Research Article |
Corresponding author: David J. Goyder ( d.goyder@kew.org ) Academic editor: Peter Bruyns
© 2023 David J. Goyder, Nina Davies, Manfred Finckh, Amândio Gomes, Francisco Maiato P. Gonçalves, Paulina Meller, Alan J. Paton.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Goyder DJ, Davies N, Finckh M, Gomes A, Gonçalves FMP, Meller P, Paton AJ (2023) New species of Asclepias (Apocynaceae), Baphia (Leguminosae), Cochlospermum (Bixaceae) and Endostemon (Lamiaceae) from the Kalahari sands of Angola and NW Zambia, with one new combination in Vangueria (Rubiaceae). PhytoKeys 232: 145-166. https://doi.org/10.3897/phytokeys.232.110110
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Four new species are described from central and eastern Angola and adjacent NW Zambia. All occur in Kalahari sand savannas rich in endemic and more widely distributed geoxylic suffrutices. Despite being known from very few collections, the conservation status of one of these new species is assessed as Least Concern, as these grasslands are nutrient-poor, are in remote sparsely populated areas, and are not threatened with conversion to agriculture. The remaining three are treated as Data Deficient. In addition, one new combination is provided for Ancylanthos rubiginosus Desf. under Vangueria as V. rubiginosa (Desf.) Lantz is an illegitimate later homonym. We also make orthographic corrections to specific epithets commemorating Ilse von Nolde, a collector who made important collections from Quela in Malange in the 1930s.
Ancylanthos, Casearia, Cochlospermaceae, geoxyle, geoxylic suffrutices, Ilse von Nolde
Plant diversity in Angola is poorly documented with very uneven geographic coverage – much of the eastern half of the country and some northern provinces are largely devoid of georeferenced plant collections (
Two initiatives have contributed data to the current publication. Firstly, a series of expeditions under the auspices of the National Geographic Okavango Wilderness Project (NGOWP) focussed initially on the headwaters of the Cuito and Cuanavale tributaries of the Okavango system in central Angola, then on river catchments further to the east in 2019. Most of these surveys were in Moxico Province, but neighbouring areas of Cuando Cubango with similar rainfall and vegetation were also documented. In parallel, The Future Okavango programme (TFO) of the University of Hamburg initiated a series of ecological studies in miombo and grassland sites around Chitembo, Bié Province, just to the west of the core NGOWP study area.
The NGOWP surveys culminated in a Checklist of the Cuito headwaters, documenting 417 species from the region (
Much of eastern Angola is overlain with deep deposits of Kalahari sand, which in the Cuito headwater zone are white, highly leached, and largely devoid of nutrients (
Tributaries of the Cuito River lie in steeply incised valleys around 150 m below the level of the surrounding plateau and their headwaters generally take the form of a wetland or seepage area surrounded by a narrow zone of exposed sand with a herbaceous or suffrutescent flora maintained free of woody vegetation by a combination of fire, frost and high water table (
Frequently, a little further downstream, a headwater lake occurs with deep open water and a fringing mat of peat, with its own suite of plants capable of surviving in this low pH environment. Wetlands are not generally known for high levels of plant diversity or endemism, although some species such as the insectivorous Genlisea angolensis R.D.Good appear to be restricted to peaty wetlands on Kalahari sand (
Further downstream, with the confluence of additional tributaries, the valleys generally become wider and the seasonally burned grassland zones more extensive. This is the habitat that is most significant botanically with many range-restricted and endemic species, including most of the species new to science described below. One notable valley grassland formation consists of extensive sand platforms perched above the level of the current watercourse; it is dominated by grasses and by geoxylic suffrutices, plants with considerable underground woody biomass and seasonal above-ground shoots. These lifeforms are adapted to above-ground disturbances such as fire and frost and were highlighted by
Several globally rare species were noted in the high rainfall Kalahari sand savannas of Gabon’s Batéké Plateau (
Morphological descriptions are based principally on examination of herbarium specimens at Kew, with smaller organs examined under a Leica MZ12.5 stereo-dissecting microscope fitted with a 10 mm graduated eyepiece graticule. Relevant type material not available for direct observation at K or BM was consulted through the JSTOR Global Plants portal www.plants.jstor.org, or the websites of individual herbaria such as COI www.uc.pt/en/herbario_digital or LISC https://actd.iict.pt/collection/actd:BIOHERB. Herbarium abbreviations follow Index Herbariorum (Thiers continually updated) except for the Collections Unit of Angola’s National Institute for Biodiversity and Protected Areas which is not listed by Thiers, and is here referred to as INBAC.
