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Language editing: Will Simonson (Cambridge), and Proofreading Pal Translation of abstracts to Portuguese: Ana Filipa Guerra Silva Gomes da Piedade Page desing & layout: Marit Arnold, Klaus A. Hess, Ria Henning-Lohmann Cover photographs: front: Thunderstorm approaching a village on the Angolan Central Plateau (Rasmus Revermann) back: Fire in the miombo woodlands, Zambia (David Parduhn) Cover Design: Ria Henning-Lohmann

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Suggestion for citations: Volume: Revermann, R., Krewenka, K.M., Schmiedel, U., Olwoch, J.M., Helmschrot, J. & Jürgens, N. (eds.) (2018) Climate change and adaptive land management in southern Africa – assessments, changes, challenges, and solutions. Biodiversity & Ecology, 6, Klaus Hess Publishers, Göttingen & Windhoek.

Articles (example): Archer, E., Engelbrecht, F., Hänsler, A., Landman, W., Tadross, M. & Helmschrot, J. (2018) Seasonal prediction and regional climate projections for southern Africa. In: Climate change and adaptive land management in southern Africa – assessments, changes, challenges, and solutions (ed. by Revermann, R., Krewenka, K.M., Schmiedel, U., Olwoch, J.M., Helmschrot, J. & Jürgens, N.), pp. 14–21, Biodiversity & Ecology, 6, Klaus Hess Publishers, Göttingen & Windhoek.

Corrections brought to our attention will be published at the following location: http://www.biodiversity-plants.de/biodivers_ecol/biodivers_ecol .php Biodiversity & Ecology

Journal of the Division Biodiversity, Evolution and Ecology of Plants, Institute for Plant Science and Microbiology, University of Hamburg

Volume 6:

Climate change and adaptive land management in southern Africa

Assessments, changes, challenges, and solutions

Edited by

Rasmus Revermann1, Kristin M. Krewenka1, Ute Schmiedel1, Jane M. Olwoch2, Jörg Helmschrot2,3, Norbert Jürgens1

1 Institute for Plant Science and Microbiology, University of Hamburg 2 Southern African Science Service Centre for Climate Change and Adaptive Land Management 3 Department of Soil Science, Faculty of AgriSciences, Stellenbosch University

Hamburg 2018

RPlease cite the article as follows:

Chisingui, A.V., Gonçalves, F.M.P., Tchamba, J.J., Camôngua Luís, J., Rafael, M.F.F. & Alexandre, J.L.M. (2018) Vegetation survey of the woodlands of Huíla Province. In: Climate change and adaptive land management in southern Africa – assessments, changes, challenges, and solutions (ed. by Revermann, R., Krewenka, K.M., Schmiedel, U., Olwoch, J.M., Helmschrot, J. & Jürgens, N.), pp. 426-437, Biodiversity & Ecology, 6, Klaus Hess Publishers, Göttingen & Windhoek. doi:10.7809/b-e.00355 Vegetation survey of the woodlands of Huíla Province

António Valter Chisingui1*, Francisco M. P. Gonçalves1,2, José João Tchamba1, José Camôngua Luís1, Marina Filomena F. Rafael1, and José L. M. Alexandre1 1 Instituto Superior de Ciências de Educação, ISCED-Huíla, Rua Sarmento Rodrigues, 230 Lubango, Angola 2 Institute for Plant Science and Microbiology, University of Hamburg, Ohnhorststr. 18, 22609 Hamburg, Germany * Corresponding author: [email protected]

Abstract: We conducted a vegetation survey in the woodlands of Huíla Province, Angola, with the aim of investigating woodland tree communities and species associations. Vegetation sampling was conducted using vegetation plots of 1 000 m2 Biodiversity where all tree species with or above 5 cm trunk diameter (DBH) were measured. A total of 456 vegetation plots were assessed and a total of 32 080 individual trees measured. Vegetation classifi cation using the ISOPAM algorithm resulted in 13 distinct tree communities. The most dominant family was Fabaceae, subfamily Caesalpinioideae, followed by Combretaceae and Euphorbiaceae. The classifi cation resulted in seven tree communities belonging to the miombo woodlands, two tree com- munities from Mopane, two from the Baikiaea-Baphia-Terminalia woodlands, and two other distinct tree communities. In general, the miombo communities were the most diverse. The study represents the fi rst plot-based vegetation survey for the region, and will provide the basis for the elaboration of the fi rst vegetation map of Huíla Province.

Resumo: O levantamento da vegetação foi realizado nos bosques da Província da Huíla, Angola, com o objectivo de inves- tigar as comunidades arbóreas e associações de espécies. O processo de amostragem da vegetação foi realizado em parcelas de 1 000 m2, onde todas as árvores com diâmetro à altura do peito (DBH) igual ou acima de 5 cm foram medidas. Avaliámos no total 456 parcelas de vegetação, que resultou num total de 32 080 indivíduos medidos. A classifi cação da vegetação foi feita com recurso ao algorítimo ISOPAM que resultou em 13 comunidades distintas. A família mais dominante foi Fabaceae, subfamília Caesalpinioideae, seguida da família Combretaceae e Euphorbiaceae. A classifi cação da vegetação resultou em sete comunidades de miombo, duas comunidades de Mopane, duas comunidades de Baikiaea-Baphia-Terminalia, e duas outras comunidades distintas. Em geral, as comunidades de miombo foram as mais diversas. O estudo representa o primeiro levantamento feito na região com uso de parcelas, e poderá proporcionar as bases para a elaboração do primeiro mapa de vegetação da Província da Huíla.

