Two new species of Romulea ( Iridaceae : Crocoideae ) from the west ern Karoo , Northern Cape and notes on infrageneric classification and range extensions

Two new species of Romulea are described from Northern Cape, raising the number of species in southern Africa to 76. R. collina J.C.Manning & Goldblatl is endemic to the Hantamsberg near Calvinia. It is distinguished in subgenus Spatalanthus by its clumped habit, yellow flowers with dark markings in the throat, and short papery bracts. A re-examination of rela­ tionships within the subgenus suggests that section Cruciatae is not monophyletic and it is accordingly no longer recog­ nized as separate from section Spatalanthus. R. eburnea J.C.Manning & Goldblatt is a distinctive species of subgenus Spatalanthus from the Komsberg near Sutherland. It is distinguished by its golden yellow flowers with the apical third of the tepals coloured pale creamy apricot, bracts with broad, translucent margins and tips, and an unusually long perianth tube, 10-13 mm long. The genus Romulea (Iridaceae: Crocoideae) compris­ es ± 90 species in sub-Saharan Africa, the Mediterranean and Near East. The centre of diversity of the genus lies in the winter rainfall region of southern Africa, where 73 species are currently recognized (Manning & Goldblatt 2001). The most recent revision of the sub-Saharan species divides the genus into two subgenera and six sec­ tions, based largely on characters of the corm (Manning & Goldblatt 2001). The fine structure of the corm pro­ vides essential information for accurate identification of many species, since the flower structure, with few excep­ tions, is conservative (Goldblatt et al. 2002). Five new species of Romulea have been described from South Africa in the twenty years since the publica­ tion of De Vos’s (1972) monograph of the genus in south­ ern Africa (De Vos 1983; Manning & Goldblatt 2001). The subsequent discovery of an undescribed species from the summit plateau of the Hantamsberg near Cal­ vinia in Northern Cape, described here as Romulea colli­ na, and a second from the Klein Roggeveld south of Sutherland, described as R. ebumea. now raises the total number of species in southern Africa to seventy six. Both new species are members of subgenus Spatalanthus, defined by the outer corm tunics splitting below into prominent, recurved or straight teeth without fibrous tips (Manning & Godblatt 2001). The Roggeveld Escarpment is well known for the diversity of its endemic geophyte flora (Manning et al. 2002) and is the centre of diversity for subgenus Spatalanthus. Over half of the twenty five known species of the subgenus are endemic to the Rog­ geveld and adjacent Bokkeveld Escarpments (Manning & Goldblatt 2001) and the description of two new spe­ cies brings the total number of endemic Romulea species in this centre to fifteen. * Compton Herbarium, National Botanical Institute. Private Bag X7. 7735 Claremont, Cape Town. * * B A. Krukofl Curator of African Botany. Missouri Botanical Garden. P.O. Box 299, St. Louis, Missouri 63166, USA MS. received: 2003-11-27. The discovery of Romulea collina raises doubts as to the validity of the distinction between the two sections Cruciatae and Spatalanthus of subgenus Spatalanthus, recognized by Manning & Goldblatt (2001) in their recent account of the genus. This distinction is re-exam­ ined and we conclude that the recognition of section Cruciatae is no longer justified. Romulea collina J.C.Manning & Goldblatt, sp. nov. TYPE.—Northern Cape. 3119 (Calvinia): summit of Hantamsberg at base of radio mast, (-BD). August 2002. flowered in cult.. July 2003, IBSA 5 (NBG195425, holo.). Plantae 50-100 mm altae caespitosae, caule subterraneaneo ad 4-ramoso, ramis ad 10 mm supra terram pro­ duces, cormo subgloboso asymmetrico base rotundo. tunicis infra divisis cuspis acuminatis recurvatis supra fibris grossis 20-40 mm longis. foliis 7-9 laminis ad 50-100 mm longis, floribus solitariis, bracteo extemo subobtuso pallide papyraceo ad apicem marginibus apiceque latis translucentibus brunneo vittato 10-12 mm longo intemo obtuso pallide papyraceo. floribus profunde cupuliformibus pallide flavis cupulo luteo. tubo perianthii infundibuliforme 4 mm longo parte inferiore ± 1 mm longo. tepalis lanceolatis ± 20 x 7 mm. staminibus flavis filamentis ± 6 mm longis dense pubescentibus in dimidio inferiore. antheris 2.5-3.0 mm longis apiculatis, ramis styli 1.5-2.0 mm longis, capsulis subglobosis ± 8 mm oblongo-ovoideis pedicelibus recurvatis. Plants 50-100 mm high, growing in clumps; stem subterranean, with up to four branches reaching 