Asclepias minutissima appears most similar to A. aurea (Schltr.) Schltr. but differs in the campanulate rather than rotate to reflexed corolla, the ascending disposition of the corona rather than radiating from the column in A. aurea, the absence of the well-developed distal tongue to the corona of the latter species, and the shorter peduncles (1–3 cm rather than (3)5–14 cm in A. aurea).
Angola. Moxico Province: Mussuma plains, 50 km NE of Lumbala, Zambezi drainage, 13°45'49"S, 021°43'25"E, fl. 7 December 2019, D.Goyder & F.Maiato 9204 (holotype: K (K001334259); isotypes: INBAC, LUBA, PRE).
Perennial herb with a single erect stem arising annually from a small napiform tuber, latex white; stems 8–15 cm long, minutely pubescent along two lines. Leaves sessile, 3–7 × 0.05 cm, filiform with inrolled margins, glabrous. Inflorescences terminal or extra-axillary, umbelliform, with 4–5 erect flowers; peduncles 1–3 cm long, minutely pubescent; pedicels c. 1 cm long, minutely pubescent. Sepals 1–1.5 mm long, narrowly to broadly triangular, glabrous. Corolla campanulate, lobes 3.5–4 × 1.5 mm, oblong, green or white, glabrous on both faces. Corona lobes 2–3 mm long, cucullate, lacking an apical tongue, pinkish cream or white. Anther wings 1 mm long. Stylar head flat. Follicles not seen. (Fig.
Known from a single collection in eastern Angola and one in NW Zambia. The Angolan population consisted of several scattered individuals on a broad open sandy plain just above the water table. Both the Angolan and the Zambian localities are on Kalahari sand deposits, and the Angolan collection was associated with common geoxylic suffrutices of the region such as Parinari capensis Harv., Syzygium guineense subsp. huillense (Hiern) F.White, Eugenia malangensis (O.Hoffm.) Nied., Lannea gossweileri subsp. gossweileri and Cryptosepalum sp. aff. mimosoides Welw. ex Oliv. Altitude 1100–1300 m. (Map
Asclepias minutissima is known from two localities some 350 km apart, but is inconspicuous and easily overlooked and is likely to be more common than the herbarium records suggest. Both localities are in nutrient-deficient sandy environments unsuitable for agriculture, and with little threat of habitat transformation as human settlements are few and far between. The new species is therefore provisionally assessed as Data Deficient.
Zambia. Mwinilunga District, 16 km along road from Matonchi Farm, 11°39'S, 24°03'E, fl. 17 November 1962, Richards 17269 (K).
Only three species of Asclepias were reported from Angola by
Asclepias randii is a much more robust plant than the other species and has pubescent stems and leaves. A. aurea and A. minor are glabrous and are slender herbs. Asclepias minor has a corona which is much reduced, not even reaching the base of the anther wings and has a short ventral appendage. So the species most similar to our new collection appears to be the highly variable A. aurea, which occurs across Namibia, southern Angola, Zambia, the Katanga region of the D.R.Congo, Zimbabwe, northern provinces of South Africa, Eswatini (Swaziland) and Lesotho (
Asclepias aurea has rotate to reflexed corolla lobes, corona lobes which radiate from the column and are extended into a long distal tongue, and longer peduncles. In addition to the rather subtle morphological characters that distinguish the new species from A. aurea, its ecological requirements, close to the water table on leached Kalahari sand, are probably also significant. Asclepias aurea occurs on richer soils. The new taxon was mentioned by
Cochlospermum adjanyae differs from all African species of the genus in possessing palmatisect rather than palmatifid or lobed leaves, with discrete leaflets rather than partially connate lobes.
