Studies in the Ericoideae . III . The genus Grisebachia

A revision of the genus Grisebachia Klotzsch in which eight species are recognized is presented. The genus belongs to the Ericaceae-Ericoideae and is endemic in the south-western part of the Cape Province. The work revealed a high degree of variability among the species, necessitating the reduction of seven species to infraspecific rank, seven species to synonomy and the rejection of one species as imperfectly known. One new species G. secundiflora E. G. H. Oliver is described.


HISTORICAL OUTLINE
W hen K lotzsch reclassified the subfam ily Ericoi deae in 1838, he described the genus Grisebachia consisting o f eight species: G. ciliaris sensu K lotzsch, G. hispida K lotzsch, G. involuta K lotzsch, G. zeyheriana K lotzsch, G. incana (B artl.)K lotzsch, G. hirta K lotzsch an d G. plum osa K lotzsch.O f these, only G. incana an d G. plum osa are retained in the present revision.L ater in the sam e w ork he described Eremia parviflora, w hich is now recognized as a species o f Grisebachia.
The follow ing year B entham (1839), w hen revising the whole fam ily E ricaceae, retained Grisebachia but in an enlaig ed fo rm .He upheld K lo tzsch 's eight species and ad d ed G. dregeana and G. serrulata, both o f which have now been reduced to synonom y.He also included F inkea K lotzsch w ith tw o species, as a section.N .E. Brow n (1906) correctly placed the latter in the genus Acrostem on o f K lotzsch.
In 1876 B entham again revised the fam ily and included a fu rth e r tw o o f K lotzsch's genera, Acros temon and Comocephalus, und er Grisebachia.The genus was then divided up in to three sections based on the shape o f the corolla, the hairiness o f the filam ents and the ovary com plem ent.
In Die N atiirlichen Pflanzenfam ilien D rude (1897) to o k a very conservative view o f the E ricoideae and placed Grisebachia as defined by B entham as one o f four sections in the genus Erem ia D .D o n .
N .E. Brow n (1906) in F lo ra Capensis changed th s system o f the earlier w orkers and adop ted Grisebacl.iias originally co n stru ed by K lotzsch.He retained Acrostem on as a distinct genus an d reduced Como cephalus and F inkea to synonom y u nder it.He retained all eight o f K lo tzsch 's species and placed Eremia parviflora correctly in Grisebachia, but as G. eremioides w hich h ad been nam ed by M acO w an in 1890.He also a d d ed ten o f his ow n species o f which only three are upheld as distinct species in the present revision, nam ely G. rigida, G. nivenii and G. minutiflora. Phillips (1926) accepted Brow n's w ork in its entirety in the first edition o f his G enera.In 1944 he put forw ard his proposals fo r a reclassification o f the family in S outh A frica fo r the second edition o f his G enera (1951).He presum ably based his ideas on the very conservative views o f D rude.He recircum scribed all o f the genera an d placed Grisebachia u n d e r Erem ia together with seven other genera, som e o f w hich are quite unrelated.
The genus has been retained in D yer's G enera (O liver 1975) in the sam e form as adopted by N .E. Brow n in F lo ra C apensis.
W hen the present revision was un d ertak en the genus Grisebachia consisted o f 21 species.As a result o f finding num erous variations and overlapping o f characters the num ber o f species has been reduced to seven.One new species, G. secundiflora, has been added.

MORPHOLOGY
In hab it m ost o f the species o f Grisebachia are erect, often form ing com pact shrublets.G. parviflora is usually sparse an d spreading am ong rocks an d vegetation and G. secundiflora is com pact b u t rath e r spraw ling.
The branches o f all species are never entirely glabrous.M ost have pubescent to pilose or tom entose branches w hen young, som etim es w ith sim ple to plum ose sto u t hairs interm ingled.These m ay be gland-tipped.
T he leaves are all typically ericoid w ith no openbacked form s and are m ostly 3-nate.In G. plum osa subsp.hispida they are always 4-nate, while in subsp.pentheri they can be occasionally 4-nate inbetw een 3-nate.M ost leaves are adpressed with one exception, G. ciliaris subsp.multiglandulosa, where they are recurved spreading.The indum entum o f the leaves is very variable and disjunctions have been used for taxonom ic division.S tout simple or plum ose hairs m ay o r m ay n ot be present on the leaves and m ay be confined to the m argins or occur on the adaxial surface as well.In m any cases these stout hairs m ay fall off and rem ain only as short stubs, w hich can easily be overlooked.
T he flowers o f all species are term inal either at the ends o f the m ain branches or m ore often at the ends o f sh o rt lateral branchlets.In G. parviflora these sh o rt branchlets m ay be aggregated together to form a loose pseudospike and in G. secundiflora the pseu d o spike is com pacted an d secund.The 3-bracteolate flowers are usually 4-12 in a head or as m uch as 36 in G. minutiflora.The bracteoles vary considerably in shape and size, particularly in G. plum osa and G. ciliaris where they are o f taxonom ic im portance.The bracteoles m ay be equal to very unequal w ith the m edian bracteole being considerably enlarged w ith an expanded base.
T h e variation also occurs within a single inflorescence w here the bracteoles m ay be very unequal in the ou ter flowers to equal in the inner flowers (Fig. 1).
The calyx in all species is 4-lobed o r 4-partite.In G. ciliaris, G. incana, G. rigida an d G. nivenii the sepals are free o r very slightly jo in ed a t the base, whereas in G. plumosa, G. parviflora, G. minutiflora an d G. secundiflora they are joined for q u a rte r to three quarters o f their length.In all species the sepals are m ore or less equal.Som etim es the lateral pair m ay be slightly narro w er th an the ab-an d adaxial pair.The indum entum o f the calyx is very variable and is used as a taxonom ic character.In m ost species there are some sto u t sim ple to plum ose h airs on the m argins o f the sepals and som etim es also on the abaxial surface.These may be gland-tipped in dif ferent taxa or in th e same taxon in the young stages.
The corolla is o f tw o basic shapes in the genus.Five o f the species, G. ciliaris, G. plum osa, G. rigida, G. incana and G. nivenii, have a corolla sim ilar to th a t in the section Cyatholom a o f the genus Erica.T he corolla tube is m ore o r less distinctly con stricted in the m iddle with an ovoid or o bovoid base and cyathiform upper p o rtio n including the large lobes.This is evident in the fresh state a n d in m ost o f the species in the pressed m aterial.But care m ust be taken w ith m aterial o f G. ciliaris subsp.ciliaris in w hich the small flowers easily loose this shape when pressed.O ccasionally, w hen the constriction is not very m arked, a cam panulate shape occurs.
In the three rem aining species, G. parviflora, G. minutiflora and G. secundiflora, the corolla has no distinct constriction.In the first tw o species it is usually funnel-shaped o r obconic an d in the th ird it is tu b u lar or tu b u lar with an inflated m iddle poition.In the m ajority o f species especially those w ith the constricted corolla the corolla is pubescent to pilose in the middle region outside and also inside aro u n d the point o f constriction.
The num ber o f stam ens in all specim ens exam ined was constantly 4 and is im p o rtan t in the generic classification.The stam ens are free an d have pilose filam ents.
The anthers are m ostly m anifest being arranged ju st above the constriction in the corolla.In G. secundiflora they are m anifest to included.The m ajority o f anthers are characteristically bipartite.In G. secundiflora t.iere is a tendency foi them to be bilobed.Awns are present in several species, b u t may be absent in anthers o f the sam e flower.This ch arac ter, used in the past for specific recognition, is o f no use taxonom ically.T he an th ers are all dorsally attached w ith an expanded apex to the filament.
The pollen in all the m aterial exam ined occurs as single grains, which are found in several o f the o ther m inor genera o f the Ericoideae.The grains are tricolporate, the furrow s being deeply chanelled and alm ost as long as the cell.In shape the grains are m ostly ellipsoid with flattened apices.In a few cases they are oblate as in G. nivenii an d some form s o f G. plumosa and G. parviflora.The sculpturing o f the surface in the first six species is either scabrate or m icroscabrate with a tendency for the elem ent rods to becom e fused at their distal ends to form tecta.In the last tw o species the fusion is com plete giving an alm ost sm ooth appearance to the pollen surface (Fig. 2).
The ovary is m ostly 2-celled w ith a single pendulous subapical ovule in each cell.Very occasionally 3-celled ovaries occur, notably in G. parviflora subsp.pubescens.In G. secundiflora the ovary is constantly slightly obliquely 1-celled.
M ature fruits th a t were found in a few species were hard-w alled nuts w ith the walls often verrucose.They are apparently indehiscent a n d contain one or tw o very soft juicy seeds.Some fruits on Levyns 1367 (G.ciliaris subsp.multiglandulosa) collected in 1925 still contained soft juicy seeds.
As in m ost genera o f the E ricoideae, the ovary in Grisebachia is seated on a nectariferous disc which, in some cases, is very conspicuous.This suggests th a t all the species are insect pollinated.O n a few occasions it was noted in the field th a t plants were visited by bees.
The stigm a varies from sim ple to capitellate, which is in accordance w ith the insect pollination.

DELIMITATION OF THE GENUS
As defined in the present revision the genus Grise bachia is characterized by having 3 bracteoles, 4 sepals, which are free or p artly fused, a 4-lobed corolla, 4 free stam ens w ith bipartite anthers and an ovary w ith 2, rarely 3 o r 1, cells and a single ovule in each cell.The im p o rta n t characters arfe the stam en num ber and bipartite anthers.
The uniform ity o f the genus has until now been recorded as very constant.It has been relatively easy to assign m aterial to the genus when identifying Ericaceae.The 2-celled ovary, 4 stam ens and bipartite anthers served to be a distinctive com bination o f characters.
N only know n from the type collections.A n exam ination o f a few flowers did n o t confirm this an d w ithout investigating num erous flowers it was decided to accept Brow n's observations.A num ber o f flowers w ith 3-celled ovaries was, how ever, fo u n d in G. parviflora subsp.pubescens.
There are several genera in the E ricoideae w hich have 2-celled ovaries an d 4 stam ens, i.e.Simocheilus, Acrostem on, Thoracosperma, Sym pieza, Aniserica an d Coilostigma, but none o f them has the distinctive bipartite anthers found in Grisebachia.They all have a different appearance from the ra th e r uniform Grisebachia.
W ith the discovery o f G. secundiflora, this uniform ity was slightly changed.This species was difficult to place satisfactorily in any o f the genera due to its 1-celled ovary.The species was clearly allied to som e species o f Eremia and Grisebachia, but could not be included in the form er on the g rounds o f having only 4 stam ens and in the latter for having a 1-celled ovary.As the genus Erem ia has recently been em ended (Oliver, 1976) to include ovary v ariatio n s from 4-celled to 1-celled but w ith constantly 8 stam ens, it was decided to place this new species u n d er Grise bachia next to G. parviflora an d em end th a t genus to include the 1-celled ovary ra th e r th an a lte r Erem ia even furth er to include 4 stam ens.The 1-celled ovary b ro u g h t the species cl >se to Anom alanthus and Syndesmanhus, neither o f which it resembles.
The close sim ilarity betw een Grisebachia and Eremia has been m entioned under th a t genus (Oliver, 1976).G. parviflora is superfi;ially sim ilar to Erem ia curvistyla in flower form and habit.G. secundiflora looks very m uch like Erem ia totta but, in b o th cases, the 4 stam ens serve to distinguish them as species belonging to the tw o separate genera.The an th ers o f all species o f Eremia except E. curvistyla are only bilobed and not distinctly bipartite as occurs in Grisebachia.In Eremiella outeniquae (Oliver, 1976) the anthers are also bipartite, b u t the rest o f the floral characters are very different from Grisebachia.
Grisebachia and Erem ia are sym patric to a great extent in the area from the C edarberg to the C old Bokkeveld.U ndoubtedly they are very closely allied and have possibly envolved from som e ancestral stock, which in tu rn arose from the genus Erica by reduction.

