Grana

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Pollen Morphology of Salvia and Some Related

Genera

D. M. Henderson , H. Prentice & I. C. Hedge

To cite this article: D. M. Henderson , H. Prentice & I. C. Hedge (1968) Pollen Morphology of
Salvia and Some Related Genera, Grana, 8:1, 70-85, DOI: 10.1080/00173136809427461

To link to this article: https://doi.org/10.1080/00173136809427461

Published online: 01 Sep 2009.

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 POLLEN MORPHOLOGY OF SrlLVIA AND
SOME RELATED GENERA
D. 31. HESDEIiSOS, I I . 1’IiI:NTICI.: antl I. C. IIEDGI:
d1.S received 1 S . I \’.t7

Snloiu is a vast genus with about 900 spccics \vidcly distrihiitctl

t h ~ ~ i i ~ htlic~ l iOld
t and Ncw \\‘orltls lvitli centres of high spccics
concentrations in Central Aiiicrica antl South \\‘est Asia. Tlicrc is
grcat variation in hahit, lcnf shape, indumcntuiii, calys, corolla
and fruit striicturc throughout the gcniis antl with tlicse c1inr:ictcrs
it can reatlily ~ >split e into several suI)-(livisions. In stamina1 structure
it is possihlc to tracc a scqucncc from the most primitive types (siicli
as S . picroc(i1y.r and S. scfinicilfiaiuei~ii),ivlicre tlie thccnc of c a d i
anthcr I)otli contain fertilc pollcn and tlic conncctivc is short, to
thosc species (such as S. fillgetis) wlicrc one thecn is fcrtilc. thc othcr
is reduced to awl-shaped sterile tissuc antl the connective is wry
clongatctl. ’I’liis staminal variation is iiniqiic in the Labintac and
1) rol)a 1)Iy t h roil gho ti t t li c CII t ire pla n t li i ngt1o m t h crc is nothing quit c
compara1)le. For thc most part, tlic generic limits of Saloia arc clcar
and uncqiiivocal allhougli thcre arc a fcw problems \vitli a group
of Xortli Anicrican species \vliicIi inay constitutc a scparatc but w r y
closely allied genus.
Allliough it is possible to tracc cvolutionary tlevclopmcnts in Sriluirc
staminal structure arid to delimit the gcnus fairly distinctly, it is
w r y difficult to sag \vliat :ire its gcncric allies antl its progcnitors.
There arc a few, small, isolatctl gciicra such a s I’crouskia, dleriandrci,
Dorgsioeclicis and Horniiriiiiii, sharing tlic two staminate character
lvith Snluia, which Iiavc bccn considered as possible rclativcs hiit on
general rnorphology thcy are quite distinct ant1 apparcntly rclic
gc n cra .
Tlic ohjcct of this siirvcy was to dctcrniinc \vlicllicr there \vas any
correlation hctwecn pollcn morphology, general morphology and
 P O LL E s .\I 0111’1 I 0 LO GT 0 I: s h LVI‘\ 71

geographic tlistrihtions and \vlictlier or not it \vas a useful :idtlitional

tnsonoinic tool in assessing relationsliips and delimiting tasa both
in Snluitr ntitl its putativc allics prior to a niorc extensive taxonomic
revision of the Old ll‘orld species 1)y one of 11s(I. C. 11.). A selection
of the species for examination \vas made in order to givc a fairly
\vide reprcscnlntion of tliffcrcnt species throughoiit the range of
Snlrtirr. Other genera csaminctl were Srilr&istriim, Horniinrrni, Bory-
sfoeclins, Z’crouskirr, Mcrinnrlrrci antl Rosninrinus. The study was
critircly carried out \villi dried matcrial in thc Etlinburgh hcrt)ariurn
(E). Although appropriate material for pollen morphological studics
is present on most Iicrharium material, a survey of it as an aid to
tasonomy is seldom undert:ilien. \\‘ortli\vhilc data is tliiis often ovcr-
loolted and one of the great ricetls of systematic palynology today-
a n account of range of structure \vitliin genera-is not fulfillcd.
‘I’licre is as great n need for lino\vledgc of uniformity ns of diversity
of pollen structurc \vitliin tam. A prcvioiis paper (IIentlcrson 1965)
clcmonstratetl great diversity within the genus dfeconopsis of an othcr-
wise rather stcnopalynous family, the I’npavcraccac.