Angola. Moxico Province: Lungué-Bungo valley, 50 km S of Munhango, near Lungué-Bungo bridge, 12°36'50"S, 018°47'59"E, fl. 20 November 2019, D.Goyder & A.Gomes 9002 (holotype: K (K001334241); isotypes: INBAC, LUBA, PRE).
Geoxylic suffrutex forming diffuse but discrete patches several metres across; above-ground stems 10–20 cm tall, sub-erect, glabrous, burned off in the dry season. Leaves palmate with (4–)5 leaflets; stipules c. 3 mm long, narrowly triangular; petioles (2–)4–6 cm long, glabrous except for a minute rusty pubescence at the junction with the leaflets; leaflets reducing in size from the central leaflet to the lateral and basal ones, central leaflet 3–5 cm long, 1.2–2 cm wide, elliptic to slightly obovate, acute or occasionally obtuse apically, the base somewhat cuneate, margins serrate at least in the upper half, glabrous except for a minute rusty pubescence at the junction with the petiole adaxially. Inflorescences minutely rusty-puberulent, terminal on the leafy shoots, with 1–3 flowers; peduncles 2.5–3 cm long; sepals subequal, 11–15 × 7–8 mm, broadly ovate or elliptic, rounded apically, minutely rusty-puberulent and with occasional dark streaks, the outer pair more deeply coloured than the inner three; petals c. 3 × 2–2.3 cm, obovate, rounded or slightly emarginate, bright yellow with linear red streaks. Stamens numerous (80+), yellow; anthers c. 5 mm long, straight or weakly curved, apical pore 0.5–1 mm long. Ovary c. 2 mm in diameter, glabrous. Fruit not seen. (Figs
Found only once in flower in grassland rich in geoxylic suffrutices on deep Kalahari sand. Material in bud had been encountered, but not collected, the day before in a similar grassland some 15 km to the NW along with the new species of Baphia described below and other geoxyles such as Sclerocroton oblongifolius (Müll.Arg.) Kruijt & Roebers, Parinari capensis, Entada arenaria Schinz and Englerophytum magalismontanum (Sond.) T.D.Penn. The fact that this conspicuously flowered species was seen only on the November 2019 expedition and not on earlier ones through the same valley system (late rainy season; early and mid-dry season) suggests that populations are highly localised and that the flowering period is short, at the end of the dry season, and perhaps dependent on rainfall following fire. Altitude 1285–1340 m. (Map
The specific epithet honours Adjany Costa who was part of the core National Geographic Okavango Wilderness Project headwaters team from the first expedition in 2015 until she left to pursue academic studies at Oxford University in 2019. Initially, she supported freshwater fish specialists Paul Skelton and Ben van der Waal, recording and preserving freshwater fish diversity from the Cuito source to the Delta, before developing the communities programme with Chris and Steve Boyes. She was a National Geographic Young Explorer, starred in the National Geographic documentary film “Into the Okavango”, and was awarded the United Nation’s Young Champions of the Earth Prize for Africa in 2019.
While Cochlostemum adjanyae is known from a single locality, these geoxyle-rich grasslands are not currently threatened as this nutrient-poor sandy environment is not conducive to agricultural development. The environment does not support many human settlements, which are few and far between. The species is probably best assessed as Data Deficient.