Introduction remained unstudied in terms of species is no doubt that conservation actions are composition and distribution of vegeta- needed, aimed at preventing and mitigat- Angola harbours an enormous variety tion communities, as most of the previous ing ecologically harmful consequences of habitats, with the miombo woodlands documentation of tree species diversity caused by habitat modifi cations and land covering about 47% of the land area of conducted in the country was based com- use change (Simila et al., 2006). To do so the country (Huntley & Matos, 1994). monly on fl oristic itineraries and not on eff ectively fi rst requires knowledge of the Within Huíla Province, miombo is also detailed vegetation surveys (dos Santos, present ecosystems as well as plot-based the dominant vegetation type. Other im- 1982). inventories that document the fl oristic portant vegetation types include Afro- Huíla Province, like many parts of An- diversity and species composition of the montane forests and grasslands in the gola, faces challenges such as land deg- woodlands. area of the escarpment, and Baikiaea radation and deforestation, and as a result Due to the long period of civil war plurijuga and Colophospermum mopane thereof, loss of biodiversity (Cabral et al., experienced by Angola, information woodlands in the southwestern parts of 2011). The main drivers of these process- on vegetation is scarce and is generally the province. Although the region is prob- es are demand for agricultural land, fi re based on early botanical work, such as ably one of the most botanically studied frequency, fuelwood extraction, charcoal the phytogeographic map of Angola parts of the country and hosts one of the production, and rapid urban develop- (Barbosa, 1970). For the woody vegeta- largest botanical collections in Angola at ment (Röder et al., 2015; De Cauwer et tion of Angolan miombo, only a single the Herbarium of Lubango (LUBA), it al., 2016; Schneibel et al., 2017). There study is known, which resulted in the fi rst

426 C A Biodiversity

Figure 1: Map of Southern Africa with the location of the study area corresponding to Huíla Province (in red) and its administrative division into 14 municipalities. Plot locations are coloured by their corresponding woodland community types, derived from the ISOPAM algorithm.

provincial map of the Bié Province based Materials and methods Vegetation sampling on 144 plots (Monteiro, 1970). Recently, We used the preliminary classifi cation a few initiatives have started to docu- Study site of a time-series (2001-2013) of MODIS ment the fl oristic diversity of the region, The woody vegetation was assessed in satellite imagery and derived phenol- though attention has been paid mostly to the woodlands of Huíla Province, located ogy metrics to identify major vegetation areas of particular botanical interest, such in southwestern Angola. The province units (Stellmes et al., 2013). The map so as the Angolan escarpment (Barker et al., occupies an estimated area of about 78 obtained was used to create a random 2015; Gonçalves & Goyder, 2016; Gon- 879 km2, divided into 14 municipalities stratifi ed plot sample design across the çalves et al., in prep.). Studies addressing (Fig. 1). The climate of the region is con- vegetation units of the study area. A total fl oristic diversity and woody species re- sidered tropical, with dry and cold winters of 456 vegetation plots of 20 m x 50 m covery following disturbance of the mi- and temperate rainy summers; the mean were used to assess the vegetation; within ombo woodlands in south-central Angola annual temperature varies from 18°C in each plot, all woody species reaching a have also been conducted (Gonçalves et the highlands of Humpata to 20°C in the diameter at breast height (DBH) ≥ 5 cm al., 2017; Revermann et al., 2018). How- eastern parts of the province (Köppen- were measured. The taxonomy of woody ever, much remains to be done in terms Geiger, 1936). Annual precipitation in- species followed the Angolan checklist of vegetation assessment to characterise creases from 700 mm in the southwest of of vascular plants (Figueiredo & Smith, the main vegetation types and species as- Huíla Province to 1 000 mm in the east 2008). sociations of Huíla Province. Thus, this (Azevedo et al., 1972). According to Bar- study aims to provide an initial classifi ca- bosa (1970), Huíla Province comprises at Data analysis tion of the woodland plant communities least eight vegetation types. The wood- To understand variation in tree species of the region. This will ultimately lead to lands include the miombo, Angolan Mo- and diversity within each of the derived the creation of vegetation maps that are pane and the Zambezian Bakiaea-Baphia tree communities, we calculated species urgently needed as a tool for conservation woodlands. richness (S) and Shannon diversity in- planning and forest management. dex (H’). The number of individual tree

B E 6 2018 427 species belonged to subfamily Caesal- pinioideae, 9% to Detarioideae, and 5% to Papilionoideae, while the subfamilies Cercidoideae and Dialioideae were each represented by one species only. Other abundant families included Combretaceae (10%), Euphorbiaceae (9%), Burseraceae (7%), and other smaller families (Fig. 2). The species accumulation curve shows that the 456 plots used in this study were suffi cient to cover much of the variation observed and species diversity encoun- tered in the studied area. At 400 plots, the graph has not yet reached its asymptotic level but is starting to converge (Fig. 3). Biodiversity

Vegetation classifi cation The vegetation classifi cation of the ISO- PAM algorithm resulted in a dendrogram in which the tree species communities can be seen. At the second hierarchi- cal level of the dendrogram, fi ve major fl oristic groups were diff erentiated. The Figure 2: Number of tree species per botanical family found in Huíla Province; only families dendrogram featured 13 terminal clusters with more than two species are shown in the pie chart. corresponding to the 13 tree communities diff erentiated from the fourth hierarchi- species per plot was subject to a vegeta- Results cal level (Fig. 4). tion classifi cation using the ISOPAM al- The tree species communities and spe- gorithm in hierarchical capacity, which Tree species richness cies associations can be detailed as seen is based on the ordination scores from Within the 456 vegetation plots surveyed in Table 2. The fi rst tree communities isometric feature mapping and partition- in the woodlands of Huíla Province, we (Communities 1, 2, 3, 5, 6, 7, and 8), with ing around medoids (Schmidtlein et al., recorded a total of 32 080 tree individu- the exception of Community 4 [Com- 2010). These were performed in R Ver- als corresponding to 176 tree species of bretum collinum-Pericopsis angolensis sion 3.2.3 (R Development Core Team, 94 genera and 43 botanical families. The woodlands], constitute typical miombo 2017) with the package ISOPAM. We Fabaceae family was the most abundant woodlands with open, dense, or medium selected the third hierarchical level of the in the study, reaching 34% of the total dense tree canopy, sometimes with dense dendrogram to describe tree communities number of species. According to a recent understorey development (Figs. 7, 8, 9, and determined the diagnostic species us- classifi cation, the Leguminosae/Fabaceae 11, 12, 13, and 14). These communities ing the phi coeffi cient with a threshold of family comprises six subfamilies (Azani were dominated by multiple key miombo 30 and a p-value of p < 0.05. et al., 2017). Of all Fabaceae, 20% of species such as Brachystegia boehmii,

Figure 3: Species accumulation curve for trees with a diameter at breast height ≥ 5 cm measured within the sampling plots Figure 4: Community dendrogram from the ISOPAM algorithm classifi cation with tree com- of the study area. munities of Huíla Province; the numbered clades indicate the diff erent tree communities.

