2003 75ndash92

Fig 1 Morphology of Merwilla and Spetaea A Merwilla plumbea (Lindley) Speta t 1355 in Lindley (1830) as Scillaplumbea Lindley B Merwilla natalensis (Planchon) Speta t 365 in Phillips (1930) as Scilla natalensis Planchon CSpetaea lachenaliiflora t 1006 in Lewis (1947) as Scilla plumbea Lindley D Spetaea lachenaliiflora WW 1527 photo-graphed at the natural locality at Bainrsquos Kloof in Dec 1987

76

ing 12 species in these three genera Several phylogenetic studies in which South African

members of Hyacinthaceae were analyzed already indi-cated the heterogeneity of a large genus Scilla sl (egSpeta 1998a b Stedje 1998 Pfosser amp Speta 1999)Moreover our previous studies on seed morphology andphylogeny of Massonieae (Wetschnig amp al 2002a bPfosser amp al in press) showed that Scilla plumbea sensuKunth (1843) and Lewis (1947) from the Bainrsquos Kloofand Dutoitskloof area represents an independent taxondistinct from other members of Massonieae

As already shown by Speta (1998a) and Pfosser ampSpeta (1999) it is difficult to find morphological charac-ters yielding synapomorphies for groups of taxa withinHyacinthaceae For example Speta (1998a) mentionedldquoIm Bereich der Morphologie ist festzustellen daszlig inallen Subfamilien beinahe alle machbaren Abaumlnderungenstattgefunden haben was zeigt daszlig die Mutationen ingewissen lsquoBahnenrsquo verlaufen Merkmalsgleichheit mehr-mals und immer wieder entstehen kann Stabile Merk-male die so wenig veraumlnderlich sind daszlig sie fuumlr denSystematiker besonders wertvoll sind sind kaum welcheaufzufindenrdquo This holds true also for genera withinMassonieae Besides seed morphology only combina-tions of characters can be used to delimit genera

The aim of the present study was to determine thephylogenetic position of this taxon in relation to othermembers of Hyacinthaceae previously included withinthe genus Scilla Based on DNA sequence data seedmorphology and karyology we conclude that this taxonis sufficiently different from other genera of Massonieaeto warrant generic status We describe the new genus andspecies Spetaea lachenaliiflora (Fig 1C D) in honor ofFranz Speta who has contributed so much to disentan-gling the confused relationships within Hyacinthaceae

MATERIALS AND METHODSTaxa sampled mdash This analysis is based on mate-

rial of subfamily Hyacinthoideae from South AfricaEurasia and East Asia In total 151 individual plantscomprising 45 genera of Hyacinthaceae have been ana-lyzed The majority of genera came from subfamilyHyacinthoideae (37) three from subfamily Urgineoi-deae four from subfamily Ornithogaloideae and onefrom Oziroe from subfamily Oziroeoideae that has beenused as outgroup to root the phylogenetic trees Voucherinformation for all plant accessions geographic originand EMBL database accession numbers are provided in


Fig 2 Morphology of Schizocarphus and Pseudoprospero A Schizocarphus nervosus (Burch) Merwe t 904 in Merwe(1943) as Scilla rigidifolia var nervosa (Burch) Baker B Pseudoprospero firmifolium (Baker) Speta t 926 in Merwe(1944) as Scilla firmifolia Baker

77

Appendix 1 Nomenclature follows that of Speta (1998ab)

DNA extraction mdash Total genomic DNA wasextracted from lyophilized and powdered leaf or seedmaterial in 700 microl CTAB buffer (2 CTAB 100 mMTris 14 M NaCl 20 mM EDTA 02 mercaptoethanolpH 80) for 30 minutes at 60deg C 500 microl chloroformiso-amylalcohol (241) were added and the extraction mixwas incubated for 15 minutes at 4degC After centrifuga-tion the DNA was precipitated with 500 microl isopropanolThe pellet was washed with 70 ethanol and dissolvedin 100 microl TE buffer

DNA sequencing mdash Two non-coding regions ofthe plastid genome were sequenced The trnL(UAA)intron and the intergenic spacer (IGS) between thetrnL(UAA)-3acuteexon and the trnF(GAA) gene were ampli-fied together in a single PCR reaction (Pfosser amp Speta1999) Amplified double-stranded DNA fragments weresequenced directly on an ABI377 automated sequencer(Perkin Elmer UK) following the DYEnamicET cyclesequencing protocol (Amersham Pharmacia USA)Both strands were sequenced using the nested sequenc-ing primers as described for the trnL-F region (Pfosser ampSpeta 1999) On average less than 1 of data matrixcells were scored as missing data

Phylogenetic analysis mdash Sequence manipula-tions were performed on a Digital Alpha 1000A 5400server under the operating system Digital Unix V40DDNA sequences were pre-aligned using the PileUp pro-gram of the GCG software package (Genetics ComputerGroup 1994) Final alignment of DNA sequences wasdone visually The sequences have been trimmed on bothends to exclude ambiguous positions in close proximityto the sequencing primers All sequences have beendeposited in the EMBL database (for accession numbersrefer to Appendix 1) Twenty-five indels in the datamatrix were coded as additional characters and treesearches were performed using the nucleotide datatogether with the indel data Phylogenetic analysis usingmaximum parsimony (MP) neighbor joining (NJ) andmaximum likelihood (ML) methods were performedwith the computer program PAUP version 40b10(Swofford 2000) MP analyses were performed eitherwithout or with successive character weighting (rescaledconsistency index) until tree lengths remained the samein two successive rounds Most parsimonious trees wereobtained by 1000 replicates of random sequence additionusing tree bisection-reconnection (TBR) branch swap-ping under the Fitch criterion (Fitch 1971) Ten thousandfast bootstrap replicates (Felsenstein 1985) were used toassess confidence limits for the resulting tree topologiesPhylogenetic reconstructions using Bayesian inferenceof phylogeny (BI) were performed with the computerprogram MrBayes (Huelsenbeck amp Ronquist 2001)

Three Markov Monte Carlo chains were run simultane-ously for 200000 generations Trees were sampled every200 generations Trees sampled after reaching stationaryphase (after c 51000 generations) were collected andfinally a majority rule consensus tree was constructedTree manipulations were performed using MacClade ver-sion 306 (Maddison amp Maddison 1992)

Morphological analysis mdash All examinations ofseed morphology were carried out on fully developed dryseeds Weight and size (arithmetic means) have beendetermined from at least ten seeds Dried seeds weremounted on aluminum stubs with ldquoLeit-Tabsrdquo and coat-ed with gold in an Agar sputter coater Electron micro-graphs were obtained with a Philips XL 30 ESEM scan-ning electron microscope operating at 20 kV Seeds wereoriented with the chalazal pole pointing left and themicropylar pole pointing right Lateral views were ori-ented with the raphe facing upwards

Karyological analysis mdash Root tip meristems ofplants grown in soil were used for studying chromosomemorphology Root tips were pretreated for 12 hours in a0002 M solution of 8ndashhydroxyquinoline at 5deg C Afterpretreatment the root tips were fixed in ethanolaceticacid 31 After staining with carmine acetic acid squashpreparations of the meristems were performed All meas-urements for constructing idiograms were obtained fromdrawings made with the drawing equipment of a ZeissAxioplan 2 imaging microscope Digital photographswere taken using the Axiocam camera (Zeiss) Ten well-scattered metaphase plates were used to produce idio-grams Calculation of symmetry values (Gi Si) ratio ofshortlong arm (r) relative length of the chromosome(cr) and -values of long and short arms and the draw-ing of idiograms were done using the computer programCHROM (Wetschnig 1992) Nomenclature of chromo-somes and centromeric positions follows Levan amp al(1964)

RESULTSPhylogenetic position of Spetaea mdash Based on

chloroplast DNA data a clear separation among the foursubfamilies of Hyacinthaceae is visible (Figs 3 4) Aclassification of genera of Hyacinthoideae into the twotribes Hyacintheae and Massonieae is also reflected inthe molecular data although tribe Massonieae appearsparaphyletic due to the basal position of PseudoprosperoDNA sequence data suggest this genus to be sister to therest of Hyacinthoideae Basal within Hyacintheae is theEast Asian Barnardia scilloides species complex (Fig 3)The remaining taxa of Hyacintheae fall into three majorclades Clade HI consists of W MediterraneanWEuropean taxa (64 bootstrap support) The Linnaean


78


Fig 3 Majority rule phylogenetic consensus tree (maximum parsimony) of tribe Hyacintheae Bootstrap support valuesafter character weighting (first number) and percent consensus values calculated from Bayesian inference of phylo-geny (second number) are indicated above branches Nodes collapsing in the strict consensus tree are marked by opencircles The highly supported Scilla sstr clade is highlighted in bold Vertical bar indicates a synapomorphic indelMajor clades within Hyacintheae consisting of more than a single genus are indicated by background shadingRepresentatives of subfamilies Urgineoideae Ornithogaloideae and Oziroeoideae (used to root the tree) have beenincluded in the analysis