Materials and Methods

l’he pollen material \vas obtainctl from specimens in the Ediii-
h r g h hcrbarii~mand is listed in the appendix. Anthers of ~ l i o l e
flo\vcrs \Yere acctolysed according to Brdtmm’s (10GO) routine and
the pollen mounted in glycerinc jclly. Light p1iotomicrogr:iptis \Yere
tnlicn \\it11 n Lcitz Ortholux 2 microscope. For electron microscopy,
pollen- was cnihcdtlctl in Epon, cut on a C:inibridgc Ultrotome,
stained in 1 % aqucoiis potassium pcrrnanganatc, and examined and
pliotograplictl with an E.1I.G electron microscope. \l’e are intlebtctl
to I’rofcssor I\. Isrown antl 1lr. .I. Tulett Departtn. of Ilotany,
Iidint)urgh University, for use of the electron microscope and as-
sistance in ninny other ways.

General Morphology
The grains of Snlvici and ik allies are sphcroitlnl or prolate and
(i-colpatc cscept for :I fe\v cultivars where, for csamplc, in Saloia
splentlens, it has been sho\vn that incrcasc in ploidy leads to pro-
portionate iiicrcasc i n iiuiiibcr of colpi (Naurizio 1956). I n polar
view the sis mcsocolpia are either sut)cqual (Fig. 1:A) or two arclarger
52 I). ) I . 11k:SDEIlSOS, 11. P l l E S T l C E A S I ) 1. C. l l l ~ I ~ G l

than tlic rciiiaining four (Fig. 1: 13, C, 1)). I n n few spccics tlicsc two
larger iiicsocolpin also diner in csinc striicturc from the foiir smaller
mcsocolpia 1'1. 2: C. 'i'lic csincs of a11 the tasn csairiirictl arc to sonic
tlegrcc rcticiilatc. Apart from tlic cqiinlity a n d incqiiality of the
nicsocolpin tlic only usal)le dil'fercnlial characters \vc hare fount1 lie
in thc fine tlclails of tlic rcticulum. Thc rcticulum is citlicr uniform
in diameter over; thc grain surface (esclutling the colpi; 1'1. 3: A, 1j)
or is up to half the diarnctcr in the polar regions compared with tlie
cqiiator (1'1. 2: C). The \valls (miiri) of tlic rcticiiluni arc usually
proritlcd with n single scrics of 1)aciila (Fig. 1: I), 1)ut a few species
hare t\vo scrics of 1)ncala (tlu~~lil~:iculatc, Fig. 1: J). I n sonic instanccs
the tliamctcr of tlic meshes is qiiilc siriall (< 1.25 p). In tlicsc cases
the ltriiiina are not closcd. I3ut in many the ltriiiina are rclativcly
\vide ( > 1.5 11). Each lunicn is then closed by a perfornted incinbranc
at tlie lcvcl of tlic top of the hacula. 'I'lic niiinbcr of pcrforntions per
liinicn varies l)et\vccn foiir a n d thirty. Thcy niay 1)c crenly distri-
hutctl ovcr tlic cntirc incmbranc (Fig. 1: F) or may 1)c marginal only
:ind thus irninedintcly adjoining the supporting bnciiln (Fig. 1: E).
This nicml)ranc is not supportctl hy 1)aciila csccpt nroiintl the pcri-
phcry of tlic luiiien. Many tasn linrc a rclatircly thin membrane,
h i t in sonic the solid striictiirc of tlie mciiil~raricis iicnvicr ant1 tlic
whole tlicn appears as n secondary rcticuluin \vithin tlic primary
one--usually at a slightly lower lercl of focus.
I n light microscopy it is also apparcnt that in sonic spccics bacula
also occur on tlic floor of the lunicn, reaching 111) towards thc rcticiilar
iIicni1)raiic hiit not reaching it (Fig. 1: L, Al). 'I'licsc tlctnils of reticular
structure arc just rcsolrable by light microscopy hut tlic critical
details hare been confirmed by clcctron inicrograplis wliicii arc
dcscribcd in a later section.