428 C A Table 1: Overview of the tree communities of Huíla Province, their diversity values, number C. mubango, and Ochna pulchra (Fig. 16). of indicator species, and total number of plots surveyed per tree community Communities of Baikiaea-Baphia also

ůƵƐƚĞƌ DĞĂŶ EŽ͘ŽĨ EŽ͘ŽĨ cover large parts of Bicuar National Park DĞĂŶ dƌĞĞŽŵŵƵŶŝƚŝĞƐ ŚŝĞƌĂƌĐŚŝĐĂů &ŽƌŵĂƚŝŽŶ ^ŚĂŶŶŽŶ ŝŶĚŝĐĂƚŽƌ ƉůŽƚƐƉĞƌ ƌŝĐŚŶĞƐƐ (BNP). Here, these woodlands formerly ůĞǀĞů ŝŶĚĞdž ƐƉĞĐŝĞƐ ĐŽŵŵƵŶŝƚLJ appeared associated with Schinziophyton ϭ͗ƌĂĐŚLJƐƚĞŐŝĂƐƉŝĐŝĨŽƌŵŝƐͲ KƉĞŶǁŽŽĚůĂŶĚƐ ϭ͘ϵϭ ϭϱ͘ϲϮ ϭϵ Ϯϵ WĂƌŝŶĂƌŝĐƵƌĂƚĞůůŝĨŽůŝĂ ϭ͘ϭ͘ϭ rautanenii, though at present, we found Ϯ͗:ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂͲ few and sparse trees at the woodland edg- KƉĞŶǁŽŽĚůĂŶĚƐ ϭ͘ϭϴ ϭϭ͘Ϯϯ ϭϬ ϯϵ ƌĂĐŚLJƐƚĞŐŝĂƐƉŝĐŝĨŽƌŵŝƐ ϭ͘ϭ͘Ϯ es of the park and also in the municipal ϯ͗ƌĂĐŚLJƐƚĞŐŝĂůŽŶŐŝĨŽůŝĂͲ KƉĞŶǁŽŽĚůĂŶĚƐ ϭ͘ϴϴ ϭϲ͘ϰϭ ϭϳ Ϯϵ limits of Quipungo and Chicomba. ŝŽƐƉLJƌŽƐŬŝƌŬŝŝ ϭ͘ϭ͘ϯ Community 11 [medium open wood- ϰ͗ŽŵďƌĞƚƵŵĐŽůůŝŶƵŵͲ DĞĚŝƵŵĚĞŶƐĞ Ϯ͘Ϭϳ ϭϴ͘Ϯϵ Ϯϰ ϰϵ WĞƌŝĐŽƉƐŝƐĂŶŐŽůĞŶƐŝƐ ϭ͘ϭ͘ϰ ǁŽŽĚůĂŶĚƐ lands] Patches of Baphia-Terminalia ϱ͗:ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂͲ KƉĞŶĚĞŶƐĞ were also found in the municipality of ŝƉůŽƌŚLJŶĐŚƵƐ ϭ͘ϲ ϵ͘Ϯϯ ϲ ϯϱ ǁŽŽĚůĂŶĚƐ ĐŽŶĚLJůŽĐĂƌƉŽŶ ϭ͘Ϯ͘ϭ Matala, and partially along Bicuar mov- ϲ͗:ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂͲ DĞĚŝƵŵĚĞŶƐĞ ing south on the way to Mulondo (Fig. ϭ͘ϰϯ ϴ ϯ ϱϯ ŽŵďƌĞƚƵŵĐŽůůŝŶƵŵ ϭ͘Ϯ͘Ϯ ǁŽŽĚůĂŶĚƐ 17). The understorey was commonly