Zagrosia persica 1 Zagrosia persica 2

Pfosseria bithynicaHyacinthus orientalis Hyacinthus orientalis var alba

Fessia vvedenskyiFessia puschkinioides

Hyacinthus litwinowii Fessia greilhuberi

Othocallis spPuschkinia scilloides

Prospero elisae 1 Prospero elisae 2

Prospero haritonidaeProspero obtusifolium

Hyacinthella dalmaticaHyacinthella heldreichii

Scilla cf bulgaricaScilla spetana

Scilla sieheiScilla nana

Scilla albescensScilla subnivalis

Scilla cydonia 1Scilla cydonia 2

Bellevalia brevipedicellataBellevalia trifoliata

Muscari botryoides Muscari comosum

Muscari parviflorum Muscari macrocarpum Schnarfia messeniaca

Nectaroscilla hyacinthoidesChouardia litardierei

Hyacinthoides non-scriptaHyacinthoides hispanica

Hyacinthoides vincentina Hyacinthoides reverchonii

Hyacinthoides lingulata Hyacinthoides aristidis

Hyacinthoides italica Autonoe haemorrhoidalis 1

Autonoe haemorrhoidalis 2Autonoe latifolia

Oncostema dimartinoi Oncostema villosa

Oncostema peruvianaTractema lilio-hyacinthus

Tractema monophyllosBrimeura amethystina

Barnardia scilloides 1Barnardia scilloides 2

Pseudoprospero firmifoliumCharybdis undulata

Charybdis aphyllaCharybdis hesperia

Rhadamanthus mascarenensis Bowiea volubilis

Bowiea spStellarioides longebracteata Stellarioides sp

Pseudogaltonia clavataDipcadi cf heterocuspe

Ornithogalum kochiiOziroe biflora

Oziroe acaulis

5 changes

100100

5460

Massonieae (Fig 4)

1001006087 8995

100100

100100100100

100100

100100

81100

80100

94100

6485

9498

-80

--

100100

-67

99100

96100

85100

100100

--92100

82100

98100

100100

5786

100100

961005599

83100

96100

86100

99100

-63

98100

86100

-91

86100

5899

MASSONIEAE

OZIROE-OIDEAE

ORNITHOGAL-OIDEAE

URGINE-OIDEAE

HYACINTHEAE HYACINTH-OIDEAE

100100

HIII

HII

HI

79


Fig 4 Majority rule phylogenetic consensus tree (maximum parsimony) of tribe Massonieae Bootstrap support valuesafter character weighting (first number) and percent consensus values calculated from Bayesian inference of phylo-geny (second number) are indicated above branches Nodes collapsing in the strict consensus tree are marked by opencircles The genus and species Spetaea lachenaliiflora is highlighted in bold Vertical bars indicate synapomorphicindels Major clades within Massonieae consisting of more than a single genus are indicated by background shadingRepresentatives of subfamilies Urgineoideae Ornithogaloideae and Oziroeoideae (used to root the tree) have beenincluded in the analysis

Lachenalia pallida 1Lachenalia aloides Lachenalia pallida 2

Lachenalia rubida 2Lachenalia spLachenalia rubida 1

Lachenalia pusilla Lachenalia cf pusilla 1Lachenalia cf pusilla 2

Lachenalia contaminata Polyxena ensifolia 6Polyxena ensifolia 5Polyxena ensifolia 1Polyxena ensifolia 4Polyxena ensifolia 3Polyxena ensifolia 2

Polyxena spPeriboea oliveri

Periboea paucifolia Polyxena calcicola

Massonia depressa 5Massonia sp 2 Massonia depressa 4

Massonia tenella Massonia pustulata 2Massonia pustulata 1

Massonia depressa 3Massonia cf sessiliflora

Massonia echinata 3Massonia sp 1Massonia echinata 2Massonia echinata 1Massonia depressa 2Massonia depressa 1

Whiteheadia bifoliaWhiteheadia etesionamibensis

Veltheimia bracteata 1Veltheimia bracteata 3Veltheimia bracteata 2Veltheimia bracteata 4

Amphisiphon stylosa 1Amphisiphon stylosa 3Amphisiphon stylosa 2Massonia marginata Massonia zeyheri 1Massonia zeyheri 2Androsiphon capense

Daubenya aurea Spetaea lachenaliiflora

Eucomis zambesiaca Eucomis vandermerwei

Eucomis montana Eucomis punctata Eucomis bicolor

Ledebouria hyacinthina 2Ledebouria somalensis

Ledebouria hyacinthina 1Resnova humifusa Resnova maxima

Drimiopsis maculata Drimiopsis spDrimiopsis kirkii

Drimiopsis botryoides Drimiopsis barteri

Ledebouria socialis Ledebouria socialis agg

Ledebouria cf concolor Ledebouria sp 2

Ledebouria cordifolia Ledebouria revoluta

Ledebouria sp 3 Ledebouria sp 4


Ledebouria sp 7 Ledebouria sp 8 Ledebouria floribunda

Ledebouria sp 1 Schizocarphus nervosus 2Schizocarphus nervosus 1

Merwilla sp 2 Merwilla sp 1

Merwilla natalensis 2Merwilla natalensis 1Merwilla kraussii

Pseudoprospero firmifolium Charybdis undulata

Charybdis aphylla Charybdis hesperia



Pseudogaltonia clavata Dipcadi cf heterocuspe

Ornithogalum kochii Oziroe biflora

Oziroe acaulis

5 changes

Hyacintheae (Fig 3)

100100

100100

5460

100100

60878995

100100

100100

100100

100100

8110080100

5599

86100

6499

84100100100

89100

100100

---96

709793100

84100

6493

100100

--

-86

9798

100100

87100

100100

5887

--

5252100100

921006090

100100

94100

6095

6498

66100

98100

99100

MASSONIEAE

OZIROE-OIDEAE

ORNITHOGAL-OIDEAE

URGINE-OIDEAE

MASSONIEAE

HYACINTH-OIDEAE

MIa

MIb

MII

MIIIa

MIIIb

80

type of the genus Scilla falls together with BellevaliaMuscari Schnarfia Nectaroscilla and Chouardia intothe highly supported clade HII (92 bootstrap support)All taxa within this clade are characterized by a synapo-morphic indel found only here and in no other taxon ofHyacinthaceae The third highly supported clade (HIII)contains genera with a predominant Asia MinorEMediterranean distribution

Twenty-four indels in the data matrix have beenfound to be synapomorphic for genera or clades ofMassonieae (Fig 4) If Pseudoprospero is excluded therest of Massonieae forms a monophyletic clade (86bootstrap support) In this clade genus Merwilla (=Scilla natalensis relationship) occupies an isolated basalposition whereas all other species previously included inScilla show other affinities A high bootstrap support forthe clade combining the remaining genera of Massonieae(89) underlines the isolated position of MerwillaSchizocarphus nervosus (= Scilla nervosa) is looselyrelated to a well-supported clade consisting of Lede-bouria Drimiopsis and Resnova (100) Spetaealachenaliiflora (= Scilla plumbea sensu Kunth andLewis) is basal in a clade with Daubenya AndrosiphonAmphisiphon Massonia marginata and M zeyheri(97) However long branches and 100 bootstrap sup-port for the taxa from Daubenya to Amphisiphon do notsuggest a direct relationship of Spetaea with this clade Itprobably should be better viewed as the sister group Thegenus Eucomis appears as a well-supported monophylet-ic group Veltheimia is sister to a clade consisting ofWhiteheadia the remaining species of MassoniaPeriboea Polyxena and Lachenalia High bootstrap val-ues in this group exclude the possibility that Spetaeamight be related to any of these genera

Different tree-building algorithms performed on thetrnL-F dataset all yielded similar tree topologies (Table1) In most cases highly supported clades (gt70 boot-strap support) received high values irrespective of themethod applied for phylogenetic reconstruction A nar-rowly defined genus Scilla sstr received sup399 supportin all analyses The enclosing clade HII consisting ofNectaroscilla Chouardia Schnarfia Muscari Belle-valia and Scilla was also highly supported (sup377 sup-port in all analyses) This already excludes the possibili-ty that any other taxon outside of this group could be amember of the genus Scilla unless a highly polyphyleticgenus is accepted Within Massonieae clade MII(including Spetaea) is highly supported (sup390 supportin all analyses) A sister group relationship betweenSpetaea and the rest of clade MII is also supported frommultiple analyses (100 support)

Seed morphology of Spetaea and selectedgenera of Massonieae mdash Seed morphological datahave been determined for Spetaea Daubenya

Schizocarphus and Merwilla1 Spetaea (Figs 5A 6A B) Weight 00048 g

Lengthwidthheight 2617519 mmThe shape of the seed of Spetaea lachenaliiflora is

ellipsoidal to ovoid and slightly compressed (Fig 6A B)Depending on the number of seeds developed per ovule[(1)ndash2ndash(3)] their shape is regular or somewhatdeformed Sometimes a slight flattening in the area of thechalaza appears but normally a compressed conicalswelling of the seed coat is seen in this as well as in themicropylar region The endosperm is not as soft as that ofMerwilla and not as hard as in members of theDaubenya-clade Color of the endosperm has a slightlyolive touch The embryo is plusmn straight and has a relativelength of about 82 of the length of the endosperm Theplusmn broadened micropylar pole of the embryo slightlyexceeds the outline of the endosperm The seed coat ishard wrinkled black and shining (Fig 5A) The hilum isa distinctive character of the seeds of Spetaea lachenali-iflora It is positioned at the raphal side of the conicaloutgrowth of the seed coat at the micropylar region andextends from almost the tip of this conus approximatelyhalf way down the seed length The form of the hilum islanceolate The white or slightly brownish color of thehilum contrasts with the black seed coat (Fig 5A) Theraphe forms a ridge extending from the chalazal end ofthe hilum to the chalazal region of the seed