Electron Micrographs
l'lic intinc has not ijccn wcll 1)rcscrvctl in oiir preparations and is
not considereti. '1'11~ cndcsinc forms R thin ( c . 300 A) r:itlicr tlis-
continuous layer. Tlic irrcgular contour of the csinc in relation to
the intinc-with irrcgiilar proccsscs into tlic intinc, is w r y marked
although it may 1)c a n artifact of scctioiiing. Thc cktcsinc is rclatirely
thick. 1'1. 1: A of thc csinc of Snluia nrinria rar. ctrlophyfa in scctioii
is typical of tIic grcat majority of thc species csarriiiictl in this study.
T h e witlc mcslics are supportccl by rnarginal baciila. 'I'lie foot layer
 i

Fig. 1. Diagrams of range of pollen slructurc in Suluia and allies.-A-D, shape and
relative size oI niesocolpin in polar view; A. subequal inesocolpia; I3,C,D, unequal nicso-
colpia; D, LNO rncsocolpia will1 complele tcctuni (Saloia npianu). E-Ii, details of rcli-
culum; E-G, with holes in rcticuluin mcmbranc; 11, uniform small reticulum without
membrane (Iiorininuni-type); I-J. sirnplibaculate (I) ant1 duplibncirlale (J) reliculum;
I<-11, floor of reticulum at low focus: I<, partial 1)acula absent; L, 11, partial bnculo
prcsent.
74 D . .\I. III:SI)EI1SOS, 11. I’llliSTICE A S D 1. C. IIEDGE

is continuous on tlic floor of tlic luniina. ?‘lie lumina a r c closet1 hy

pcrforatc nicniI)raiics forniing a “tcctuin” at tlic level of thc apices
of the baciila. ‘I’he planc of the section passes througli n region of
tlircc perforations in this rncnibrnnc. Ininicdiatcly to thc riglit four
perforations arc scctionctl. Xo frcc standing bncula are prcscnt
within thc luniina of thc rcliculum in these sections nlthough a fctv
occur in thc pollen of this species. 1’1. 1: 13 of Sdniccstninl [c.t-(iii[i
sho\vs similar features but tlic plane of section lies along thc inuri
so that n inore continuous ro\v of 1)ncul:i and less obvious pcrforatc
tcctuni is prcscnt. ’I’he two contrasting types of scgnicnts of Sduici
cipinnn pollen arc illustrated i n 1’1. 1: C & I); C is a section
through tlic largc, more or less continuously tcctatc segnicnts; D is
of one of tlic four discontinuously tcctatc segments. l‘liis tlilTercntial
patterning of scgnicnts presents an intcrcsting prohlcm i n niorplio-
gcnctic control in this group of t h e e spccics invcstigatctl and is
comparatively rarc in pollen morphology.

Pollen Types of Saluia and Its Allies

\\‘c have found it advisahlc to divide thc I)ollcn \vc have stutlictl
into somewlint arbitrary groups. ‘l’hcsc tlcscribc belo\v in terms
of light microscopy. \\‘c Iiavc adopted \\‘ittniann & \\’alkcr’s (1 96.5)
suggcstion of describing the appcnrnncc at various lcvcls of focus but
have also iriscrtetl :I short diagnosis of tlic pollen types \vc rccognisc.
It \viIl hc nt once apparent that we arc tlcaling with a rclntivcly
stcnopnlynoiis group hut nevertheless thc tlivcrsity in fine niorplio-
logical tletnil is still considcrahlc an d forms tlic Insis of tlic sclicnic
\vc propose.

1. S‘iLvI‘i-‘rYr~E
I’ollcn grains G-colpatc, prolate to splicroitlal (30-40 p ) , nicso-
colpia unequal in size, four sinnller and two largcr. I n polar view
the smallcr niesoco1pi:i form tlic grc:itcr, sonicwhat flattcnctl \vall,
the largcr niesocolpia forming thc iiiorc acute cntl (Fig. 1: IS; 1’1.
2: U). Esinc about 1.5 ,u thick, clctesinc much thiclter than cndcsinc.
cktcsinc with tcctum, reticolntc, muri siniplibaculatc, often with free-
standing b:iciiIa within the luniina; rcticular loculi \vitli a inorc or
lcss per foratc ~ n cmb ran c.
Focus A. Rcticulatc. AIeshcs at pole markctlly smaller than in
niesocolpia (at pole: 1 p in tlinmclcr, in nicsocolpin 3 p ) - 1’1. 2: C.
 P O L L E S J l o l i l ' I l o L o G Y O F SAL\'I,\ 7.i

I.'ocus 13. Ikticulate wit11 1iiciii1xxiic ixt\vccn tlic muri \\.it11

scattered or marginal pores in the nicmbrane (1'1. 3: A, 13). Sonic
species form a secondary reticulum \vithin the primary reticuloin.
I:ocus C. Reticulate with reticulum ~ ~ 1 1 1ofs one row of bacula.
I.'ocus I). A s Focus C but with some ccntr:il, free-standing 1)aciila
\vitIiin the lumiiia or in sonic species in only a proportion of the
lu niina.