KƉĞŶǁŽŽĚůĂŶĚƐ Biodiversity ϳ͗ƌĂĐŚLJƐƚĞŐŝĂƐƉŝĐŝĨŽƌŵŝƐͲ ǁŝƚŚĚĞŶƐĞ ϭ͘ϰ ϲ͘ϳϴ ϯ ϯϲ dominated by Mundulea sericea, Vitex WƚĞůĞŽƉƐŝƐĂŶŝƐŽƉƚĞƌĂ Ϯ͘ϭ͘ϭ ƵŶĚĞƌƐƚŽƌĞLJ mombassae, Ximenia americana and KƉĞŶǁŽŽĚůĂŶĚƐ ϴ͗:ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂͲ ǁŝƚŚĚĞŶƐĞ ϭ͘Ϯ ϲ͘Ϭϳ Ϯ ϱϰ X. caff ra, while in the herbaceous layer ƵƌŬĞĂĂĨƌŝĐĂŶĂ Ϯ͘ϭ͘Ϯ ƵŶĚĞƌƐƚŽƌĞLJ we documented Scadoxus multifl orus, ϵ͗,ĞdžĂůŽďƵƐŵŽŶŽƉĞƚĂůƵƐͲ DĞĚŝƵŵĚĞŶƐĞ ϭ͘ϳϰ ϵ͘ϱϯ ϲ ϱϱ Grewia monticola, and Erithrina baumii. WƚĞůĞŽƉƐŝƐĂŶŝƐŽƉƚĞƌĂ Ϯ͘ϭ͘ϯ ǁŽŽĚůĂŶĚƐ Community 12 [open woodlands] ϭϬ͗ĂŝŬŝĂĞĂƉůƵƌŝũƵŐĂͲ ůŽƐĞĚĚĞŶƐĞ Ϭ͘ϱϲ ϯ Ϯ ϭϵ ĂƉŚŝĂŵĂƐƐĂŝĞŶƐŝƐ Ϯ͘Ϯ͘ϭ ǁŽŽĚůĂŶĚ This community represents woodlands ϭϭ͗ĂƉŚŝĂŵĂƐƐĂŝĞŶƐŝƐ DĞĚŝƵŵŽƉĞŶ dominated primarily by Colophosper- ƐƵďƐƉ͘ŽďŽǀĂƚĂͲdĞƌŵŝŶĂůŝĂ Ϭ͘ϳ ϯ͘ϰ Ϯ ϭϬ ǁŽŽĚůĂŶĚ ƐĞƌŝĐĞĂ Ϯ͘Ϯ͘Ϯ mum mopane and normally occurs at ϭϮ͗ŽůŽƉŚŽƐƉĞƌŵƵŵ low altitudes below 1 000 m, as in the ŵŽƉĂŶĞͲ^ƉŝƌŽƐƚĂĐŚLJƐ KƉĞŶǁŽŽĚůĂŶĚƐ ϭ͘ϭϭ ϱ͘ϯϴ ϱ Ϯϵ ĂĨƌŝĐĂŶĂ Ϯ͘ϯ͘ϭ municipalities of Chibia, Gambos, and ϭϯ͗ŽůŽƉŚŽƐƉĞƌŵƵŵ DĞĚŝƵŵĚĞŶƐĞ Quilengues. Here, C. mopane constitutes ŵŽƉĂŶĞͲWƚĞƌŽĐĂƌƉƵƐůƵĐĞŶƐ ϭ͘ϳϵ ϵ͘ϱϯ ϭϭ ϭϵ ǁŽŽĚůĂŶĚƐ ƐƵďƐƉ͘ĂŶƚƵŶĞƐŝŝ Ϯ͘ϯ͘Ϯ the most dominant tree species (Fig. 18), and appears occasionally associated with B. fl oribunda, B. longifolia and B. spici- bretum engleri, and Buxus benguellensis, Spirostachys africana, Acacia nilotica, formis. The unique exception is Julber- the latter being endemic to the region. and Pterocarpus rotundifolius. The shrub nardia paniculata found in four of the Community 10 [closed dense wood- layer is dominated by Commiphora afri- identifi ed communities. lands] In the study area, this community cana, Grewia welwitschii, Bolusanthus Community 4 [medium dense wood- occurs in southeastern parts of the mu- speciosus, and Pseudomussaenda mon- lands] This community, despite holding nicipality of Gambos, where it forms teiroi, mostly in the Mopane woodlands few key miombo species, can be consid- closed dense woodlands dominated by of Quilengues. In some of drier areas of ered a successional stage of miombo due Combretum celastroides, C. engleri, Hip- the region, we found Commiphora mol- to the high dominance of Combretum pocratea parvifolia, and sparsely trees lis, Commiphora multijuga, Terminalia species (Fig. 10). These woodlands are of Baikiaea plurijuga. In the herbaceous prunioides, Schrebera alata, and Rhig- normally accompanied by medium-sized layer, we documented Adenium boehmi- ozum obovatum, also associated with tree species around 3-4 m in height, and anum, Gloriosa superba, and Hibiscus Colophospermum mopane woodlands. hardwood trees of Pericopsis angolensis. phoeniceus, among others. These commu- Community 13 [medium dense wood- Community 9 [medium dense wood- nities give way to shrublands dominated lands] lands] This community occurred in low- by Baphia massaiensis subsp. obovata. In These communities cover areas in the er-altitude areas covering large patches reality, most of these areas are occupied southwestern parts of the region around of the northwest of the region (Fig. 15). by private farmers, moving towards more Chibia and Gambos. Colophospermum Pteleopsis anisoptera is the dominant tree open woodlands in the municipality of mopane is sparsely distributed along with species, and is a generally small tree. Occa- Quipungo, where Baikiaea plurijuga con- Pterocarpus lucens spp. Antunesii; other sionally it was found associated with other stitutes the tallest canopy tree, occasion- tree species include Commiphora ango- tree species such as Cassia angolensis and ally associated with Phylenoptera nelsii, lensis, Kirkia acuminata, Peltophorum Commiphora mollis. In the highlands of Combretum apiculatum subsp. apicula- africanum, Ptaerxylon obliquum, and En- Humpata, communities of P. anisoptera tum, C. collinum, C. psidioides, and C. tada abyssinica. The generally closed un- generally formed dense and impenetrable zeyheri. Below the tree canopy, the veg- derstorey is dominated by thorny species thickets where we also documented Di- etation is dense, being dominated again such as Acacia nilotica, A. ataxacantha, chrostachys cinerea, Tarchonanthus cam- by Baphia massaiensis subsp. obovata, A. welwitschii, A. tortilis, and Commi- phoratus, Haplocoelum foliolosum, Com- Bauhinia urbaniana, Croton gratissimus, phora africana (Fig. 19).

B E 6 2018 429 Biodiversity Figure 7: Brachystegia spiciformis-Parinari curatellifolia woodlands Figure 8: Julbernardia paniculata-Brachystegia spiciformis wood- [open woodlands] (photo: F. Gonçalves). lands [open woodlands] (photo: F. Gonçalves).

Figure 9: Brachystegia spiciformis-Diospyros kirkii woodlands [open Figure 10: Combretum collinum-Pericopsis angolensis woodlands woodlands] (photo: F. Gonçalves). [medium dense woodlands with dense understorey] (photo: F. Gonçalves).

Figure 11: Julbernardia paniculata-Diplorhynchus condylocarpon Figure 12: Julbernardia paniculata-Combretum collinum woodlands woodlands [open dense woodlands] (photo: F. Gonçalves). [medium dense woodlands] (photo: F. Gonçalves).