Spetaea lachenaliiflora is easily distinguished fromall other members of Massonieae by its unique seed mor-phology A somewhat similar wrinkled seed coat and aconspicuous large hilum is found only in Veltheimia andWhiteheadia (Fig 5C D 6D) However size and shapeof the seeds and color and shape of the hilum differ fromSpetaea Seeds of the Daubenya clade have a smoothseed coat and a more globular shape of the seed (Fig6C) The hilum is much smaller and of different shape(Fig 5B) than that of Spetaea although it is easily visi-ble because of its contrasting color

2 Daubenya (Figs 5B 6C) Weight 00114 gLengthwidthheight 262524 mm

The shape of the seed of Daubenya aurea is globu-lar rarely slightly ellipsoidal (Fig 6C) Sometimesdeformations occur due to space limitations of the devel-oping young seeds within one locule A slight flatteningat the chalazal region may occur Occasionally a conicalswelling and folding of the seed coat can be found in thisarea as well as at the hilum The endosperm is hard andcolorless-hyaline to whitish The straight embryo has arelative length of about 80 of the length of theendosperm The somewhat broadened micropylar pole ofthe embryo slightly exceeds the outline of theendosperm The chalazal flattening and the micropylarswelling are not in the longitudinal axis of the embryobut angular above the axis at the raphal side The seed


81

coat is hard brownish-black or black The seed coatappears smooth in well-developed seeds The surface is

shining and a polygonal cellular pattern with a diameterof about 20 microm can be observed The hilum is small and


Table 1 Tree scores and support values gt50 ( bootstrap consensus) of clades within Hyacinthoideae as obtainedfrom different phylogenetic reconstruction methods MP maximum parsimony (10000 fast bootstrap replicates)MP+SW maximum parsimony with successive character weighting (10000 fast bootstrap replicates) BI Bayesian infer-ence of phylogeny (consensus of 740 trees collected after burn-in of Markov chains) NJ neighbor joining (10000 repli-cates) ML maximum likelihood (100 bootstrap replicates) Values gt70 and clades receiving gt70 support in allanalyses are printed in bold

MP MP+SW BI NJ MLClade bootstrap bootstrap consensus bootstrap bootstrap

Hyacinthoideae 68 81 100 62 65HyacintheaeMassonieae (exc Pseudoprospero) - 55 99 - -Hyacintheae - 80 100 - 52Barnardia 89 94 100 93 87HIHIIHIII - - - - -HI - 64 85 - -TractemaOncostema - - 80 60 -AutonoeHyacinthoides - - 67 - -Autonoe 100 100 100 100 100Hyacinthoides 95 99 100 97 90Hyacinthoides italicaaristidis lingulata 97 96 100 97 97H reverchoniivincentinahispanicanon-scripta 81 85 100 85 71HII 79 92 100 81 77SchnarfiaMuscariBellevaliaScilla 58 82 100 52 65Muscari 92 98 100 98 94BellevaliaScilla - 57 86 - -Scilla 99 100 100 100 100HIII 94 96 100 98 100Hyacinthella 82 83 100 53 65Prospero 85 86 100 78 94OthocallisFessiaHyacinthusPfosseriaZagrosia 92 98 100 91 77FessiaHyacinthus litwinowii - 58 99 - 55FessiaHyacinthusPfosseriaZagrosia 87 86 100 92 81Hyacinthus orientalisPfosseriaZagrosia - - 91 55 -Massonieae (excl Pseudoprospero) 66 86 100 53 71Merwilla 58 64 99 85 65SchizocarphusMIEucomisMIIVeltheimiaMIII 76 89 100 64 77MI 99 100 100 100 100MIa 51 - 96 55 -MIb 67 70 97 65 61Drimiopsis 77 84 100 72 71ResnovaLedebouria hyacinthina L somalensis 64 64 93 67 65Eucomis 100 100 100 100 100MIIVeltheimiaMIII - - 86 - 52MII 96 97 98 90 97MII excl Spetaea 100 100 100 100 100AmphisiphonMassonia zeyheriM angustifolia 87 87 100 86 71VeltheimiaMIII 58 58 87 - 61Veltheimia 100 100 100 100 100MIII 89 92 100 84 90MIIIa - - - - -MIIIa excl Whiteheadia etesionamibensis 54 52 52 - 52MIIIa excl Whiteheadia 99 100 100 98 100MIIIb 100 100 100 100 100PeriboeaPolyxena 60 60 95 59 55Lachenalia 65 66 100 88 61

Tree length 720 456995 719 725 721Consistency index (CI) 0704 0874 0704 0699 0703Retention index (RI) 0917 0965 0917 0915 0917Rescaled consistency index (RC) 0646 0844 0646 0640 0645Homoplasy index (HI) 0296 0126 0296 0301 0297

82

elliptic or ovate (Fig 5B) In most seeds its pale browncolor contrasts with the dark brown or black seed coat

Seeds of the monotypic Daubenya show strong sim-ilarities with those of Amphisiphon stylosa Androsiphoncapense Massonia marginata (= M angustifolia L ffide Manning amp van der Merwe 2002) and Massoniazeyheri (Wetschnig amp al 2002a b) Differences occur insize and color of the seeds the extent and size of theswellings of the testa and the surface of the testaHowever it is difficult to distinguish these taxa by seedmorphology although they are unique among Masson-ieae and therefore easily distinguishable from other taxaof this tribe

3 Schizocarphus (Fig 6E) Weight 00022 gLengthwidthheight 2315ndash(17)14ndash15 mm

Seeds of Schizocarphus nervosus are ellipsoidal toovoid (Fig 6E) with occasional flat facets These facetsare caused by deformations of the seed coat and not bydeformations of the endosperm The shape of theendosperm is ellipsoidal colorless-hyaline to whitishand not as soft as that of Merwilla natalensis Thestraight embryo is positioned in the longitudinal axis ofthe endosperm measuring 80 of the endosperm lengthAt the micropylar end the embryo slightly exceeds theoutline of the endosperm The color of the seed coat isbrownish-black It is soft and breaks into small pieces

when damaged The basic structure of the seed coat is asystem of brain-like wrinkles The slightly shining sur-face of these wrinkles (primary structure of the seedcoat) shows a polygonal to orbital pattern with a diame-ter of the fields of 30ndash50 microm The hilum can be found inmost seeds but is small and inconspicuous The raphe iseither invisible or can be observed only as a vertex with-in the wrinkles of the seed coat

The seeds of Schizocarphus nervosus are easy to dis-tinguish from all other genera of Massonieae Similarityin the primary structure of the seed coat exists only withthat of Merwilla

4 Merwilla (Fig 6F) Weight 00045 g Lengthwidthheight 6434 (at the broadest position)1ndash23mm

The seeds of M natalensis appear obtuse-triangularto slightly crescent in lateral view (Fig 6F) The shape ofthe transverse section is compressed-elliptic flat-ellipticwith a distinct center rib or flat-triangular That seeds areflat or triangular in transverse section is caused by theirposition within the locule The seed coat forms narrowwings at the edges and above the center rib of theendosperm The endosperm is narrow-obovate to lanceo-late Frequently a rib is formed at the raphal broadsideThe endosperm is yellow and softer than that of othermembers of Massonieae The straight embryo is com-


83

Fig 5 Seed morphology of Spetaea lachenaliiflora (A hilum) Daubenya aurea (B hilum) Veltheimia bracteata (Chilum) and Whiteheadia bifolia (D hilum and raphe) Bar = 05 mm

pressed and only slightly shorter than the endospermThe color of the testa is light-brown to beige in dry seedsThe seed coat is soft and easily detachable with a needleand breaks into small pieces when damaged The surfaceof the unwrinkled seed coat (primary structure of theseed coat) shows a polygonal to plusmn orbital pattern of50ndash79 microm in diameter The edges of these polygons aresomewhat raised and most likely correspond to cell

walls The seed surface is dull In most seeds the raphe isnot visible The vascular bundle leads through the moreor less narrow-winged edge of the seed coat

Seeds of Merwilla natalensis are the most distinctiveamong Massonieae The shape of the seeds the color ofthe seed coat and the yellow and soft endosperm areunique in this tribe Furthermore the seeds germinatefast (the epigaeal germination starts already 5ndash9 days


84

Fig 6 Scanning electron micrographs of seeds of Spetaea lachenaliiflora and relevant genera of Massonieae A BSpetaea lachenaliiflora (B with hilum and raphe) C Daubenya aurea D Veltheimia bracteata E Schizocarphus ner-vosus F Merwilla natalensis AndashE lateral view F raphal view Bars = 1 mm

after sowing) The primary structure of the seed coatshows similarity with that of Schizocarphus nervosus