PHYTOGEOGRAPHY
T he genus Grisebachia is endem ic in the so u th w estern and western parts (Fig. 3   Com pact erect shrubs up to 0 ,5 m, rarely 1 m, high.Branches num erous erect pubescent to tom entose with longer stou t hairs inbetween, som etim es glandtipped, som etim es becom ing glabrous and grey.Leaves 3-or 4-nate inbricate and adpressed to spread ing recurved, up to 4 m m long, linear-oblong to ovate, acute to obtuse, glabrous o r puberulous to tomentose and canopubescent when young an d with short to long stou t plum ose or simple eglandular or glandular hairs on the m argins only or also on the abaxial surface becom ing glabrous on the abaxial surface an d scabrid w ith the sto ut hairs falling off leaving short trun cate stubs, rarely only crisped pubescent w ithout any stout hairs.Flowers (1) 6-12 (16)-nate in term inal erect or nodding heads; pedicels short, 0 ,5 -1 ,5 m m long, puberulous som etim es w ith longer stouter hairs inbetween, som etim es glandtip p ed ; bracteoles m edian to adpressed, m arkedly unequal to subequal, the m edian from lanceolate to broadly ovate from an expanded base, 1 ,5 -3 , 5 x 2 , 7 m m , the laterals usually sm aller and narrow er, m ostly oblong-elliptic, all ciliate with short to long stou t plum ose to simple hairs which may be glandtipped, w ith or w ithout an even to sparse covering o f sim ilar shorter or equally sized hairs on the abaxial surface, som etim es ju st crisped pubescent.C alyx jo in ed for one th ird to two thirds o f its length, cam panulate som etim es 4-angled at the base, pink, glabrous to pubescent w ith the lobes ciliate w ith sh o rt to long simple to m arkedly plum ose stou t to soft hairs which m ay be gland-tipped, occasionally w ith an even to sparse covering o f sim ilar shorter or equally sized hairs on the abaxial surface; lobes narrow ly to broadly deltoid, slightly sulcate a t the apex.Corolla 4-lobed up to 4 m m long, very co n stricted half to two thirds the way up above an ovoid to obovoid base cyathiform above, often 4-angled the angles alternating w ith the calyx segm ents, pubescent to sparsely so in the m iddle region and pilose on the inside at the m outh o r constrictio n ; lobes broadly ovate to deltoid, obtuse erect to slightly spreading, glabrous or slightly pubescent dow n the centre outside.Stam ens 4, free; filam ents narrow ly linear broadened at the point o f attach m en t, pilose, w hite; anthers exserted o r m anifest, 0 ,5 -1 ,3 m m long obovate, dorsally attached, scabrous, m uticous or rarely m inutely aw ned the awns occurring only in a few flowers or an th ers; pore a b o u t h a lf the length o f the cell; pollen grains single.Ovary 2-celled, com pressed, ovoid to oblate, obtuse to subacute sm ooth to verrucose rarely pilose at the apex otherw ise glabrous, seated on a disc; style exserted; stigm a small, subcapitate; fruit hard verrucose.Figs 5-8.
A species form ing erect shrublets up to 0 ,5 m occurring on sandy coastal flats in the w estern Cape Province from Cape T ow n to G raafw ater an d on m o untain slopes in the C lanw illiam area, flowering from June to Septem ber.
G. plumosa is characterized by the calyx being joined for one q u arter to th ree-quarters o f its length, the corolla-tube being distinctly constricted in the m iddle and the habit being erect.
In his treatm ent o f the genus B row n recognized five species in the group with joined calyces, basing the separation on the form o f the sto u t hairs on the calyx.The five species were G. plum osa, G. hirta, G. pilifolia, G. pentheri an d G. solivaga.O n the small am o u n t o f m aterial available to him this classification was feasible.But since F lo ra Capensis num erous collections o f these species have been m ade.A n exam ination o f all this m aterial show ed a degree o f variation in the diagnostic characters sufficient to w arran t the five species being regarded as one single com plex o f taxa with discontinuities occurring only in one character an d with partial separation in oth er characters between the constitu en t taxa.T he oldest nam e applicable to this com plex is G. plumosa K lotzsch.
It was also found th a t v ariation in the n um ber o f leaves per w horl in som e specimens o f G. pentheri overlapped with the num ber in the very sim ilar G. hispida which had been separated off from the rest o f the species in the genus on this character.G. hispida therefore had to be included in the G. plumosa Com plex.
The above six taxa were then exam ined as one com plex group.It was found th a t the g roup could be divided into two form series on the position o f the stout hairs on the leaves, one w ith the hairs only on the m argins the other w ith the hairs also scattered on the abaxial surface.The first series co ntained G. plum osa and G. solivaga, the second co ntained G. hispida, G. pentheri, G. pilifolia an d G. hirta.
In the first form series the variatio n in calyx hairs from the very plum ose m aterial o f G. plum osa in the M am re area to the alm ost sim ple-haired specim ens in the A u ro ra area show ed an overlap w ith the type and only collection o f G. solivaga from ju st west o f Clanwilliam .M aterial which had been nam ed as G. hirta was found to constitute a distinct new taxon m ore closely allied to G. plumosa on the leaf character.F urtherm ore, a collection m ade in the G if berg (Oliver 4951) was found to be nearest to G. plumosa and, although som ew hat anom alous, was referred to this series which then consisted o f G. plumosa (in cluding G. solivaga) and the tw o new taxa.
In the second form series G. pilifolia showed considerable variation w ith an overlap in the dis tinguishing characters with G. pentheri thus necessitat ing its reduction to synonom y.A close relationship with G. hispida was show n to exist with only a partial separation on the num ber o f leaves per w horl an d a distinct separation in the plum osity o f the hairs.G. pentheri and G. hirta appeared to be very sim ilar w ith only one character show ing any disjunction, i.e. the position o f the stout hairs on the abaxial surface o f the calyx.This series therefore consisted o f G. hispida, G. pentheri (including G. pilifolia) and G. hirta.
W ithin these two form series recognizable on the single character difference o f leaf hairs, several m ore or less distinct taxa could be distinguished again on various single character differences.The com plex occurs in tw o m ain d istribution centres, the M am re area in the south and the m ountains west o f the O lifants River in the n orth.Regional separation o f the two series in the com plex is only partial.The " plum osa" series is concentrated in the south with some outliers in the far no rth and the " hispida" series in the n orth with outliers in the south.It was decided fo r reasons o f expediency to regard the com plex as one species w ith six subspecies based on a single ch aracter d isjunction occurring w ith some degree o f regional sep aratio n over a wide part of the d istrib u tio n range o f the species (Fig. 4).This classification, th o u g h n o t final, attem pts to show the type o f variatio n th a t occurs in this complex.Only a th o ro u g h biosystem atic study o f the populations will solve the problem s an d should either confirm o r correct the above classification I have given.
In floral an d foliage characters this is a very variable taxo n in w hich six subspecies are recognized.Branches pubescent to tom entose w ith sh o rt stout plum ose hairs inbetween, rarely ju st tom entose.Leaves 3-nate m ostly erect and adpressed, pubescent to canopubescent and ciliate w ith short, occasionally long, sto u t plum ose hairs, becom ing glabrous an d often scabrid edged w ith the cilia falling off leaving sh ort truncate setae.Bracteoles pubescent, rarely glabrous, ciliate w ith sh o rt stout plum ose hairs, rarely also w ith sim ilar hairs on the abaxial surface.Sepals pubescent, rarely glabrous, ciliate w ith stou t plum ose, rarely subplum ose hairs, with a few shorter ones scattered over the abaxial surface.Subsp.plumosa is characterized by having m ostly plum ose, occasionally simple, cilia on the leaf m argins an d no glands on the calyx.The cilia are present at least in the young stages as they often fall off leaving m inute truncate setae which can easily be overlooked.This latter character was used by B entham in creating his G. serrulata.Branches pubescent becom ing glabrous w ith sh o rt sto u t plum ose hairs inbetw een w hen older.Leaves 3-nate, adpressed but som etim es slightly recurved, pubescent becoming glabrous on the abaxial surface, ciliate w ith short sto u t plum ose hairs, occasionally gland-tipped when young, som etim es falling off w hen older and rem aining as sh o rt tru n cate setae.Bracteoles and sepals pubescent m ainly at the apex, ciliate w ith simple to sparsely plum ose gland-tipped hairs on the m argins and slightly sh o rter ones on the abaxial surface.Subsp.irrorata is recognizable by the gland-tipped hairs on the calyx and lack o f sto u t hairs on the abaxial surface o f the leaves.

K ey to the subspecies
M aterial o f this tax o n had been, until now, placed under G. hirta K lotzsch due to the key ch aracter o f a glandular calyx w ithout taking into account the significant differences in the leaf indum entum .In subsp.irrorata the leaves are typical o f the " plum osa" kind in which the sto u t hairs are confined to the m argins o f the leaves and do n o t occur on the abaxial surface as they occur in subsp.hirta (G.hirta Klotzsch).
The collection, Compton 9530, is interm ediate between subsp.irrorata and subsp.plumosa in having glands only on the abaxial surface o f the calyx.There appears to be no interm ediate between this taxon an d subsp.hirta.H ow ever, after a biosystem atic study this taxon m ay be show n to be a hybrid between subsp.plumosa and subsp.hirta.
The three taxa, subsp.plumosa, subsp.irrorata and subsp.hirta are sym patric on the sandy flats o f the M am re area.These flats have unfortunately been decim ated by alien vegetation and hum an activity.The pressure on the area from industrial and u rban developm ent is now very great.
A thorough biosystem atic study o f the populations from this area was not possible and probably never will be possible for a m ore objective assessm ent o f their relationships.
Branches pubescent w ith sim ple hairs only.Leaves 3-nate adpressed, lanate w ith simple hairs, occasionally with a few tufts o f sto u ter hairs on the m argins.Bracteoles pubescent and lanate at the apex.Sepals very sparsely pilose, ciliate w ith irregularly and sparsely plum ose hairs, rarely with a few sim ilar hairs on the abaxial surface.Subsp.eciliata is distinct in the species for having leaves which do n ot possess stout hairs on either the m argins or abaxial surface.The pubescence is very short, lanate an d crisped.There is, however, an occasional tuft o f hairs on the m argins o f the leaves b ut n ot sim ilar to those found in the rest o f the species.
This taxon is som ew hat anom alous in th a t it has a sim ilarity to G. plum osa com plex in w hich the closest affinity is with the m aterial form erly know n as G. solivaga N .E .Br. now form ing p a rt o f G. plumosa subsp.plumosa.T he crisped pubescence, lack o f distinct stout hairs and the tufts on the leaves are sim ilar to the condition found in G. ciliaris subsp.ciliaris which occurs in the same area.The broad calyx lobes are sim ilar to the broad sepals in the m aterial form erly know n as G. dregeana Benth.The sepals o f the latter are, how ever, free or very slightly joined a t the base.
It was decided to place this taxon under G. plumosa on the basis o f the fused calyx segments and to leave G. ciliaris to be characterized by its free sepals.The taxon is, however, a close link between these tw o species and points to the need for a thorough biosystem atic study o f all the O lifants River taxa to understand their relationships.Subsp.hispida m ay be distinguished by its 4-nate leaves and calyx thickly covered w ith densely plum ose hairs.In all the m aterial exam ined the leaves were 4-nate.In subsp.pentheri som e specim ens have been found to possess 4 -nate leaves on branches w ith mostly 3-nate leaves.T he subspecies has the largest leaves, flower-heads an d flowers in the species.Branches pubescent, rarely glabrous, w ith long stiff plum ose eglandular or gland-tipped hairs i.ibetw een.Leaves 3-nate, rarely also 4-nate, pubescent w hen young, ciliate w ith sto u t hairs and clothed w ith a few sim ilar hairs on the abaxial surface, the hairs being sim ple and gland-tipped to plum ose an d eglan d u lar, often breaking off and rem aining as sh o rt tru n cate setae.Bracteoles and sepals m ostly glabrous, rarely sparsely pubescent, ciliate w ith sim ple to plum ose cilia w hich may be gland-tipped, occasionally clothed w ith a few sim ilar shorter hairs on the abaxial surface dow n the middle.Subsp.pentheri is characterized by its 3-nate leaves, w hich are very rarely 4-nate on the same branch, its calyx w hich is m ore sparsely hairy and less plum ose th an in subsp.hispida a n d in the glandular form by having m ost o f the hairs o n the m argins o f the sepals.This subspecies is very variable in the form o f the hairs on the leaves an d calyx.In F lo ra Capensis Brow n recognized tw o separate species based on these hairs, his ow n G. pilifolia w ith its sim ple to plum ose eglandular hairs and later in the ad d en d a Z ah lb ru ck n er's G. pentheri w ith its gland-tipped simple to subplum ose hairs.Since F lo ra Capensis, m ore m aterial o f this g roup has been collected and has exhibited a com plete range betw een the two extrem es thus necessitating a red u ctio n o f G. pilifolia to synonom y under G. pentheri w hich itself h ad to be reduced to subspecific ran k in the G. plum osa com plex.
T he relationship betw een the glan d u lar form s o f subsp.pentheri and subsp.hirta is very close an d it is only w ith some careful exam ination th a t they can be distinguished.The only ch aracter w hich shows any discontinuity is the d istrib u tio n o f the hairs on the calyx.In subsp.pentheri the sto u t hairs are m ostly confined to the m argins o f the calyx lobes w ith the hairs on the abaxial surface being few an d shorter.In subsp.hirta the hairs are m ore o r less evenly distributed over the calyx an d are o f the sam e length.This relationship is interesting because the two tax a are widely separated.B ranches pubescent to tom entose w ith long stiff plum ose gland-tipped hairs inbetween.Leaves 3-nate, pubescent w hen young becom ing glabrous, ciliate w ith long stout simple to plum ose gland-tipped hairs and clothed w ith sim ilar hairs on the abaxial surface, erect to spreading-recurved.Bracteoles an d sepals puberulous sometimes sparsely so, ciliate an d evenly clothed on the abaxial surface w ith num erous short sto u t simple to sparsely plum ose gland-tipped hairs.Subsp.hirta is characterized by its glandular stout mostly simple hairs evenly distributed on the calyx.It differs only in this respect from subsp.pentheri and from subsp.irrorata in having gland-tipped hairs on the m argins and abaxial surfaces o f the leaves.
The relationship betw een subsp.hirta and subsp irrorata is very close w ith the flowers being alm ost identical.The difference in the leaves, how ever, is distinct.In subsp.irrorata the leaves are typical o f subsp.plumosa w ith short stout plum ose cilia.A s all three taxa are sym patric o n the sandy flats near M am re there is a possibility th a t hybridisation an d introgression m ay occur.A th o ro u g h biosystem atic study o f populations from the area will have to be carried o ut to ascertain the relationships o f these taxa.