430 C A Figure 13: Brachystegia spiciformis-Pteleopsis anisoptera Figure 14: Julbernardia paniculata-Burkea africana woodlands Biodiversity woodlands [open woodlands with dense understorey] (photo: F. [open woodlands with dense understorey] (photo: F. Gonçalves). Gonçalves).

Figure 15: Hexalobus monopetalus-Pteleopsis anisoptera wood- Figure 16: Baikiaea plurijuga-Baphia massaiaiensis subsp. obovata lands [medium dense woodlands] (photo: F. Gonçalves). woodlands [closed dense woodlands] (photo: F. Gonçalves).

Figure 17: Baphia massaiensis subsp. obovata-Terminalia sericea Figure 18: Colophospermum mopane-Spirostachys africana wood- woodlands [medium open woodlands] (photo: F. Gonçalves). lands [open woodlands] (photo: F. Gonçalves).

B E 6 2018 431 Tree species diversity The inspection of diversity of the main fl oristic groups showed that in general, miombo woodlands with Brachystegia spiciformis-Parinari curatellifolia, B. longifolia-Diospyros kirkii, and Julber- nardia paniculata-Diplorhynchus con- dylocarpon [Communities 1, 3, 5] have Figure 19: the highest species richness (Tab. 1), Colophospermum followed by Hexalobus monopeta- mopane-Pteocarpus lucens subsp. antunesii lus-Pteleopsis anisoptera woodlands [medium dense wood- [Community 9] and Colophospermum lands] mopane-Pterocarpus lucens subsp. an- (photo: F. Gonçalves). tunesii woodlands [Community 13]. The tree community with the highest Biodiversity tree species richness was Combretum collinum-Pericopsis angolensis wood- lands [Community 4]. Communities of Julbernardia paniculata-Brachystegia spiciformis [Community 2], Baikiaea plurijuga-Baphia massaiensis subsp. obovata [Community 10], Baphia mas- saiensis ssp obovata-Terminalia sericea [Community 11], and Colophospermum mopane-Pterocarpus lucens subsp. an- tunesii [Community 12] showed the low- est species richness (Fig. 5). The communities of the fi rst major cluster, Communities 1-5, were the most diverse as measured by the Shannon diversity index, with the exception of Community 2. These communities also showed the highest species richness and greatest number of indicator species of Figure 5: Box plots depicting the tree species richness in the 13 identifi ed communities. communities. The second most diverse community was represented by tree com- munities 6 to 9 and 13 (Fig. 6). Commu- nity 9 showed also the highest number of plots sampled. The second highest number of plots sampled was found in Community 8. Despite the high sampling eff ort in these communities, they dem- onstrated very low species richness as well as low numbers of indicator species (Tab. 1)

Discussion

Diversity of the woodlands of Huíla Province This study represents the fi rst plot-based woody vegetation assessment of Huíla Province in Angola, aiming to charac- terise and describe the woodlands of the Figure 6: Shannon index of the plots of the 13 tree communities. region. A total of 456 plots were surveyed

432 C A in the region, suffi cient to document as areas heavily impacted by agriculture almost intact, towards the southeastern major fl oristic groups occurring in the are reported to hold tree species with high and western parts of the region, where the study area, as evidenced by the species light demand and fast growth, like many woodlands are generally sparser (Com- accumulation curve. This pattern may Combretum species (Jew et al., 2016). munities 1, 2, 3, 5, 6, 7, and 8). Miombo imply that any further increase of sam- These species recruit and establish eas- woodlands show variations in terms of pling eff ort would be expected to lead ily following disturbance and in this way density and species composition through- to inclusion of additional rare species, additional species are added to the oth- out the region, with diff erences in spe- as the sample size of 456 was high and erwise typical miombo species (Banda cies composition being more evident at may capture almost all woody species et al., 2006; Gonçalves et al., 2017). local scale. Local abiotic conditions and occurring in the region, and can be use- The presence of standing trees of the changes from Brachystegia spiciformis ful to characterise the species diversity hardwood species Pericopsis angolensis to Julbernardia paniculata communities, and relationship between woody species was also documented in early succession together with various other woody spe- and site conditions. As mentioned before, of the south-central Angolan miombo cies, may also infl uence the stand struc- most of the previous studies carried out woodlands (Gonçalves et al., 2017). A ture and composition of the woodlands in the country date to the colonial era. threshold of H’ = 2 has been mentioned (Revermann & Finckh, 2013). Signs of These studies characterised the vegeta- as the minimum value above which an tree damage caused by fi re, woodcut- Biodiversity tion types based on general aspects or ecosystem can be regarded as medium to ting, and/or agriculture activity were also were limited to dominant species only high diversity (Giliba et al., 2011). The documented; this is a stark reminder that (but see Revermann et al., 2018). As with mean Shannon diversity index found in fi re frequency, together with other human other parts of Angola, there is still a great the Mopane woodlands of the study area disturbance, plays a major role in miom- deal of work to be done to describe the was relatively low compared to similar bo woodland dynamics, aff ecting physi- vegetation of Huíla Province. Barbosa habitats in northern Botswana; this is cal structure, composition of species, and (1970) recognises about eight vegetation attributed to anthropogenic disturbance, also woodland recovery following distur- types for the Huíla region, including the suggesting that non-protected woodlands bance (Chidumayo, 2002; Furley et al., most important woodlands surrounding are negatively impacted by human activ- 2008). the study area. With the present study, ity, with a direct eff ect on the composi- Community 4, Combretum collinum- we are able to document 13 distinct tree tion and diversity of species (Teketay Pericopsis angolensis woodlands, was communities within the region; most of et al., 2018). Within the study area we found associated with key miombo and them constitute fairly typical miombo found H’ values below 1; this may not non-miombo species, mainly in the mu- woodlands, dominated by the Fabaceae only be related to the minimum number nicipalities of Caconda, Caluquembe, family, of which the subfamily Caesal- of plots surveyed, but also to the mono- and Chicomba. Similar patterns of key pinioideae is the largest. The dominance dominance of single tree species such as miombo species occurring together with of the Fabaceae family within the mi- Baikiaea plurijuga-Baphia massaiensis- other woody species have also been doc- ombo eco-region has been widely re- Terminalia sericea. The Shannon index umented in the Tanzanian miombo wood- ported in various studies (Byers, 2001; may be strongly infl uenced by the occur- lands (Banda et al., 2008). These fi ndings Munishi et al., 2011). Within the study rence of rare species such as Entandro- contrast with the pattern usually consid- area of Huíla Province, miombo wood- phragma spicatum, which can be found ered common for miombo woodlands, lands occupy large areas, being relatively in the area associated to the woodlands. suggesting that on a larger spatial scale, dense with canopy tree species around E. spicatum was reported to be very rare the species composition of miombo is 12-15 m in height in the municipalities of in the study area and probably in risk of very high, and common genera as Brach- Chipindo and Cuvango. The woodlands local extinction due to its high value as ystegia, Julbernardia, and Isoberlinia are become open with increasing altitude, as timber (Barbosa, 1970). not always dominant at the local scale observed in Caconda, Caluquembe, and (Mwakalukwa et al., 2014). patches of the Humpata plateau, and due Vegetation classifi cation Community 9, Hexalobus monopeta- to high land use pressure caused by agri- The Angolan miombo woodlands cover lus-Pteleopsis anisoptera woodlands, culture, the woodlands appear also rela- an extensive area of central Angola, ex- was widely distributed across the region. tively open in the municipality of Jamba. tending into Democratic Republic of Stands of Hexalobus monopetalus are re- The mean Shannon index values found Congo (Burgess et al., 2004). Most of this ported to occur mainly at low altitudes in in the study area were generally low, with eco-region is found at elevations between southern Africa (Coates Palgrave, 2005), the exception of Community 4 [Com- 1 000-1 500 m above sea level and in- but in the study area, these species also bretum collinum-Pericopsis angolensis cludes the highlands of Huíla, Huambo, appeared to be very common in disturbed woodlands] with a mean H’ equal to 2.07. and Bié (Barbosa, 1970; Huntley, 1974a). woodlands. This community was also The highest value of Shannon diversity Within our study area, miombo wood- found on rocky outcrops and mountain- found in this community supports the lands covered large areas, ranging from ous areas of the Quilengues and Humpata view that these woodlands may represent high rainfall sites in the north, where the plateau. Small trees of key miombo spe- regrowth of typical miombo woodlands, woodlands seem to be much denser and cies such as Brachystegia spiciformis and

