Botanical Journal of the Linnean Society Volume 60 Issue 383 1968 [Doi 10.1111%2Fj.1095-8339.1968.Tb00087.x] a. EL-GAZZAR; L. WATSON; W. T. WILLIAMS; G. N. LANCE -- The Taxonomy of

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J . Linn.Soc. (Bot . )60, 383, p . 237W i t h 5 jlyuresPrinted in arcat Bri tainPebnuc.ly, 1968

237

The taxonomy of Sulwia: a test of two radically differentnumerical methodsBY A. EL-GAZZAR, L. WATSON, F.L.S.,Botany Department, So ut hm pt m Universityw. T. WILLIAMS, F.L.S. AND a. N. LANCE

C.S.I.R.O. Computing Research Section, Canberra, Australia(Acceptedfor publication September, 1967)

Forty-one variations in calyx morphology and venation, stamen form, trichomes, style&ape and aspects of habit and gross vegetative morphology were recorded for a cosmopolitansample of 100 Salvia species. Comparative data were subjected to computation under twofundamentally different programs, MLJLTCLAS a mixed-data method) and CENTCLAS(taxonomically crude, strictly qualitative). he resulting classifications were essentiallysimilar, but not identical; and inspection suggests that the differences result largely fromtaxonomic distortion inherent in the CENTCLAB method.A classification of these Salviaa is suggested, taken from the MULTCLAS hierarchy, which isfundamentally different from anything yet proposed by taxonomists. Of the five subgenera,only Cabsphace has gained support; and of the twelve sections only Hymenosphace is homo-geneous. The new scheme seems taxonomically reasonable, and is informative in that thegroupings are consistent with observations other than those on which it was based. The mainsubdivision represents a distinction between Central and South American Salvias and therest, and reflects variation in stylar morphology, though neither geography nor stylar formwere subjected to computation. It is concluded that even CENTCLAB, used alone, could havegreatly enhanced the taxonomic understanding of this well-known genus.

INTRODUTOTIONSalvia is t)he argest genus of Labiutae and one of he best known, taking in some of themost spectacular species. Bentham 1832-6) was unhappy about the quality of his

specimens and confessed dissatisfaction with his classification; but, as in so many othercaws is account represents the most recent attempt a t a world monograph, notwith-standing a threefold increase in the number of described species since his day. The treat-ment of Briquet 1897) is in essence a nomenclatural variant of Bentham's, and latertaxonomic accounts of Salvia are restricted to regional floras. The most important ofthese is the work of Epling 1938, 1939), who has given descriptions for 600 species,distributed among 91 sections. However, he has dealt only with American species, themajority of which were grouped by Bentham under the subgenus Cahp?um. A morpho-logical study of Salvia stamens by Zalewska 1928) has never been incorporated into thetaxonomy of the genus.

ClearlySalvia is due for a taxonomic reappraisal, and interested as we are in the applica-tion of numerical techniques to Angiosperm taxonomy, this genus appealed to us as asuitable subject for investigation.OBSERVATIONS

The accepted classification of Salvia reflects Bentham's obsession with stamens, andthere is precious little comparative information on record. However, these plants have16



238 A. EL-GAZZAR,.WATSON,. T. W ~ ~ ~ z a a a sLND . N. LANCEreadily provided a splendid catalogue of variations. Mostof the observations are novel;but they are all of the kind that can be made rapidly without sophisticated techniques.

The calyxM ~ P h O k l YThe Salvia calyx conforms with the usual Labiate pattern in seeming to represent fivefused sepals. However, the calyces of our sample can be arranged under five headings asfollows:

(1) Calyx &dentate, with all the teeth moreor less equal (Fig.1 ).2) Calyx bilobed, dorsal lobe entire, and two ventral teeth which may be very small(Fig. 1b). That of S. leucophyUa Greene seems best regarded as a modification of thistype, in which the ventral teeth are absent.(3) Calyx bilobed, with the dorsal lobe bicuspidate, and two ventral teeth. Thedorsal lobe may be notched in the middle (Fig.1c), or not (Fig. Id).(4) Similar to type 3), but having the dorsal lobe tricuspidate (Fig. le). In somespecies (e.g.S. sylvatria L,),he two lateral spines of the dorsal lobe overlap the middleone (Fig. If) .( 5 ) Calyx generally much larger in size than in types 1) o 4), the teeth more or lessequal, obtuse and expanded (Fig. 1g).Annuluethe annulus, and have simply recorded its presenceor absence.Some species manifest a dense ring of simple hairs within the calyx throat:we call this

VenationThe number of main veins in the calyx is remarkably constant within the species, butamong them varies from8 to 17or more. The vein number is not a mere reflectionof calyxshape: for example, our calyx type 2)may have 8,9,11 ,13,14 or 15 veins; and the samevein numbers occur in different calyx types.

Cell-walls of inner epidermisIn 70 yoof the species examined, the cell-walk of the inner epidermisof the calyx tubeare heavily thickened with many oanal pits traversing them. There am some species wherethe calyx does not show this condition at all, and some where the thickened walls areconfined to the basal region.