Blaeria ciliaris L.f., Suppl. 122 (1782).
Shrublets m ostly low grow ing an d com pact or erect up to 75 cm high.Branches subglabrous to pubescent, occasionally arach n o id , the hairs thick and m atted, erect or retrorse or very sparse and erect, som etim es with stouter longer hairs inbetw een w hich are either plum ose or sim ple and gland-tipped.Leaves 4-nate, erect and adpressed, som etim es im bri cate to spreading recurved, 1 -4 ,5 m m long w ith the petiole very short to 0 ,5 mm long, from linear to ovate to obovate, very variable in the indum entum , pubescent w ith dense crisped retrorse hairs or p uberu lous with erect hairs, eciliate or rarely with sm all com pound tufts on the m argins o r som etim es with stout plum ose cilia with few to m any spreading plum e branches, the cilia often falling off and rem ain ing as short setae, all becom ing m ore or less glabrous with age, som etim es subglabrous to glabrous and shiny on the abaxial surface and ciliate and clothed on the abaxial surface w ith sto u t sim p b gland-tipped hairs.Flowers 3-12 in capitate, som etim es nodding, heads a t the ends o f lateral b ranchlets; bracteoles 3 subequal to m arkedly unequal in the outer flowers o f the inflorescences to equal in the inner flowers, mostly m edian to rem ote, adpressed or recurved, the m edian 1 ,3 -5 x 0 ,4 5 -2 ,3 m m , sm all an d oblong to narrow ly ovate w ith a relatively large keel-tip and no m arkedly expanded base to broadly elliptic or ovate with a broad flat base and relatively sm all b u t distinct keel-tip, from alm ost glabrous to puberulous all over, som etim es w ith a distinct apical tu ft o f lanate hairs, sometimes ciliate w ith short to long stout sim ple to plum ose eglandular o r gland-tipped hairs, rarely with a few sim ilar hairs on the abaxial surface at the keel-tip; the pedicel 1 ,0 -2 ,5 m m long, puberulous to sparsely glan d u lar pilose.C alyx 4-partite som etim es slightly joined a t the base, 1 , 5 ^, 3 x 0 , 4 -2 , 3 mm, very variable in size an d indum entum , sm all narrow ly oblong to oblong-ovate to broadly elliptic or large oblong-elliptic to b roadly elliptic and ovate, slightly keel-tipped occasionally w ith a knoblike apex, glabrous to pubescent som etim es w ith a distinct apical tu ft o f lan ate o r straight hairs, ciliate with long stout sim ple to plum ose crooked or straight hairs which are eglandular o f gland-tipped, plum e branches long an d spreading or short and erect, sometimes clo th ed w ith sim ilar hairs on the abaxial surface, the apex devoid o f cilia o r ciliate, the cilia as long as, m ostly longer th an , the w idth o f the sepals.Corolla 4-lobed, 2 ,5 -7 m m long, constricted in the m iddle to tw o -th ird s o f the way up, som etim es inconspicuously so, inflated below in the lower part and often 4-angled, the angles altern atin g w ith the sepals, cyathiform above the constriction, pubescent outside m ainly in the m iddle region, pubescent to pilose, rarely su b g lab ro u s inside aro u n d the constric tion; lobes erect to slightly spreading, broadly to narrow ly deltoid, sm o o th to slightly crenulate, obtuse, occasionally em arginate.Stam ens 4, free; filam ents mostly linear, expanded at the apex a t the po in t o f attachm ent to th e an th er, sparsely pilose to villous; anthers m anifest, b ip artite, 0 ,8 -1 ,5 m m long, m ostly oblong, scabrid to long scabrid, m uticous or aristate; awns up to h a lf th e length o f the cell; pore up to half the length o f the cell; pollen grains single.Ovary 2-celled w ith a single ovule in each cell, m ostly compressed, ovoid to oblate, obtuse, glabrous to pilose at the apex; style filiform , glabrous, far exserted; stigma subsim ple to capitellate.Figs 9-16.
A species form ing low com pact sem ispreading to erect shrublets up to 0 ,5 m occuring in sandy areas in m ountains betw een Porterville an d N iew oudtville in the w estern C ape, flowering from A ugust to November.
G. ciliaris is characterized by having the calyx segments free o r only very slightly jo in ed at the base, the corolla-tube m ore o r less distinctly constricted in the m iddle an d the cilia on the calyx longer than the w idth o f the sepals.
G. ciliaris is one o f the oldest described species among the m inor genera o f the Ericoideae.Strangely the species is very isolated and far-rem oved from Cape Tow n w here o th er species m ore accessible existed, but were overlooked fo r so long.Despite its long standing, the species has been very much confused until now.L innaeus, the younger, stated in the protologue th a t the species had 3-nate leaves based undoubtedly on a T hunberg specim en.T h u n berg him self later published a fuller description from his own specim en stating th a t the leaves were 4-nate.This error was subsequently repeated by num erous authors until R ach (1853) corrected this.
A sim ilar situ atio n exists with G. ciliaris as occurs in G. plumosa.In F lo ra Capensis Brow n recognized six species w hich he grouped on the ch aracter o f a It w ould app ear th a t we are dealing w ith an aggre gate species o f spatially separated noninterbreeding populations which are in the first stages o f evolving into a num ber o f distinct entities which may eventually becom e sufficiently distinct to be regarded as separate species.A t present, sim ilarities are too close to justify this latter classification.
A n im p o rtan t feature and character o f use in delim iting the subspecies is the nature o f the cilia on the calyx, som ething which is easily observable an d yet som ew hat difficult to define (Fig. 11) p a rti cularly in regard to the plum e sidebranches.