B E 6 2018 433 Julbernardia paniculata were commonly as the surrounding areas of Matala and (Barbosa, 1970). Further indicator spe- found associated to the community (Gon- Quipungo were largely depleted, most cies of the community were Commiphora çalves, 2009). likely by timber over-exploitation and species, and shrubs of Croton mubango, Community 10, Baikiaea plurijuga- clearance of large areas for agriculture common in the understorey. Baphia massaiensis subsp. obovata purposes in the two municipalities. woodlands, belongs to the Zambezian To the south, moving towards Mulon- Baikiaea woodlands eco-region, which do in the municipality of Matala, we were Conclusion forms a mosaic of Baikiaea plurijuga- able to characterise woodland as Baphia dominated forest, woodlands, thickets, massaiensis-Terminalia sericea wood- Information on Angolan vegetation is and secondary grassland in Angola, Na- lands (Community 11). This community scarce, as most studies conducted in the mibia, Botswana, Zambia, and Zimba- also covers large areas of Bicuar Na- country date back to the colonial era. bwe (Burgess et al., 2004). These dense tional Park. Shrubby species within the This study, carried out in Huíla Province, woodlands, described in the southeastern community were generally very few, and represents the fi rst plot-based vegetation parts of Gambos, appear similar to the included Vitex mombassae and Grewia survey in this province and provides the dense Baikiaea-Burkea woodlands fi rst spp. The previous vegetation studies re- fi rst vegetation classifi cation for this re- Biodiversity described for the Okavango basin along fer to this community as Terminalia seri- gion. A high sampling eff ort guaranteed the Cubango River, characterised by a cea, Acacia nilotica, A. tristis, or Hip- that most tree species expected to occur closed canopy and thicket-like understo- pocratea-Baphia-Croton-Combretum in the Huíla were recorded and that rep- rey (Revermann & Finckh, 2013; Wallen- spp. shrublands, where Baikiaea may or resentative tree communities were identi- fang et al., 2015). Further north of Gam- may not occur (Teixeira, 1968; Barbosa, fi ed. As such, this survey constitutes the bos, Baphia massaiensis subsp. obovata 1970). In fact, T. sericea may also appear basis for the fi rst detailed vegetation map dominates the landscape; this attracts pri- associated to these species in our study of the province. Additionally, a deeper vate farmers to these areas, as this species area; however, we were not able to assess analysis of environmental drivers of veg- is of high nutritional value for livestock these shrublands, as they generally form etation patterns will be needed in order to (Maiato & Sweet, 2011). The Baikiaea dense and almost impenetrable thicket, explain the distribution and variation of woodlands become more open and con- and standing trees rarely occur. species composition and diversity in the stitute one of the dominant vegetation Mopane woodlands cover an estimat- woodlands of Huíla Province. types of Quipungo within the adminis- ed area of 55 000 km2 in southern Africa trative division of Bicuar National Park (Makhado et al., 2014). The woodlands (BNP). A detailed vegetation account of stretch between Angola and Namibia in Acknowledgements this area points to about six vegetation the southwest, from the marginal moun- communities occurring within the BNP, tain chain at the base of Serra da Chela The research was carried out in the including woodlands, shrublands, and to more open and sub-desert habitats framework of SASSCAL and was spon- grasslands with aquatic and semi-aquatic (Barbosa, 1970; Menezes, 1971). The sored by the German Federal Ministry of vegetation (Teixeira, 1968). We docu- Mopane woodlands in Angola grow over Education and Research (BMBF) under mented Baikiaea plurijuga-Baphia mas- vast areas, and are typically associated promotion number 01LG1201M. We also saiensis subsp. obovata woodlands dur- with ferralitic and black clay soils. With- thank the Angolan Ministry of Science ing the fi eld survey as one of the major in these areas, C. mopane appears associ- and Technology through the Angolan woody vegetation components in the re- ated with tree species, which mirrors the escarpment fl oristic diversity project for gion, which is not clearly described in this much drier climate previously document- partially funding the fi eld surveys. Ad- previous study. Barbosa (1970) described ed between the municipalities of Car- ditionally, we thank the vice governor of these woodlands as occupying large areas aculo and Virei in the Namibe Province Huíla Province, Mrs Maria João Tchipa- of the BNP. The Baikiaea-Baphia wood- (Maiato & Sweet, 2011). In our study, lavela, for her personal commitment and lands here can be considered part of an two communities (Communities 12 and support given; the municipal administra- extensive area of dry tropical woodland 13) with C. mopane as a character species tors of Huíla Province; and traditional in southern Africa; in Angola, it has its were identifi ed with only slightly diff er- authorities and the local population for southeast limit in deep Kalahari sands ing species composition. Community 13, their collaboration during fi eld surveys. along the Angolan-Namibian border. Colophospermum mopane-Pterocarpus We thank our colleagues from the Uni- Here the canopy is dominated by tree spe- lucens subsp. antunesii woodlands was versity of Hamburg, Rasmus Revermann cies such as Schinziophyton rautanenii, mostly characteristic of the drier areas and Manfred Finckh, for supporting us Guibourtia coleosperma, and Pterocar- of Huíla Province. This community may with methodology and fi eldwork, and pus angolensis; pure stands of Baikiaea represent a vegetation of contact between John Lister Godlee at the University of plurijuga only rarely occur (Gonçalves medium open miombo woodlands of Edinburgh, who kindly revised the Eng- et al., 2018). The woodlands in the BNP Julbernardia paniculata, Brachystegia lish of the manuscript. may represent the most intact unit of this spicifomis and B. boehmii of Chibia and vegetation community in Huíla Province, typical Mopane woodlands further south