Calcium oxalate crystals are found in the calyces of many species ofSalvia.Their shapesand sizes are so variable tha t we have contented ourselves with recording only theirpresence or absence. They are consistently absent among certain African species.The atamem

We have been unable to recognize all the stamen types delimited by Zalewska (1928),who contemplated them from a unotional and adaptive standpoint. It seems to ua thatfour basic types can be recognized using as oriteria the nature of the two ends of the articu-lated connective, as well as other morphological features:



239

Fig. 1. Calyx morphology of S a l k : a) 8. uvanduhefolia; b ) S. apecioaa; c) S. iaidica;d)S. oemeriama; e)S. oaaefolia; f)8. ylvealria; 8)S. cetabu2oaa. Venation is omittedonlyin g). For descriptionsaee text. a) to f), x 6; g), x 2.6.



240 A. EL-GAZZAR,. WATSON,W. T. WILWS AND G. N. LANCE(1) The two parts of the connective unequal, the longer being fertile while theshorter is sterile and reduced to a small process receiving no vascular supply from thestaminal filament Fig. 2 a).2) Similar to the preceding, but the sterile part of the connective is much longer andfurnished with a vascular strand from the staminal filament. The sterile parts of the

two stamens in the flower are often in contact via the minute glandular hairs on theadjacent margins Fig. 2b).(3) Both arms of the connective Worm, more or less equal, and fertile, but the twomain pollen-producing cells of the stamen are morphologically distinct Fig. 2 c) Weplace the stamens of 8. urduacea and S . columbariaeunder this heading although theylack the articulation between the staminal filaments and the connectives.4) Only the outer end of the connective is fertile while the inner is typically sterile,short, broad and flattened. The flat parts of the two stamens in the flower are usuallycontiguous via their abortive half-anthers Fig. 2d).

C

Fig. 2. Type8 of stamens in Salvia: a) . vel.ticillala; b) . ombrophila; c ) S. hegelmai t r i :d)S. dominica. All x 6.



Numerical uxorunny of SalviaForma of hairs

241

Leaves, bracts and calycesof Salvia species are commonly invested with hairs, of which(1) Simple, multicellular and eglandular, with the apical cell shortest and pointed2) Similar to type ( l ) ,but with the apical cell in the form of a globular secretory

(3) Similar to type (1),but with the apical cell in the form of a very long and narrow4) Long, undulate and intermingled to form a dense compact mat covering the leaf

( 5 ) Uniseriate branched (Fig. 3e), confined to certain species in Central and South

five types are easily distinguishable :

(Fig. 3a) .gland (Fig.3b)appendage, its lumen occluded (Fig. 3c).surface. The cell junctions are noticeably swollen (Fig. 3d).America.

Fig. 3. Diagrams of the hair types found on leaves, bracts and calyces ofSulvia species.Seetext for details. x 13.All these types occur on leaves, bracts and calyces, although the combinations vary

from species to species. Furthermore, in any species the indumentum of leaves, bractsand calyces may be different.



242 A. L-GAZZAR,.WATSON,. T. Wmmius ANDG.N. LAXWThe tyle

The fldvvia;style is always unequally bXd and borne underneath the upper lip of thecorolla.In somespecies he upper arm of the style ismuch longer than the lower (Fig.4a)elsewhere the opposite is the case (Fig. 4b). The style is more or less hairy in some species,glabrous in others.

Fig. 4. a) Style of S. anguelqoZolia with the upper arm longer than the lower, (b) that ofS. iinnata with the lower arm longer than the upper. x 13.

Vegetative w p b b g yWe have made use of a few gross morphological characters, easily recorded fiom herb-arium material but seemingly ignored by taxonomists. Leaves may be aimpleor dissected;

narrowly linear and sessileor broad and petiolate; smooth or rugose;glabrousor denselyhairy. Bracts may be reflexed or surrounding a glomerule of flowers.Table 1. m m a y of compra.tive observathns recordedfor 100 Salviaspec;@. Characters 1 to 15 were subje.cted to m ~ hh mctere16and 17 were left out

Qualitative charactera:h8VeS: 1. Simple/compound2. eeseile/petiolate3. smooth/rugoeecalyx : 4. calp-oq&&, present/absent6.6.

cell-wallsofinnerepidermis, thickened/not thickenedannulusofhaim within calyx-throat, present/abbsentBracts: 7. reflexed/not reflexedStamens: 8. staminel filamentsor connectives, hairy/glaborusStyle: 9. hairy/glaborue

Calyx : 10. number of main veins, 9 categoriesStamens: 12. type of tamen, 4 categories

Multistate charactera:11. type ofcalyx, 6 categorim

Hairs : 13. tyPeS Ofh 8 h n CdJX, 4 C a h g O r i e S14.15. types ofhairs on eaves, 4 categorieetypes ofhaim on bracts, 4 cat0goriesAdditional characters :Stamens: 16. innerend of connective, pointing upwarda/pointbgd o w n dStyle: 17. upper arm onger/lower arm onger