10 mm above-ground, these relatively stout and flushed maroon. Corm subglobose. asymmetric, base rounded, tunics split into curved acuminate teeth below, drawn into coarse fibres above, these 20-40 mm long. Cataphylls 3, flushed maroon above ground. Leaves 7-9. all basal, sheaths flushed deep purple, blades two to four times as long as flowering stems, narrowly 4-grooved, 50-100 x 1.0-1.5 mm. Inflorescence: up to 4 solitary flowers; 18 Bothalia 34,1 (2004) outer bracts subobtuse, pale and papery throughout with veins in centre pale green, with broad, translucent mar­ gins and tip flecked pale brown, 10-12 mm long; inner bracts obtuse, pale and papery throughout with only main veins pale green, with broad translucent margins flecked pale brown, about as long as outer. Flowers deeply cup-shaped, cup ± 11 mm deep, pale canary yel­ low with golden yellow cup; outer tepals with dark median blotch or transverse zone in throat, reverse marked with dark maroon longitudinal streaks; inner tepals with narrow brownish maroon median line and diffuse transverse zone in throat, unscented, ± 30 mm diam.; perianth tube funnel-shaped, 4 mm long with lower narrow portion ± 1 mm long, tepals lanceolate, ± 20 x 7 mm. Stamens yellow; filaments inserted at base of cup, free, densely hairy in lower half, 6 mm long; anthers apiculate, parallel, 2.5-3.0 mm long. Style dividing opposite upper third of anthers, branches 1.5-2.0 mm long, divided for about two thirds of their length. Capsules subglobose, ± 8 mm long, pushed onto soil sur­ face or slightly underground by strongly recurved pedicels which later become erect and 10—15 mm long. Seeds subglobose or angled by pressure, reddish brown, 2.5-3.0 mm diam. Flowering time: late June to mid-July, possibly to late July. Figure 1. FIGURE 1.— Romulea collina. A, whole plant; B, flower; C, outer bract; D, inner bract; E, adaxial markings on outer tepal; F, adaxial markings on inner tepal; G, stamens and style; H, seed; I, t/s leaf. Scale bars: A-F, 10 mm; G, H, 2 mm. Artist: John Manning. Distribution and ecology This clump-forming species is known from a single, very localized colony on the summit plateau of the Hantamsberg overlooking Calvinia on the Bokkeveld Escarpment (Figure 2). The plants grow in seasonally moist, dolerite clays in open renosterveld, invariably in small clumps that appear to originate from seeds that are shed directly at the base of the parent plant. The short peduncles curve downwards immediately after flowering but become erect when the fruits mature. They do not, however, elongate appreciably so that the mature cap­ sules dehisce 10-15 mm above the ground at the base of the plant. Flowers open around midday and close in midafternoon. They are short-lived, lasting only two days. At up to 3 mm diam., the seeds of R. collina are among the largest recorded in the genus, where seeds are mostly 1-2 mm diam. (De Vos 1972). Romulea collina is yet another of several species of Iridaceae that are endemic to the Hantamsberg, an isolated, flat-topped massif representing a northern outlier of the Roggeveld Escarpment. Iridaceae endemic to the slopes and summit of the Hantamsberg include Hesperantha hantamensis Schltr. ex Foster, H. oligantha (Diels) Goldblatt Bothalia 34,1 (2004) 19 FIGURE 2. —Distribution of Romulea collina, O; and Romulea eburnea, • Moraea reflexa Goldblatt and Romulea hantamensis (Diels) Goldblatt, among others (Van Wyk & Smith 2001). Diagnosis and relationships Romulea collina is readily distinguished by its clumped habit rounded corm with the outer tunics splitting into acumi­ nate teeth, yellow flowers with dark markings in the throat, and short bracts, 10-12 mm long, that are entirely pale and papery at flowering. The corm with tunics splitting into acuminate teeth at the base are a defining characteristic of Romulea sub­ genus Spatalanthus but within the subgenus, the relationships of R. collina are more difficult to define. The rounded corm with curved basal teeth is characteristic of section Spatalanthus but in other respects, particularly the relatively short more or less entirely membranous bracts and the long filaments, the species closely resembles R. membranacea M.P. de Vos, anoth­ er species from the Roggeveld Escarpment Conservation status: Vulnerable (B1 + 2c, D). Apart from the difference in the corm shape, R. collina is separated from R. membranacea by the longitudinal dark streaks on the outside of the outer tepals and by the pro­ nounced dark blotches on the inside. Despite their different corm morphology, the overwhelming similarity between the two