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B E 6 2018 435 Table 2: Vegetation classifi cation of the woody vegetation of Huíla province based on 1 000 m2 vegetation plots. The 465 table displays the indicator species of 466 every vegetation community identifi ed in the analysis. Species are ordered according to Pearson’s phi coeffi cient of association p <= 0.05. The phi value 467 ranges from 0 to 100, where phi > 30 is regarded as diagnostic of tree species. Species can be associated with more than one group or group combinations. 468 Only species with p ≤ 0.05 are shown in the table.

sĞŐĞƚĂƚŝŽŶŽŵŵƵŶŝƚŝĞƐ ϭ Ϯ ϯ ϰ ϱ ϲ ϳ ϴ ϵ ϭϬ ϭϭ ϭϮ ϭϯ ϭнϮ ϭнϰ Ϯнϯ Ϯнϱ Ϯнϲнϳ ϭϮнϭϯ ƉͲǀĂůƵĞ ŽŵŵƵŶŝƚLJϭ͗ƌĂĐŚLJƐƚĞŐŝĂƐƉŝĐŝĨŽƌŵŝƐͲWĂƌŝŶĂƌŝĐƵƌĂƚĞůůŝĨŽůŝĂǁŽŽĚůĂŶĚƐ ƌĂĐŚLJƐƚĞŐŝĂďŽĞŚŵŝŝ ϵϬ ϳϭ Ϭ͘ϬϬϭ WĂƌŝŶĂƌŝĐƵƌĂƚĞůůŝĨŽůŝĂ ϴϲ ϲϵ Ϭ͘ϬϬϭ ƌĂĐŚLJƐƚĞŐŝĂƐƉŝĐŝĨŽƌŵŝƐ ϴϯ ϴϮ ϭϬϬ Ϭ͘ϬϬϭ hĂƉĂĐĂŬŝƌŬŝĂŶĂ ϳϲ ϰϰ Ϭ͘ϬϬϭ ƌĂĐŚLJƐƚĞŐŝĂĨůŽƌŝďƵŶĚĂ ϳϮ Ϭ͘ϬϬϭ ƌĂĐŚLJƐƚĞŐŝĂůŽŶŐŝĨŽůŝĂ ϳϮ Ϭ͘ϬϬϭ ^LJnjLJŐŝƵŵŐƵŝŶĞĞŶƐĞ ϲϵ ϱϭ Ϭ͘ϬϬϭ ŶŝƐŽƉŚLJůůĞĂďŽĞŚŵŝŝ ϲϮ ϰϭ Ϭ͘ϬϬϭ &ĂƵƌĞĂƌŽĐŚĞƚŝĂŶĂ ϲϮ ϯϲ Ϭ͘ϬϬϭ ŽďŐƵŶŶŝĂŵĂĚĂŐĂƐĐĂƌŝĞŶƐŝƐ ϲϮ ϰϵ Ϭ͘ϬϬϭ

