A brief botanical survey into Kumbira forest, an isolated patch of Guineo-Congolian biome
expand article infoFrancisco Maiato Pedro Gonçalves, David John Goyder§
‡ Herbarium of Lubango, ISCED-Huíla (Lubnago, Angola) and University of Hamburg, Hamburg, Germany
§ Herbarium, Royal Botanic Gardens, London, United Kingdom
Open Access


Kumbira forest is a discrete patch of moist forest of Guineo-Congolian biome in Western Angola central scarp and runs through Cuanza Norte and Cuanza Sul province. The project aimed to document the floristic diversity of the Angolan escarpment, a combination of general walk-over survey, plant specimen collection and sight observation was used to aid the characterization of the vegetation. Over 100 plant specimens in flower or fruit were collected within four identified vegetation types. The list of species includes two new records of Guineo-Congolian species in Angola, one new record for the country and one potential new species.


Kumbira forest, Guineo-Congolian, floristic diversity


Angola lies almost wholly within the southern zone of tropical grassland, bounded by tropical rain forest of the Congo in the north and by the Kalahari Desert in the south (Shaw 1947). In general the country comprises four main physiographic components: a coastal plain from the Atlantic seaboard of approximately 200 m above sea level, and from 12 to 200 km wide; a narrow steep escarpment, from 200 to 1000 m altitude; an interior plateau, occupying nearly 80% of the country lying between 1000 and 1600 m, and a mountain belt rising above the escarpment and plateau of about 2620 m in central plateau (Huntley and Matos 1994).

Between the Karoo-Namib phytochorion of the coastal belt and the Brachystegia dominated Zambesian phytochorion of the interior plateau, a discontinuous series of moister vegetation type extends southwards from the Guinea forest and Congo savanna systems following the escarpment (Huntley 1974, White 1983). The Angolan escarpment is dominated by semi-deciduous forest and a mosaic of forest-savanna and gallery forest of Guineo-Congolian affinity, which is restricted to the interior of Cabinda province and large but discontinuous patches of forest in Zaire, Uíge, Cuanza Norte and Cuanza Sul provinces (Barbosa 1970, Huntley 1974).

These vegetation formations, as referred above, cover large areas of Cabinda with a tree strata of about 30 to 40 m height, while in the south are restricted to extensive “Coffee forests” in Dembos, Cazengo and Gabela regions (Huntley 1974). In this latter Kumbira forest, located in Conda, Cuanza Sul province is no doubt the most important and, probably the most southerly and most isolated patch of this biome, with various elements of Congo basin and West African affinities, dominated by genera Celtis, Morus, Albizia, Bombax and Pterocarpus (Barbosa 1970).

To these formations can be added the afro-montane forests, also of great biogeographic interest (Hall 1960a), restricted to small and isolated patches of forest in Benguela, Huambo and Huíla provinces. The total area of these forest patches was estimated to be approximately 200 ha, the best known of which is Mount Moco in Huambo province which provides habitat for its flora and avifauna (Huntley 1974, Olmos 2008 unpub., Maiato 2009, Mills 2010). The Angolan escarpment with approximately 1000 km of extent is unique, beautiful and constitutes the section of the Great Escarpment of Southern Africa poorly known in terms of its biodiversity, but high level of endemism (Huntley and Matos 1994, Figueiredo 2008, Clark 2011).

Despite the recent published checklist of Angolan vascular plants (Figueiredo and Smith 2008), data on the Angolan flora is mostly limited to the literature of pre-independence era, and current knowledge on Angolan plants is poor and restricted to isolated and focused studies carried out by individuals or institutions. Increasing interest and efforts are being made in order to document and obtain baseline data of areas with high socio-ecological importance (for example The Future Okavango Project – and the Okavango Wilderness Project – which focus on the Cubango and Cuito river catchments respectively). However large parts of the country, such as the Angolan escarpment, remain to be studied biologically.