species in other particulars makes it difficult to accept that they are not immediately related. R. membranacea is currently placed in section Cruciatae, which is defined by a corm pointed at base and with straight basal teeth (Manning & Goldblatt 2001) but the relationships of this species have always been problematic. De Vos (1972) placed it in her subsection Atrandrae on the basis of the broad membranous margins of the bracts but pointed out that it was anomalous here in its pointed corm and lack of marginal veins in the leaves. Manning & Goldblatt (2001), placing greater emphasis on corm structure, preferred to ally it with the other species with straight basal teeth, which it matches in leaf anatomy. The chromosome number 2n = ± 24 is, how­ ever, inconsistent with this alliance and accords better with section Atrandrae. The discovery of the new species R. col­ lina, with the rounded corm of section Spatalanthus but the le

The genus Romulea (Iridaceae: Crocoideae) compris es ± 90 species in sub-Saharan Africa, the Mediterranean and Near East.The centre of diversity of the genus lies in the winter rainfall region of southern Africa, where 73 species are currently recognized (Manning & Goldblatt 2001).The most recent revision of the sub-Saharan species divides the genus into two subgenera and six sec tions, based largely on characters of the corm (Manning & Goldblatt 2001).The fine structure of the corm pro vides essential information for accurate identification of many species, since the flower structure, with few excep tions, is conservative (Goldblatt et al. 2002).
Five new species of Romulea have been described from South Africa in the twenty years since the publica tion of De Vos's (1972) monograph of the genus in south ern Africa (De Vos 1983;Manning & Goldblatt 2001).The subsequent discovery of an undescribed species from the summit plateau of the Hantamsberg near Cal vinia in Northern Cape, described here as Romulea colli na, and a second from the Klein Roggeveld south of Sutherland, described as R. ebumea.now raises the total number of species in southern Africa to seventy six.Both new species are members of subgenus Spatalanthus, defined by the outer corm tunics splitting below into prominent, recurved or straight teeth without fibrous tips (Manning & Godblatt 2001).The Roggeveld Escarpment is well known for the diversity of its endemic geophyte flora (Manning et al. 2002) and is the centre of diversity for subgenus Spatalanthus.Over half of the twenty five known species of the subgenus are endemic to the Rog geveld and adjacent Bokkeveld Escarpments (Manning & Goldblatt 2001) and the description of two new spe cies brings the total number of endemic Romulea species in this centre to fifteen.

Romulea collina
Plants 50-100 mm high, growing in clumps; stem subterranean, with up to four branches reaching 10 mm above-ground, these relatively stout and flushed maroon.Corm subglobose.asymmetric, base rounded, tunics split into curved acuminate teeth below, drawn into coarse fibres above, these 20-40 mm long.Cataphylls 3, flushed maroon above ground.Leaves 7-9.all basal, sheaths flushed deep purple, blades two to four times as long as flowering stems, narrowly 4-grooved, 50-100 x 1.0-1.5 mm.Inflorescence: up to 4 solitary flowers; outer bracts subobtuse, pale and papery throughout with veins in centre pale green, with broad, translucent mar gins and tip flecked pale brown, 10-12 mm long; inner bracts obtuse, pale and papery throughout with only main veins pale green, with broad translucent margins flecked pale brown, about as long as outer.Flowers deeply cup-shaped, cup ± 11 mm deep, pale canary yel low with golden yellow cup; outer tepals with dark median blotch or transverse zone in throat, reverse marked with dark maroon longitudinal streaks; inner tepals with narrow brownish maroon median line and diffuse transverse zone in throat, unscented, ± 30 mm diam.; perianth tube funnel-shaped, 4 mm long with lower narrow portion ± 1 mm long, tepals lanceolate, ± 20 x 7 mm.Stamens yellow; filaments inserted at base of cup, free, densely hairy in lower half, 6 mm long; anthers apiculate, parallel, 2.5-3.0 mm long.Style dividing opposite upper third of anthers, branches 1.5-2.0mm long, divided for about two thirds of their length.Capsules subglobose, ± 8 mm long, pushed onto soil sur face or slightly underground by strongly recurved pedicels which later become erect and 10-15 mm long.Seeds subglobose or angled by pressure, reddish brown, 2.5-3.0 mm diam.Flowering time: late June to mid-July, possibly to late July.Figure 1.