Numerical taxonomy of Salvia 243We have examined a cosmopolitan sample of 100 species including representatives fromthe questionable genera Audibertia Benth. and Salviastrum Scheele; also SchraderiuMedic., which is Benthams section Hymemsphace promoted to generic rank. The samplethus takes in about 10 yoof Sdv ia semu 2ato. Table 1 summarizes the characters recorded,which were classified as qualitatives and multistates for a mixed-data analysis (see

Lance Williams, 1967). Characters 13 to 15 are multistates of the non-exclusivetype,where there is more than one possibility for a single species (i.e. one plant may exhibitmore than one of the hair types on any of the organs studied). In certain species havingcaducous bracts we were unable to record characters7 nd 15,which were thus inapplicablein five cases. Characters 16 and 17 were recorded, but were not incorporated in thenumerical analyses: they were kept in reserve, along with geographical and cytologicalinformation,for use in evaluating the results.

THE PROGRAMSThe data were analysed as follows:1 ) Under the mixed-data program MULTCLAS, which uses the non-metric coefficientand has provision for flexible sorting, in which reversals are impossible and the in-tensity of clustering can be varied by changing the parameter /3. Experience suggests thata suitable value of Pfor most purposes is .25 and this was used here. MULTCLAS catersfor qualitatives, inapplicables and multistate characters including non-exclusives, andalso for quantitative characters, which were absent from our data. For details of theprogram, see Lance Williams 1967);MULTCLAS is the program therein referred to as aversion of MULTIST using flexible strategy.(2) Under the program CENTCLAS. This is the program referred to in Lance Williams1967) as CENTBET; it caters for qualitative or continuous data , but not for multistateattributes or mixed data, and uses a conventional centroid sorting strategy. It has atpresent no provision for missingor inapplicable values. We have used it in its qualitativeform with an information statistic, which involved dichotomizing all the information,including that for the multistate attributes. The theory of the statistic used is given inLance Williams 1966).

COMPARING THE RESULTS O F TEE TWO ANALYSESThe total information content, as given on the computer print-out, has been usedas ameasure of level when plotting the CENTCLAS hierarchy (Fig. 5). For the MULTCLAS

Table 2. Com parison between the alternative groups of Salvia species under MULTCL~SandCENTCLAS schemes in t e r n of the species they contain. Ast er ie b indicate w s of migration

BfULTCLAS O E N T O L A SGroup 18. erticillata, S. bwnmuelleri, S. sprucei*, S. S. verticillata, S . bwnmuelleri, S. clevelandi,macroetaohya*, S. clevelandi, S. apiana, S. S . apiana, S. aonomenaia, S. mellijera, S.aonomenaia, S. leucophylla, S. nzellijera, S. argenteo*, S . crassifolia, S. acetabuloea S.spathacea, S . craaaijololia. S. acetabubaa, S. dracocephaloides,S. cryptantha, S. ajricanu, S.cryptantha, S. ajricana, S. aurea, S . nivea, S.cana&nsia, S . cmpresaa.Group 2S. verbenaca, S. patensis, S. horminum, S. S. verbenaca, S. prdenaia, S. horminum, S.hnata, S . montbretii, S. viridia, S. v i r g a t a , S. lanata S. montbvetii, S . viridis, S. v i rgah S.glutinoaa, S. argentea, S. sylvestria. S. aclarea glutirwaa, S. sclarea, S. hkT080 ? 3TW,S.S. pandiflwa*, S. hieroaolymitana,S. bicolor,S. bicolor,S. aethiopia,S.bahnaae*, S. aminiana*.aethiopia.

aurea, S. nivea, S. canarieneis, S. cmpreaea.



244 A. EL-GAZZAR,. WATSON,W. T. wIlm4 M~NDG. N. IANCIETable 2--continued.

Group 3S. lavandulaefolia,S. aromdica, S. aucheli , S.h q e l m a i e l i ,S. candelabrum.s lxdim s ibanotim,s riloba,s kinalia,Group 4S. m d m c e a , S. columbariae, S. texana, S.plebeia,S.d l t i o r h i z a ,S. aponica,S. orakohlei,S. h l l e y a n a , S. lyrata,S. pinnata. S. campanu-lata S. roemeriana, S. roaaefolia*, S. hetero-tricha+,S. irablitziana,S. dracocephaaloidea*.Group 6S. runcinata, S. monticola, S. cooperi, S.dominim, S. balanaae, S. controveraa, S.jaminiana S. atenophylla, S. deaerti. S.aegyptiaca.Group 6S. aplendene S. lavandubidea, S. coocinea, S.g r a h i , S. lasiocephala, S. srcbinciaa. S.hyptoidtw,S. corrugates S.pocurrena, S. lachno-ata;ehya, S. remota, S. m e x i m n a , S. azurea, S.npecioaa, S. benthamiana. S. farinucm, S.fu lgena, S. lanceolata, S. roamarindda, 8.tortuoaa, S. p r a n e n a w , S. ornbrophila, S.conferti) ra, S. meliaaaeflora,S. involucrda.