In terms of biodiversity, only the avifauna has been investigated in any detail (Hall 1960a, Cagan and Riley 2005, Mills 2010, Cáceres et al. 2015). Recommendations have being made to undertake botanical surveys in the Kumbira area (Barbosa 1970, Huntley 1974). The ecological importance of Kumbira forest in maintaining the highest number of Angolan endemic avifauna highlighted the area as important for conservation of biodiversity. But at the same time concerned due to the increasing human pressure (Huntley 2011 unpub.). To fill in the gap in terms of botanical diversity, we undertook a botanical survey into Kumbira forest. Here we document our current knowledge of the plant diversity and phytogeographical affinities of Kumbira Forest, the findings are based primarily on a rapid botanical assessment conducted between 10th and 18th June 2014.


Study area

The central escarpment of western Angola on which Kumbira forest is part of, runs through Cuanza Norte and Cuanza Sul provinces. Barbosa (1970) recognized three types of moist high forest along this escarpment occurring in discrete geographic and climatic zones. North to south, these are:

i. forests of subtype Cazengo to the north of the Cuanza River;

ii. forests of subtype Amboim, between the Cuanza and Keve (Queve) Rivers, of which the most significant is the area of forest around Gabela;

iii. forests of subtype Uku (Vila Nova do Seles) to the south of the Keve River;

Kumbira forest (11°07.00'S; 014°17.00'E) is a discrete patch of moist forest vegetation in this third zone, SW of Conda (Figure 1), where cloud gathers under the knife-edge ridge of the Njelo mountain which reaches around 1500 m in elevation and prevents the cloud from moving further inland. The forest forms part of about 200 000 ha of semi-deciduous moist forest (Cagan and Riley 2005) and occurs on middle altitude slopes at around 700–900 m. Below this altitude dry open vegetation predominates, and at higher elevation the forest gives way to woodland and ultimately open rocky mountain slopes into rocky gorges in Njelo mountain.

Figure 1. 

Map of Angola and its provinces, Cuanza Sul province highlighted and Kumbira in Conda municipality (black dot). Kumbira forest (top), the forest with Njelo mountain in the background (bottom).

Kumbira forest is considered to be part of Gabela Important Bird Area (Dean 2001), and holds unique and threatened bird community (Cagan and Riley 2005). Although its ornithological importance the area is not protected by law, the proposed implementation of Gabela Natural Reserve, recently reinforced (Huntley 1974; Huntley 2011 unpub.) is unfortunately far from being achieved.

Sampling strategy

The botanical team consisted of staff from the Royal Botanic Gardens Kew, UK and from the Herbarium of Lubango, Angola. The team used a combination of general walk-over survey (Filgueiras et al. 1994), plant specimen collection and sight observations to aid the characterization of the vegetation and the compiling of an inventory of the plant species present. Over 100 herbarium specimens were collected, mostly of flowering and fruiting plants, but some sterile collections (lacking flowers or fruits) were made of the more common or important species which were not flowering or fruiting at the time of our visit.

Due to the limited time available, no plot-based surveys to quantify the composition of the different vegetation types were conducted. Plant specimens were collected in duplicate or triplicate, with one set deposited in Lubango for incorporation into the main collection, the remaining set(s) being taken back to the UK where they were identified by comparison with material housed in the Kew herbarium. A range of botanical specialists were consulted to help in the identification of difficult plant groups.

Due to the paucity of useful literature on Angolan plants, and to the limited previous botanical work in the region, it was not possible to name all collections made, particularly sterile material, but we were able to name the large majority. Those named only to genus are nevertheless included within the checklist in Appendix 1. We have not included the additional species listed by Gossweiler and Mendonça (1939) and Barbosa (1970) from this forest subtype in the Appendix 1, as it is not possible to say which forest patch they are from.

The recent Angolan plant checklist by Figueiredo and Smith (2008) was used as the baseline for assessing species records and distributions in the country. The African Plants Database ( and the World Checklist series ( for a number of plant families were used as the standards for up-to-date taxonomy of African plant species.


From the vegetation survey carried out in Kumbira forest, four main vegetation types were identified. Here we provide a very brief overview of the forest types and their dominant species as recorded by the rapid botanical survey. The habitat types outlined here are also assigned to each of the species listed in (Appendix 1).

