Massonia bakeriana M.Pinter, Mart.

Pinter, Michael, Martínez-Azorín, Mario, Crespo, Manuel B. & Wetschnig, Wolfgang, 2015, Massonia bakeriana (Asparagaceae, Scilloideae), a new pustulate species from the Northern Cape Province (South Africa), Phytotaxa 222 (1), pp. 51-60 : 52-57

Massonia bakeriana M.Pinter, Mart. -Azorín & Wetschnig sp. nov. ( Figs. 1 − 6 ).

Species notabilis ab Massonia echinata et M. mimetica primo aspectu aemulans, sed eis propria combinatione characterum facile distinguitur foliis pallide viridulis, pustulis 0.6−1 mm in diam.; perigonii tubo ad faucem valde aperto, egibboso laevique (in illis 6 gibbis munito, quae ovarium occultans); filamentis albidis a basi breve connatis (ca. 1 mm long.); et antheris purpureo-violaceis. A priore insuper etiam differt foliarum pustulis majoribus et trichomatibus apicalis destitutis (in illa 0.3−0.4 mm diam. et trichomate unico apicale munitis). A posteriore insuper multo discrepat etiam foliis supra non cinnamomeis, pustulis foliarum minoribus (in illa 1−2 mm diam.); et tubo filamentorum non purpureo maculato.

Type:— SOUTH AFRICA. Northern Cape, Williston (3120): Slangberge, south of Williston (- DB), southwest slopes of kloof, common between and under rocks, 1220 m elevation ; flowers whitish to pink, 19 May 1976, M. F. Thompson 3155 (holotype PRE0549801 -0!, isotypes PRE0488986 -0!, PRE0513174 -0!)

Herbaceous perennial plant. Bulb ovoid to subglobose, tunicate, 20 − 30 × 20 − 28 mm, inner tunics fleshy and white, outer tunics leathery and brownish. Leaves 2, deciduous, opposite, spreading and appressed to the ground, synanthous, ovoid to suborbicular with obtuse apex, with a short apicule <1 mm long, (4 −)6 − 8(− 10) × (5 −)6 − 8(− 10) cm, with entire to minute papillose margins; adaxial side glabrous, pale green, with symmetrical, cone-like, dark green, (11 −)15 − 30(− 42) emergences/cm 2, 0.6 − 1 mm in diam., with a small apical papilla; abaxial side smooth, green. Inflorescence a dense, subcapitate raceme, up to 20 − 30 mm long, with 15 − 20 flowers, shortly overtopping ground level. Bracts lanceolate, long acuminate, 15 − 24(− 30) × 5 − 10(− 14) mm, membranous, green in the upper half and white below, with slightly darker green venation, glabrous with entire margins. Pedicels 10 − 14 mm long. Flowers tubular, funnel-shaped above. Perigone white, free segments (6 −)7 − 8 × 1.2 − 2 mm with a short greenish central band at the tip, first straight and erect, later spreading and finally at anthesis strongly reflexed with a slight curve at the base. Perigone-filaments tube 9 − 13 × (2 −) 2.5 − 4 mm, cylindrical, funnel-shaped above, bearing a wide open mouth that shows the ovary in apical view. Filaments white, (12 −)14 − 16(− 18) mm long, in young flowers sometimes unequal in length, connate at the base for less than 1 mm forming a short filaments-tube above the perigone-filaments tube, spreading, straight, aligned at the same angle as the funnel-shaped tube, attenuate; anthers 2 − 3 mm long when closed, oblong, violet-purple, dorsifixed, pollen yellow. Ovary narrowly oblong, greenish with a violet to purplish tinge, 5 − 6 × 1.7 − 2 mm, slightly contracted at the joint with the style. Style white, slender, gradually tapering to the apex, 16 − 20 mm long, curved, with a bend at the apex. Capsule loculicidal, 10 − 12 × 7 − 10 mm, valves splitting down to the base, oblong in lateral view and trigonous with blunt edges in apical view. Seeds globose, glossy black, ca. 1.4 − 1.6 × 1.3 − 1.6 mm, smooth.