3-
ssp involuta ssp bolusii ssp ciliaris ssp ciliciiflora ssp multiglandulosa 1 2 B ranches pubescent w ith retrorse simple hairs, som etim es arachnoid.Leaves adpressed, 1 -2 ,5 m m long, narrow ly ovate to oblong to obovate, hairy w ith dense crisped retrorse hairs, often becom ing glabrous on the abaxial surface.Bracteoles equal to slightly unequal, m edian to rem ote, often recurved, the m edian 1 ,3 -1 ,8 x 0 ,4 5 -0 ,8 mm, narrow ly ovate to elliptic to oblong with a relatively large keel-tip and no m arkedly expanded base, m ostly pilose w ith crisped hairs, very rarely ciliate w ith a few sh ort sto ut hairs; pedicel up to 1,5 mm long.Sepals 1 ,5 -2 ,2 x 0 ,4 -1 1 m m, narrow ly oblong to oblong to oblong-ovate, often with a swollen knob-like apex, very slightly keel-tipped, pilose at the base and som etim es in the upper half, the apex clothed with a tu ft o f lanate hairs, ciliate with simple to plum ose sto ut hairs with irregular short and long spreading plum e branches, eglandular or gland-tipped, cilia often crooked, as long as or longer than the w idth o f the sepal, occasionally with some sim ilar hairs on the abaxial surface, the apex usually devoid o f cilia, rarely w ith cilia.Corolla up to 3 ,5 mm long, distinctly or indistinctly constricted in the m iddle, the constriction som etim es n o t visible in dried m aterial, pubescent outside in the m iddle region, pilose to alm ost glabrous inside around the constric tion.Anthers 0 ,8 -0 ,9 m m long, scabrous, aristate rarely m uticous; awns up to half the length o f the cell.Ovary glabrous to pilose at the apex.Fig. 13.This subspecies is distinguished by the can o p ubescent leaves, the hairs being crisped an d n o t glandular, the crisped pubescence on the bracteoles, w ith rarely short stout hairs, an d the relatively sh o rt calyx hairs, the hairs being sim ple and straig h t to crooked and irregularly plum ose w ith spreading branches (Fig. 12).
V ariation within the subspecies occurs in the w idth o f the sepals where the b ro ad form (Lavis 19811) merges into w hat was G. dregeana recorded as a single collection from the G ifberg.Sim ilarly variation in the pubescence on the ovary apex and the presence o r absence o f a n th e r awns provided a g rad atio n with G. dregeana.Branches puberulous to pilose w ith simple hairs, som etim es w ith sh o rt sto u t plum ose hairs adm ixed.Leaves extrem ely variable, 1 ,2 -3 ,5 mm, linearlanceolate to ovate or obovate, pubescent, sometimes with dense crisped hairs o r alm o st lanate becom ing glabrous, ciliate with sto u t plum ose hairs or ju s, com pound tufts, rarely only pubescent, cilia variablet m ostly straight w ith few to m any long spreading plum e branches, eglandular, often falling off and rem aining as stubs.Bracteoles sub-equal to very unequal, adpressed to the calyx, the m edian elliptic to broadly elliptic o r ovate, 2 , 0 -3 , 2 x 1 ,0 -1 ,7 mm, with broad flat base and relatively sm all but distinct keel-tip, the laterals oblong-elliptic to obovate som etim es oblique, glabrous to puberulous, ciliate in the upper half w ith stout plum ose hairs.Sepals 2 ,1 -3 ,0 x 0 ,9 -2 0 m m , oblong-elliptic to very broadly elliptic, pubescent rarely subglabrous, ciliate with short to long sto u t plum ose hairs, rarely subplum ose, eglandular, plum e branches spreading, relatively long, rarely sh o rt and erect, w ith sim ilar hairs on the abaxial surface, ciliate at the apex and w ith an apical tu ft o f straig h t not lanate hairs.Corolla 3-4 m m long, pubescent to pilose inside and outside in the middle region.Anthers m uticous rarely m inutely awned.Ovary glabrous.This subspecies is distinguished by having a large median bracteole m ore th a n 2 x 1 mm but less th an 3,2 mm long w ith a b ro ad base an d relatively small keel-tip and w ith m arginal cilia, leaves m ostly ciliate with stout plum ose hairs at least when young, sepals less th an 3 m m long w ith very plum ose sto u t hairs with spreading branches.This is a very variable taxon particularly as to the leaves, som e being close to subsp.ciliaris w ith the short crisped in d u m en tu m an d only a few tufted cilia.The m ajority o f specim ens has awnless anthers whereas those o f subsp.ciliaris are m ostly awned except in the so u th ern populations.The bracteoles and leaves w ith distinct cilia serve to distinguish subsp.bolusii.T he larger linear-leaved form tends tow ards subsp.involuta.A superficial sim ilarity exists between subsp.bolusii and G. plumosa subsp.pentheri which occurs on the west side o f the O lifants River valley.The free, as opposed to jo in e d , sepals with num erous o r few abaxial hairs serve to distinguish the two taxa.Branches pubescent with sim ple hairs.Leaves adpressed, up to 4 ,5 mm long, m ostly lanceolate, straight, sparsely p uberulous becom ing glabrous, ciliate with sh o rt sto u t plum ose hairs w hich becom e setae.Bracteoles unequal, m edian, adpressed to the calyx, the m edian 4 -5 x 1 , 7 -2 , 3 mm long elliptic to ovate with an expanded flat base and distinct keel-tip, the laterals a b o u t 3 m m long oblong w ith a slight keel-tip, all g labrous except for a few hairs on the keel-tip, ciliate with long slightly plum ose straight hairs, the plum e branches very small and pointing tow ards the apex o f the cilium.Sepals 4 , 3 x 1 , 9 -2 , 3 mm broadly elliptic, slightly keeltipped, glabrous, with apical tuft o f short straig h t hairs, ciliate with long straight slightly plum ose hairs, plum e branches very small and pointing tow ards the apex o f the cilium.Corolla 6-7 mm long, constricted tw o-thirds o f the way up, pubescent in the m iddle region outside, villous inside at the constriction.Anthers ab o u t 1,5 mm long, m uticous, scabrous.Ovary glabrous.This subspecies is characterized by its overall larger flowers and inflorescence, the bracteoles being longer th an 4 ,2 mm and the sepals longer th a n 4 mm, b oth w ith a broad base and relatively small keel-tip.
The larger size is the only differentiating c h aracter betw een this subspecies and some form s o f subsp.bolusii.It also has close sim ilarities with som e form s o f subsp.ciliciiflora but, again, the size difference is pronounced and the bracteole shape slightly different.The sepal cilia (Fig. 12) are m ore closely related to those o f subsp.ciliciiflora th a n those o f subsp.bolusii.
Subsp.involuta is very restricted in its d istrib u tio n possibly occurring in only one or two p o p u lations on the western side o f the K rakadouw range.N o recent collections have been made.These populations are allopatric to those o f subsp.bolusii and subsp.ciliciiflora m aking interchange o f genetic m aterial highly im probable.Branches pubescent with erect to retrorse sh o rt sim ple hairs.Leaves adpressed, 1 ,5 -3 ,0 mm long, linear to ovate, m ostly pubescent w ith adpressed crisped hairs becom ing som ew hat glabrous on the abaxial surface, rarely glabrous when young, occasion ally ciliate with short stout gland-tipped hairs or ju st apiculate.Bracteoles rem ote or m edian, unequal to subequal, slightly recurved, 1 ,4 -2 ,3 x 0 ,5 -0 ,8 mm , oblong to elliptic, the laterals linear to linearelliptic, with a distinct keel-tip and flat base, pubescent to subglabrous but with an apical tu ft o f crisped hairs, ciliate with long stout subplum ose h airs; pedicel long pilose.Sepals 1,6 -2 ,5 x 0 ,6 -1 ,3 m m , oblong to broadly elliptic with a slight keel-tip pubescent to glabrous with or w ithout an apical tuft o f straight hairs, ciliate with long straight plum ose hairs, plum e branches small forw ard pointing, rarely subplum ose, rarely gland-tipped.Corolla 2 ,5 -3 ,0 mm long, distinctly constricted and 4-angled T his subspecies is characterized by the small usually subequal narrow -based bracteoles w ith co n spicuous long plum ose cilia and by the leaves, when glandular, with the glands apical or m arginal only and usually short-stalked to sessile and by the sepals with long plum ose hairs having very sm all erect branches.
Relationships with the oth er subspecies are in three directions and are som ew hat difficult to explain in the case o f subsp.ciliaris due to the geographical isolation o f the latter.The relationships with subsp.involuta and subsp.multiglandulosa are understandable due to the reasonably close proxim ity o f the p o p u la tions.
T here is considerable variation w ithin this su b species which necessitated the inclusion o f G. apiculata and G. zeyheriana in the synonom y.The typical form o f subsp.ciliciiflora possesses leaves with a crisped, som etim es retrorse, indum entum and calyx with num erous long sparsely plum ose hairs with plume branches forw ard pointing.In some form s the leaves possess sessile or subsessile m arginal glands and a large apical gland.Branches puberulous to subglabrous with stouter gland-tipped hairs adm ixed.Leaves recurved-spreading sometimes straight and adpressed, 1 ,5 -4 ,5 mm long with petiole 0 ,5 mm long, lanceolate to ovate m ostly glabrous and shiny on the abaxial surface and sparsely puberulous on the adaxial surface, rarely entirely glabrous, occasionally puberulous all over when young, ciliate and clothed on the abaxial surface with short to long stout simple gland-tipped hairs, sometimes those on abaxial surface falling off in erect leaves.Bracteoles m edian to rem ote, subequal to unequal, 0 ,8 -2 ,5 mm long the laterals mostly 1,0 mm long, linear to oblong-ovate to oblong-elliptic with an enlarged keel-tip, glabrous to pilose in the lower half, som etim es crisped at the apex, ciliate with stout gland-tipped simple hairs with a few on the keel-tip; pedicel up to 2 ,5 mm long, sparsely puberu lous with simple and gland-tipped hairs.Sepals 1 ,9 -2 ,5 x 0 ,5 -0 ,8 mm oblong to oblong-ovate with a slight keel-tip, glabrous, rarely with a few scattered hairs, ciliate with long straight subplum ose to simple hairs with sim ilar hairs on the abaxial surface, hairs often gland-tipped.Corolla ab o u t 4 ,5 mm long, distinctly 4-angled, pubescent below the con striction som etim es sparsely so and confined to the angles, villous inside.Anthers ab o u t 0 ,8 mm long, scabrous, m uticous.Ovary glabrous.Fig. 16 This subspecies m ay easily be recognized by its leaves w hich are erect to recurved-spreading m ostly glabrous b u t w ith distinct long gland-tipped hairs on the m argins an d abaxial surface, by its small bracteoles up to 1 , 8 x 1 , 0 m m which have stout subplum ose to sim ple hairs on the m argins and abaxial surface o f th e keel-tip and by the sim ple to subplum ose eg landular o r gland-tipped hairs on the sepals.
The m aterial available varies som ew hat in floral and foliage characters.T he leaves are alw ays glandciliate w ith long sto u t hairs on the m argins and abaxial surface an d are m ostly distinctly recurvedspreading.But a few specim ens have erect adpressed leaves like those o f subsp.ciliciiflora.The calyx cilia m ay be sim ple o r occasionally plum ose with plume branches like those in subsp.ciliciiflora.It was found th a t the only distinguishing ch aracter is the presence o f abaxial hairs on the leaves and bracteoles in subsp.multiglandulosa.Small com pact shrublets to 30 cm high.Branches pubescent to tom entose w ith reflexed hairs, occa sionally with sto u t plum ose to gland-tipped sub plum ose hairs in between.Leaves 3-nate adpressed, 1 ,5 -2 ,0 m m long, elliptic to oblong-obovate, p u b e scent becom ing glabrous on the abaxial surface, ciliate with a few very short stout plum ose hairs or w ith short stout gland-tipped hairs; petiole very short, pubescent, som etim es with gland-tipped hairs.Flowers in small term inal heads of 3-6 (9) on the ends o f lateral branchlets, pink, occasionally w hite; pedicel up to 1 ,0 m m long pubescent; bracteoles subequal to unequal, m edian b u t adpressed, 1 ,0 -1 ,4 m m long, narrow ly oblong to elliptic oblong often w ith an enlarged keel-tip, acute or obtuse, the laterals linear, pubescent, ciliate with short stout plum ose hairs or subplum ose gland-tipped hairs.C alyx 4 -p artite; lobes 1 ,2 -1 ,8 x O ,3 ^),6 5 m m , m ostly narrow ly oblong, occasionally elliptic-oblong or linear, acute, pubescent w ith longer straight hairs at the apex, ciliate w ith stout plum ose eglandular hairs to ciliate w ith sto u t subsim ple gland-tipped hairs, all shorter th an the w idth o f the lobe, som e tim es w ith sim ilar b u t shorter hairs on the abaxial surface.Corolla 2 ,4 -2 ,7 mm long, constricted at the m iddle, ellipsoid below, urceolate above, pilose to villous in the m iddle region and slightly up the back o f the lobes, pilose inside around the co n stric tio n ; lobes very broad, obtuse, erect-spreading.Stam ens 8, free; filam ents linear, m uch dilated a t the p o int o f attachm ent, sparsely pilose; an th ers m anifest, ab o u t 0 ,7 mm long with oblong parallel to spreading cells, scabrid edged, aristate; aw ns sm all to obsolete, arising from the apex o f the fila m ents, scabrid; pore relatively small ab o u t one q u arte r the length o f the cell.Ovary 2-celled with a single pendulous ovule in each cell, com pressed, broadly ovoid, pubescent on to p and seated on a distinct nectariferous disc; style filiform, glabrous, far exserted; stigm a sim ple to capitellate.Fruit a hard verrucose nut.G. incana may be distinguished from related taxa by its small flowers, sm all narrow sepals less than 2 ,0 x 0 ,6 5 mm w hich have slender straight cilia as long as but m ostly shorter th an the w idth o f the sepal and by the straight hairs form ing the apical tuft on the bracteoles and sepals.
The species affords a good exam ple o f geographical vicarism with its closely related species which are well separated spatially, e.g.G. ciliaris subsp.ciliaris, G. rigida an d G. nivenii.
Difficulty was experienced in distinguishing G. incana from G. ciliaris subsp.ciliaris.The form er occurs only on the sandy coastal flats adjacent to the Cape Peninsula, w hereas the latter is confined to the sum m its o f m ountains a ro u n d V anrhynsdorp and N iew oudtville.
N. E. Brow n unfortunately m isinterpreted the corolla shape in G. ciliaris subsp.ciliaris and so isolated it from G. incana in his revision.Several characters were exam ined in detail and found to have a certain degree o f disjunction and, when used in com bination, served to distinguish the two taxa.In G. incana the leaves possess hairs which are m ostly erect as opposed to the crisped retrorse hairs in G. ciliaris subsp.ciliaris.The leaves are usually edged with short stout plum ose hairs or gland-tipped hairs, whereas in G. ciliaris subsp.ciliaris this rarely occurs.The calyx in G. incana is m ostly pilose with a tuft o f longer straight hairs at the apex and w ith cilia shorter than the w idth o f the sepal.In G. ciliaris subsp.ciliaris the calyx is very sparsely puberulous with a distinct apical tuft o f long interwoven crisped hairs and with cilia longer th a n the w idth o f the sepal.
To the east there occur two closely related species, G. rigida and G. nivenii, both in restricted separate areas.G. incana differs from b oth these species in the size o f the sepals, which are less th an 2 ,0 x 0 ,6 5 mm, and in the texture and indum entum o f the leaves.In the glandular form o f G. incana the leaves are very sim ilar to those in G. rigida but are not so inflated, are m ore pubescent and have the glands confined to the m argins.
The sepals in this species are the sm allest and narrow est in the genus; in one specim en being only 0 ,3 mm wide.T his feature m akes the corollas m ore easily visible th a n in o th er species.The anthers are unique in the genus in having the sm allest pores relative to the size o f the cell.
Two fairly distinct form s occur in the m aterial so far collected.The specim ens from the north around M am re have bracteoles an d sepals with m ore plum ose cilia w hich are eglandular.Those from the south in the K raaifontein area have sub plum ose gland-tipped hairs on the sepals, bracteoles and on the leaves.This variation is, however, clinal with no distinct disjunction betw een the tw o extremes.
N. E. Brow n described G. alba from a single collection m ade by A dm iral G rey an d based it on the single ch aracter o f white flowers including the anthers.W hite-flow ered form s o f G. incana have been collected, b u t these have h ad pale brow n anthers.In all o th er characters, G. alba is identical to the glandular form o f G. incana and is presum ed to be only an a b e rra n t albino o f this species.
G. incana is fairly restricted in its distribution occurring only on the recent sand deposits on the coastal flats a t Sir Low ry's Pass, Eerste River, K raai fontein, below T ygerberg and near M elkboschstrand.The records from Sim onstow n, K leinm ond, du T o it's K lo o f an d Vogelvlei are very doubtful.