Group 7S. anguatifolia, S. paeudo-coccinea, 9 revoluta,S. acoparia, S. leucocephala, S. flooculoeo, S.w r i i a n a .

S.piaidiea,S. libanotiea,S. tribba.S. o&inalia,S. grandiflora, S. aromatica, S. lyrdaf, S.lavandulmfolia. S. aucheri, S. hegelmaieri, S.candelabrum.S. carduacea, S. c o l u d a r i a e , S. tezana, S.plebeia, S. miltiorhiza, S.jap onic a, S.forakohlei,S. bulleyana, S. pinnakz. S. eanyxmulokr S.roemeriano, S. habliteiana, S. roeaefolia+, S.aylveatria+,S. atenophylla+.S. runcinata, S. mont icob, S. cooperi, S.dominica,S.controveraa,S. deaerti,S. aegyptiaca.

S. applendena. S. lavandubidea S. coccinea, S.grahwni , S. lasiocephala, S. aubinciaa, S.hyptoidea,S. c m g a t a , S.procurrena, S. lachno-atachya, S. r e w t a , S. mexicana, S. azurea, S.a p o w a a , S. benthamiana, S. f a r i m S.fulgena, S. lanceolalcr, S. roanw7inoidea, S.tortuoaa, S. a p w e i , S. mucroatachya, S.paranen&, S. ombophi la , S. confertiflora, S.m l k a w f l o r a , S. involucrata, S. M e r o t r i c h , S.anguatifolia, S. paewlo-coccinea. S. acoparia, S.apathaoea+.S. leucocephala. S. revoluta, S. moriana S.flocculoaa, S. leucophylla*.

Table 3. Diugm8tic features for group8 C and D in Fa 5 , picked out byeye from the data-matrices as rearranged under MULTCUS and CENTOLASrespectivelyC E N T C L A B

Group C 37 species)1. Leaves dissected in 10 species.2. 6 species with type 1)stamens.3. 25 species with type (4) stamens.4. 6 species with type (3)stamens.5. 3 species with annulus ofhairs within calyx-throat.6. 9 species with type 6) calyx.7. No species with type (1) calyx.8 . 34 species with 13-veined calyx.

Group D (26 species)1. Leaves dissected in 12 species.2. No species with type (1) stamens.3. One species with type 4) stamens.4. 26 species with type 3) stamens.6. 4 species with annulus of

hairs within calyx-throat.6. No species with type 6) calyx.7. 8 species with type 1)calyx.8. 6 pecies with 13-veinedcalyx.M ULTCLAB

Group C (42 species)1. Leaves diseected in 4 species.2. 8 species with type 1) stamens.3. 20 species with type 4) stamens.4. 12 species with type (3) stamens.6.No species with 'annulus' ofhairs within calyx-throat.6. 8 species with type (6) calyx.7. 8 species with type 1)calyx.8. 29 species with 13-veinedcalyx.

Group D (26 species)1. Leaves dissected in 18 species.2. No species with type 1) stamens.3. 6 species with type 4) stamens.4. 19 species with type (3) stamens.6. 7 species with annulus of6. One species with type (6) calyx.7. No species with type 1) calyx.8 . 13 species with 13-veined calyx.

hairs within cdyx-throat.



Numerical t u x o n m y of Salvia 2462 5

2 2

2 0

1.7

1.51

12

O(

0.7i

C

III:

MULTCLAS

l hbb l1 25 7 6

C

A

CENTCLAS 1600

1400

1200

1000

800

600

400

IFig. 6. The MULTCLAS and CENTCLAS hierarchiesfor 100Sulwia.species seeTable 2), baaed onthe characters listed in Table 1 . Numbers of species in the low-levelgroupings re indicated.

hierarchy, where no exactly comparable measure exists, the non-metric measure a t thepoint of fusion has been used to define the level. For purposes of discussion, we had tochoose stopping levels beneath which further subdivision of groups seems taxonomicallyunprofitable. This does not mean t hat we consider the very low-level aggregations meaning-less : merely tha t the similarities between adjacent groups are so high that we have beenunable to discover external criteria to support them.

The two hierarchies coincide closely regarding the two main groups ofSalvia species, Aand B in Fig. 5 . They also agree to a marked extent over the low-level groupings 1 to 7 : infact the small group 7 is the only one showing a correspondenceof ess than 70 yoamong thespecies (Table 2), and the only major difference between the two hierarchies concern therelative positions of the two groups 3 and 5 in C and D . In order to evaluate taxonomic-ally the divergent versions of C and D , i t is necessary to reconsider the data-matr ix asrearranged by CENTCLASand MULTCILASespectively. Table 3 summarizes the correlatingcharacters which pick out the groups: for the sake of simplicity we have left out thosecharacters where there are no obvious correlations in this context. Taking the CENTCILASarrangement first, we we t ha t there is no criterion here for an absolute distinction, but thetwo groupsexhibit markedly different tendencies regarding stamen types 2,3and 4), and