Eponymy:—The specific epithet ‘ bakeriana’ honours John Gilbert Baker (1834–1920), for his leading work on Massonia and influential treatment of the genus in Flora Capensis (1897).

Biology:—In wild populations leaves are found from April to May. Massonia bakeriana flowers around May in the natural habitat, whereas in cultivation in a greenhouse in the Northern Hemisphere the leaves appear in September and it flowers from October to November.

Distribution:—The new species is known to us from three main areas: the Kamiesberg area, in the surroundings of Kamieskroon; the surroundings of Loeriesfontein and Calvinia on the Bokkeveld Plateau; and the Jan Swartsberge and Slangberge, W and SW of Williston, N of the Roggeveld ( Fig. 7 ). The new species appears to be common in the Jan Swartsberge and the Slangberge.

Habitat:— Massonia bakeriana seems to be confined to rocky areas. In the Kamiesberg and the Bokkeveld Plateau, it grows between 700 and 1000 m of elevation, and in the Jan Swartsberge and Slangberge it occurs in elevations between 1000 and 1300 m. The Kamiesberg localities fall within the Succulent Karoo Biome, Kamiesberg Mountains Shrubland (SKn6) and also in the Fynbos Biome, Namaqualand Granite Renostersveld (FRg1). Localities on the Bokkeveld Plateau also fall within the Succulent Karoo Biome, Namaqualand Klipkoppe Shrubland (SKn1); Hantam Karoo (SKt2); and Knersvlakte Shale Vygieveld (SKk4). In the Swartsberge and Slangberge, W and SW of Williston localities are found mostly confined to the Nama-Karoo Biome, Upper Karoo Hardeveld (NKu2); they could also occur in the Western Upper Karoo (NKu1) ( Mucina & Rutherford 2006).

species.

Taxonomic relationships:— Massonia bakeriana is at first sight related to M. echinata and M. mimetica based on flower and leaf morphology. It also shares similar distribution ranges with the latter two species ( Fig. 7 ). However, Massonia bakeriana differs from both latter species by the reflexed free portions of perigone segments at anthesis, lacking a strong sigmoid curve at the base; the wide open, funnel-shape mouth of the perigone-filaments tube lacking gibbosities, where the ovary is visible in apical view; the longer filaments and ovary; and the style usually bend in the apical portion ( Figs. 2 − 3 ; Table 2). Massonia bakeriana shares with M. echinata the green leaves bearing emergences and the white flowers, but the latter differs by its leaves with less and smaller emergences bearing a hair instead of a papilla on top ( Figs. 5 , 6 ); flowers tubular with narrow mouth of the tube, usually having gibbosities; free portion of perigone segments at anthesis with a strong sigmoid curve; filaments and ovary shorter; and the style straight ( Table 2). Massonia bakeriana shares with M. mimetica the distinctly pustulate leaves, showing emergences with a papilla on top ( Figs. 5 , 6 ), but the latter differs by the leaves being orange-brown or cinnamon-coloured; the flowers cream-white to pale-yellow; free portion of perigone segments at anthesis with a strong sigmoid curve at the base; the filaments-tube tinged with reddish; the shorter filaments and ovary; and the straight style ( Martínez-Azorín et al. 2013). Other species with pustulate leaves, like M. longipes and M. pustulata , differ from M. bakeriana by the ovary truncate at the apex and clearly differentiated from the narrow style. Furthermore other differences on flower and leaf morphology are shown in Wetschnig et al. (2012). There are also important differences in ecology of M. bakeriana and its closest relatives. Whereas Massonia mimetica appears to be confined to high-altitude inland plateaus supporting deep, red sandy soils, where the peculiar leaves show mimesis of the environment, M. bakeriana occurs in adjacent mountainous regions and grows in rocky habitats. Massonia echinata grows in course-grained sandy soils and hard-stony clay.

Molecular data:—In our preliminary phylogenetic studies (Martínez-Azorín et al., unpublished data) Massonia bakeriana , M. echinata and M. mimetica form a monophyletic group within the genus which is well supported, a placement that agrees with their shared morphological characters. Moreover, they all show differences in their DNA sequences supporting segregation at the species rank. A more complete sampling of Massonia including a higher number of taxa and additional markers is ongoing.