5 Pseudoprospero No seeds of this monotypicgenus [P firmifolium (Baker) Speta] were available forthis study Fig 4c in Jessop (1975) shows a papillose pri-mary structure of the seed coat for this species The

papillae are flattened with three grooves and the cells are5- or 6-angled with a diameter of 25ndash35 microm A reductionof the height of such papillae could lead to a similar pri-mary structure as that found in Merwilla andSchizocarphus

Karyology of Spetaea and chromosome


85

Fig 7 Karyology of Spetaea lachenaliiflora A phase-contrast micrograph of a metaphase plate of Spetaea lachenaliif-lora 2n = 20 B idiogram of the diploid chromosome set Chromosomes are arranged according to their cr-values (rela-tive length of the chromosome) s short arm l long arm r ratio of ls = percentage of whole karyotype length Bar= 10 microm

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

s 08 08 19 11 13 11 12 11 11 10 09 09 08 08 08 06 08 06 06 06 microm10 10 23 14 16 14 14 13 13 12 12 11 10 10 10 08 10 08 08 08

l 147 147 53 50 38 31 12 12 11 11 10 09 10 09 09 11 08 08 07 07 microm12 14 microm

180 180 65 61 46 38 15 15 13 13 12 12 12 12 12 13 10 09 09 09 14 17

r 188 188 34 57 29 28 10 11 10 11 11 10 12 11 12 17 10 12 12 12

cr 190 190 103 92 62 52 29 28 27 25 24 23 22 22 21 21 19 17 17 17

B

counts in Massonieae mdash Spetaea lachenaliiflora hasa diploid chromosome number of 2n = 20 (Fig 7A BTable 2) Diploid chromosome numbers among relativesof Spetaea range from 2n = 10 to 2n = 56 (Table 2) Thekaryotype of Spetaea lachenaliiflora (Fig 7A) is veryasymmetric with Si = 3032 (Si = sum of the short armstimes100sum of the long arms) Differences in length amongshort chromosome arms are small ranging from 063 to191 microm Chromosomes 7ndash20 show long arms of 072 to122 microm length whereas chromosomes 1ndash6 show con-spicuous longer arms ranging from 313 microm (chromo-somes 6) to 1469 microm (chromosomes 1 and 2) Theselong arms account for the high Gi-value of 869 (Gi =total length of the smallest chromosome times100totallength of the longest chromosome) Chromosomes 1 and2 account for 38 of the total length of the diploid chro-mosome set Chromosomes 3 and 4 account for 195and chromosomes 5 and 6 for 114 The six macro-chromosomes together therefore sum to 689 of thetotal length of the karyotpye

Among members of Massonieae the same diploidchromosome number of 2n = 20 as in Spetaea lachenali-iflora have been reported for Drimiopsis barteri(Kootin-Sanwu 1969) and Eucomis vandermerwei (DeWet 1957) However these counts are dubious since allother counts of these species and of other species within

these genera provided different numbers Johnson andBrandham (1997) published 2n = 20 for someLachenalia species (L alba L comptonii L undulata Lsp and L sp nov) Chromosome numbers based on n =5 or n = 10 are frequently found among species ofMassonieae For example Schizocarphus (own counts ofS nervosus WW 1526 and WW 1528 both with 2n = 40)and Resnova (own counts of R humifusa WW 1524 with2n = 10) as well as some species of Ledebouria andLachenalia (see Venter 1993 Johnson amp Brandham1997) have chromosome numbers based on n = 5 or n =10 It is most likely that the chromosome numbers of theDaubenya clade (2n = 32 34 see Table 2) also have orig-inated from ancestors with 2n = 30 That no diploid chro-mosome number of 2n = 20 has been reported fromthose species of Massonieae previously regarded asbelonging to Scilla sl further substantiates the inde-pendent generic status of Spetaea

TAXONOMIC CONCLUSIONThe new taxon is distinct by a combination of mor-

phological karyological and molecular characters foundnowhere else among Massonieae Morphologically it ischaracterized by succulent canaliculate leaves blue to


86

Table 2 Chromosome numbers of Spetaea its closest relatives (Daubenya clade Veltheimia Whiteheadia) and SouthAfrican taxa previously included within the genus Scilla sensu Reid (1993 ie inlcuding Merwilla PseudoprosperoSchizocarphus Spetaea) Chromosome counts determined in this study are in bold Vouchers are deposited in LI (WW= W Wetschnig)

Species Voucher Chrom number ReferenceAmphisiphon stylosa WF Barker 2n = 32 Brandham (1989) Johnson amp

Brandham (1997)Androsiphon capense Schltr 2n = 34 Brandham (1990)Daubenya aurea Lindl WW 1152 2n = 32 this paperMassonia zeyheri Kunth WW 1109 2n = 34 this paperMerwilla kraussii (Baker) Speta WW 2178 2n = 40 this paperMerwilla natalensis (Planchon) Speta 2n = 32 De Wet (1957)

(= Scilla natalensis Planchon)Merwilla natalensis (Planchon) Speta 2n = 40 Gimenez Martin (1959)Merwilla natalensis (Planchon) Speta n = 20 21 Ratter amp Milne (1973)Pseudoprospero firmifolium (Baker) Speta 2n = 18 De Wet (1957) Jessop (1970)

(= Scilla firmifolia Baker)Pseudoprospero firmifolium (Baker) Speta WW 1322 2n = 18 this paperResnova humifusa (Baker) U amp D Muumlller-Doblies WW 1524 2n = 10 this paperSchizocarphus gerrardii Merwe 2n = 56 De Wet (1957)Schizocarphus nervosus (forma gerrardii) 2n = 42 Jessop (1970)Schizocarphus nervosus Merwe 2n = 28 De Wet (1957)Schizocarphus nervosus Merwe WW 1526 2n = 40 this paperSchizocarphus nervosus Merwe WW 1528 2n = 40 this paperSpetaea lachenaliiflora Wetschnig amp Pfosser WW 1527 2n = 20 this paperVeltheimia bracteata Harv ex Baker 2n = 40 Taylor (1925) Coleman (1940)

(= V viridifolia Jacq) Delay (1947)Veltheimia capensis (L) DC 2n = 40 Sato (1942)

[= V glauca (Ait) Jacq]Whiteheadia bifolia (Jacq) Baker WW 1122 2n = 40 this paper

violet tepals that are fused at the base forming a cam-panulate perigon and 2 to 3 ovules per locule Seed mor-phology (in parallel to molecular data) does not onlyyield differential characters for this taxon but also helpsto detect relationships with other genera We thereforedescribe here the new genus and species Spetaealachenaliiflora

Spetaea Wetschnig amp Pfosser gen nov ndash Typusgeneris Spetaea lachenaliiflora Wetschnig amp Pfosser

Descriptio Bulbus globosus Pedicelli adsunt pro-phylla desunt Tepala uninerva campanulata basibusconnatis tubum crateriformem formantibus obtusacaerulea ad violacea Ovarium sine stylo ovula 2 vel 3 inquoque loculo stylus 7ndash9 mm longus Semina ellip-soidea compressa leniter rugosa in statu sicco nigrasemina hilo conspicuo albido 1 spec S lachenaliiflora

Description Bulb globose Pedicels present pro-phylls lacking Tepals 1-nerved campanulate fused atthe base and forming a narrow cup obtuse blue to vio-let Ovary not stipitate ovules 2 to 3 per locule style 7ndash9mm long Seeds compressed ellipsoidal slightly wrin-kled black when dry hilum conspicuous whitish Onespecies S lachenaliiflora

Spetaea lachenaliiflora Wetschnig amp Pfosser spnov ndash Holotype [South Africa] Du Tois kloof [=Dutoitskloof] felsige Orte [= rocky places] III HJanuar [J F] Dregravege 1997 (B isotype W)

Descriptio Planta 20ndash45 cm alta Bulbus hypogaeusdiametro ca 35 cm folia bulborum solida pallium atro-fuscum non fibrosum Radices tenues Folia frondis ca6 succulenta canaliculata linearia ad lanceolata acumi-nata 15ndash25 cm longa 1ndash3 cm lata non maculata nitidaglabra Scapus 1 erectus (inflorescentia inclusa) 20ndash50cm longus teres diametro ca 4 mm glaber Racemusdiametro ca 4ndash5 cm multiflorus densiflorus non rami-ficans rachis 10ndash20 cm longa bracteae adsunt 2ndash3 mmlongae 15 mm latae recurvatae margine membranaceoPedicelli patentes 09ndash19 cm longi glabri Tepala 9ndash11mm longa 18ndash25 mm lata Filamenta basibus connatispatula caerulea ad violacea 9-12 mm longa Antheraeca 25 mm longae in statu intacto caeruleae ad vio-laceae dorsifixae pollen luteum Pistillum 9ndash12 mmlongum Ovarium 3 mm longum diametro ca 2 mmCapsula obtusa 3-lobata obovata ca 10 mm longadiametro ca 8 mm Semina ca 26 mm longa ca 17 mmlata ca 19 mm alta leniter rugosa in statu sicco nigrahilum ca 12 mm longum et 03 mm latum albidumtypus germinationis ignotus Numerus chromosomatumdiploideus 2n = 20 Africa australis

Description Plant 20ndash45 cm high Bulb hypogeousc 35 cm in diameter bulb scales firm pallium dark-brown not fibrous Roots thin Foliage leaves about 6succulent canaliculate linear to lanceolate acuminate