246 A. EL-GAZZAR,. WATSON,W. T. W m ND G. N. LANCEcalyx form and venation 6,7 and 8).The most impressive correlationhere is between type4) stamens and 13-veinedcalyces, involving 23 out of37 species in group C.The MULTCLASarrangement also picks out two series with different tendencies among calyces and stamens,but the distinctions regarding features3 ,4 , 6 and 8are less clear. On the other hand it canbe seen that MULTCLAS is also acknowledging distinctions (not absolute, but impressive)in leaf form 1) and calyx construction (6) ignored under CENTCLAS.Furthermore, theMULTCLAS treatment has placed in group C 16 out of the 17 Salvias in our sample withcalyx types (I)and (6) , nd in doing so it draws attention to a fact which we had over-looked while investigating calyx morphology. There are only 17 species in which thecalyx is not markedly 2-lipped, and of these eight constitute our calyx type I ) , the resttype (6). We take it as morphologically significant that 16 of them are aggregated in

We consider the MULTCLAS version superior, since it accounts for more aspects of theMULTCLhS groupcplants diversity, and has even taught us something about calyx morphology.

REALLOOATIONIt is in theory possible to re-examine the constitution of the groups at any chosen levelin the hierarchy, to ascertain whether any of the individuals would be more appropriatelyplaced in a different group, thus redressing errom due to t he migration problem. Themethod is necessarily iterative, but, at least in a strictly Euclidean system, the process isknown to converge rapidly (for an account, me Jancey, 1966 ; for a proof of convergence,see Needham, 1966). The problem of its extensionto non-Euclidean classificatory models(such as we have used) is under investigation, but has so far proved intractable. As acompromise solution, we have reallocated the 8dvia species on a Euclidean model withthe attribute-axes weighted accordingto their contributionsto the primary classification.The operation was carried out on the MULTULAS hierarchy,at the 7 groups level: i.e. acrossthe minor groupings 1 to 7 in Fig. 5 . It resulted in six reallocations, which we judgeto be

very reasonable. It is noticeable that five of the reallocated species were correctly placedunder UENTGZAS(cf. Tables2and 4).Table4, in ourview, represents a satisfactory arrange-ment ofSdviaspecies in the light of the available evidence, with thesix ransfers indicatedby asterisks.

Table 4. Chai cation of Salvia 8pecim. Derived f r m the MULTOLAS hierarchy (Fig. 5),but incorporating reallocations (aaterisked; 8ee text)Group AGroup 1S. uerticilhfa L., S. bornmwlkri Haueskn.,S. ckve ndi Greene, S. spalhacea:Greene, S . aphnaJepson, 8. wrophylh Greene, 8.met,?ift?m areene, 8. 8munnek Greene, 8. cetabubm L.,S. dracocephuEoides* Boias., S . cryptonthaMontbr. et Auch., S. afrioana L., S . aurea L., . n kThunb., S . ConariSnaia L.,S. compema Vent, S. m a a fo l h Ded.

S. eylveetriaL.,S. v i d d i a L., S. montbretii Benth.,S. hnutu Roxb.,S. sclareaL.,S. aethi opi aL.,S. verbena00L., . prate L., . lwrminm L., . argent- L..S. glutinoeaL., S. hieroeolynilomcBoim.,S virgataAit., S. bicolor Lam.S. pieidieo Bo k. et Heldr., S. libanoticaBobs. et Gaill., S. o f i i n a l i a L., S. grandiflora* Etling,S. tsiloboL., S . aromdiCaBoh. , S. auoheri Benth.,S. ?bqelmaieri P. R., S. candelabrumBoiss.,S. vandukwfoZbVahl, S . ro8aefolia* Sm.

8.wtvhacea Benth.,S. co2wmbariue Benth., S. tezonaGray, S miltiorhim Bunge, S. japonicoThunb.,5. orskohkiL., .plekia R. Br.,S. bzrlleyanaDiels,8 c a m p a n u bWall., S . hablitzkanaPall.,S. roemel.iana Scheele, S. pinnda L., 7. lgmtaL.

Group 2

Group 3

Group 4



Numerical t m m y f Salvia 247Group 5S.runcin t L., S. wwn&kola Benth., S. cooperi Skm.,S. balansae No6, S. dominim L., . ateno-phylla Burch., S. controversa Tenore, S. aminiana Nod. S. deserti Decne.,S. aegyptiaca L.

Group BGroup 6S. splendens Sello., S. lavanduloidee H. B. K., S. coccinea Juts.,S. grahami Benth., S. mbincisaBenth., S. lasiocephab Hook. et Am., S. h y p t o d a Mart. et Gal., S. comugda Vahl, S. lachno-siachys Benth., S. procurrene Benth., S. remota Benth., S. w x i c a n u L., S. azurea Lam., S.paranensis Dudn, S. speciosa Presl. S. ahnacea Benth., S. benthamiana Gardn. et Field., S.fulgens Cav., S. lanceolata Willd., S. tortuosa H. B . K.,S. r o m 7 i n o i d e e St. Hil., S.conferti&n-aPohl,S. lnelieaaeJlwaBenth.. S. ombrophilaDusBn, S. involucrata Cav.. S. heterotricha* Fernald.S. angustijolia Cav., S. pseudococcinea Jacq., S. acoparia Epling, S. macroatachya' H. B. K.,S. ~pruce i *Briq.. S. revolt& Ruiz et Pav., S. leucocephala, H. B. K., S. jeocculoea Benth.,S.m ri n Mart.