Cochlospermum Kunth is a pantropical genus of around 16 species of trees, shrubs and geoxylic suffrutices, or if expanded to include the herbaceous neotropical genus Amoreuxia DC., 20 species. C. noldeae Poppend. from NE Angola, C. macnamarae Hislop, K.R.Thiele & Brassington and C. arafuricum Cowie & R.A.Kerrigan from Australia (
Species of Cochlospermum subg. Cochlospermum have a single apical pore to the anthers, and in addition to growth form, can be distinguished by leaf indentation or lobing, indumentum, and the position and timing of flowers on the shoots. A collection made in the geoxyle-rich Lungué-Bungo valley grasslands of Moxico in November 2019 is unique in the African species in having palmatisect leaves, divided to the base rather than being merely lobed. The only other taxon in the subgenus with this leaf character is the nomenclaturally illegitimate C. gillivrayi subsp. gregorii (F.Muell.) Poppend., an Australian tree. The Angolan material is described as a new species. It is perhaps closest morphologically to C. wittei Robyns from the Upemba region of Katanga. Cochlospermum wittei is also associated with savanna and woodland on sand plateaux (
1 | Trees or shrubs at least 4 m tall | C. angolense Welw. ex Oliv. |
– | Geoxylic suffrutices or low shrubs with annually produced shoots less than 4 m tall | 2 |
2 | Flowering mostly near ground level after fires and before the development of leafy shoots; savanna regions north of the equator | C. tinctorium A.Rich. |
– | Flowering towards the tip of leafy shoots | 3 |
3 | Leaves 3-lobed, the apex of the lobes attenuate; anthers 7 mm long; NE Angola: Malange |
C. noldeae Poppend. |
– | Leaves mostly 5–7-lobed, the apices acute, obtuse or rounded; anthers 4–6 mm long | 4 |
4 | Leaves palmately compound with leaflets free to the base; eastern Angola: Moxico | C. adjanyae |
– | Leaves palmately lobed, the lobes connate for at least some of their length | 5 |
5 | Leaves with lobes connate for at least half their length, lobes rounded or obtuse apically; leaves generally silvery-white beneath; West African savannas | C. planchonii Hook.f. ex Planch. |
– | Leaves with lobes connate for less than half their length, lobes obtuse or acute apically; leaf indumentum variable | 6 |
6 | Leaves with lobes connate for ¼–½ of their length; Central African Republic | C. intermedium Mildbr. |
– | Leaves with lobes connate for less than ¼ of their length; DR Congo: Katanga | 7 |
7 | Leaves glabrous beneath; sepals minutely puberulous; ‘forest’ understorey | C. wittei subsp. wittei |
– | Leaves silvery-white beneath; sepals minutely tomentose, greyish; savanna on Kalahari sand | C. wittei subsp. incanum (Robyns) Poppend. |
Ilse von Nolde, who lived in Quela, eastern Malange Province where she and her husband farmed coffee, made important collections of plants from the region between 1928 and 1938 (
Differs from E. tubulascens (Briq.) M.Ashby in the little-branched habit, the sessile rather than petiolate and linear rather than elliptic leaves, the fewer-flowered (4–6-flowered rather than 6–10-flowered) verticils in the inflorescence, the pale violet rather than pinkish white flowers and the longer calyx (fruiting calyx 8–9 mm long rather than 5.5–6 mm).
Angola. Moxico Province: Confluence of Cuito River and its 1st tributary, the Kalua River, c. 65 km SSW of Munhango, 12°44'55"S, 018°21'16"E, fl. 20 Oct. 2016, D.Goyder & F.Maiato 8762 (holotype: K (K001333409); isotypes: INBAC, LUBA).
Aromatic perennial suffrutex with few stems arising from a thick woody rootstock; stems erect, 15–30 cm tall, branched near the base, square in section above, more rounded below, pubescent with both glandular and eglandular hairs. Leaves verticillate, ascending, sessile, linear and folded along the midvein, 2.5–5 cm long, 0.1–0.2 cm wide, pubescent. Inflorescence lax with 4–6-flowered verticils 1.5–2.5 cm apart; bracts lanceolate or narrowly ovate, 3.5–4.5 mm long; pedicels 1–5 mm long, longer in lower verticils. Calyx 5 mm long at anthesis, pubescent with spreading hairs and sessile glands, posterior lip purplish; fruiting calyx 8–9 mm long. Corolla pale violet, 8–10 mm long; tube 6–7 mm long, straight, parallel-sided, dilating at the throat. Filaments c. 0.5 mm long, glabrous or pilose. Ovaries pubescent at apex. (Fig.