Moist high forest (F)

Surveyed at former coffee plantation at foot of Serra Njelo, c.7 km SW of Conda [centred on 11°09.26'S, 014°17.56'E]). The canopy trees in this part of the forest were retained as shade for the coffee grown underneath (Coffea canephora, C. robusta), and there has been considerable regrowth of forest understorey since the plantations were abandoned. The canopy is c. 25-30 m high, and the commonest tree is the seasonally deciduous Albizia adianthifolia, Trema orientalis, Markhamia zanzibarica, Antidesma venosum and several species of Ficus are common elements. We also encountered Anthocleista schweinfurthii, Cola welwitschii, Pteleopsis diptera, Synsepalum cerasiferum, Turraea vogelii and Vitex welwitschii in some areas. An arborescent Dracaena and a species of Erythrina were also noted (sight records only).The understorey was rich in Rubiaceae, and the herbaceous flora included many ferns and occasional epiphytes, Justicia paxiana is recorded from Angola for the first time.

Figure 2. 

Turraea vogelii Hook.f. ex Benth, Cola welwitschii Exell & Mendonça ex R. Germ., Pancovia golungensis (Hiern) Exell & Mendonça, Cochlospermum angolense Welw. ex Oliv., Inga vera Willd. subsp. vera, Pavetta gossweileri Bremek, Pittosporum viridiflorum Sims, Clerodendrum poggei Gürke

Submontane forest/woodland (W)

(Upper margins of Moist High Forest). The moist high forest gives way to deciduous woodland at higher elevations (above c. 900 m). Characteristic trees and shrubs include Harungana madagascariensis, Dombeya rotundifolia, Cochlospermum angolense, Grewia flavescens, Pittosporum viridiflorum and Hymenocardia acida. We also encountered Podocarpus milanjianus, an afromontane element, in the gullies at high altitude.

Inga-dominated former coffee plantation (Inga)

(Surveyed at: mostly at Monte Belo estate, former coffee plantation at foot of Serra Njelo, c. 11 km SW of Conda [centred on 11°10.68'S, 014°16.36'E]). The exotic, evergreen Inga trees (Inga vera from northern South America) planted initially as shade for the coffee are now overgrown and let little light through the canopy, seriously reducing the development of a shrub or herb layer except in gaps. Inga vera was recently referred as one of the most impressive and worrying invasive species of western Angola, forming dense stands in localized sites as observed in Kumbira forest (Rajmánek et al. 2016). Nevertheless, we did record a variety of ferns and other herbaceous plants in this area, in addition to the very large fig Ficus saussureana, the first record of this Guineo-Congolian species from Angola. We observed some regeneration of Inga from seed, demonstrating its potential as an invasive species. A second exotic mimosoid legume tree was encountered in this area – Leucaena leucocephala – also native to the New World.

Ruderal and secondary habitats (Sec)

There is extensive encroachment of agriculture in the vicinity of villages, roads and tracks, and extraction of timber trees in some areas of forest. We did not survey this beyond making occasional observations. The commonest subsistence crops grown in the region are manioc (cassava) and plantain. The oil palm Elaeis guineensis is widely planted. We also encountered dense stands of the pigeon pea Cajanus cajan in one area. Disturbed areas of forest, recently cleared, had large patches of the invasive shrub Solanum mauritianum.

Plant diversity

Our records are a brief snapshot, based on a visit of just six collecting days, at just one time of the year (June). They are by no means comprehensive. Nevertheless, we recorded 92 species from the forest.

New plant records for Angola

We report two new records of Guineo-Congolian forest species for Angola – the tree Ficus saussureana and the herb Justicia paxiana. In addition, Tarenna pavettoides is newly recorded for the country, and may represent an undescribed subspecies. We also report a potential new species in Rytigynia sp. nov. (Appendix 2).


Phytogeographical context

Coastal regions of Angola are strongly influenced by the Benguela cold-water current which reduces surface evaporation and rainfall. Although the current moves off-shore north of Namibe, corresponding with the northern limit of the Namib Desert, coastal vegetation north of the desert remains dry, and is dominated by dry scrub woodland and succulents such as aloes, euphorbias and baobabs.