Distribution and ecology
This clump-forming species is known from a single, very localized colony on the summit plateau of the Hantamsberg overlooking Calvinia on the Bokkeveld Escarpment (Figure 2).The plants grow in seasonally moist, dolerite clays in open renosterveld, invariably in small clumps that appear to originate from seeds that are shed directly at the base of the parent plant.The short peduncles curve downwards immediately after flowering but become erect when the fruits mature.They do not, however, elongate appreciably so that the mature cap sules dehisce 10-15 mm above the ground at the base of the plant.Flowers open around midday and close in midafternoon.They are short-lived, lasting only two days.At up to 3 mm diam., the seeds of R. collina are among the largest recorded in the genus, where seeds are mostly 1-2 mm diam.(De Vos 1972).
Romulea collina is yet another of several species of Iridaceae that are endemic to the Hantamsberg, an isolated, flat-topped massif representing a northern outlier of the Roggeveld Escarpment.Iridaceae endemic to the slopes and summit of the Hantamsberg include Hesperantha hantamensis Schltr.ex Foster, H. oligantha (Diels) Goldblatt

Diagnosis and relationships
Romulea collina is readily distinguished by its clumped habit rounded corm with the outer tunics splitting into acumi nate teeth, yellow flowers with dark markings in the throat, and short bracts, 10-12 mm long, that are entirely pale and papery at flowering.The corm with tunics splitting into acuminate teeth at the base are a defining characteristic of Romulea sub genus Spatalanthus but within the subgenus, the relationships of R. collina are more difficult to define.The rounded corm with curved basal teeth is characteristic of section Spatalanthus but in other respects, particularly the relatively short more or less entirely membranous bracts and the long filaments, the species closely resembles R. membranacea M.P. de Vos, anoth er species from the Roggeveld Escarpment Conservation status: Vulnerable (B 1 + 2c, D).
Apart from the difference in the corm shape, R. collina is separated from R. membranacea by the longitudinal dark streaks on the outside of the outer tepals and by the pro nounced dark blotches on the inside.Despite their different corm morphology, the overwhelming similarity between the two species in other particulars makes it difficult to accept that they are not immediately related.R. membranacea is currently placed in section Cruciatae, which is defined by a corm pointed at base and with straight basal teeth (Manning & Goldblatt 2001) but the relationships of this species have always been problematic.De Vos (1972) placed it in her subsection Atrandrae on the basis of the broad membranous margins of the bracts but pointed out that it was anomalous here in its pointed corm and lack of marginal veins in the leaves.Manning & Goldblatt (2001), placing greater emphasis on corm structure, preferred to ally it with the other species with straight basal teeth, which it matches in leaf anatomy.The chromosome number 2n = ± 24 is, how ever, inconsistent with this alliance and accords better with section Atrandrae.The discovery of the new species R. col lina, with the rounded corm of section Spatalanthus but the leaf anatomy and bract morphology of R. membranacea, suggests that the earlier placement by De Vos (1972) of R. membranacea in subsection Atrandrae (= series Atrandrae of Manning & Goldblatt 2001) is preferable.Within series Atrandrae, R. collina and R. membranacea are probably most closely related to R. diversiformis M.P.de Vos, with which they share yellow flowers, fruiting peduncles that are curved at maturity rather than coiled, and leaves that lack marginal veins along the furrows.