15ndash25 cm long 1ndash3 cm broad unspotted shiningglabrous Scape 1 erect 20ndash50 cm long including theinflorescence terete about 4 mm in diameter glabrousRaceme 4ndash5 cm in diameter many-flowered denseunbranched rachis 10ndash20 cm long bracts present 2ndash3mm long 15 mm broad recurved apex membranousPedicels horizontally spreading 09ndash19 cm longglabrous prophylls lacking Tepals 9ndash11 mm long18ndash25 mm broad Filaments fused at the base spread-ing blue to violet 9ndash12 mm long Anthers c 25 mmlong violet to purple when still closed dorsifixed pollenyellow Pistil 9ndash12 mm long ovary 3 mm long 2 mm indiameter Capsule obtusely 3-lobed obovate about 10mm long about 8 mm in diameter Seeds c 26 mm longc 17 mm broad c 19 mm high slightly wrinkled blackwhen dry hilum about 12 mm long and 03 mm widewhitish type of germination unknown Diploid chromo-some number 2n = 20 South Africa Western CapeProvince Endemic in a small area in the vicinity ofDutoitskloof and Bainrsquos Kloof

Eponymy mdash The generic epithet is chosen tohonor Franz Speta born 22 Dec 1941 in LinzUrfahrAustria who drew the authorsrsquo attention to the peculiari-ties of this plant and morphological differences with theiconotype of Scilla plumbea Lindl The specific epithetwas chosen to indicate overall flower morphology whichresembles that of members of the genus Lachenalia

Key to relevant genera of Massonieae inSouth Africa mdash [Largely based on Speta (1998b) gen-era regarded until recently as belonging to Scilla are indi-cated in bold]1 Tepals free or slightly connate 21 Tepals connate

Daubenya sl Veltheimia Whiteheadiaand all other gen-era of Massonieae

2 Perigone plusmn stellate (without a pseudotube) 32 Perigone plusmn campanulate (with a short or long pseudo-

tube) 63 Raceme without an apical tuft of conspicuous sterile

green bracts 43 Raceme with a conspicuous apical tuft of sterile large

green bracts Eucomis4 Locules 2-ovulate tepals pink or whitish

Pseudoprospero4 Locules with several ovules 55 Tepals blue ovary white or bluish seeds flattened

Merwilla5 Tepals light blue to whitish or greenish white ovary

bluish green seeds not flattened Schizocarphus6 Ovary stipitate tips of tepals distinctly reflexed

Ledebouria6 Ovary not stipitate tepals not or only slightly reflexed 7 7 Inner tepals nearly closed at anthesis tepals succulent


87

and persistent in fruit Drimiopsis7 Inner tepals not nearly closed at anthesis tepals not suc-

culent and persistent in fruit 88 Tepals many-nerved (3ndash7) pink-purple Avonsera8 Tepals 1-nerved 9 9 Tepals greenish whitish pink-purple or bluish with a

conspicuous vitta style to 25 mm long Resnova9 Tepals vivid blue to violet without a conspicuous vitta

style 7ndash9 mm long Spetaea

ACKNOWLEDGEMENTSA grant from the Republic of South Africa to WW in 1987 to

collect most of the material for this study is highly appreciated Wethank Rod and Rachel Saunders for their efforts to provide seedsand other plant material We are grateful to E StabentheinerInstitute of Plant Physiology Karl-Franzens-University Graz foruse of the SEM laboratory G Prenner made most of the micro-graphs Sincere thanks go also to C Scheuer for his help with theLatin description N Kilian from the Botanic Garden andBotanical Museum Berlin-Dahlem provided scans of originalplates

LITERATURE CITEDBaker J G 1897 Liliaceae Pp 253ndash528 in Thiselton-Dyer

W T (ed) Flora Capensis vol 6 Reeve LondonBrandham P E 1989 123 Amphisiphon stylosa

Hyacinthaceae Kew Mag 6 58ndash61Brandham P E 1990 155 Androsiphon capense

Hyacinthaceae Kew Mag 7 124ndash128Coleman L C 1940 The cytology of Veltheimia viridifolia

Jacq Amer J Bot 27 887ndash895Delay C 1947 Recherches sur la structure des noyaux quies-

cents chez les Phanerogames Rev Cytol CytophysiolVeacuteg 9 169ndash222 10 103ndash229

De Wet J M J 1957 Chromosome numbers in the ScilleaeCytologia 22 145ndash159

Felsenstein J 1985 Confidence limits on phylogenies anapproach using the bootstrap Evolution 39 783ndash791

Fitch W M 1971 Toward defining the course of evolutionminimum change for a specific tree topology Syst Zool20 406ndash416

Gimenez Martin G 1959 Nuacutemero cromosoacutemico en especiesde Scilla Geneacutet Ibeacuter 11 97

Goldblatt P amp Manning J 2000 Cape plants A conspectusof the Cape flora of South Africa Strelitzia 9 1ndash743

Huelsenbeck J P amp Ronquist F R 2001 MrBAYESBayesian inference of phylogenetic trees Bioinformatics17 754ndash755

Jessop J P 1970 Studies in the bulbous Liliaceae 1 ScillaSchizocarphus and Ledebouria J S African Bot 36233ndash266

Jessop J P 1975 Studies in the bulbous Liliaceae in SouthAfrica 5 Seed surface characters and generic groupingsJ S African Bot 41 67ndash85

Johnson M A T amp Brandham P E 1997 New chromo-some numbers in petaloid monocotyledons and in othermiscellaneous angiosperms Kew Bull 52 121ndash138

Kootin-Sanwu M 1969 IOPB chromosome number reportXXII Taxon 18 436

Kunth C S 1843 Enumeratio Plantarum Omnium HuiusqueCognitorum IV J G Cottae Stuttgardiae amp Tubingae

Levan A Fredga K amp Sandberg A A 1964Nomenclature for centromeric position on chromosomesHereditas 52 201ndash220

Lewis G J 1947 Scilla plumbea Fl Pl Africa 26 t 1006 +2 pp

Lindley J 1830 Scilla plumbea Lead-coloured ScillaEdwardacutes Bot Reg 16 t 1355 + 2 pp

Maddison W P amp Maddison D R 1992 MacCladeAnalysis of Phylogeny and Character evolution Version30 Sinauer Associates Sunderland Massachusetts

Manning J C amp Van Der Merwe A M 2002 Hyacinthac-eae A new combination in Daubenya Bothalia 32 63ndash65

Merwe F Z van der 1943 Schizocarphus nervosus Fl PlS Africa 23 t 904 + 2 pp

Merwe F Z van der 1944 Scilla firmifolia Fl Pl S Africa24 t 926 + 2 pp

Pfosser M amp Speta F 1999 Phylogenetics of Hyacinthaceaebased on plastid DNA sequences Ann Missouri BotGard 86 853ndash875

Pfosser M amp Speta F In press From Scilla to Charybdis ndashis our voyage safer now Pl Syst Evol

Pfosser M Wetschnig W Ungar S amp Prenner G In pressPhylogenetic relationships among genera of Massonieae(Hyacinthaceae) inferred from plastid DNA and seed mor-phology J Pl Res 117

Phillips E F 1930 Scilla natalensis Fl Pl S Afr 10 t 365+ 2 pp

Ratter J A amp Milne C 1973 Some angiosperm chromo-some numbers Notes Roy Bot Gard Edinburgh 32429ndash438

Reid C 1993 Hyacinthaceae (Part B) Pp 146ndash155 inArnold T H amp De Wet B C (eds) Plants of SouthernAfrica Names and Distribution [Mem Bot Surv SAfrica 62] National Botanical Insitute Pretoria

Sacircto D 1942 Karyotype alteration and phylogeny in Liliaceaeand allied families Jap J Bot 12 57ndash161

Speta F 1998a Systematische Analyse der Gattung Scilla L(Hyacinthaceae)Phyton (Horn Austria) 38 1ndash141

Speta F 1998b Hyacinthaceae Pp 261ndash285 in Kubitzki K(ed) The Families and Genera of Vascular Plants vol 3Springer-Verlag Berlin

Stedje B 1998 Phylogenetic relationships and generic delim-itation of sub-Saharan Scilla and allied African genera asinferred from morphological and DNA sequence data PlSyst Evol 211 1ndash11

Swofford D L 2000 PAUP Phylogenetic Analysis UsingParsimony (and other methods) Version 4 SinauerAssociates Sunderland Massachusetts

Taylor W R 1925 The chromosome morphology ofVeltheimia Allium and Cyrtanthus Amer J Bot 12104ndash115

Venter S 1993 A revision of the genus Ledebouria Roth(Hyacinthaceae) in South Africa PhD Thesis Univ ofNatal Pietermaritzburg

Wetschnig W 1992 CHROM ein neues Computerprogramm


88

zur Darstellung chromosomenmorphologischer DatenPhyton (Horn) 31 251ndash256

Wetschnig W Pfosser M amp Prenner G 2002a ZurSamenmorphologie der Massonieae Baker 1871(Hyacinthaceae) im Lichte phylogenetisch interpretiertermolekularer Befunde Stapfia 80 349ndash379