1 A. Species with many free-standing baciiln and scattered per-

lorations in the tcctoni of each lumen ( P I . 3: A).
l(osninriniis lavnndiilticeiis S . grcindillorri S. plebein
I(. officiiinlis S. liydrnngeci S . ptcrocalys
S. lasiciiillici S. rosifolia
Sri liiici (i tie xi opli y lln S. laoct i i clii loides S. sic/Trulicosci
S . n egyp 1 icicn S . leriifolin S . lriclioclnda
S. cnbirlicci S . mcicroclilcci~igs S. Iriciispitlei l a
S. cciesl)ilosci S. nie.z.icaiia S. lriloba
S. ciii i i 11 b nr iti ri S. Iiioiilbreiii S . oerlicillaln
S. clinopotlioitles S. p il iferei
s. f i l lye11s S . p isidiccc

1 I). Species \\.it11 one or two h c u l a occasionally, o r nonc in each

re h i Iar 1o culu s .
Snlvirr nrinriri var. S . 1ini bn in S. scli inn 111 nusen ii
cn lop11 y 1Cl S. rhylidea: S. sclnrcn
S . bracliycinlli(i S. edelbergii S . iiirgcitci
S . glufiiiosri
S . liorniiriiini

1 C. Species with only marginal pcrfor:itions in tlic meiii1)rane of

each reticular loculus.
Snloici kecrlci s. pollJslctclilJc1
s. le11ctr I1 ill (1 S. piirpiirc(i

1 I). As 1 X hut \vitli cncli major reticular loculus \\it11 secondary

rcticoliini. (Solid parts of the reticular meiiibranc thick liltc the
iliain rcticiilum h i t not regularly supported by h a c u l : ~ )
Saloici drobovii S. eiieiiisiaiici S . forskci li 1ei
2. SA ~ , Iv.\ I. I:L' co 1'1 1T LLA-'I'T I' I:
Pollen grains 6-colpatc, ohlate (32-5G t i in diani.). hlcsocolpia
ant1 csiiic ns in type 1.
Focus r\. Ilcticdatc. hIcshcs at polcs (1.25 p in tlinni.) 1iiarkctl1~-
snialler than those at mesocolpia (3.5 t i in diani.).
I'ociis 1). Iieticulatc \vitIi mesh walls of a single ro\v of hacula, \vitIi
inany smnllcr and shorter lmciila \vithin the reticular loculi.

3. s ;\ 1,VIA A PI'\ S,\-'l.T I> I:

I'ollcn grains G-colpntc, oblate (45-55 p in diaiiicter). 'I'he meso-
colpia unequal ns in Salvin-type 1)ut muri tliiplibaciilatc. Esine 1 ti
t hiclc .
Focus A. Iieticulatc pattern on the four sinaller mesocolpia;
larger mcsocolpia not rcticulnte.
P'ocus 13. Reliciilntc pattern of small iiicsocolpia l>rolic11into t\vo
ro\vs of hncula (PI. 2: 1,) with many free-standing hacula within the
locu li .
Sn loici cipin i i (1 S . soiioiiieiisis

4 . SA LVI A J A Po sI cA -'I s I>E

As Salvia-type h i t with uniform reticulum.
(a) Snloin cidinn I ifol in S . jiirisicii
S . cixillaris s. I'nIens
S. jnpoiiicci S . tiiberiferci
( b ) A s ( ( 1 ) hut \\it11 inarginal p r f o r a t i o n s only in c a d i liiminal
t c c111I l l .
Sn 1i )in 11isp n 11 ica