The reassessment of the relationships of R. collina and R. membranacea implies that the pointed corm with straight basal teeth has arisen more than once within the genus.If this interpretation is correct then the heterogeneity evident within the section assumes a new significance.Section Cruciatae is currently divided into two series.Series Cruciatae, with the removal of R. membranacea, contains three species, two of which are clearly sister taxa [R.cruciata (Jacq.)Baker and R. eximia M.P.de Vos] that share the unusual chromosome number 2n = 18.On the basis of their cytology, R. cruciata and R. eximia are better allied with R. rosea (L.) Eckl.(section Spatalanthus, series Roseae), the only other species in the genus with In = 18.The third species in this series, the poorly known R. vlokii M.P.de Vos has bracts that more closely resemble those found in series Atrandrae of section Spatalanthus, which is where De Vos (1983) placed it.Series Tubiformes contains a single highly derived species, R. hantamensis (Diels) Goldblatt, with a chromosome number In = 30, which is unique within sub genus Spatalanthus and its relationships remain unclear.If the pointed corm is interpreted as a convergent character state, then it is likely to have evolved three times in section Cruciatae and a fourth time in R. membranacea.Section Cruciatae as currently construed is probably not monophyletic and it seems more appropriate that the species within it be removed to section Spatalanthus.Accordingly, we no longer recognize section Cruciatae, and subgenus Spatalanthus should no longer be subdivided into sections.Romulea eburnea J.C. Manning & Goldblatt, sp. nov.TYPE.-NorthernCape, 3220 (Sutherland): 28 km along Komsberg Pass road from southern end, foot of Smoushoogte, alluvial wash along Meintjiesplaasrivier, (-DC), 28 August 2003, J. Manning 2886 (NBG, holo.; MO, iso.).
Plants 100-150 mm high; stem subterranean, with up to two branches reaching 30 mm above ground, these relatively stout and flushed maroon.Corm subglobose, asymmetric, base rounded, tunics split into curved acuminate teeth below, drawn into coarse fibres above, these up to 2 mm long.Cataphylls 3, flushed maroon above ground.Leaves 2-4, sheathing portion pale with adhering sand grains, blades of two longest leaves up to six times as long as flowering stems, narrowly four-grooved, 150-200 mm long, when more than two leaves present, then uppermost one or two with blades less than half as long, 10-100 x 1 mm.Inflorescence: up to 2 soli tary flowers; outer bracts pale reddish brown with broad, translucent margins and broad, membranous tip flecked with pale brown, 18-22 mm long; inner bracts bifid, pale reddish brown with broad, translucent margins and broad, membra nous tip flecked with pale brown, about as long as outer or slightly longer.Flowers deeply cup-shaped, cup ± 12 mm deep, pale creamy apricot with lower two thirds of tepals and cup deep yellow; outer tepals with reverse flushed dull olive in centre in lower half, unscented, ± 30 mm diam.; perianth tube funnel-shaped, 10-13 mm long with lower narrow por tion 6-8 mm long, tepals lanceolate, 23-27 x 7-9 mm.Stamens yellow; filaments inserted at base of cup, free, pubes cent at base, 6-7 mm long; anthers suberect or lightly incurved, ± 8 mm long.Style dividing opposite tips of anthers, branches 3 mm long, divided for about half their length.Capsules oblong-ovoid, 10-11 x 5.5-6.0 mm, pushed onto soil surface or slightly underground by strongly recurved pedicels which later coil up when dry. Seeds subglobose or angled by pressure, reddish brown, 1.8-2.0mm diam.Flowering time: late August to early September.Figure 3.

Distribution and ecology
This distinctively coloured species is known from a single colony below Smoushoogte Pass south of Sutherland at the foot of the Klein Roggeveldberge (Figure 2).Romulea ebumea grows in an alluvial wash along the Meintjiesplaasrivier in seasonally moist, deep sandy loam.The veg etation is open scrub dominated by Asparagus capensis, Chrysocoma ciliata, Eriocephalus eximius, and Dimorphotheca cuneata, and quite distinct from the adjacent renosterveld (Elytopappus rhinocemtis) community that occu pies fine-grained clays derived from shale.