4.
Grisebachia rigida N .E .Br. in FI. C ap. 4 , 1 :343 (1906) Shrublets com p act and low to erect up to 50 cm high.Branches pubescent with sim ple recurved hairs, very occasionally gland-tipped, rarely w ith stout plum ose hairs adm ixed.Leaves 3-nate up to 3 ,O x 1,0 m m , m ostly elliptic to narrow ly elliptic, occasionally ovate o r oblong-obovate, thick and fattened, pubescent or m inutely scabrous on the abaxial surface, rarely glabrous and shiny, pubescent on adaxial surface, ciliate with 7-9 short sto u t gland-tipped hairs and with some scattered over the abaxial surface, petiole very short, shortly g landular pubescent.Flowers 1-8-nate on the ends o f lateral branchlets, pink, rarely w hite; pedicels a b o u t 1 mm long, pubescent with some plum ose hairs a t the apex; bracteoles equal to slightly unequal with the m edian slightly broader, adpressed to the calyx, ovate to oblong-ovate to elliptic to narrow oblong-elliptic, obtuse, glabrous or sparsely pubescent, ciliate w ith stout simple to very slightly plum ose hairs which are m ostly gland-tipped, rarely sparsely pilose inside, keel-tipped.C alyx 4-lobed, slightly joined at the base; lobes ovate-elliptic to oblong-elliptic to broadly elliptic up to 2 , 1 -2 , 8 x O ,9 -2 ,0 mm, often w ith incurved m argins, subacute, keel-tipped, glabrous or sparsely pubescent m ostly in the lower half, ciliate with broadly based stout hairs alm ost fim briate in places an d with sim ilar hairs up the centre o f the abaxial surface, hairs m ostly simple or very slightly plum ose, rarely gland-tipped, often crooked.Corolla up to 4 ,4 mm long often oblique, distinctly c o n stricted in the m iddle; tube up to 3 mm long globose ellipsoid, spreading above the constriction, 4-angled, puberulous outside w ith glabrous patches opposite the sepals, pilose on the inside mainly at the p o in t o f co n striction; lobes erect-spreading slightly crenulate and em arginate a b o u t 1 mm long, very b ro adly obtuse, pubescent at the base in the middle.Stam ens 4; filam ents linear, sparsely to densely pilose, up to 2 ,5 m m long; anthers m anifest, attached dorsally one th ird the way up, variable in size, 0 ,7 -1 ,1 mm long w ith oblong to obovate parallel or spreading cells, alm ost glabrous to scabrous, occasionally with som e long tra n sp aren t hairs on the edges, m uticous \ v ' " S/J (I/ </ \ \ \ s ■ / or arista te ; awns up to 0 ,4 m m long o r one th ird the length o f the cell, arising from the filam ent apex, spreading laterally to descending, m inutely scabrous; pollen grains single.Ovary 2-celled w ith a single pendulous ovule in each cell, 0 ,6 x 0 ,8 -0 ,9 x 0 ,9 m m , broadly ovoid to ellipsoid, com pressed, obtuse, variously pilose at the apex, very unevenly w rinkled; style up to 4 mm long, glabrous, exserted; stigm a slightly capitellate.Fruit a h ard verrucose n u t.G. rigida is characterized by its free o r slightly jo in ed sepals which are m ore th a n 2 , 0 x 0 , 6 5 m m w ith very bro ad flattened subplum ose o r sim ple cilia and by its fattened leaves which are ciliate w ith short gland-tipped hairs and w ith a few sim ilar hairs on the abaxial surface.
The species differs from G. nivenii in the leaf cilia, less plum ose calyx and glabrous inner surface o f the sepals.It is closely related to G. incana from which it is easily distinguished by its b ro ad er sepals w ith their broad cilia and by the leaves.
G. rigida varies in the size o f the sepals w here in the type, Bolus 5193, they m ay be as m uch as 2 ,7 x 2 , 0 mm.The anthers also vary in size, shape an d in the occurrence o f awns.A few specim ens have an th ers w ith long colourless hairs, a feature very rarely seen in the Ericoideae.
The species occurs on the recent sandy alluvial flats at the eastern base o f the Stettyns M ountains betw een W orcester and V illiersdorp where isolated pockets o f fynbos grow (Fig. 20).T he surrounding area possesses m ountain renosterveld on the shales and W ittenberg quartzites.In this area the species is very susceptible to extinction due to encroaching agriculture and burning.B olus's record between French H oek an d Villiers d o rp has not been reconfirm ed.A lthough som ew hat rem oved from the m ain populations an d com ing from a com pletely different valley system, this record could be correct due to the num erous sandy alluvial patches in the area.