Group 7

Table 5. Comparison between the orthodox c2assiJications of Salvia (includingAudibertia,Salviastrum and Schraderia) and groups 1 to 7 of Table 4 . Name-s are t h e of Bentham(1832-6, 1848), Bentham S Hooker (1876) and Briquet (1897) for mbgenera and ( i nparen theses) sectiom. Numerals indicate numbers of species

Group 117 species): S a l k (E ua ph ace), 2 ; Salt& (H ywn oep hac e; Schraderia), 6 ; S c h e a ( A et hi op ie ),1;Leonia (Hemisphace), 1 Audibertia, 6.14 species): Salwia (Drymosphace), 1; Sclara (Hormunum), 2; Sclarea (Aethwpia), 6; Sclccrea(Pleithioephace),5.11 species) Salwia (Euapha ce),11.13 species) Salvia (Eusphace), 1;S a l k ( Dr ym o ep ha ce ), 4; Leonia (P ycnoqha ce) , 1 Leonia(Echinoephace), 1 eonia (Notwuphace).2; Leonia (Heteroephace),2; Salvia9trum, 1.10 species) Sclarea (Pleithioephuce), 3 ; Leonia (Heteroephace),2; Leonia (Notiouphuce),2.

( 2 6 specks): Calosphace (Caloephme), 26.9 species) Caloaphace (Caloephace),7.

Group 2

Group 3

Group 4

Group 6Group 8Group 7

Table 6. Diagnostic features of group8 A and B in Table 4 . Features 1 o 7 were picked outby eye from the rearranged data-m atrix; features 8 to 10 are additional, i . e . not subjected tocomputation

Group A (85species)2. Stamen types l ) , 3) and 4).3. Stamens glabrous in 60 pc iee .4. Style glabrous in 59 species.5. Leaves dissected in 22 species.6. 36 species with glandular hairs.7. No branched ha& on calyx, bracts, or8. Lower lobe of style longer than upper.9. Inner end of connective points upwards

1. All types of calyx.

leaves except in S. lewophylla.

in 61 species.American species.10. European, Asian, Afrioan and North

Group B 36 species)1. Calyces of type 2) only.2. Stamen type 2) only.3. Stamens hairy except in S. lanceolata.4. Style densely hairy in 28 species.5. Leaves always simple.6. 27 species without glandular hairs.7. 11 s p i e s with branched heirs on all orany of calyx, b r a t s or leaves.8. Lower lobe of style shorter han upper.9. Inner end of connective pointsdownwards.

species.10. Only Central and South American



248 A. EL-GAZZAR,. WATSON,W. T. WILLT~MSND G. N. LANCETable 7. Dkqm8tic featurm of groups C and D in Table 4. Features 1 to 6 were picked out byeye from the rearranged data -ma trix; eatures 7 and 8 are additional

Group A , C 42 species) Group A , D 23 species)1 . Leaves simple in 36 species.2. Stamen types 1) 8 species), 3 ) 14

species) and 4) 20 species).3. No species with annulus of hairs withincalyx-throat in this group.4. Calyx types 1 ) and 5) account for 17species i.e. calyx often moreor leasregular).

1. Leaves dissected in 16 species.2. Stamen types 3) 17 species) and 4)3. All species with the annulus 7 ) fall4. Calyx types 1 ) and 6) absent: calyx

6 species).

always 2-lipped.5. 41 species with crystals in calyx.6. Leaves sessilein only 3 species.7. No species with flowers less than 1 cm long.8. Species mainly European and

6. 8 species without crystals in calyx.6. Leaves sessile in 12 species.7. 7 species with flowers leas than 1 cm long.8. North and South African, East AsianMediterranean. and North American species.

T A X O N O W O DISCUSSIONOur treatment of Salvia species bears little resemblance to any orthodox classification.Comparison of Table 4 with Table 5 shows that of the five subgenera only Cahphuceemerges in tac t and of the 12 sections only Hymenosphace appears to be homogeneous.The species from Audibertia, Sa lv bt ru m and Schraderiu have all been submerged amongthe Salvias.We intend to carry out further computations in order to investigate the feasibility ofextracting diagnoses by these means. In the meantime, Tables6 and 7 serve to show tha tour scheme not only accounts for the wide range of comparative observations listed inTable1,but that i t is also in harmony with evidence ha t was not subjected to computation.Thus, the recognition of two main groupsof Salvia species( A and B) is supported by the