Kumbira Forest is a very localized patch of forest between this dry coastal vegetation and the moist savannas of the plateau. Barbosa (1970) regarded the forest as typically Zambezian, the canopy dominated by Albizia species that lose their leaves in the dry season. Barbosa also quoted from Gossweiler and Mendonça (1939) who commented on the high percentage of Ficus species. However, we found that most of the species within the forest are Guineo-Congolian rather that Zambezian in floristic affinity sensuWhite (1983) - the species recorded are generally widespread within the Guineo-Congolian phytogeographical region but often rare or previously unrecorded in Angola.

And it must be noted that Albizia adianthifolia which we encountered in the forest, is distributed widely across tropical Africa, and not just in the Zambezian region. Kumbira Forest must be one of the most southerly areas in western Angola with significant Guineo-Congolian vegetation. The upper limits of the forest contain occasional afromontane elements, and the forest merges into moist woodland characterized by widespread species.


We surveyed key sites in a small portion of Kumbira Forest in Cuanza Sul Province, reporting on diversity and floristic affinities of the flora. The vegetation types were identified: moist high forest, submontane forest & woodland, and Inga-dominated former coffee plantation. In addition, ruderal or disturbed areas also occur.

Species composition is overwhelmingly Guineo-Congolian, and this forest represents one of the most southerly areas of such vegetation along the Angolan escarpment. Most species have a wide geographic distribution, but some such as Pavetta gossweileri are more restricted, and are reported from just Cuanza Sul and Cuanza Norte. Deinbollia laurifolia is restricted to riverine lowland habitats from Cuanza Sul to Cameroon.

Much of the area surveyed had good regeneration of the understorey since coffee cultivation ceased, but we observed large trees being taken out of the forest, and evidence of village agriculture encroaching in a number of places. Areas formerly planted with New World legume species, particularly Inga vera, were in less favourable condition as the overgrown Inga shades out the understorey. There is some evidence of this species reproducing and spreading naturally, and we observed other potentially invasive species such as Solanum mauritianum. In addition, some coffee estates are being reestablished, which will reduce the extent of undisturbed or recovering forest further. Three species are newly recorded for Angola, Ficus saussureana, Justicia paxiana and Tarenna pavettoides. We also report a potential new species of Rytigynia.


The authors are extremelly grateful to the Angolan Ministry of Science and Technology for funding this project, under a bilateral cooperation programme with South African Department of Science and Technology. F.M.P.G thanks the staff of the Herbarium of Lubango for all support given and José Camôngua Luís for preparing the map of the study area. D.J.G thanks also the botanists consulted for identification of difficult plant groups.


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Appendix 1

List of the vascular plants collected and observed in Kumbira Forest, Cuanza Sul, Angola. Habitat types in Kumbira forest relate to the phytochoria recognised by White (1983) as follows: Forest = I. Guineo-Congolian; Woodland = VIII. Afromontane. Most species found in secondary forest and Inga-dominated forest are also Guineo-Congolian in distribution and affinity.