Although known from a single locality, Endostemon palustris occupies a habitat that is extensive within the upper catchment of the Cuito and its tributaries. The area is not threatened with agricultural development being both nutrient poor and many kilometres from any human habitation, but with so little information the species is provisionally assessed as Data Deficient.
Endostemon N.E.Br. is an isolated genus of 20 species within the tribe Ocimeae, with two centres of endemism – Angola and the Horn of Africa. It can be recognised within the Orthosiphon Benth. group of genera by its short, villous staminal filaments, an expanded shield-like base to the style, and pollen with alternating wide and narrow mesocolpia (
Most similar morphologically to B. massaiensis Taub., from which it can be readily distinguished by its geoxylic lifeform, flowering and fruiting at ground level on short, prostrate above-ground shoots, and the villous suture of the keel petal.
Angola. Moxico Province: tributary of the Lungué-Bungo River 42 km SSE of Munhango, 12°31'34"S, 018°40'13"E, fl. 22 October 2016, D.Goyder & F.Maiato 8772 (holotype: K (K001333933); isotypes: INBAC, LUBA).
Geoxylic suffrutex forming large patches; above-ground shoots prostrate, 5–10 cm long, pubescent, arising from extensive woody below-ground stems. Leaves unifoliolate; stipules linear, 3–4 mm long, densely pubescent with silvery hairs; petiole 3–5 mm long, pulvinus barely apparent; leaflet narrowly obovate-oblong and generally folded along the midvein, 4–6 cm long, 1–2 cm wide, obtuse apically, tapering somewhat towards the base, ± glabrous except for the major veins beneath. Inflorescences with an indumentum of greyish or golden hairs; flowers 1–3 in sessile or subsessile axillary fascicles; pedicels 15–20 mm long; bracteoles 1–2 mm below the calyx, caducous. Calyx 7–10 mm long, spathaceous. Petals white, the standard with a yellow triangular mark towards the base; standard 10–12 × 8–10 mm, emarginate; wings c. 10 × 2 mm; keel 10–12 × 2.5–3 mm, lower suture villous distally. Stamens 10, free. Ovary 6–9 mm long, villous, with a glabrous upturned style. Legume c. 7 cm long, c. 1.5 cm wide, brown. Seeds c. 10 mm long, dark brown or black. (Figs
Flowering in October and November in the late dry season or at the onset of the rains; fruiting in February. Known from three sites, the first in the Lungué-Bungo river system of western Moxico Province at an elevation of around 1350 m, the second in the Cusseque River system of Bié Province some 200 km to the SW of the Moxico site at around 1530 m, and the third just over the watershed into the Cacuchi River valley which drains into the Rio Cuchi and is around 1540 m. The three sites are similar topographically, with broad fossil river terraces and sandy alluvial deposits rich in geoxylic suffrutices (
Baphia arenicola is known from three localities, two of which have vouchered herbarium collections. TFO project made many unvouchered observations of the plant in the Cusseque and Cacuchi River valleys and it is clear that there are extensive populations of this species. There are no significant threats to these nutrient-poor grasslands rich in geoxylic suffrutices as the environment is not conducive to agricultural development. B. arenicola is therefore provisionally assessed as Least Concern.
Angola. Bié Province: Cusseque, TFO core site relevé 23324, 13°41'53"S, 017°06'43"E, fl. 29 October 2011, Finckh 132753 (HBG, K); Cusseque, TFO core site relevé 23349, 13°41'53"S, 017°07'48"E, fl. 2 November 2011, Revermann 132895 (HBG, K); Cusseque village, Chitembo, 13°43'21"S, 017°05'53"E, fr. 17 February 2014, Gomes & Maiato 161 (LUBA); Cusseque 15 September 2019, Finckh 145383A (HBG, LUBA); Cusseque, 28 January 2020, Finckh 145352B (HBG, LUBA).