5. SALVIA SIIVOSIC.~-?'YPE
As Salria-type b u t with iiicsocolpia all equal in size.
salu iu 11 ippO i i icn
G. SA ~~vI A I I I IITI.:I. IA-TTI' E
As Salvia-type b u t \vitIi reticulum on larger mcsocolpia much
smaller than rcticuluni on smaller mesocolpia.
Snlvin liirlclla
5. HoI~>IIsu.\I-‘~’YI,I:
l’ollcii grains G-colpatc, pro1:ite (20-34 p). llesocolpia eqiial i n
size. llembra11c al)sent, luiiiina small (Fig. 1 H ant1 1’1. 3: C). Esinc
1.25 p thick.
Focus A . 1’ol:ir ant1 mcsocolpial ~neslieseqiial in size and small
(usually c. 0.8 p).
1:ocus 13. Ileticulatc; lower part of muri consisting of a single
ro\v of h a c ~ l a .
jlorniiniiiii p~rericiiciini dlcriciiitlra slrobilifcrti

s. I) o I1 T S T O 1:c I I AS -‘h
I:
Pollen grains 6-colpatc, prolate (26-31 11). hIcsocolpia eqiial in
size, iiicslies small, :iricl uniform.
I)orystocchris linsta tci Z+roi)skia n f riplicifol in 1’. ti brolci I ioitlcs

9. S~\LvI’\sTllu~l-‘l’Tl~l~

As in Salvia-type h u t with each of tlic two larger iiicsocolpia

iiiore t1i:in t\vicc as hrontl ;is each o f the four smaller iiiesocolpia.
(See 1’1. ‘1: 13.) k i n e 1.2.5 ,LL thick.
Scilviaslruni lc.~.-ti~iii~ii

Taxonomic Conclusion
I h m the foregoing descriptions of pollen structure, i t is clear that
the most rc\vartling aspect of tlie investigation has 1)ecn tlic interest
of tlic fine structure in itself and that the rewards for the taxonomist
arc less ohviolis. Despite this, sonic general remarlis of a taxoiioiiiic
nature arc a n appropriate adjunct to tlic plynological account a s
several interesting sniall points have crncrgctl.
\\’itliin most of the nine pollen groups there is little correlation
l)et\veen the sI)ecics-groupings and their niorphology or geography.
For cxaiiiplc, New \\rorid species of Stiluia arc represented in four
of tlic six groups that include Snlvias ant1 several esaniples could
he cited to sho\v kick of correlation I)et\veen morphologically
coiiipara1)lc species ant1 the ~ ~ o l l c n - g r o u ~ ~ i nGroup
gs. 1 A, for in-
stance, contains several dilTerent unrelated sections’ of the Old

Xllliougli lliere is no up to d a l e and coiiiprehcnsivc account of the sub-generic

tnsa in Strluin, tliosc rclerrcd to herc arc thosc 01 Epling (1939) for tlic Xmcricnn
species, Pobctliinora (195 1) lor the Soviet species and Briquet (1895) lor the remainder.
78 D. JI. l I I ~ X I ~ l ~ l ~ S11.
O SI’IIIISTICII
, A S D I. C. 11III)GII

\\’orltI: Sect. I’liysospliacc ( S . plcroccilu.~),Sect. ISuspliacc (S. grrrji(/i-