5.
Grisebachia nivenii N .E .Br. in FI. Cap. 4 , 1 :343 (1906).Shrublets com pact, erect, up to 50 cm high.Branches m inutely pubescent with retrorse hairs.Leaves 3-nate, adpressed, up to 1 ,7 x 1 , 2 m m , broadly ovate to elliptic, very rounded and thick, obtuse or acute, glabrous on the abaxial surface, pubescent on the adaxial surface, ciliate w ith 5-6 sh o rt sto ut plum ose cilia; petiole very sh o rt and bro ad , ciliate.Flowers in term inal globose heads o f 3-8 on the ends o f lateral branchlets, pink, rarely w hite; pedice's very short, sparsely pubescent, som etim es w ith sto u t plum ose hairs a t the apex; bracteoles adpressed, about 1,3 m m long, subequal, rarely m arkedly unequal, mostly oblong-elliptic som etim es the laterals obliquely so, the m edian rarely angular-ovate, all keel-tipped, obtuse or acute, puberulous outside and inside, ciliate w ith stout plum ose hairs.C alyx 4-lobed, sometimes slightly joined a t the base; lobes broadly elliptic to narrow ly oblong-elliptic, 2 ,1 -2 ,9 x 0 ,6 5 -1 ,3 mm, acute, keel-tipped, puberulous on the abaxial surface m ainly tow ards the base, sparsely puberulous on the inside, ciliate with sto u t b ro a d plum ose hairs with sim ilar hairs over the adaxial surface tow ards the centre, rarely gland-tipped.Corolla up to 4 ,0 mm long, som etim es oblique, distinctly constricted in the m iddle; tube up to 3 m m long, globose ovoid to ellipsoid, urceolate above the constriction, pube scent to pilose in the m iddle an d lower p a rt and up the back o f the lobes and inside the tube a ro u n d the constriction; lobes broadly obtuse, erect or slightly spreading.Stam ens 4; filam ents linear, sparsely to densely pilose; an th ers m anifest, a b o u t 0 ,8 mm long, oblong, dorsifixed one th ird o f the way up, papillate, aristate; pore a b o u t a th ird o f the length o f the cell; awns sm all, spreading to deflexed, arising from the filam ent at the p o in t o f attach m en t to the an th e r; pollen grains single.Ovary 2-celled with a single pendulous ovule in each cell, 0 ,7 -0 ,6 mm, ovoid com pressed, obtuse sparsely pilose at the apex; style exserted, up to 4 mm long; stigm a capitel late or subsim ple.Fruit verrucose, hard.abaxial surface an d have 5 -6 very sh o rt stout plum ose cilia an d by its m ore o r less free sepals which have short stout flattened plum ose cilia.
The species is closely allied to G. rigida and G. incana.It differs from G. rigida in having glabrous shiny leaves, even w hen young, w ith plum ose cilia and no gland-tipped cilia.The calyx cilia are m ore plu mose th an in G. rigida and there is a sparse pubescence on the inside o f th e sepals.F ro m G. incana it differs in leaf details, the la tte r having pubescent nonfattened leaves.The sepals in G. nivenii are bro ad er, m ore than 2 ,0 x 0 ,6 5 m m .G. nivenii is geographically isolated as it occurs only on the sandy flats in the hilly co u n try so u th east o f Sw ellendam w here it is far rem oved from its closest allies, G. rigida an d G. incana.Like nearly all the other species in th e genus it is confined to a few small patches o f alluvial sand.T he m ain m orphological difference between these tw o tax a lies in the form o f the inflorescence.In A the flowers are generally 1-4 -n a te a t the ends o f sh o rt axillary branchlets w hich are often clustered together in a pseudospike along the m ain an d lateral branches.In B the flowers are term inal on the ends o f sh o rt branchlets w ith up to 36 flowers form ing a cluster.These clusters usually hang dow nw ards m aking the plan ts less conspicuous th an those of tax o n A. Except for one vicariad, plants o f taxon B are glan d u la r particularly on the m argins o f the calyx.T he glands term inate plum ose cilia.In taxon A the few glands are confined to the short simple cilia on the calyx.
T aking into account the hab itat and m orphological differences, I have decided to regard the tw o tax a A & B as closely related species referred to G. parvi flo ra and G. minutiflora respectively and to recognize several subspecific taxa.
Low com pact to spreading shrublets up to 20 cm high, rarely up to 50 cm.Branches erect or spreading often entw ining am ong the surrounding vegetation, w ith num erous short branchlets, pubescent som etim es w ith glands adm ixed.Leaves 3-nate up to 3 m m long w ith the petiole, erect to spreading, straight to m arkedly recurved, linear to lanceolate rarely ovate, acute, flat to trigonous, glabrous or a t first puberulous, ciliate som etim es with gland cilia or sessile glands, som etim es glan d -ap icu late: petiole adpressed ciliate.
Flowers 1-^-nate at the ends o f extrem ely short branchlets arranged in a spike-like m anner along the branches; pedicels very short, less th a n 0 ,5 mm long; bracteoles 3, equal to subequal, adpressed to the calyx or slightly spreading, very variable in size, up to 1,5 m m long, in shape from linear to elliptic-oblong to ovate, acute to obtuse, glabrous or pubescent ciliate w ith o r w ithout sessile o r stalked glands white.C alyx 4-lobed for its length, cam panulate 0 ,6 -1 ,8 mm long an d 0 ,3 -0 ,8 m m wide, glabrous or pubescent, w hite; lobes erect variable in shape from elliptic-oblong to ovate to su b q uad rate, the apex acute cuspidate or su btruncate with an apiculus, with or w ithout a distinct keel-tip and w ith or w ithout a distinct m edian ridge, ciliate, with or w ithout sessile o r stalked glands.Corolla 4-lobed, up to 3 x 1 , 7 mm long, m ostly 2 x 1 , 2 m m, funnel-shaped sometim es broadening m ore above the m iddle, nowhere constricted, occasionally tubularellipsoid; tube glabrous inside som etim es glabrous outside o r puberulous in lower half to fully pubescent; lobes erect to incurved, rarely slightly spreading, obtuse, broader th an long, glabrous entire.Stam ens 4 included or m anifest; filam ents filiform g lab ro u s; anthers up to 1,1 mm long w ith oblong parallel separate cells, basal, m inutely scabrous, a ristate; awns the length o f the cell to rud im en tary , ciliate; pore the length o f the cell, pollen grains single.Ovary 2-celled with a single ovule per cell in one subspecies, rarely 2 ovules per cell and som etim es 3 cells, ovoid to ellipsoid, up to 1 x 0 , 7 m m , obtuse glabrous to puberulous on the top, som etim es thickly pubescent; style far exserted, straight or curved, glabrous up to 4 mm long; stigm a m inutely capitate.Figs 22 & 23.
A species generally low, sparse an d spraw ling in habit, occasionally com pact, occurring frequently on dry stony slopes and flats on m ountains from the C edarberg to D u T o it's K lo o f eastw ards to near Swellendam, flowering from as early as July to as late as January.
A very variable taxon in which three subspecies are recognized.
G. parviflora is characterized by the flowers being 1-4 -n a te in small heads clustered along the branches in a congested spike-like m anner, the corolla tube n ot contracted in the middle and the simple cilia on the calyx lobes.The species is closely allied to G. minutiflora and has a rem arkable superficial resem blance to Eremia curvistyla (N .E. Br.) E. G. H. Oliver differing basically in the num ber o f stam ens.All three species are sym patric.
The true identity o f this species has been over looked until now as it has nearly always been referred to the later G. eremioides M acO w an.K lotzsch in describing it placed the species in D o n 's genus Eremia using as his type an Ecklon & Zeyher collection from " Hills between Puspas Valley and K ogm ansk loof M o u n tain s" .W ithout seeing the type, N. E. Brown followed K lotzsch in keeping the species in Eremia.But later in his w ork on Grise bachia he stated th a t he had seen the type and found th at the species was conspecific with G. eremioides which he proceeded to retain under the Kew Rule.This was picked up by D ruce in his search th ro u g h F lo ra Capensis for new com binations but never applied by subsequent botanists.
U nfortunately the holotype in Berlin is no longer extant and all the Ecklon & Zeyher m aterial distributed as Eremia parviflora K lotzsch tu rn s out to be Anomalanthus scoparius K lotzsch.N. E. Brown on exam ining the type sent to him on loan stated " The description o f K lotzsch is very erroneous, as the calyx is n o t subequal to the corolla, but considerably shorter th a n it, and the stam ens are 4, not 8 as K lotzsch states.It is identical w ith Z eyher 1117, except th at the leaves are straighter, like those o f Schlechter 10091" .This statem ent clears up the discrepancies in the type description an d also paves the way for the typification o f the species.W ithout any authentic duplicate m aterial available, I consider th a t it is justifiable to rely on N. E. B row n's com parison and therefore select Zeyher 1117 in the herbarium at Kew as the neotype.The above published note by N. E. Brown also appears on the sheet a t Kew.
Five duplicates o f Zeyher 1117 are located in various h erbaria an d all com e from " Flats between the W itsenberg and Skurfdeberg" .The locality o f H ouw H oek on one o f the sheets in SAM as cited by M acO w an in his protologue is erroneous.
To date no additional m aterial o f G. parviflora has been collected in the sam e area as th a t visited by Ecklon & Zeyher.But in the variatio n pattern, I have found in the calyx th a t the broadly elliptic-acum inate lobes o f Zeyher 1117 could well have occurred in the holotype.The leaf difference noted by N. E. Brown is not vitally significant as variatio n in this ch aracter has been noted even on the same plant.N .E. Brown described (b) an d (c) w ith their narrow elliptic-oblong acute lobes in com bination with the possession o f erect straight leaves as a separate variety, var.grata.I have found th a t neither the calyx lobe shape n o r the leaf arran g em en t show any significant differences.There is a definite intergrading between the broad based apiculate and the narrow elliptic-oblong acute sepals as seen in Compton 6624 from R oodesandberg, Stokoe 6067 from the W ellington m ountains and Esterhuysen 11057 from Stettynsberg.In leaf arrangem ent variatio n from erect straight leaves through to curved slightly spreading leaves can occur on the same plant.T here were therefore no grounds for keeping the tw o species separate.Small shrublet, com pact to spreading.Leaves stiffly trigonous, m arkedly spreading-recurved when m ature.C alyx lobes sub q u ad rate, subtruncate with a thickened apiculus, rarely very broadly angularovate, closely ciliate w ith sh o rt simple hairs occasionally with a few subsessile glands, otherw ise glabrous, rarely sparsely puberulous.Corolla glabrous, rarely sparsely puberulous.Ovary thickly pubescent on the upper half.There is, then, in the C edarberg a reasonable discontinuity in several characters coupled w ith a spatial separation.This should w arrant recognition at specific level, but as there are definite similarities with certain elem ents to the south, I feel th a t recognition is only justifiable a t subspecific level.
On two collections N. E. Brow n described this vicariad as var.eglandula.A n exam ination o f all the collections showed th at sm all subsessile glands are present on the m argins o f the calyx.Low com pact to sem i-spreading shrublet up to 20 cm high.Branches long when spreading and som etim es rooting a t the nodes, pubescent, w ith glandular hairs interm ingled when young, 3-angled when young.Leaves 3-nate, up to 3 ,5 m m long, the petiole 0 ,5 m m long, ovate to narrow ly oblong, straight erect or slightly spreading, im bricate or sho rter th a n the internodes, subobtuse, thick, pubescent o r glabrous w hen young w ith a few to num erous sessile glands on the m argins, becom ing glabrous except on the adaxial surface, sometimes term inating in a sessile gland.Flowers in term inal globose heads o f up to 36 flowers on sh o rt branchlets, not form ing congested spikes, w hite; pedicels alm ost none up to 0 ,8 m m long; bracteoles 3, adpressed or recurvedspreading, equal or very unequal in som e o u ter flowers in inflorescences, m ostly 0 ,5 -1 ,7 m m long, if equal th en linear acute or subacute and m inutely keel-tipped, if unequal then the m edian one large an d leaflike an d well keeled, glabrous or pubescent, ciliate tow ards the base with simple hairs and a m ixture o f sim ple and larger plum ose and glandtipped hairs tow ards the apex.C alyx 4-lobed from 4 its length, up to 1,9 mm long, pubescent, the pubescence sh o rt over the whole surface or in zones w ith som etim es longish hairs; tube obconic o r tu b u lar w ith spreading lobes; lobes ovate-oblong to very broadly ovate up to 0 ,9 mm long and 1 m m bro ad , erect or spreading, som etim es with pubescence on the inside a t the top, ciliate w ith sh o rt o r long cilia w hich are sim ple o r variously plum ose fro m base to apex, m ostly gland-tipped, keel-tipped an d thickened, acute to subobtuse.Leaves pubescent w hen young, occasionally w ith a few sessile glands on the m argins.Pedicels up to 0 ,8 mm long; bracteoles usually adpressed and recurved tow ards the apex, m ostly up to 1,7 m m long.C alyx pubescent in zones or evenly w ith pubescence also on the inner surface in the u p per quarter, cilia plum ose to the apex and not glandtipped.G. minutiflora is characterized by the globose inflorescences o f 6-36 flowers scattered along the branches, the corolla-tube n o t contracted in the middle, the plum ose cilia on the calyx lobes and the general glandular condition o f the flowering branches.It is closely allied to G. parviflora.
W hen Brown revised the genus he had tw o collections o f Schlechter to exam ine and justifiably described them as two new species, G. minutiflora and G. nodiflora, basing them on the prescence or absence o f gland-tipped plum ose or simple cilia on the calyx lobes.
Since then no further m aterial referrable to G. nodiflora has been collected, but there have been 12 collections of G. minutiflora.The type o f G. nodiflora possesses no glands on the calyx lobes, whereas in the collections o f G. minutiflora there are always some glands p esent on the cilia, but these may be extremely small in some flowers.The m aterial was then exam ined for o th er characters.Brown used the differences in the size o f the inflorescence heads, but this is invalid as the heads in Oliver 4105 are m uch larger th a n in Schlechter 10188.He also used the degree o f feathering on the cilia, but this is very variable in G. minutiflora from simple to alm ost fully plumose.
Slight differences were found in the length o f the pedicel which is absen t to 0 ,4 m m long in G. m inuti flora and up to 0 ,8 m m long in G. nodiflora.The bracteoles o f the form er are usually adpressed whereas in the latter they are approxim ate b u t curvedspreading.I decided to reduce G. nodiflora to sub specific level u n d er G. minutiflora, because o f the difference in the g lan d u lar state o f the cilia and the slight discontinuities in the bracteoles and pedicel characters coupled w ith the allopatric distribution.
A t first exam in atio n I found th a t there was a distinct difference in the type o f pubescence on the calyces with G. m inutiflora having zones th a t were pubescent and glab ro u s and w ith G. nodiflora having an evenly pubescent calyx.Close exam ination o f Schlechter 10188 d u plicates show ed there to be two distinct form s. In one th e pubescence is zoned as in G. minutiflora, in th e o th e r it is evenly distributed over the calyx, a co n d itio n n o t found in G. m inuti flora.This variability suggests a closer relationship between the tw o tax a th a n was previously accorded to them .
The m aterial o n sheets o f Schlechter 10188 can easily be separated in to tw o form s, A & B, on the type o f pubescence an d the shape o f the calyx lobes.In form A the pubescence is zoned an d th e calyx lobes are oblong to elliptic-oblong acute.In form B the pubescence is denser an d evenly d istrib u ted over the calyx the lobes o f w hich are transversely broadly elliptic obtuse.These variations can only be recognised as form s at present as they presum ably cam e from one population.T his has n o t been rediscovered and until such tim e as it is, no fu rth er statu s can be given to this v ariatio n .
The specim ens exam ined in detail have been assigned to the tw o form s as follow s: N. E. Brown did n o t designate a holotype but labelled one sheet in BO L (form A only) and one in K (form A & B) as types.F rom the protologue two characters can be pinp o in ted to determ ine which form he used to describe his species, i.e. " ca yx lobes oblong, a c u te " .T hese undoubtedly refer to form A. I have th erefore chosen the Schlechter sheet labelled as the type in BO L as the lectotype.
The species was first seen by me at the Ceres W ildflower Show in O ctober 1974 w hen I was doing the nam ing o f specimens.The collector and locality unfortunately could n ot be traced.A few weeks later the m aterial collected the previous year in the Sw artruggens by D r J. M acG regor was sent to me for identification.
A n exam ination o f the above collections showed th a t they constituted a new an d very distinct species which I was n o t able to place satisfactorily in any know n genus.Follow ing D r M acG regor's directions, I visited the Sw artruggens an d located the species in three disjunct sparse populations.A range o f m aterial was collected and exam ined for variations in the critical character, the 1-celled uniovulate ovary.A n exam ination o f num erous flowers produced only a few with unequally 2-celled ovaries.
The species is rem arkably sim ilar to Eremia totta (Thunb.)G. D on in the ou tw ard appearance o f the flowers and leaves but can n o t be placed near th at species which has 8 stam ens a n d a 4-celled ovary.
Eremia has been considerably am ended to include the 1-celled Eremia curvistyla (N .E .Br.) E. G .H. Oliver but still retains the co n stant character of eight stam ens (Oliver, 1976).
To a lesser extent the species is sim ilar in outw ard appearances to Grisebachia parviflora (Fig. 22) and G. minutiflora (Fig. 24) both o f which it grew with in the Swartruggens.They all possess four stamens.U ntil this revision, all species o f Grisebachia possessed 2-celled ovaries w ith very few exceptions having 3-celled ovaries.
There were thus four ways o f dealing with the new species: (1) placing it under Eremia and having to am end the generic circum scription even fu rther to include the 4-stam ened condition, thus causing a breakdow n in the distinction between Eremia and Grisebachia' , (2) placing it under Grisebachia and am ending the generic circum scription to include this 1-celled species; (3) placing it in either Anomalanthus or Syndesm anthus, genera w ith 4 stam ens a nd a 1-celled ovary, b u t w hich have no resem blance to it; (4) describing the species as a separate m onotypic genus.
T aking into acco u n t the im plications o f the above, it was decided to b ro ad en the circum scription of Grisebachia to include the 3-celled variations p a rti cularly in G. parviflora subsp.pubescens an d the 1-celled, rarely 2-celled, condition occurring in the new species.
In the Sw artruggens G. secundiflora was found in three separate p o p u latio n s consisting o f only a few scattered plan ts each.A t the lower altitude the plants were grow ing on sandy flats w ith a p opulation o f G. minutiflora nearby.H igher up the m ou n tain they were grow ing on sandy, rocky slopes together with some plan ts o f G. parviflora subsp.pubescens.
In all cases G. secundiflora form ed decum bent yet com pact shrubs up to 0 ,5 m high a n d up to 1 m across w ith n u m erous ascending to decum bent branches.T he branches were strikingly bare and devoid o f leaves except tow ards their ends.The white conspicuous secund inflorescences were subterm inal.This co ntrasted strongly w ith the very com pact low shrublet o f G. minutiflora o r the sparse spreading procum bent to sem i-erect plants o f G. parviflora ssp.pubescens w hich were all in fruit.Some variatio n in floral characters occurs.The pubescence on the calyx may be present o r absent on different twigs in the collections Oliver 6105, 6107 and 6115 an d is present in M acGregor s.n.It is absent in Oliver 5044 from the flower show.The corolla tube in the collections o f O liver and M acG regor from the Sw artruggens are sh o rt, tu b u la r and inflated in the low er h alf w hereas in the m aterial from the flower show it is distinctly longer and narrow er.As this latter m aterial is unlocalized no further com m ents on its statu s can be given.K lotzsch w rongly ascribed the W illdenow specim en to the species which had, up until then, been cited as Blaeria ciliaris L.f.In his description he stated th a t the leaves were 4-nate, possibly repeating the slip m ade by T hunberg and copied by m ost su b sequent authors.The W illdenow specimen has in fact 3-nate leaves.R ach (1855) in exam ining K lotzsch's specim en and th a t o f T hunberg stated that they were n o t o f the sam e species.Brown noted th at T h u n b erg 's specim en was identical to the type in the L innaean H erbarium .

I
have been able to exam ine the W illdenow specim en in the Berlin H erbarium .The m aterial certainly does not belong to G. ciliaris (L.f.)K lotzsch subsp.ciliaris and is only in young bud stage from which it is n ot possible to identify it with any certainty.