Wetschnig W Pfosser M amp Prenner G 2002bSamenmorphologische und molekulare Untersuchungenzur Sippengliederung der Massonieae (Hyacinthaceae) P153 in Bundesanstalt fuumlr alpenlaumlndische LandwirtschaftGumpenstein (ed) Bericht uumlber das 10 Oumlsterr Botaniker-treffen BFA Irdning [Abstr]

Williams R 2000 Hyacinthaceae Pp 610ndash619 in LeistnerO A (ed) Seed plants of Southern Africa families andgenera Strelitzia 10 610ndash619


89


90

Appendix 1 List of taxa investigated in this study with vouchers citation information and EMBL accession numbersAll vouchers are deposited in LI unless otherwise indicated Two accession numbers for the trnL-F locus indicate thatthe sequences of the trnL intron and the trnL-F spacer have been deposited separately

Species Voucher Origin EMBL accession no

Amphisiphon stylosa WF Barker 1 Wetschnig 1160 S Africa AJ507957Amphisiphon stylosa WF Barker 2 Wetschnig 1144 S Africa AJ507959Amphisiphon stylosa WF Barker 3 Wetschnig 1145 S Africa AJ507958Androsiphon capense Schltr Wetschnig 1129 S Africa AJ507955Autonoe haemorrhoidalis (Webb amp Berth) Speta 1 Klenner H160 Spain AJ232518AJ2326411

Autonoe haemorrhoidales (Webb amp Berth) Speta 2 Vitek 1993 Spain AJ508001Autonoe latifolia (Willd) Speta Ehrendorfer H015 Morocco AJ232517AJ2326401

Barnardia scilloides Lindl 1 Pfosser H599 Korea AJ507998Barnardia scilloides Lindl 2 Pfosser H638 Japan AJ507999Bellevalia aff brevipedicellata Turrill Jahn H214 Greece AJ232547AJ2326701

Bellevalia trifoliata Kunth Speta H052 Greece AJ232548AJ2326711

Bowiea volubilis Harv ex Hook f Pfosser H222 S Africa AJ232454AJ2325771

Bowiea sp Pfosser H600 Madagascar AJ507922Brimeura amethystina (L) Chouard Pfosser H225 cult ex B G Tallinn AJ232510AJ2326331

Charybdis aphylla (Forskaringl) Speta Strauss H878 Jordan AJ4261132

Charybdis hesperia (Webb amp Berth) Speta Speta H764 Spain AJ4260882

Charybdis undulata (Desf) Speta Raus M117 Israel AJ507924Chouardia litardierei (Breistr) Speta Pfosser H230 Croatia AJ232541AJ2326641

Daubenya aurea Lindl Wetschnig 1162 S Africa AJ507956Dipcadi cf heterocuspe Baker Pfosser H603 Madagascar AJ507926Drimiopsis barteri Baker Bjoslashrnstad 1158 Tanzania Z99137Z991383

Drimiopsis botryoides Baker ssp botryoides Nordal 1600 Tanzania Z99139Z991403

Drimiopsis kirkii Baker Pfosser L121 cult B G Vienna AJ507952Drimiopsis maculata Lindley Speta H002 cult LI AJ232502AJ2326251

Drimiopsis sp Pfosser H642 Madagascar AJ507953Eucomis bicolor Bak Schnabel M162 S Africa AJ507933Eucomis montana Compton Schnabel M130 S Africa AJ507932Eucomis punctata (Thunb) LacuteHerit Speta H221 cult LI AJ232500AJ2326231

Eucomis vandermerwei Verdoorn Wetschnig 2171 S Africa AJ507935Eucomis zambesiaca Baker Wetschnig M163 ex Manchester Univ AJ507934Fessia greilhuberi (Speta) Speta Pfosser H013 cult B G Vienna AJ232534AJ2326571

Fessia puschkinioides (E Regel) Speta Fritsch H211 Tadschikistan AJ232536AJ2326591

Fessia vvedenskyi (Pazij) Speta Speta H234 Uzbekistan AJ232535AJ2326581

Hyacinthella dalmatica Chouard Gutermann H297 Croatia AJ232526AJ2326491

Hyacinthella heldreichii (Boiss) Chouard Markus H210 Turkey AJ232527AJ2326501

Hyacinthoides aristidis (Coss) Rothm Schneider H065 Algeria AJ232521AJ2326441

Hyacinthoides hispanica (Mill) Rothm Voglmayr H338 Spain AJ232525AJ2326481

Hyacinthoides italica (L) Rothm Pfosser H300 France AJ232519AJ2326421

Hyacinthoides lingulata (Poir) Rothm Koenen H066 Tunisia AJ232520AJ2326431

Hyacinthoides non-scripta (L) Chouard ex Rothm Pfosser H235 France AJ232524AJ2326471

Hyacinthoides reverchonii (Degen amp Hervier) Speta Voglmayr H307 Spain AJ232523AJ2326461

Hyacinthoides vincentina (Hoffmanns amp Link) Rothm Scheiblreiter H305 Portugal AJ232522AJ2326451

Hyacinthus litwinowii Czerniakowska Akhani H573 Iran AJ508689Hyacinthus orientalis L Speta H067 Romania AJ232539AJ2326621

Hyacinthus orientalis L var alba Plass M178 Syria AJ508002Lachenalia aloides (L f) Engl Pfosser H159 cult B G Vienna AJ232508AJ2326311

Lachenalia cf pusilla Jacq 1 Wetschnig 1146 S Africa AJ507988Lachenalia cf pusilla Jacq 2 Wetschnig 1164 S Africa AJ507987Lachenalia contaminataAiton Pfosser M141 S Africa AJ507985Lachenalia pallida Aiton 1 Wetschnig 1168 S Africa AJ507981Lachenalia pallida Aiton 2 Pfosser H021 cult B G Vienna AJ232507AJ2326301

Lachenalia pusilla Jacq Wetschnig 1115 S Africa AJ507986Lachenalia rubida Jacq 1 Wetschnig 1131 S Africa AJ507982Lachenalia rubida Jacq 2 Pfosser M140 S Africa AJ507983Lachenalia sp Wetschnig 1110 S Africa AJ507984Ledebouria cf concolor (Bak) Jessop Wetschnig 1412 S Africa AJ507946Ledebouria cordifolia (Baker) Stedje amp Thulin Nordal amp Stedje 2409 Malawi Z99143Z991442

Ledebouria floribunda (Baker) Jessop Wetschnig 1433 S Africa AJ507937Ledebouria hyacinthina Roth 1 Jha H838 India AJ507944Ledebouria hyacinthina Roth 2 Jha H839 India AJ507945


91

Appendix 1 (continued)Ledebouria revoluta (L f) Jessop Nordal 2082 Zimbabwe Z99146Z991473

Ledebouria socialis (Baker) Jessop Pfosser H014 cult B G Vienna AJ232501AJ2326241

Ledebouria socialis agg WW 2170 S Africa AJ507951Ledebouria somaliensis (Baker) Stedje amp Thulin Nordal 2296 Ethiopia Z99150Z991513

Ledebouria sp 1 Pfosser H641 Madagascar AJ507938Ledebouria sp 2 Pfosser H795 Guinea AJ507941Ledebouria sp 3 Wetschnig 1421 S Africa AJ507939Ledebouria sp 4 Wetschnig 1419 S Africa AJ507940Ledebouria sp 5 Mucina LM2010015 S Africa AJ507947Ledebouria sp 6 Mucina LM2010019B S Africa AJ507948Ledebouria sp 7 Mucina LM2010016 S Africa AJ507949Ledebouria sp 8 Mucina LM2010019A S Africa AJ507950Massonia depressa Houtt 1 Wetschnig 1142 S Africa AJ507980Massonia depressa Houtt 2 Wetschnig 1143 S Africa AJ507977Massonia depressa Houtt 3 Wetschnig 1140 S Africa AJ507972Massonia depressa Houtt 4 Wetschnig 1134 S Africa AJ507978Massonia depressa Houtt 5 Wetschnig 1139 S Africa AJ507975Massonia echinata L f 1 Wetschnig 1161 S Africa AJ507967Massonia echinata L f 2 Wetschnig 1138 S Africa AJ507974Massonia echinata L f 3 Wetschnig 1135 S Africa AJ507971Massonia marginata Willd ex Kunth Wetschnig 1101 S Africa AJ507960Massonia pustulata Jacq 1 Wetschnig 1148 S Africa AJ507970Massonia pustulata Jacq 2 Wetschnig 1132 S Africa AJ507969Massonia cf sessiliflora (Dinter) U amp D Muumlller-Doblies Wetschnig 1137 S Africa AJ507973Massonia sp 1 Wetschnig 1136 S Africa AJ507968Massonia sp 2 Wetschnig 1133 S Africa AJ507979Massonia tenella Soland ex Baker Wetschnig 1145 S Africa AJ507976Massonia zeyheri Kunth 1 Wetschnig 1153 S Africa AJ507961Massonia zeyheri Kunth 2 Wetschnig 1109 S Africa AJ507962Merwilla kraussii (Baker) Speta Wetschnig 2178 S Africa AJ507929Merwilla natalensis (Planchon) Speta 1 Wetschnig 1534 S Africa AJ507931Merwilla natalensis (Planchon) Speta 2 Wetschnig M278 S Africa AJ507930Merwilla sp 1 Puff H219 cult B G Vienna AJ232499AJ2326221