Family Species Habitat type Collectors Voucher number(s)
Dennstaedtiaceae Blotiella currorii (Hook.) R.M.Tryon Forest Goyder & Maiato 7701, 7719, 7720, 7722
Dryopteridaceae Bolbitis gemmifera (Hieron.) C.Chr. Forest Goyder & Maiato 7702, 7726
Lomariopsidaceae cf. Lomariopsis Forest Goyder & Maiato 7718
Polypodiaceae Platycerium sp. Forest Sight record
Pteridaceae Pteris catoptera Kunze var. catoptera Inga Goyder & Maiato 7783
Thelypteridaceae Christella dentata (Forssk.) Brownsey & Jermy Forest Goyder & Maiato 7721, 7727
Podocarpaceae Podocarpus milanjianus Rendle Woodland Goyder & Maiato 77800
Araceae Culcasia angolensis Welw. ex Scott Forest Goyder & Maiato 7802
Asparagaceae Asparagus africanus Lam. var. puberulus (Baker) Sebseb Forest Goyder & Maiato 7751
Dracaena sp. Forest Sight record
Commelinaceae Aneilema beniniense (P.Beauv.) Kunth Forest Goyder & Maiato 7741
Palisota cf. schweinfurthii C.B.Clarke Forest Goyder & Maiato 7715
Dioscoreaceae Dioscorea praehensilis Benth. Forest Goyder & Maiato 7758
Marantaceae Marantochloa leucantha (K.Schum.) Milne-Rendh. Forest Goyder & Maiato 7778
Acanthaceae Acanthus montanus (Nees) T.Anderson Forest Goyder & Maiato 7707
Justicia flava (Vahl) Vahl Secondary forest Sight record
Justicia paxiana Lindau Forest Goyder & Maiato 7717
Apocynaceae Motandra guineensis (Thonn.) A.DC Forest Goyder & Maiato 7804
Oncinotis sp. Forest Goyder & Maiato 7777
Bignoniaceae Markhamia zanzibarica (Bojer ex DC.) K.Schum. Forest Goyder & Maiato 7710
Bixaceae Cochlospermum angolense Welw. ex Oliv. Woodland Goyder & Maiato 7796
Cactaceae Rhipsalis baccifera (J.S.Muell.) Stearn Forest/Inga Sight record
Cannabaceae Trema orientalis (L.) Blume Forest Goyder & Maiato 7765
Cleomaceae Cleome sp. Forest Goyder & Maiato 7713
Combretaceae Combretum angolense Welw. ex M.A.Lawson Forest Goyder & Maiato 7712
Combretum collinum Fresen. Woodland Goyder & Maiato 7792
Combretum platypetalum Welw. ex M.A.Lawson Forest Goyder & Maiato 7757, 7766
Pteleopsis diptera Engl. & Diels Forest/woodland Goyder & Maiato 7725, 7793
Connaraceae Rourea thomsonii (Baker) Jongkind Forest Goyder & Maiato 7703
Dichapetalaceae Dichapetalum crassifolium Chodat Forest Goyder & Maiato 7704
Ebenaceae Diospyros heterotricha (Welw. ex Hiern) F.White Forest Goyder & Maiato 7704
Euphorbiaceae Acalypha paniculata Miq. Forest Goyder & Maiato 7762
Antidesma venosum E.Mey. ex Tul Forest Goyder & Maiato 7768
Croton gratissimus Burch. Forest Goyder & Maiato 7801
Mallotus oppositifolius (Geiseler) Müll. Arg. Forest Goyder & Maiato 7734
Gentianaceae Anthocleista schweinfurthii Gilg. Forest Goyder & Maiato 7774
Hypericaceae Harungana madagascariensis Lam. ex Poir. Woodland Goyder & Maiato 7791
Lamiaceaee Clerodendrum poggei Gürke Forest Goyder & Maiato 7736
Clerodendrum volubile P.Beauv. Forest Goyder & Maiato 7714
Vitex welwitschii Gürke Forest Goyder & Maiato 7770
Leguminosae: Mimosoideae Acacia pentagona (Schumach. & Thonn.) Hook.f. Forest Goyder & Maiato 7748
Albizia adianthifolia (Schumach.) W.Wight Forest Goyder & Maiato 7740, 7764
Inga vera Willd. subsp. vera Inga Goyder & Maiato 7759, 7764
Leucaena leucocephala (Lam.) de Wit Inga Goyder & Maiato 7780
Leguminosae: Papilionoideae Cajanus cajan (L.) Millsp. Secondary forest Goyder & Maiato 7784
Dalbergia saxatilis Hook.f.var. saxatilis Forest Goyder & Maiato 7769
Desmodium repandum (Vahl) DC Forest Goyder & Maiato 7742
Erythrina sp. Forest Sight record
Millettia drastica Welw. ex Baker Forest Goyder & Maiato 7787
Neonotonia wightii (Widht & Arn.) J.A.Lackey Forest Goyder & Maiato 7723