Fig
Fig. 1.-Variation in the size of the median bracteole from outer to inner flowers in a single inflorescence o f Grisebachia plumosa subsp.plumosa.Drawn x l 6 from Thompson 791 (STE).

F
i g .4 .-Di s t r i b u t i o n o f Grisebachia plumosa: ® s u b s p .plumosa; © s u b s p .hirta; O s u b s p .irrorata\ % s u b s p .eciliata; (5 s u b s p .hispida; © s u b s p .pentheri.sepal, x 16, drawn from Schlechter 8480 (PRE).
This subspecies is very sim ilar to som e form s o f subsp.pentheri, som e o f w hich used to constitute part o f w hat was form erly G. pilifolia N .E .Br.In these form s the calyx and leaves are eglandular a n d the hairs very plum ose.Subsp.hispida appears to be very restricted in its distribution, occurring only on the sandy hills and flats near Paleisheuwel associated w ith dry fynbos scrub which, according to A cocks's m ap o f Veld Types (1953), is classified as True Fynbos an d n o t C oastal M acchia.(e) subsp.pentheri (Zahlbr.)E. G. H. Oliver, com b, et stat nov.G. pentheri Zahlbr. in Ann.Naturh.Mus.Wien.20:42 (1905); N.E.Br. in FI.Cap. 4 ,1 : 1128 (1909).Type: Elandsfontein, Clanwilliam, Aug. 1894, Pent her 2925 (BM!; C a p e .-3218(Clanwilliam): Uitkomst, Graafwater, 427 m, (-BA ), Compton 4945 (BOL; NBG); 4949 (BOL); Compton 6789 (NBG; STE); Compton 24218 (NBG; STE); Kanovlei, east of Graafwater, 396 m (-BA), Oliver 3869 (STE) Subsp.pentheri occurs frequently in scattered p opulations in sandy areas on the m ountains o n the west side o f the O lifants River near Clanw illiam .U nfortunately m uch o f the h ab itat o f this tax o n has been lost to farm ing practices an d all th a t rem ains is in the rocky unusable areas.The fynbos in w hich the plants grow m ay w ithout hum an a n d anim al intervention becom e quite tall an d erect plants o f this taxon have been seen up to 1 m high.(f) subsp.hirta (K lotzsch) E. G. H. Oliver, com b et stat.nov.

F
i g .9 .-Di s t r i b u t i o n o f Grisebachia ciliaris: A s u b s p .ciliaris; O s u b s p .bolusii', © s u b s p .involuta-, # s u b s p .ciliciiflora; © s u b s p .multiglandulosa.calyxdivided to the base an d m uticous anthers.He then separated the species on the nature o f the indum entum on the leaves, the sepal size an d the length and form o f the sepal hairs.These species were G. bolusii, G. apiculata, G. involuta, G. velleriflora, G. dregeana and G. zeyheriana.A seventh species, G. thunbergii (G.ciliaris), he characterized incorrectly by placing it with those species n ot having a distinctly constricted corolla.On the small am o u n t o f m aterial available to Brown the recognition o f these tax a as distinct species was feasible but n um erous subsequent collections have provided a considerable degree o f variation which broke dow n m any o f the existing discontinuities in the m edian bracteoles(Figs 10 & 11), sepal hairs (Fig.12) an d leaf glands.

Fig. 11 .
Fig. 11.-Grisebachia ciliaris, scatter diagram showing the variation in the median bracteole of the outer 3 flowers in a single inflorescence.Each dot represents the mean measurement of each inflorescence.
Compton 20884 (BOL; NBG; STE); Hutchinson 763 (BM; BOL; K; PRE); 790 m, Oliver s.n.(STE); 762 m, F ig .13.-Grisebachia ciliaris subsp.ciliaris.1, flower, x 8 ; 2, corolla, x 8 ; 3, bracteoles, x l6 : a, laterals; b, median; c, median inside view; all drawn from the fragment of the holotype (K); 4, flower, x 8 ; 5, corolla, two views, x 8 ; 6, sepal, x 16; 7, anther, side, front and back views, x 16; 8, ovary, x 16; 9, leaf, x 16; all drawn from Oliver 3860 (STE); 10, sepal, x 16; 11, ovary, x 16; both drawn from Marloth 7646 (STE); 12, median bracteoles, x 16, a from Marloth 7646 (STE) and b from Middlemost 1594 (NBG); 13, anther, back, front and side views, x 16, drawn from Drêge 7803 T h ere are three groupings o f popu latio n s w ithin the subspecies.The n o rth ern group occurs on the N iew oudtville Plateau an d has no plum es on the leaves n o r cilia on the calyx apices.The two sou th ern p o p u latio n s on the G ifberg and a t L ockenberg are interm ed iates between subsp.ciliaris and subsp.bolusii in occasionally having plum es on the cilia and a few cilia on the sepal apices.D ue to the large spatial separation, a hybrid origin is ruled out.A nother line o f relationship occurs with subsp.ciliciiflora o f the C itrusdal area.F ro m this latter it differs in the type o f plum ose sepal cilia and in having sh o rt sepal cilia.Som etim es the leaves o f these tw o subspecies are rem arkably sim ilar in having a crisped retrorse indem entum an d no cilia.G. ciliaris (G.thunbergii R ach) was incorrectly assessed by Brown, w ho ju d g ed the corollas to be w ithout any distinct constriction in the middle.This condition is ap p a re n t in the dried m aterial which, when thoroughly boiled, som etim es shows a slight constriction.However, all fresh m aterial exam ined possessed distinctly constricted corollas.This subspecies is m ost closely related to G. incana from the flats near C ape T ow n-an unusual distri butional relationship.It is distinguished by the calyx hairs being as long as o r longer th an the w idth o f the sepals and, if equal, w ith a distinct tu ft o f apical crisped hairs.(b) subsp.bolusii (N .E .Br.) E. G. H. Olive com b, et stat.nov.G. bolusii N.E.Br. in FI.Cap. 4 ,1 : 340 (1906).Type: Moun tains near Pakhuis Pass, Bolus 8681 (BOL, holo!; K!; N H!; PRE!; STE!; Z!).

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i g . 1 5 .-Grisebachiaciliaris s u b s p .ciliciiflora.1, c o r o l l a , x 8 ; 2, le a f, xl6; b o t h d r a w n f r o m Oliver 4 0 1 7 (STE); 3, la te r a l b r a c te o le , x 8 ; 4 , m e d ia n b r a c te o le , x 8 ; 5 , s e p a l, x 8 ; 6, a n t h e r , f r o n t , s id e a n d b a c k v ie w s , x 1 6 ; 7 , o v a r y , x 1 6 ; a ll d r a w n f r o m th e h o lo ty p e , Masson s .n .(BM); 8, le a f, x 16, d r a w n f r o m Stokoe in SAM 5 4 8 4 7 (STE); 9, le a f, x 16, d r a w n f r o m Schlechter 4 9 6 9 (STE).at the base, pubescent to pilose outside and inside in the middle region.Anthers a b o u t 1 m m long, m uticous long scabrous; pore h alf the length o f the cell.Ovary glabrous.Fig. 15.Compton 16128 (NBG; STE); Compton 20965 (BOL; NBG; PRE; STE); Leipoldt in BOL 21655 (BOL); Stokoe in SA M 54847 (SAM; STE); Elandskloof Pass, (-CA ), Hafstrom & Acocks 1043 (PRE; S; STE); Waterfall between Citrusdal & Elandskloof, (-CA), Stokoe 7712 (BOL; NBG; N H ; PRE); Williams sub Baker 1821 (BM); Kleinfontein east of Citrusdal 7 6 2 m, (-CA), Oliver (E; K; MO; P; PRE; STE); Allandale, south-east of Citrusdal, 5 4 8 m (-CA), Oliver 5007 (BM; G; S; STE); near Citrusdal (-CA), Rust s.n.
Subsp.ciliciiflora occurs in the C itrusdal area mostly at lower altitudes on sandy open patches and on the m ountains north-w est o f the tow n.The locality near W uppertal, Compton 24264, is unusual and inexplicable.(e) subsp.multiglandulosa E. G. H. Olive subsp.nov., similis subspecie ciliciiflorae, sed distinguitur piliis longis glandulis m arginibus et paginis abaxialibus foliorum bracteolarum que.T y p e.-C ape, O lifants River Valley above T oorgat on the farm G rootfontein, Oliver 3972 (STE, holo.!; K !; M O !; N B G !; PR E!).
Subsp.multiglandulosa is confined to the m ountains at the south ern end o f the m ain O lifants River valley w here it occurs m ainly o n sandy open flat areas.The m ajority o f p o p u latio n s is allopatric to those o f subsp.ciliciiflora, occurring at high altitude only.In the region o f P iekenierskloof there is, however, an overlap.The specim en, Levyns 1367, is an interm ediate very sim ilar to the type and only collection o f G .apiculata (subsp.ciliciiflora).U n fortunately all th e p o p u latio n s a p p ear to have been removed in this a re a by agriculture thus m aking a study o f the p o p u latio n s im possible.
Figs 17 & 18.A species form ing small com pact shrublets occurring in sandy places on the flats between Sir Low ry's Pass, K raaifo n tein and M am re, flowering early from A pril to July.
Fig. 21.A species form ing com pact erect shrublets up to 50 cm high occurring in a very restricted area o f sandy flats south-east o f Swellendam , flowering from July to Septem ber.
The surviving p o p u la tions in the area now lie w ithin the b o undaries o f the B ontebok N atio n a l Park.There is som e con fu sio n a b o u t the collector o f the lectotype o f the species.It was labelled as " C.B.S. M asson" b ut N. E. Brow n changed this to Niven.The handw riting is definitely n o t N iven's, b u t m atches th at on the type o f Erem ia brevifolia Benth.w hich Brown cited as collected by M asson.N either o f these labels exactly m atches the handw riting o f M asson in the Kew A rchives.B row n labelled this sheet as the type.V ariation w ithin th e species is very slight.The sepals in Z eyher 3330 are som ew hat n arro w er th an in the other specim ens a n d have som e gland-tipped hairs.The G. parviflora/G.minutiflora group A long the m ain ch ain o f m o u n tain s a n d high level plateaux ru n n in g in a n o rth -s o u th direction from the C edarberg th ro u g h the C old Bokkeveld to the W orcester D istrict, there is a series o f vicarious taxa.Superficially they are very sim ilar in th eir low com pact to spreading habit, small w hite subcalycine to calycine flowers, m anifest an thers and ciliate calyx lobes.One very variable taxon (A) is w idespread from the C edarberg to the m ountains south o f W orcester an d eastw ards to near Swellendam, occurring on dry stony slopes w ith short dry restiad/ericoid vegetation.It usually form s a low com pact to spraw ling sh rublet w ith branches spreading am ongst the restiads.It consists o f three distinct allopatric vicariads a n d has had the nam es G. parviflora (K lotzsch) D ruce (G .ermioides M acO w an) and G. similis N .E. Br. applied to it.The second taxon (B) is m uch m ore restricted, occurring in the central a n d southern Cold B okkeveld on dry open sandy flats an d form s a low co m p act shrublet som etim es slightly spraw ling and ro o tin g a t the nodes.T his has had the nam es G. m inuti flora N .E. Br. a n d G. nodiflora N.E.Br. applied to it.As w ith the taxon A, there are two distinct allo p atric vicariads in this taxon.T o m y know ledge the two tax a only grow in reasonably close proxim ity in the H artebeeskloof and W inkelhaak areas where I have observed them .In the form er locality the plants o f taxon A (G. parviflora) were few and were outliers of the larger po p u latio n s higher up the rocky slopes.The plants o f tax o n B (G. minutiflora) were locally com m on on open sandy patches.In the latter locality taxon B w as flowering three m onths later th an the early flow ering vicariad o f tax o n A.