Merwilla sp 2 Puff H218 cult B G Vienna AJ232498AJ2326211

Muscari botryoides (L) Mill Kleesadl H011 Austria AJ232545AJ2326681

Muscari comosum (L) Mill Neuner H056 Italy AJ232546AJ2326691

Muscari macrocarpum Sweet Fritsch H212 Turkey AJ232544AJ2326671

Muscari parviflorum Desf Plass M138 Syria AJ508003Nectaroscilla hyacinthoides (L) Parl Scheiblreiter H016 Portugal AJ232542AJ2326651

Oncostema dimartinoi Raf Vezda H178 Italy AJ232514AJ2326371

Oncostema peruviana (L) Speta Pfosser H198 Portugal AJ232516AJ2326391

Oncostema villosa (Desf) Raf Gruber H217 Tunisia AJ232515AJ2326381

Ornithogalum kochii Parl Speta H717 Slovenia AJ507927Othocallis sp Pasche H179 Turkey AJ232533AJ2326561

Oziroe acaulis (Baker) Speta Weigend sn Peru AJ507921Oziroe biflora (Ruiz amp Pavon) Speta Chase 793 (K) Chile AJ232453AJ2325761

Periboea oliveri U amp D Muumlller-Doblies Wetschnig 0005 S Africa AJ507989Periboea paucifolia (W F Barker) U amp D Muumlller-Doblies Wetschnig 1154 S Africa AJ507990Pfosseria bithynica (Boiss) Speta Vasak H232 Bulgaria AJ232540AJ2326631

Polyxena calcicola U amp D Muumlller-Doblies Muumlller-Doblies H216 S Africa AJ232506AJ2326291

Polyxena ensifolia (Thunb) Schoumlnl 1 Wetschnig 1113 S Africa AJ507994Polyxena ensifolia (Thunb) Schoumlnl 2 Wetschnig 1157 S Africa AJ507992Polyxena ensifolia (Thunb) Schoumlnl 3 Wetschnig 1150 S Africa AJ507996Polyxena ensifolia (Thunb) Schoumlnl 4 Wetschnig 1104 S Africa AJ507995Polyxena ensifolia (Thunb) Schoumlnl 5 Wetschnig 1156 S Africa AJ507993Polyxena ensifolia (Thunb) Schoumlnl 6 Wetschnig 1147 S Africa AJ507991Polyxena sp Saunders M061 S Africa AJ507997Prospero elisae Speta 1 Speta H068 Greece AJ232530AJ2326531

Prospero elisae Speta 2 Ehrendorfer H155 Greece AJ232531AJ2326541

Prospero haritonidae Speta Speta H027 Greece AJ232528AJ2326511

Prospero obtusifolium (Poiret) Speta Speta H053 Morocco AJ232529AJ2326521

Pseudogaltonia clavata (Masters) Phillips Speta H220 cult B G Vienna AJ232475AJ2325981

Pseudoprospero firmifolium (Baker) Speta Wetschnig 1322-01 S Africa AJ507928Puschkinia scilloides Adams var libanotica Pfosser H229 Lebanon AJ508688


92

Appendix 1 (continued)

Resnova humifusa (Baker) U amp D Muumlller-Doblies Wetschnig 1524 S Africa AJ507942Resnova maxima Merwe Hankey M275 S Africa AJ507943Rhadamanthus mascarenensis Baker Pfosser H610 Madagascar AJ507923Schizocarphus nervosus (Burch) Merwe 1 Saunders M132 S Africa AJ507936Schizocarphus nervosus (Burch) Merwe 2 Stedje 9415 Zimbabwe Z99157Z991583

Schnarfia messeniaca (Boiss) Speta Pfosser H640 cult B G Vienna AJ508004Scilla albescens Speta Speta H237 Greece AJ232553AJ2326761

Scilla cf bulgarica Speta Speta H158 Romania AJ232555AJ2326781

Scilla cydonia Speta 1 Jahn amp al H215 Greece AJ232549AJ2326721

Scilla cydonia Speta 2 Speta H489 Greece AJ232550AJ2326731

Scilla nana (JA amp JH Schultes) Speta Speta H238 Greece AJ232552AJ2326751

Scilla siehei (Stapf) Speta cv ldquoPink Giantrdquo Speta H010 cult LI AJ232551AJ2326741

Scilla spetana Kereszty Speta H227 Austria AJ232556AJ2326791

Scilla subnivalis (Halacsy) Speta Speta H240 Greece AJ232554AJ2326771

Spetaea lachenaliiflora Wetschnig amp Pfosser Saunders M164 S Africa AJ507954Stellarioides longebracteata (Jacq) Speta Pfosser H407 cult B G Vienna AJ232471AJ2325941

Stellarioides sp Pfosser H608 Madagascar AJ507925Tractema lilio-hyacinthus (L) Speta Hoog amp Hoog H298 Spain AJ232511AJ2326341

Tractema monophyllos (Link) Speta Raus H049 Spain AJ232513AJ2326361

Veltheimia bracteata Harv ex Baker 1 Wetschnig 1535 S Africa AJ507965Veltheimia bracteata Harv ex Baker 2 Wetschnig 1525 S Africa AJ507964Veltheimia bracteata Harv ex Baker 3 Speta H060 cult LI AJ232503AJ2326261

Veltheimia bracteata Harv ex Baker 4 Saunders 2009 S Africa AJ507963Whiteheadia bifolia (Jacq) Baker Wetschnig 1130 S Africa AJ507966Whiteheadia etesionamibensis Muumlller-Doblies Lavranos amp Pehlemann H444 Namibia AJ232504AJ2326271

Zagrosia persica (Hausskn) Speta 1 Leep H440 Turkey AJ232537AJ2326601

Zagrosia persica (Hausskn) Speta 2 Stevens H500 Turkey AJ232538AJ2326611

Sequences first reported in 1Pfosser amp Speta (1999) 2Pfosser amp Speta (in press) and 3Stedje (1998)


Fig 1 Morphology of Merwilla and Spetaea A Merwilla plumbea (Lindley) Speta t 1355 in Lindley (1830) as Scillaplumbea Lindley B Merwilla natalensis (Planchon) Speta t 365 in Phillips (1930) as Scilla natalensis Planchon CSpetaea lachenaliiflora t 1006 in Lewis (1947) as Scilla plumbea Lindley D Spetaea lachenaliiflora WW 1527 photo-graphed at the natural locality at Bainrsquos Kloof in Dec 1987

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Oziroe acaulis

5 changes

100100

5460

Massonieae (Fig 4)

1001006087 8995

100100

100100100100

100100

100100

81100

80100

94100

6485

9498

-80

--

100100

-67

99100

96100

85100

100100

--92100

82100

98100

100100

5786

100100

961005599

83100

96100

86100

99100

-63

98100

86100

-91

86100

5899

MASSONIEAE

OZIROE-OIDEAE

ORNITHOGAL-OIDEAE

URGINE-OIDEAE


100100

HIII

HII

HI

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Oziroe acaulis

5 changes

Hyacintheae (Fig 3)

100100

100100

5460

100100

60878995

100100

100100

100100

100100

8110080100

5599

86100

6499

84100100100

89100

100100

---96

709793100

84100

6493

100100

--

-86

9798

100100

87100

100100

5887

--

5252100100

921006090

100100

94100

6095

6498

66100

98100

99100

MASSONIEAE

OZIROE-OIDEAE

ORNITHOGAL-OIDEAE

URGINE-OIDEAE

MASSONIEAE

HYACINTH-OIDEAE

MIa

MIb

MII

MIIIa

MIIIb

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

s 08 08 19 11 13 11 12 11 11 10 09 09 08 08 08 06 08 06 06 06 microm10 10 23 14 16 14 14 13 13 12 12 11 10 10 10 08 10 08 08 08

l 147 147 53 50 38 31 12 12 11 11 10 09 10 09 09 11 08 08 07 07 microm12 14 microm

180 180 65 61 46 38 15 15 13 13 12 12 12 12 12 13 10 09 09 09 14 17

r 188 188 34 57 29 28 10 11 10 11 11 10 12 11 12 17 10 12 12 12

cr 190 190 103 92 62 52 29 28 27 25 24 23 22 22 21 21 19 17 17 17

B







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Oziroe acaulis

5 changes

100100

5460

Massonieae (Fig 4)

1001006087 8995

100100

100100100100

100100

100100

81100

80100

94100

6485

9498

-80

--

100100

-67

99100

96100

85100

100100

--92100

82100

98100

100100

5786

100100

961005599

83100

96100

86100

99100

-63

98100

86100

-91

86100

5899

MASSONIEAE

OZIROE-OIDEAE

ORNITHOGAL-OIDEAE

URGINE-OIDEAE


100100

HIII

HII

HI

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Oziroe acaulis

5 changes

Hyacintheae (Fig 3)