R. ebumea was first noticed almost a decade ago by local bulb enthusiasts who reported the occurrence of a pale-flow ered species on the Komsberg but in the absence of speci mens it was impossible to verify its identity.The possibility that it represented an outlying population of R. membranacea was suggested by Manning & Goldblatt (2001) but it is now clear that the plants represent a distinct species.It is another of several species in series Atrandrae that are endemic or near-endemic to the Roggeveld Escarpment around Sutherland, including R. hallii M.P.de Vos, R. komsbergensis M.P.de Vos and R. multifida M.P.de Vos.

Diagnosis and relationships
The rounded corms with tunics fragmenting into curved teeth, place Romulea ebumea firmly in subgenus Spatalanthus.It is immediately recognized by its largely golden yellow flowers with the apical third of the tepals coloured pale creamy apricot, bracts with broad, translu cent margins and tips, and the unusually long perianth tube.This is funnel-shaped and 10-13 mm long with the lower narrow portion 6-8 mm long.In most other species of Romulea with a similar-shaped perianth, the tube is 4-6(-10) mm long with the lower narrow portion no more than 2 mm long.Within subgenus Spatalanthus, the presence of vascular bundles along the margins of each of the ribs in the leaves, the broad membranous margins and tips to the bracts, and fruiting peduncles that coil when dry. are all consistent with series Atrandrae.Within series Atrandrae, R. ebumea is possibly most closely allied to R. diversiformis, with which it shares yellow flowers lacking dark markings in the throat and a long style that divides at or beyond the tips of the anthers.In an extraordinary coincidence, both R. diversi formis, described just over Fifty years ago (De Vos 1952), and R. ebumea share the same type locality and we have in the past collected R. diversiformis in flower in late September within a few dozen metres of the R. ebumea population, then long past flowering, without being aware of its existence.R. diversiformis was until recent ly thought to be restricted to the Klein Roggeveld area but is now known to occur also on the Hantamsberg near Calvinia, a considerable distance away at the northern end of the Roggeveld Escarpment (Manning & Goldblatt 2001).R. diversiformis is distinguished from R ebumea by its greater number of leaves, 6 or more versus 2^4, uniformly golden yellow flowers with short perianth tube, 4-6 mm long versus 10-13 mm long and fruiting peduncles that do not coil up when dry.The two species differ also in leaf anatomy, R. diversiformis lacking rib marginal veins but possessing secondary veins in the ribs.R. diversiformis grows in Fine-grained, clay soils that are seasonally waterlogged or actually shallowly inundated, whereas R. ebumea is known only from sea sonally moist, sandy soils.
This species was based on a single collection made on the summit of the Flaminkberg just southeast of Nuwerus in southern Namaqualand.The recent discov ery of a second collection antedating the type by almost twenty years, substantially increases the known range of the species.This collection matches the type in all respects, including the large white flowers, bracts with broad, brown-spotted membranous margins and the white-spotted cataphyll and there is no doubt that it rep resents this species.Romulea maculata is now known from near Nuwerus in the south of Namaqualand to Komaggas in the north and may be expected to occur through most of the higher ground of Namaqualand.The species is restricted to seasonally moist, sandy loam on granitic slopes.Romulea multisulcata M.P.de Vos in Journal of South African Botany.Suppl.9: 139 (1972).

Additional material examined
This species was described from seasonal pools on the Bokkeveld Mountains near Nieuwoudtville in Northern Cape (De Vos 1972) but later collections from the foot of the Gifberg-Matsikamma massif nearby (Manning & Goldblatt 2001).clearly represent a second set of popu lations differing from the type only in their white rather than yellow flowers.However, populations from Hondeklip Bay in central Namaqualand that were also assigned to this species on the basis of a single herbari um collection (Manning & Goldblatt 2001) prove, on examination of living plants, to lack the characteristic multisulcate leaves of R. multisulcata and accord more closely with R. tabularis Eckl.ex Beg.This species of waterlogged coastal flats is widely distributed along the west coast of South Africa from Cape Agulhas in the south to northern Namaqualand.The Hondeklip Bay populations thus fall within its recorded range.Romulea multisulcata thus remains endemic to southern Namaqua land.where it is restricted to a few scattered seasonal pools near Vanrhynsdorp and Nieuwoudtville.