additional criterion that it represents an absolute distinction between species ofour samplewith the lower arm of the style longer than the upper (groupA), and those (groupB)wherethe opposite is the case. Geographical distributions of the species are also illuminating;for group B is Central and South American whereas group A takes in species from therest of the world. A variation in stamen form, observed by Bentham 1832-6) but notsatisfactorily accounted for in his taxonomic scheme, is also in harmony: in group B theinner end of the articulated connectiveis sterile and points downwards, while in groupAit is sometimes fertile and usually pointing upwards.Table 7 shows that groups C and D are a h geographically credible. Group C ismainly European andMediterranean, although a few species occur in Persia, the Himalayasand South Africa; group D , on the other hand, takes in the majority of North Americanand East Asian Salvias(adiscontinuity well knownto phytogeographers) as well as someexclusively North and South African species. Furthermore, even in our fragmentaryspecimens, the suggested redistribution of the speciesimposes ome pattern on variationin flower length; for 20 out of 26 species in groupA with flowers more than 2 cm long fall ingroup C, while all seven species with flowers less than1cm long are in group D .Any nomenclatural confusion arising out of problems of species delimitation could makenonsense of minor groupings in the lower reaches of a hierarchy. However, from ageneticists point of view, the low-level fusions (indicating, we hope, possibilities ofhybridization) are of interest. Thus although the restricted size ofour am pl e limits thedirect value in floristic work ofthe minor groupings1 to 7 in Fig. 5 , it is worth attempting toassess their taxonomic worth. As might be expected, it is not eaayto discover additionalfeatures to distinguish adjacent low-level groupings. In fact, we have notasyet found anycriteria to support the splitting of the Central and South American Salvias (i.e. groupB)into groups 6 and 7 other than the hair differences involving leaves, calyces and bracte



Numerical twonumy of Salvia 249which were picked out by the computer. However, the minor groups 1 to 5 certainly seemtaxonomically reasonable (see Table 8).In the first place they are geographically likely,with ten African species constituting group 5 , group 2 centred on the Mediterranean andgroups 1 and 4 exhibiting the well-known phytogeographical discontinuity betweenEast Asia and North America. The flowersofSalv iaspecies are generally 1.5to 3-0cm long,and in our entire sample only ten species had them less than 1.0 cm. It is, therefore,Table 8. Diagnosesfor the seven low-level groups of Salvia species shown n Table 4.Featuresin bold were not incorporated in the computation. Cytological data from Stewart 1939),Darlington Wylie 1955),and Cave 1958-64).Figures i n parentheses indicate numbers

of species in each group for which chromosome counts are on recordGroup Diagnostic characters

1 Calyx 13-veined,types 2)and 6).Bracts and calyces without hair types 2) and 4).Cell-wallsof inner epidermisof sepals not undulateand thickened arare situation) n 10/17 species.Leavessimple in 16/17 species. Stamen types 1) and 4) . filaments glabrous. Style glabrous in 16/17species, lower lobe longer than upper. Inner end of connective pointing upwards.Chromosome numbers 8)multiples of 8,11,12 and 13. Californian and Asian species.2 Calyx 13-veined, ype 4).Bracts and calyces with hair types 2)and 4).Leaves simple. Stamenstype a), filaments glabrous. Style glabrous in 12/14 species, lower lobe longer than upper.Inner end of connective pointing downwards. Chromosome numbers 8) multiples of8 and 11. Mediterranean and S . European species.3 Calyx 14- or 17-veined, mainly type 1). Stamens type 3), filaments glabrous. Style glabrous,lower lobe longer than upper. Inner end of connective pointingupwards. Chromosomenumbers 2)multiples of 7. Mainly European species.4 Calyx 11-veinedexcept in S. carduecea andS. c o l u m b a r k e , mainly type 2);annulusof hairswithin the throatin4 species. Leaves dissected in 7/13 species. Stamens type 3). filamentshairyor glabrous. Style often glabrous,lower lobe longer than upper. Inner end of connective

pointing upwards. Chromosome numbers 4) multiples of 8, 9, 12 and 13. Chinese,Japanese and N. American species.6 Calyx 13-veinedexcept inS. monticoh and S . cooperi, type 4).Bracts and calyces without hairtypes 2 ) and 4). Leaves dissected and sessile. Calyx-crystals n only 3 species. Stamen types3 ) and 4),filaments glabrous. Style glabrous, lower lobe longer than upper. Inner end ofconnective pointing upwards. Flowers small, sometimes minute less han 1.0 cm in5/10 species). Chromosome numbers 2) multiples of 8. N. and S. African species.6 Calyx 8-, 9-, 11-, 13-, 16- r many-veined,of type 2)except in 5. procurrens.Leaves and braatawithout hair type 6) . Leaves simple. Stamens type 2) , filaments often hairy. Style commonlyhairy, upper lobe longer than lower. Inner end of connective pointing downwards.Chromosome numbers 3) multiples of 11. Central and SouthAmerican species.7 Calyx 13- or 14-veined, type 2). Bracts, calyces and leaves with hair type 6). Leaves simple.