por(i), Sccl. IIpicnospIince (S. hgclrciligea), Scct. Homalospllncc
(S.I i i o n 1brelii), Scc 1. 1 5c ni 0spli nc c (S.( I cgup 1iticci), Scct . 1I cm is 1) 11a c
( S . oerlicillala) ant1 Scct. Notiosplincc (S. plcbcia); Group 1 13 ~ 0 1 1 -
tnins siihgcnus Jfncrosphcrce ( S . sclrn~tillicrusenii).sect. Gongrospliacc
( S . rhytitlctr), Sect. I’lctliiosplincc (S. uirgnltr), Scct. Horminunl
( S . lioriii in r i n i ) , nnd S ect . I) ry i n ospli R Cc (S. g 1ii 1in osu) . ‘I’hc (I ivis i o n
mntlc lict\vccn 1 A ant1 1 I3 lio\vcver is of inlercst in that for tlic most
part 1 A contains spccics \vliicli in floral structwc arc little spccinlisctl
\\*hcrens tliosc of 113, again for tlic most part. arc spccics with the
most ndv:inccd staininn1 structure, i.c. Sccls. I’hysosplincc, Euspliacc,
IIyncnosplinc‘c arc thc iiiorc priinitiw scclioris \vlicrcas Sccts.
Gongrosplincc, I’lctliiospliacc niitl 1)ryiiiosplincc arc tlic 1110re
ntlvniicctl sections. Xnothcr intcrcsting featiirc of typc 1 is the prc-
scncc i n i t (in 1 A) of S . plebeirr; in gcncrnl fncics this is n iiiost ntypicnl
Srililiri n n t l is gcncrnlly rcgnrdctl :is n very nnoinnloiis incmbcr of tlic
gCllUS.
‘hvo intcrcsting features cmcrgcd nniong the spccics from tlic
Anicricns. ‘I’lic first is tlic recognition of Groups 2-S. Icacopliylln
tyl)c--:iiid 3-S. :ipian:i-typc-to :iccoiiimorlntc scvcral s1)ccics wliich
at oiic tiinc \vcrc regnrdccl :is iiicliibcrs of tlie scpnratc gcnus diitli-
bcrlicr. l‘hc sccoiitl is thc dislinctivc pollcn of Srrli~irrslriini lc.z.trniini
\\.liicli puts it in :i group of its o\vn. Snlvitisfriini is :i gcniis of n fe\v
spccics in ‘I’esns nntl Mcsico which 1i:is a similar stnniin:il structnrc
to tlint foiiiitl in soiiic Srrlvin spccics but is tlistinguislictl hy tlif-
fercnces in tlic calyx nntl fruit. A s is thc case in tlic Xudibcrtia
types of Sriliiici, Stilvirr.slritni is a nnrro\v SCgregntC of Srrluirc n n t l the
tlilTcrcnccs in pollcn t)*pcs arc n most intcrcsting nt1tlition:d distin-
guishing fcnturc.
Aiiiong tlic rclalctl gencrn considcrctl, tlic clcnr scparation from
S~tloiciof its possiblc dlics, I)orgstocchtis, I’eror1sliici. illcririntlrm arid
Ilorriiiriiirii (in groups 7 nnd ,8) is an ndtlitional reason for not
regarding tlicin ns closely conncctctl hut rntlicr as dislinct iinrclatctl
gcncra.

List of Specinlens Esaniined (E)