100100

100100

5460

100100

60878995

100100

100100

100100

100100

8110080100

5599

86100

6499

84100100100

89100

100100

---96

709793100

84100

6493

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--

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9798

100100

87100

100100

5887

--

5252100100

921006090

100100

94100

6095

6498

66100

98100

99100

MASSONIEAE

OZIROE-OIDEAE

ORNITHOGAL-OIDEAE

URGINE-OIDEAE

MASSONIEAE

HYACINTH-OIDEAE

MIa

MIb

MII

MIIIa

MIIIb

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

s 08 08 19 11 13 11 12 11 11 10 09 09 08 08 08 06 08 06 06 06 microm10 10 23 14 16 14 14 13 13 12 12 11 10 10 10 08 10 08 08 08

l 147 147 53 50 38 31 12 12 11 11 10 09 10 09 09 11 08 08 07 07 microm12 14 microm

180 180 65 61 46 38 15 15 13 13 12 12 12 12 12 13 10 09 09 09 14 17

r 188 188 34 57 29 28 10 11 10 11 11 10 12 11 12 17 10 12 12 12

cr 190 190 103 92 62 52 29 28 27 25 24 23 22 22 21 21 19 17 17 17

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Oziroe acaulis

5 changes

100100

5460

Massonieae (Fig 4)

1001006087 8995

100100

100100100100

100100

100100

81100

80100

94100

6485

9498

-80

--

100100

-67

99100

96100

85100

100100

--92100

82100

98100

100100

5786

100100

961005599

83100

96100

86100

99100

-63

98100

86100

-91

86100

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MASSONIEAE

OZIROE-OIDEAE

ORNITHOGAL-OIDEAE

URGINE-OIDEAE


100100

HIII

HII

HI

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Oziroe acaulis

5 changes

Hyacintheae (Fig 3)

100100

100100

5460

100100

60878995

100100

100100

100100

100100

8110080100

5599

86100

6499

84100100100

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709793100

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9798

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5887

--

5252100100

921006090

100100

94100

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98100

99100

MASSONIEAE

OZIROE-OIDEAE

ORNITHOGAL-OIDEAE

URGINE-OIDEAE

MASSONIEAE

HYACINTH-OIDEAE

MIa

MIb

MII

MIIIa

MIIIb

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

s 08 08 19 11 13 11 12 11 11 10 09 09 08 08 08 06 08 06 06 06 microm10 10 23 14 16 14 14 13 13 12 12 11 10 10 10 08 10 08 08 08

l 147 147 53 50 38 31 12 12 11 11 10 09 10 09 09 11 08 08 07 07 microm12 14 microm

180 180 65 61 46 38 15 15 13 13 12 12 12 12 12 13 10 09 09 09 14 17

r 188 188 34 57 29 28 10 11 10 11 11 10 12 11 12 17 10 12 12 12

cr 190 190 103 92 62 52 29 28 27 25 24 23 22 22 21 21 19 17 17 17

B







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Oziroe acaulis

5 changes

100100

5460

Massonieae (Fig 4)

1001006087 8995

100100

100100100100

100100

100100

81100

80100

94100

6485

9498

-80

--

100100

-67

99100

96100

85100

100100

--92100

82100

98100

100100

5786

100100

961005599

83100

96100

86100

99100

-63

98100

86100

-91

86100

5899

MASSONIEAE

OZIROE-OIDEAE

ORNITHOGAL-OIDEAE

URGINE-OIDEAE


100100

HIII

HII

HI

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Oziroe acaulis

5 changes

Hyacintheae (Fig 3)

100100

100100

5460

100100

60878995

100100

100100

100100

100100

8110080100

5599

86100

6499

84100100100

89100

100100

---96

709793100

84100

6493

100100

--

-86

9798

100100

87100

100100

5887

--

5252100100

921006090

100100

94100

6095

6498

66100

98100

99100

MASSONIEAE

OZIROE-OIDEAE

ORNITHOGAL-OIDEAE

URGINE-OIDEAE

MASSONIEAE

HYACINTH-OIDEAE

MIa

MIb

MII

MIIIa

MIIIb

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

s 08 08 19 11 13 11 12 11 11 10 09 09 08 08 08 06 08 06 06 06 microm10 10 23 14 16 14 14 13 13 12 12 11 10 10 10 08 10 08 08 08

l 147 147 53 50 38 31 12 12 11 11 10 09 10 09 09 11 08 08 07 07 microm12 14 microm

180 180 65 61 46 38 15 15 13 13 12 12 12 12 12 13 10 09 09 09 14 17

r 188 188 34 57 29 28 10 11 10 11 11 10 12 11 12 17 10 12 12 12

cr 190 190 103 92 62 52 29 28 27 25 24 23 22 22 21 21 19 17 17 17

B







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Oziroe acaulis

5 changes

Hyacintheae (Fig 3)

100100

100100

5460

100100

60878995

100100

100100

100100

100100

8110080100

5599

86100

6499

84100100100

89100

100100

---96

709793100

84100

6493

100100

--

-86

9798

100100

87100

100100

5887

--

5252100100

921006090

100100

94100

6095

6498

66100

98100

99100

MASSONIEAE

OZIROE-OIDEAE

ORNITHOGAL-OIDEAE

URGINE-OIDEAE

MASSONIEAE

HYACINTH-OIDEAE

MIa

MIb

MII

MIIIa

MIIIb

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

s 08 08 19 11 13 11 12 11 11 10 09 09 08 08 08 06 08 06 06 06 microm10 10 23 14 16 14 14 13 13 12 12 11 10 10 10 08 10 08 08 08

l 147 147 53 50 38 31 12 12 11 11 10 09 10 09 09 11 08 08 07 07 microm12 14 microm

180 180 65 61 46 38 15 15 13 13 12 12 12 12 12 13 10 09 09 09 14 17

r 188 188 34 57 29 28 10 11 10 11 11 10 12 11 12 17 10 12 12 12

cr 190 190 103 92 62 52 29 28 27 25 24 23 22 22 21 21 19 17 17 17

B







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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

s 08 08 19 11 13 11 12 11 11 10 09 09 08 08 08 06 08 06 06 06 microm10 10 23 14 16 14 14 13 13 12 12 11 10 10 10 08 10 08 08 08

l 147 147 53 50 38 31 12 12 11 11 10 09 10 09 09 11 08 08 07 07 microm12 14 microm

180 180 65 61 46 38 15 15 13 13 12 12 12 12 12 13 10 09 09 09 14 17

r 188 188 34 57 29 28 10 11 10 11 11 10 12 11 12 17 10 12 12 12

cr 190 190 103 92 62 52 29 28 27 25 24 23 22 22 21 21 19 17 17 17

B







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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

s 08 08 19 11 13 11 12 11 11 10 09 09 08 08 08 06 08 06 06 06 microm10 10 23 14 16 14 14 13 13 12 12 11 10 10 10 08 10 08 08 08

l 147 147 53 50 38 31 12 12 11 11 10 09 10 09 09 11 08 08 07 07 microm12 14 microm

180 180 65 61 46 38 15 15 13 13 12 12 12 12 12 13 10 09 09 09 14 17

r 188 188 34 57 29 28 10 11 10 11 11 10 12 11 12 17 10 12 12 12

cr 190 190 103 92 62 52 29 28 27 25 24 23 22 22 21 21 19 17 17 17

B







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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

s 08 08 19 11 13 11 12 11 11 10 09 09 08 08 08 06 08 06 06 06 microm10 10 23 14 16 14 14 13 13 12 12 11 10 10 10 08 10 08 08 08

l 147 147 53 50 38 31 12 12 11 11 10 09 10 09 09 11 08 08 07 07 microm12 14 microm

180 180 65 61 46 38 15 15 13 13 12 12 12 12 12 13 10 09 09 09 14 17

r 188 188 34 57 29 28 10 11 10 11 11 10 12 11 12 17 10 12 12 12

cr 190 190 103 92 62 52 29 28 27 25 24 23 22 22 21 21 19 17 17 17

B







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84







85


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

s 08 08 19 11 13 11 12 11 11 10 09 09 08 08 08 06 08 06 06 06 microm10 10 23 14 16 14 14 13 13 12 12 11 10 10 10 08 10 08 08 08

l 147 147 53 50 38 31 12 12 11 11 10 09 10 09 09 11 08 08 07 07 microm12 14 microm

180 180 65 61 46 38 15 15 13 13 12 12 12 12 12 13 10 09 09 09 14 17

r 188 188 34 57 29 28 10 11 10 11 11 10 12 11 12 17 10 12 12 12

cr 190 190 103 92 62 52 29 28 27 25 24 23 22 22 21 21 19 17 17 17

B







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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

s 08 08 19 11 13 11 12 11 11 10 09 09 08 08 08 06 08 06 06 06 microm10 10 23 14 16 14 14 13 13 12 12 11 10 10 10 08 10 08 08 08

l 147 147 53 50 38 31 12 12 11 11 10 09 10 09 09 11 08 08 07 07 microm12 14 microm

180 180 65 61 46 38 15 15 13 13 12 12 12 12 12 13 10 09 09 09 14 17

r 188 188 34 57 29 28 10 11 10 11 11 10 12 11 12 17 10 12 12 12

cr 190 190 103 92 62 52 29 28 27 25 24 23 22 22 21 21 19 17 17 17

B







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the scilla plumbea puzzle-present status of the genus scilla sensu lato in southern africa and...

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