Psophocarpus scandens (Engl.) Verdc. Secondary forest Goyder & Maiato 7785
Loranthaceae Tapinanthus constrictiflorus (Engl.) Danser Forest Goyder & Maiato 7789
Malvaceae Cola welwitschii Exell & Mendonça ex R. Germ. Forest Goyder & Maiato 7730, 7747
Dombeya rotundifolia (Hochst.) Planch. var. rotundifolia Woodland Goyder & Maiato 7795
Grewia flavescens Juss. Woodland Goyder & Maiato 7794
Grewia floribunda Mast. Forest Goyder & Maiato 7744
Urena lobata L. Secondary forest Goyder & Maiato 7776
Meliaceae Turraea vogelii Hook.f.ex Benth Forest Goyder & Maiato 7728, 7756
Moraceae Ficus conraui Warb. Inga Goyder & Maiato 7761
Ficus saussureana DC. Inga Goyder & Maiato 7782
Ficus sp. Forest Goyder & Maiato 7739
Trilepisium madagascariense DC. Forest Goyder & Maiato 7733
Passifloraceae Adenia lobata (Jacq.) Engl. subsp. lobata Forest Goyder & Maiato 7752
Phyllanthaceae Hymenocardia acida Tul. Woodland Goyder & Maiato 7798
Thecacoris trichogyne Müll. Arg. Forest Goyder & Maiato 7716
Pittosporaceae Pittosporum viridiflorum Sims Woodland Goyder & Maiato 7797
Primulaceae Maesa welwitschii Gilg. Forest Goyder & Maiato 7709
Rhamnaceae Gouania longipetala Hemsl. Forest Goyder & Maiato 7724
Rubiaceae Bertiera orthopetala (Hiern) N.Hallé Forest Goyder & Maiato 7771
Coffea canephora Pierre ex A.Froehner Forest/Inga Goyder & Maiato 7786
Empogona glabra (K.Schum.) Tosh & Robbr. Forest/Inga Goyder & Maiato 7755
Mussaenda erythrophylla Schumach. & Thonn. Forest Goyder & Maiato 7803
Pavetta gossweileri Bremek Forest/Inga Goyder & Maiato 7760
Psychotria nigropunctata Hiern Woodland Goyder & Maiato 7799
Rothmannia longiflora Salisb. Forest Goyder & Maiato 7745
Rothmannia whitfieldii (Lindl.) Dandy Forest Goyder & Maiato 7746
Sherbournia bignoniiflora (Welw.) Hua Forest Goyder & Maiato 7705
Tarenna pavettoides (Harv.) Sim?subsp. nov. Forest/Inga Goyder & Maiato 7775
Rytigynia?sp. nov. Forest Goyder & Maiato 7788
Sapindaceae Deinbollia laurifolia Baker Forest Goyder & Maiato 7743
Pancovia golungensis (Hiern) Exell & Mendonça Forest Goyder & Maiato 7772
Paullinia pinnata L. Forest Goyder & Maiato 7767
Sapotaceae Synsepalum cerasiferum (Welw.) T.D.Penn. Forest Goyder & Maiato 7754
Solanaceae Solanum anomalum Thonn. Forest Goyder & Maiato 7711
Solanum mauritianum Scop. Secondary forest Goyder & Maiato 7738
Solanum terminale Forssk. Forest Goyder & Maiato 7749
Umbelliferae Steganotaenia araliacea Hochst. Inga Goyder & Maiato 7779
Urticaceae Urera trinervis (Hochst.) Friis & Immelman Forest Goyder & Maiato 7773
Violaceae Rinorea ilicifolia (Welw. ex Oliv.) Kuntze Forest Goyder & Maiato 7750
Vitaceae Cissus aralioides (Welw. ex Baker) Planch. Forest Goyder & Maiato 7790
Leea guineensis G.Don Secondary forest Goyder & Maiato 7708

Appendix 2

List of species collected in Kumbira forest which represent new records for Angola.

Family Species Notes
Acanthaceae Justicia paxiana Lindau Widely distributed in West Africa and the Congo Basin, but not recorded before from Angola
Moraceae Ficus saussureana DC. Widely distributed in West Africa and eastern and western margins of the Congo Basin, but not recorded before in Angola.
Rubiaceae Rytigynia ?sp. nov. We were unable to match this to any known species of the genus.
Rubiaceae Tarenna pavettoides (Harv.) Sim This species has one subspecies distributed across West Africa and northern limits of the Congo Basin, and other subspecies in Eastern and Southern Africa. It has not been reported from Angola before, and our collection may represent a new subspecies.