Dorgsloechns Iitislnlct Boiss. Sr. I-Icltlr. Turkey; Antalga, Takhtnli dag,
Davis 15048.
Zforniiririni pgrcnaicuni I,, Italy; Cortina d’Ampczzo-Grail Pass,
Davis 17556.
,\lerinndrn slrobilifcrn Ueiitli.
perovskici nbrolnnoidcs I<arel.
-
p. nlriplicifolin Ueiitli.
I<osrrinriniis lnvandulaccus dc KoE.
11. officinalis L.
Salvin aderiophgllo I Icdgc & I Iuhcr-
JIoratli
S. adinnlijolin Stib.
S . crcggplinca L.
S . apinnn Jeps.
- U.S.A.; California, To1):itop:i ;\Its.,
S . nrinnn Hedge wir. cnlophgla
I-rctigc
S. tixillrrris AIoc. B ScssC
S . brciclignnllici (Bordz.) I'obed.
S . cribiilicn I3cntli.
S . cncspilosn AIoiit. L! Aucli.
S. ciriririharinci AI.! L G .
S . clinopodioidcs H. 13. I<.
S. dorii (Kcll.) Abr:iriis. s s p
gilmnrii (Epl.) Abrams.
S . tirobovii I3otsch.
S . rfiglirlcn BcntIi. (syii.: S . edel-
bergii Ilccli. f.)
S . cvansiana Haiid.-hIzt.
S. forsknlilei L.
s. fulgclls Cav.
S . gliilinosn L.
S . grnndijlorci Etl.
Indin; Kotgneli-Sutlej, Lncc 835.
Afganistnn; Barriinii, Darrnli Sinknr,
IIcdgc & \\'endclbo 4131.
Afglianistnn; Kabul, Kocic 2328.
Afghanistan; Ihbiil, Tnng-i-Ghnru,
IIcdgc B \\'cndclbo 4263.
Algeria; Oran, FnurC 39.
Turkey; Adaiin, Adnna-Kozan,
Davis 26632.
Turkey; Dciiizli, I(izil1iisar-Pesilovn,
Hubcr-AIorntli 12722.
China; I<ianglisi, Hwnngdschu-ling
\\'ring-Te-I Iui 372.
Iraq; Ilublin, tir. Kegcv. \\'hcelcr
Hairies 1 3 I\' G1.
U.S.A.; California, N. Clnremotit, Los
Angclcs, Balls 8697.
Abrnms B AlcGrcgor 1-17.
Afganistnn; Alnymann, AInyinniin-
BelEcrngli, Hedge CQ \\'endelbo 3G89.
Alcsico; I Iitl:ilgo, P:icIiucn, Pringlc
6905.
Cult. Iloynl Uotniiic Garden, ISdiii-
burgh C3003, froin seed of Davis.
B:iliicliist:in; Clinrn Butney, Lace
4008.
Turkey; AIaras, A'urilink tlag-I~lbi-
stan, St:iiiitoii & Henderson 5630.
Alesico, Oasnca, Sierra dc Sari
Fil i 1) c, I'rin glc 4 9-17.
Alcsico; hliclioncnii, I'atzcunro,
I'ringle 1258.
U.S.A.; California, N.\\'. Camp
Cnjon, S:in Bcrnnrtlino, B:ills 8.193.
Asia mctlin; I'amiro-alnj, Kyr-tan,
Grczdillo 20 vi 37.
Afgnnist:iii; Aliiijnii, Etlelberg 1300.
Yuiiiinn; \Vci I-Isi, A1cL:ircri 149.
'I'urkcy; 'I'rabzoii, Alnckn, Stniiitoii Sr
Henderson 5885.
hIesico; Sierra dc AIistun Pucbln,
Purpus 3289.
Turkey; Hamcsckcni, Un11s 499.
Turkey; Aiitnlyn, AIaiiavgnt-Akscki,
Davis 35i92.
S. hirlclla \‘ah1
S . hispanica I..
S. hormiriient L.
S . hydrangea DC.
S. jnponiccc Tliunh.
S . jiirisicii Kosanin
S. kccrlii Ilcntli.
S. lasianliict Beritli.
S . laonndicloidcs 11. 13. I<.
S. lerii/oliei Bcntli.
S . lcircarilhci Cav.
S. leucophglltr Greenc
S . linibnln C. A. ;\Icy.
S. nicierochlezniys Boiss. d Icy.
S. nicxicnrirc L.
S. moiilbrefii Bcnth.
S . niurizii Epl.
S. riipponica ~ I i q u c l
s. pnlcr1s Cav.
S . pili/ertc Alontb. B Auch.
S. pisidica Uoiss. & Heldr.
S. plebein R. Br.
S . rhylidca I3cnth. (Syn.: S . poly-
chroma Iiech. f. B I3IclB.)
S. rliylitiea Benth. (Syn.: S. poly-
clccdn Rccli. f.)
Ecuador; Tunguragua, Ikicienda
Yanayacu Alocha, I3alls 718.1.
AIcsico; Xliclioacnn, Uruapan,
I’ringlc 10115.
Turkey; Adana, Ccylian-Osnianiyc,
Davis 26703.
Iran; Azerbaijan, I<hoy, Laniond
2551.
China; Kiangsi, Kwci Tsung Sze,
Chco 353.
Cult. Royal Botanic Garden, Btlin-
biirgli, C 1487.
Alcsico; IIitlalgo, Aletcpcc Station,
I’ringIe 10312.
AIesico; Oasaca, Monte Alban,
Pringlc 4862.
Guatcmala; I3aja \‘crapaz,
Turckhcini 111532.
Afglianistan; Kabul, Slier Darwasn,
IIctlge B \\rentlelbo 2027
XIcsico; Mexico, Tcnango, Balls
8608.
U.S.A.; California, Iiancho Santa
Aria Bot. Garden, Balls 0388.
Turkey; Agri, Dogbnynzit-A&-i,
l,a11101id255-1.
Turkey; Bitlis, Iksadiyc-IJclli, 1)avis
22405.
AIesico; AIcsico, Rio IIondo Canyon,
I’ringlc 10115.
Turkey; AIardin, Alardin-Savur,
Davis 28517.
U.S.A.; California, Rancho Santa
Ana I3ot. Garden, I3alls 8181.
Japan; Sngami, Ihbotobe Pass, Ali-
zuspim 490.
AIcsico; AIcsico, Tultenango Canyon,
Pringlc 8675.
?‘url;ey; Alaray, Ahray-G6ksun road,
Stainton B Hcndcrson 5527.
Turkey; Antalya, Elninli-I<orbuteli,
Davis 35214.
Nepal; I,a\vanijuln, I~olunin,Sykcs B
\\‘illinins 674.
Afghanistan, Nuristan, AIinjan, Edel-
berg 1437.
Afghanistari; Sliibar Pass, I<oclz
13507a.
 Jlcsico, Oasaca, I’urpus 4168.
Afglianistan; Qataghan, hlirza Atbili
Pass, IIcdgc (4r \\’cndclbo 3557.
JIcsico; Orizaba, I3alls 428G.
Afghanistan; nr. Bulola, Gilli 3502.
Turkey; Coruh, Ardunu~-I(ordcvan
Da& Davis 30087.
s. scliriicrlhallsenii Rgl. Ccntral Asia; Tadzikistan, 1’1. Orient.
13ucliaricac G14.
S. sclnreci L. Iran; hlazendcran, Nczva Kuli,
\\‘cndclbo 100.1.
.q. soriontcnsis Grcenc U.S.A.; California, Rancho S:int:i
Ann Botanic Garden, Balls 0350.
s. spcilhncea Grccnc U.S.A.; California, Rancho Santn
Ann Botanic Gardcn, I3alls 8180.
s. sirfiriilicosa Jlont. B Aucli. Turkey; Elazifj, Elazik-Pertck.,
Davis 20190.
S. lriclioclada Bcntli. Kurdistan; Kuli-Sclin, Bornmullcr
1721.
s. trierrspiclnla II. (4r G . hksico; Onsaca, Sierra de Sari Filipc,
Pringlc 4845.
S. lrilobn L. f. Turkey; Izniir, Samsun Dafj, Davis
4liO0.
S . firbifern Ldvl. China; Yurinan, Chi l’su IIt.,
IIcLarcn 110b.
S . imticilirfn L. Turkey; Bitlis, Rcsndiyc-Pelli,
Davis 22407.
S . uirgalci Jacq. Turkey; Artvin, Artvin, Stainton B
IIcndcrson 5010.
Snluicislriim lcxcinum Schcclc U.S.A.; S. Texas, Iccrrvillc, Hcllcr
1635.