Baphia Afzel. ex G.Lodd is a genus of around 50 species of woody legumes which has diversified across tropical Africa (
Most floral characters invite comparison with the locally common woodland species B. massaiensis, with its spathaceous calyx split longitudinally down a single line, bracteoles longer than wide and positioned shortly below the apex of the pedicel, glabrous staminal filaments, and pubescent ovary. The keel petal which is somewhat villous along its line of fusion, however, suggests links to B. bequaertii De Wild., another miombo woodland species of the region. Preliminary molecular analyses by one of us (PM) places the new taxon close to the latter species, with estimated divergence times from B. bequaertii less than 1 mya. Divergence times from B. massaiensis, on the other hand, are estimated to be between 11 and 27 mya.
Ancylanthos fulgidus Welw. ex Hiern in Oliv. (ed.), Fl. Trop. Afr. 3: 159 (1877). Type: Angola, Huíla, Mumpulla to Lopollo, Oct. 1859, Welwitsch 3160 (lectotype: LISU (LISU208624) designated here; paralectotypes: BM, K, LISU, P, PRE). (Basionym).
Ancylanthos rubiginosus Desf., Mém. Mus. Hist. Nat. 4: 5 (1818). Vangueria rubiginosa (Desf.) Lantz, Pl. Syst. Evol. 253: 181 (2005), non V. rubiginosa K.Schum., Bot. Jahrb. Syst. 23: 457 (1897) [= Rytigynia rubiginosa (K.Schum.) Robyns, Bull. Jard. Bot. État Bruxelles 11: 209 (1928)]. Type: Angola [without locality or collector, but probably Benguela/Huila Plateau, 1785–1787, J.J. da Silva (Exell & Mendonça 1956: IX & XI, following note on specimen at P)] (holotype: P (P00138559)).
Ancylanthos ferrugineus Welw., J. Trav. Nat. Hist. 1: 29 (1868), nomen nudum.
Ancylanthos bainesii
Hiern in Oliv. (ed.), Fl. Trop. Afr. 3: 160 (1877). Type: Baines s.n. (lectotype: K (K000412071) designated by
Vangueria fulgida is a small woody species with conspicuous orange flowers. It can form single-stemmed plants to 1.5 m in height, but over much of its range it behaves as a geoxylic suffrutex, forming patches of much shorter above-ground shoots that are burned off each year. It is found mostly on Kalahari sands and is widely distributed across Angola, western Zambia, Botswana, Namibia and Zimbabwe (
Molecular studies of the tribe Vanguerieae A.Rich. ex Dumort. (
The earliest synonym listed by
Ancylanthos bainesii
AG, DJG & FMPG are most grateful to Steve Boyes and John Hilton of the Wild Bird Trust for inviting us to participate in the National Geographic Okavango Wilderness Project. The Angolan Ministry of the Environment in Luanda, in particular Adjany Costa and Dra Paula Francisco, facilitated access and permits. We are also grateful to Chris Boyes for permission to use his photograph of Cochlospermum adjanyae.
The authors have declared that no competing interests exist.
No ethical statement was reported.
No funding was reported.
Conceptualization: DJG. Investigation: DJG. Writing – original draft: DJG. Writing – review and editing: PM, AJP, FMPG, ND, AG, MF.
David J. Goyder https://orcid.org/0000-0002-3449-7313
Manfred Finckh https://orcid.org/0000-0003-2186-0854
Amândio Gomes https://orcid.org/0000-0002-7812-5490
Francisco Maiato P. Gonçalves https://orcid.org/0000-0002-8859-7491
Paulina Meller https://orcid.org/0000-0001-6711-4385
Alan J. Paton https://orcid.org/0000-0002-6052-6675
All of the data that support the findings of this study are available in the main text.