Additional specimens studied (paratypes):— SOUTH AFRICA. Northern Cape. Hondeklipbaai (3017): Kamieskroon , A. le Roux (photo!) ; Kamiesberg (3018): Bergplätze bei [mountainous areas near] Pedroskloof (- AA), 1 November 1839, Drège 2683c ( P01855936 photo!) ; Loeriesfontein (3019): Brandkraal Farm (- CC) [as M. echinata ] (photo! in Summerfield (2004: 29)) ; Loeriesfontein (3019): Loeriesfontein )) ; Loeriesfontein (- CD), grown from seeds ex Gordon Summerfield 3820, WW04971 ( GZU!) ; Calvinia (3119): Eselskop (- AA), Jiří Maule (photo!, http://www. zelenelisty.cz/clanky/werbar---cibuloviny/massonia-pustulata.html) ; Williston (3120): Jan Swartsberge ca. 30 km W of Williston (- BC), 19 April 2014 in bud, D. Human (photo!) ; Williston (3120): Slangberge, south of Williston , southwest slopes of kloof (- DB), 1220 m elevation, common between and under rocks, flowers whitish to pink, 19 May 1976, M. F. Thompson 3155 ( PRE0549801 -0!, PRE0488986 -0!, PRE0513174 -0!) .

Figures

FIGURE 1. Habit of Massonia bakeriana M.Pinter, Mart.-Azorín & Wetschnig flowering in cultivation from Loeriesfontein (WW04971). A. Bulb; B. Sprouting plant; C. Plant at anthesis. Scale bar: 1 cm.

FIGURE 2. Flower and bract morphology of Massonia bakeriana M.Pinter, Mart.-Azorín & Wetschnig flowering in cultivation from Loeriesfontein (WW04971). A. Mature flower in apical view; B. Bracts; C. Development of flowers from the bud (on the left) to full anthesis (on the right). Scale bars:A: 0.5 cm; B, C: 1 cm.

FIGURE 3. Dissected flowers of Massonia bakeriana M.Pinter, Mart.-Azorín & Wetschnig flowering in cultivation from Loeriesfontein (WW04971). A. Bud; B. Young flower; C. Flower at anthesis. Scale bar: 1 cm.

FIGURE 4. Seed morphology of Massonia bakeriana M.Pinter, Mart.-Azorín & Wetschnig. A. Seed, lateral view; B. Seed, raphal view; C. Seed, longitudinal section; D. Seed, transversal section. Scale bars: 0.5 mm.

FIGURE 5. Leaf surface and emergence morphology in the studied Massonia species. A, D. M. bakeriana M.Pinter, Mart.-Azorín & Wetschnig. B, E. M. mimetica Mart.-Azorín, M.Pinter, M.B.Crespo & Wetschnig. C, F. M. echinata L.f. A–C: apical view of leaf surface; D–F: section of leaf with emergence morphology in lateral view.A–D, F fresh leaves, E fixed leaf. Scale bars:A–C: 0.5 cm; D–F: 0.5 mm.

FIGURE 6. SEM micrographs of leaf surface showing emergence morphology in the studied Massonia species. A, D. M. bakeriana M.Pinter, Mart.-Azorín & Wetschnig. B, E. M. mimetica Mart.-Azorín, M.Pinter, M.B.Crespo & Wetschnig. C, F. M. echinata L.f. A–C: general view of leaf surface; D–F: detail of pustules morphology in apical view. Scale bars:A–C: 1 mm; D–F: 0.2 mm.

FIGURE 7. Known distribution of Massonia bakeriana M.Pinter, Mart.-Azorín & Wetschnig. (green triangles), Massonia echinata L.f. (blue squares) and Massonia mimetica Mart.-Azorín, M.Pinter, M.B.Crespo & Wetschnig (red circles) in the Northern Cape Province of South Africa. Type localities are indicated by symbols with white centres.