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he basic v ariation occurring in G. parviflora is found in the shape o f the calyx lobes, the arran g e m ent o f the leaves and the indum entum o f the flowers, the w idest range being in the shape o f the calyx lobes.In subsp.parviflora three groups occur: (a) large lobes w ith a b ro ad elliptic base and acum inate apex, sparsely gland-ciliate o r ciliate, com m on, central to southern in distrib u tio n represented by the neotype, Zeyher 1117: (b) large lobes, narrow elliptic-oblong w ith an acute apex, distinctly gland-ciliate, less com m on an d confined to the central region o f the distributional range (W interhoek to central Cold Bokkeveld).(c) sm all lobes m ostly elliptic-oblong w ith an acute apex, n o rth ern in distrib u tio n centred on the southern an d central C edarberg.
N. E. Brown also described G. similis var.publicalyx.The collection M aguire 1778 from G ydo, presum ably from one population, possesses glabrous calyces.This also occurs in the C om pton an d Esterhuysen collections from Slab Peak.The indum entum o f both the calyx and corolla occurs random ly with m ost southern collections having a glabrous calyx but the lack o f hairs and pubescence o f the collections from the northern C edarberg an d eastern Bokkeveld are significant in the delim itation o f subsp.eglandula and subsp.pubescens.(b) subsp.eglandula (N .E. Br.) E. G. H. Oliv stat.nov.Grisebachia eremioides var.eglandula N.E.Br. in FI.
Fig. 2 2 .7 , 2 2 .8 .C a p e .-3219(Wuppertal); Pakhuis (-A A ), Barker 4505 (NBG); Esterhuysen 5924 (BOL; PRE); Esterhuysen 21764 (BOL); Krakadouw, 910 m (-A A ), Bodkin s.n.(BOL); Stokoe in S A M 55129 (NBG; PRE; SAM); Stokoe in SA M 56776 (SAM); Rocklands, 790 m (-A A ), Kruger 1031 (STE); Eselbank, 1220 m (-AC), Schlechter 8818 (BM; BOL; G; K; P; PRE; STE; Z); Crevasse Peak, 1220 m (-AC ), Taylor 7459 (PRE; STE).Without precise locality: near Clanwilliam, Leipoldt 135 (BOL); Bokkeveld, 1580 m, Schlechter 8919 (7) (BM; BOL; E; G; K; MO; P; PRE; STE; UPS; W; Z).In the C edarberg, the n o rth ern part o f the distribution range o f this species, tw o reasonably distinct groupings, A & B, o f specim ens can be m ade on the shape o f the calyx lobes, the pubescence, the arrangem ent o f the leaves and the distribution.G ro u p A has small narrow acute puberulous calyx lobes with no m arked apiculus and erect straight leaves and is ascribed to subsp.parviflora.G ro u p B has generally small flattened q uadrate to subquadrate glabrous calyx lobes w ith a distinct apiculus and m arkedly recurved leaves.The pubescence on the ovary is m uch longer an d denser.There is very little overlap in these characters between the tw o groups in the C edarberg and b oth seem to be fairly distinct.The affinities o f group B appear to lie w ith the collections m uch further south.The leaf arrangem ents in A and B are very distinct in the C edarberg b ut n o t betw een B and some random southern collections o f subsp.parviflora in which the leaves can be spreading and curved eg. in Bolus 5403 from T ulbagh.T he calyx shapes o f B, although distinct in the C edarberg, have sim ilarities in some southern collections.The distributions o f the tw o groups A & B are relatively easily separable w ith A occurring west and south o f the K rakadow -W elbedacht m ountain range and B n orth an d east o f the range.A detailed investigation o f the range is necessary to ascertain w hether this separation is in fact true or ju st due to lack o f records.
N. E. Brow n an n o tated the collection Leipoldt 135 in BOL as the type.It is u n fo rtu n ate th a t it is unlocalized.The collection Schlechter 8919 given as ju st Bokke veld is undoubtedly this subspecies an d I regard the locality as an erro r.(c) subsp.pubescens E. G. H . Oliver, subsp.nov., a subspecie typica et subspecie eglandula floribus m ajoribus, tu b o corollae om nino pubescenti, calyce om nino pubescenti, d istributione et florescentia dignoscenda.T y p e .-Cape, C eres D ist: K atbakkies in the Sw artruggens (-D C ) Oliver 4310 (STE, h o lo .;B M ; B O L; E ; G ; K ; M O ; N B G ; P R E ; S).A n erect to spreading sh ru b up to 50 cm high.C alyx pubescent; lobes o b long-triangular to broadly so, ciliate w ith fine sim ple hairs an d sto u ter glandtipped hairs adm ixed.Corolla 2-3 m m long and 1 ,2 -1 ,7 m m b ro a d ; tu b e pubescent over the whole length, tubular-ellipsoid.Fig. 2 2 .9-2 2 .1 2 .Cape.-3 2 1 9 (Wuppertal): Schurweberg, east of Bokkeveld Tafelberg, 1 0 6 0 m (-C D ), Esterhuysen 20651 (BOL; K; NBG; MO; PRE; S; STE); Zuurvlakte north of Rietvlei in the Swartruggens, 1 0 6 0 m (-C D ), Oliver 6114 (PRE; STE); Kat bakkies in the Swartruggens, 1 2 2 0 m (-D C ), Levyns 1860 (CT; SAM); 1 1 8 8 m, Oliver 4310 (BM; BOL; E; G; K; MO; NBG; PRE; S; STE); Oliver 4312 (B; C ; There are several collections from the eastern p art o f the C old B okkeveld which have a different appearance an d earlier flowering tim e from the rest o f the collections o f G. parviflora.The flowers are generally m uch larger with th e corolla tube com pletely pubescent.T he Levyns collection from K atbakkies has the coro lla tu b e pubescent for th ree-q u arters o f its length.In nearly all the collections o f G. parviflora the corollas are g labrous to p u berulous and then usually below the level o f the sepals.T he collections o f M iddlem ost from Bokkerivier to th e so u th are interm ediate tending tow ards the pubescence o f the Levyns collection.The flowering tim e o f the eastern Bokkeveld collections is significant being from June to Septem ber fo r m aterial in full flower.The collections of Levyns 1860 an d Esterhuysen 20651 & 29847 and Oliver 6114 th o u g h recorded fo r Sept., Oct. and Nov. are o f fruiting m aterial.T he flow ering tim e for collections o f G. parviflora elsewhere and particularly in th e adjoining areas o f the Bokkeveld are Sept.-D ec.fo r m aterial in full flower.This means th a t cross-pollination, even if the populations were sym patric, could n o t take place.T he M iddlem ost collections from B okkerivier were collected in N ovem ber in full flower.Despite there being little m orphological disjunction between the g roup o f eastern Bokkeveld collections and the rest o f G. parviflora, I consider th a t the geographical an d reproductive isolation w arrants recognition o f this g ro u p as a distinct subspecies o f parviflora.7. Grisebachia minutiflora N .E .Br. in FI.Cap. 4 ,1 :3 4 8 (1906).T ype: C ape, near K lein Vlei in Cold Bokkeveld, Schlechter 10064 (B M !; B O L!; G !; K, h o lo !; M O !; P !; P R E !; S !; S T E !; W !).
Corolla 4-lobed, obconic to tu b u lar w ith spreading upper half, n o t constricted, glabrous inside and outside, up to 2 ,4 m m long; lobes short b road obtuse slightly crenulate, erect o r slightly spreading.Stam ens 4, m anifest o r slightly exserted; filam ents filiform, glabrous, sigm oid a t the apex; an th ers up to 0 ,7 mm long with a lm o st parallel sides, oblong, m inutely scabrous, aw ned a b o u t $ the way up the back o f the cell; aw ns up to i the length o f the cell som etim es spreading; pore a b o u t £ the length o f the cell.Ovary 2-celled w ith a single pendulous ovule per cell, ellipsoid, puberulous a t the apex; style filiform, glabrous, f a r exserted up to 2 ,2 m m long; stigm a sim ple o r slightly swollen.Fig. 24.A species form ing co m pact erect to sem ispreading low shrublets, occurring in sandy places in the C old B okkeveld n o rth o f Ceres, flowering from O ctober to January.A variable tax o n in which tw o subspecies are recognized.K ey to the subspecies Cilia on the calyx gland-tipped................... (a) subsp.minutiflora Cilia on the calyx not gland-tipped............ (b) subsp.nodiflora Leaves pubescent w hen young w ith several sessile glands on the m argins, becom ing glabrous exceps cn the adaxial surface, or glabrous w ith nu m ero u t sessile glands over the surface but pubescent on the adaxial surface.Pedicels alm ost absent or up to 0 ,4 m m long; bracteoles usually adpressed, m ostly up to 1,1 m m long, occasionally up to 1,3 m m .C alyx pubescent in zones, glabrous on inner surface, cilia plum ose o r simple, gland-tipped.Figs 2 4 .1 -24 .8& 25.

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y p e .-Cape, Ceres D istrict, Sw artruggens in the C old Bokkeveld, Oliver 6105 (STE, h o lo .;B M ; BO L; E ; G ; K ; M O ; N B G ; P; P R E : S; W).

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i g . 2 6 .-Grisebachiasecundiflora.1, f lo w e r , x 8 ; 2 , c o r o l l a , x 8 ; 3, th r e e b r a c te o le s , x 1 6 ; 4 , s e p a ls , l a t e r a l a n d a b a x i a l, x 1 6 ; 5 , a n t h e r , b a c k , f r o n t a n d s id e v ie w s , x 1 6 ; 6 , g y n o e c iu m , x 1 6 ; 7 , le a f, x 8 ; a ll d r a w n f r o m th e h o lo ty p e , Oliver 6 1 0 5 (STE); 8 , flo w e r, x 8 , a ll d r a w n f r o m Oliver 5 0 4 4 (STE).F ig . 2 7 .-Di s t r i b u t i o n o f Grisebachia secundiflora.tosh o rt tu b u lar an d m uch inflated in the low er half, finely pubescent in the middle region an d glabrous inside, w hite: lobes varying from 0 , 6 x 0 , 6 -l , 0 x 0 ,9 m m , erect to slightly spreading, obtuse, glabrous.Stam ens 4, free; filaments narrow linear, glabrous w hite; anthers up to 1 , 0 x 0 , 5 m m , included to m anifest, subbasally attached on the dorsal surface sm ooth to minutely scabrid, aristate; aw ns dorsal pointing dow nw ards, a b o u t half the length o f the cell, m inutely ciliate, w hite; pore a b o u t h a lf the length o f the cell; pollen grains single.Ovary 1-celled w ith a single pendulous apical ovule, very rarely 2-celled, slightly oblique, 0 , 7 x 0 ,6 mm, glabrous, greenish, seated on a distinct d a rk red nectariferous disc; style filiform, 4 ,0 -4 ,5 m m long, exserted, glabrous; stigm a capitellate.Figs 26 & 27.
Table M ountain Series.This is very evident in the localities o f G. rigida, G. nivenii, G. minutiflora, G. secundiflora and G. ciliaris subsp.ciliaris, subsp.bolusii, subsp.The form er is p articularly w idespread Perennial w oody shrublets, erect up to 50 cm , from M am re to the Paleisheuwel area.In the n o rth rarely 1 m, or com pact and spreading to p ro stra te it is confined to sandy pockets in the m ountains.The an d spreading.Leaves 3-nate, rarely 4-nate, erect isolated localities o f G. incana in the area from M am re im bricate to spreading an d recurved, pubescent to Sir L ow ry's Pass are ascribed to the occurrence often w ith stout subplum ose to plum ose hairs on the of suitable sandy restionaceous sites.N o species have m argins and abaxial surface.Flowers in term inal been recorded from the extensive sands o f the Cape heads usually on sh o rt lateral branchlets som etim es Flats proper.T his is pro b ab ly due to the sand over-form ing congested pseudo-spikes.
ciliciiflora and subsp.multiglandulosa.The h a b ita t o f G. ciliaris subsp.involuta is n o t know n, as details are not given on the three collections and I have n o t collected the subspecies, b u t it w ould probably fit the requirem ents.On the low lands o f the west coast the sands are recent and alternate w ith the heavier clay soils o f the M alm esbury beds.H ere the species G. plumosa and G. incana occur.surfacescabrate to m icroscabrate w ith elem ent piT^a n s o -r r v 6. 339 (1802), p ro p a rte ; T hunb., ro(js free t0 f usecj Ovary 2-celled w ith a single r l.C ap.364 ( l o i J ) , p ro p arte.subapical pendulous ovule in each cell, rarely 3-celled, Eremia K lotzsch in L innaea 12:498 (1838), pro in one species obliquely 1-celled, w ith a d istin ct parte; Phill. in J1 S. A fr. Bot.10: 70 (1944), pro nectariferous disc.S tyle exserted.Stigm a sim ple to parte; et G en. ed. 2, 560 (1951), p ro parte.capitellate.Fruit a h ard apparently indehiscent nut.K ey to the species