HEFERENCES

BRIQUET,J., 1805. Labintac. - Englcr (4r Prantl, Natiirl. Pflanzcnfam.

I\’, 3A: 183-375.
ITPLISG, C., 1038. Tlic Californian Salvias. - Ann. 310. bot. Gdti. 25:
05-152.
- 1030. Rcvision of Salvia, Sect. Calosphace. - Feddc Rcp. Bcili. 110:
1-380.
EIIDTXIAS, G., 1060. The acetolysis mctliod. - Svcnsk bot. Tidskr. 51:
561-56-1.
I~ESDEKSOS, D. II., 1065. T h c Pollcii hlorphology of Meconopsis. - G r a m
palynol. 6: 101-200.
G - 669844 Grona Pel. vol. 8: 1
82 I). .\I. I I I ~ S l ~ l ~ I I S O11.S , I’IIKSTICI: , \ S I ) I. C. IIEDGE

1956. Po1leiigest:iltung bei einigeii polyploidcn Kultur-

h I , ~ u r i r z ~ o A.,
,
pflanzen. - Gmna polgnol. 1 (2): 59-G9.
POLIEDIJIOVA, 1:. G , , 1954. Labintnc p.p. - Itomarov Flora URSS 21:
24 4 -3 G 3.
\\‘ITT>r,w, G., & WALKER, D., 19G5. Townrds Siiiiplificotioii i n Sporotlcrin
Description. - I’ollcn Sporcs 7: 442-15G.
83
s4

PI. 2. A, n. optical scction, polar view of: A, I f o r m i n i m pyrennicrim; U, Salvinslriim

lexanicm x 1250; C, Salvin apinnn, polar riew; D, Saivia sonomensis, cquatorial view
x 800,
 I’OLLES J I O I ~ P I I O L O G T01: S,\LVI:\ 85

1’1. 3. Equatorial views, upper half high focus, lower linlf lo\ver fociis; A, Saluin lerii-
/ o h ; B, Saluin rliglitlen; C , Iiorminum pyrenaiecim; D,1)orysloecha.c haslala. XI1 x 800.