Stamens type 2). filaments hairy. Style hairy, upper lobe longer than lower. Inner end ofconnective pointing downwards. No chromosome counts on record. Central and SouthAmerican species.

interesting to find an aggregation of five species with minute flowers among the ten speciesof group 5 . We have seen that tho orientation of the inner end of the connective dis-tinguishes, though not absolutely, between groups A and B. We now find that all thespecies in Group A exceptional in this respect go to form group 2. Finally, althoughchromosome counts are only sporadically available, it is noticeable tha t both the cyto-logically known species in group 3 have multiples of seven-a number unknown elsewhere.

METHODOLOOICAL CON CLUSIONSWatson, Williams Lance 1966)have noted that taxonomic groups seem quite robustin the face of variations in the numerical approach; and the present study supports this



250 A. EL-GAZZAB,. WATSON, . T. W AND G.N. LANCEsuggestion.However, their data-matrix provided a less stringent test than our informationon Salvia, which, being more extensive, constituted a far more difficult problem. Further-more, the methods compared by Watson, Williams Lance were all of the same basictype, confined to qualitative attributes with no provisionformultistates and inapplicables.Here however, we have used a qualitative method alongside a mixed-data method wheremultistates and inapplicables are cateredfor.The differences between the two programs can be illustrated in terms of the charactersused here, by considering the characters 10 (calyx vein numbers) and 15 (bract hairs).Aspecies might exhibit any one of the nine venation types, but the presence of one logicallyexcludes the possibility of h d i n g any other. Under CENTCLAS, a species with 15-veinedcalyx comesto differ from one with a 13-veinedcalyx by two characters; a stateof affairsunreasonable in giving undue weight to one aspect of calyx morphology. MULTCUeliminates this sort of error, regarding the distinction as a single character difference.Asregards character 15,we were unable to record bract details (in fact five variants) in fivespecies. Under IWLTOWB these features simply become inapplicables,but for CENTCLAS wewere obliged to treat them as negatives. The species concerned, however, have appearedin the same groups under the two sohemes.In view of the fundamental differences between the two programs, the similaritybetween the alternative hierarohies (Fig. 5 ) is remarkable. Had the level of taxonomicunderstanding of Sdvia been high, the differences would have been important. But theavailable classificationsof thia relatively well-known genus are80 primitive that CENTCLILSalone, using our quite short list of oharacters and with its illogical treatment of bractdetails, could have made a valuable aontribution.

ACKNOWLEDQEMENTSWe wish to thank the Keepers of the herbaria at Manohester Museum, EdinburghUniversity and the Botany School, Cambridgefor the loan of specimens.A. E-G. thanks the

Egyptian Government for financial support.BEFERENCES

BENTHAP,4 1832-6. L - w Uenera et Speckea. London.BENTHAM,., 1848. In De Candolle,Prodrmua, XII, Paris.BENT-, U. HOOKER,. D., 1876. UeneraPIontorum,Vol. 2. London.BBIQUET,., 1897. In Engler andPrantl,Die.N urZkhen Pfinzenfamilien, Vol. 4 3a and 3b). Leipzig.CAVE,M. S., 1068-64. ed.). .h.dex to Plan4 ~h~omoaOmember8, 2 voh. The University of NorthDABLINGTON,. D. WYLIE,. P., 1966. Chromosome A l h fFlowering Phnta, 2nd edition. London:EPLINQ,., 1938. A revisionofSaZvia:eubgenua CaloephaceI. F& Repert. Spec. Nw. eih. 110 1):LING, C., 1939. A revisiono f s a l k : subgenus Cakwphace.11. F&8 Repert. Spec.Nw. eih. 112 2):

JANCEY, . C., 1966. Mulkiimemional group analysis. A M . J . Bot. 14: 127-30.LANCE, N. WIUUMS, W. T., 1966. Computer programs for hierarchical polythetio cleesificationLANCE,U. N. WILLIAMS,. T., 1967. A general theory o f classificatorysorting strategies. 1. Hier-NEEDHAM,. M., 1966. T h e termination of certain iterative processes. Memorandum RM-6188-PR,STEWABT, . S., 1939. Chromoeomenumbere of Californian Salviae. Am.J . Bot. 26 ( 8 ) :674-6.WATSON,., WILUAMS,W. T. LANOE,0 N., 1966. Angiosperm taxonomy: a comparative study ofsome numerical techniques.J . Linn.Soo. (Bot.)S9: 491-601.ZALEWSKA, ., 1928. Recherchea su r 16volution des Btamines, considBde du point de vue de leuradaptationA la pollinisation desf l e w de laSauge (Salwia).BuZI. int. Acad.pol. Sci . Lett.3 7) 133-60.

caroliia Press.Ueorge Allen Unwin Ltd.1-160.161-383.

similarityanalyses).Cornput.J . 9: 60-4.archical systems. Cornput. J . 9: 373-80.Rand Corporation,California.

Documents

Botanical Journal of the Linnean Society Volume 60 Issue 383 1968 [Doi 10.1111%2Fj.1095-8339.1968.Tb00087.x] a. EL-GAZZAR; L. WATSON; W. T. WILLIAMS; G. N. LANCE -- The Taxonomy of