Biodiversity ůďŝnjŝĂĂŶƚƵŶĞƐŝĂŶĂ ϱϱ Ϭ͘ϬϬϭ hĂƉĂĐĂŶŝƚŝĚĂǀĂƌ͘ŶŝƚŝĚĂ ϱϱ Ϭ͘ϬϬϭ ƵƌŬĞĂĂĨƌŝĐĂŶĂ ϱϱ Ϭ͘ϬϬϭ DŽŶŽƚĞƐĂĨƌŝĐĂŶƵƐ ϰϭ Ϭ͘ϬϬϭ ƌŝĚĞůŝĂŵŽůůŝƐ ϰϭ Ϭ͘Ϭϭ WƚĞƌŽĐĂƌƉƵƐĂŶŐŽůĞŶƐŝƐ ϯϭ Ϭ͘Ϭϭ ŽŵŵƵŶŝƚLJϮ͗:ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂͲƌĂĐŚLJƐƚĞŐŝĂƐƉŝĐŝĨŽƌŵŝƐǁŽŽĚůĂŶĚƐ :ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂ ϵϮ ϭϬϬ ϵϳ ϵϮ Ϭ͘ϬϬϭ ƌĂĐŚLJƐƚĞŐŝĂƐƉŝĐŝĨŽƌŵŝƐ ϴϮ Ϭ͘ϬϬϭ ŽďŐƵŶŶŝĂŵĂĚĂŐĂƐĐĂƌŝĞŶƐŝƐ  ϰϵ Ϭ͘Ϭϭ hĂƉĂĐĂŬŝƌŬŝĂŶĂ ϰϰ Ϭ͘ϬϬϭ ^LJnjLJŐŝƵŵŐƵŝŶĞĞŶƐĞ ϱϭ Ϭ͘ϬϬϭ &ĂƵƌĞĂƌŽĐŚĞƚŝĂŶĂ ϯϲ Ϭ͘ϬϬϭ ŽŵďƌĞƚƵŵĐŽůůŝŶƵŵ ϯϲ ϯϲ Ϭ͘Ϭϭ ŽŵŵƵŶŝƚLJϯ͗ƌĂĐŚLJƐƚĞŐŝĂůŽŶŐŝĨŽůŝĂͲŝŽƐƉLJƌŽƐŬŝƌŬŝŝǁŽŽĚůĂŶĚƐ ƌĂĐŚLJƐƚĞŐŝĂůŽŶŐŝĨŽůŝĂ ϭϬϬ Ϭ͘ϬϬϭ ŝŽƐƉLJƌŽƐŬŝƌŬŝŝ ϲϲ Ϭ͘ϬϬϭ WƐĞƵĚŽůĂĐŚŶŽƐƚLJůŝƐŵĂƉƌŽƵŶĞŝĨŽůŝĂƐƵďƐƉ͘ ĚĞŬŝŶĚƚŝŝ ϲϮ Ϭ͘ϬϬϭ ŽŵďĞLJĂƌŽƚƵŶĚŝĨŽůŝĂ ϱϵ Ϭ͘ϬϬϭ ŬĞďĞƌŐŝĂďĞŶŐƵĞůůĞŶƐŝƐ ϰϱ Ϭ͘ϬϬϭ ƌŝĚĞůŝĂŵŝĐƌĂŶƚŚĂǀĂƌ͘ŵŝĐƌĂŶƚŚĂ ϯϴ Ϭ͘ϬϬϭ ƵƐƐŽŶŝĂĂŶŐŽůĞŶƐŝƐ ϯϭ Ϭ͘ϬϬϭ sŝƚĞdžŵĂĚŝĞŶƐŝƐƐƵďƐƉ͘ŵĂĚŝĞŶƐŝƐ ϯϭ Ϭ͘ϬϬϭ ŽďŐƵŶŶŝĂŵĂĚĂŐĂƐĐĂƌŝĞŶƐŝƐ ϳϮ Ϭ͘ϬϬϭ ŽŵŵƵŶŝƚLJϰ͗ŽŵďƌĞƚƵŵĐŽůůŝŶƵŵͲWĞƌŝĐŽƉƐŝƐĂŶŐŽůĞŶƐŝƐǁŽŽĚůĂŶĚƐ ŽŵďƌĞƚƵŵĐŽůůŝŶƵŵ ϴϲ Ϭ͘ϬϬϭ WĞƌŝĐŽƉƐŝƐĂŶŐŽůĞŶƐŝƐ ϲϵ Ϭ͘ϬϬϭ ŝƉůŽƌŚLJŶĐŚƵƐĐŽŶĚLJůŽĐĂƌƉŽŶ ϲϯ Ϭ͘ϬϬϭ ĂƉŚŝĂďĞƋƵĂĞƌƚŝŝ ϱϱ Ϭ͘ϬϬϭ ƌŝĚĞůŝĂŵŽůůŝƐ ϱϭ Ϭ͘ϬϬϭ ŽŵďƌĞƚƵŵnjĞLJŚĞƌŝ ϰϳ Ϭ͘ϬϬϭ WŚLJůůĂŶƚŚƵƐƌĞƚŝĐƵůĂƚƵƐ ϰϳ Ϭ͘ϬϬϭ ZŽƚŚŵĂŶŶŝĂĞŶŐůĞƌŝĂŶĂǀĂƌ͘ĞŶŐůĞƌŝĂŶĂ ϰϭ Ϭ͘ϬϬϭ ŽŵŵƵŶŝƚLJϱ͗:ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂͲŝƉůŽƌŚLJŶĐŚƵƐĐŽŶĚLJůŽĐĂƌƉŽŶǁŽŽĚůĂŶĚƐ :ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂ ϵϳ Ϭ͘ϬϬϭ ŝƉůŽƌŚLJŶĐŚƵƐĐŽŶĚLJůŽĐĂƌƉŽŶ ϴϯ Ϭ͘ϬϬϭ WƐĞƵĚŽůĂĐŚŶŽƐƚLJůŝƐŵĂƉƌŽƵŶĞŝĨŽůŝĂƐƵďƐƉ͘ ĚĞŬŝŶĚƚŝŝ ϱϳ Ϭ͘Ϭϭ ƌĂĐŚLJƐƚĞŐŝĂďŽĞŚŵŝŝ ϱϭ Ϭ͘Ϭϭ ŝŽƐƉLJƌŽƐŬŝƌŬŝŝ ϰϵ Ϭ͘ϬϬϭ ŽŵŵƵŶŝƚLJϲ͗:ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂͲŽŵďƌĞƚƵŵĐŽůůŝŶƵŵǁŽŽĚůĂŶĚƐ :ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂ ϵϮ Ϭ͘ϬϬϭ ŽŵďƌĞƚƵŵĐŽůůŝŶƵŵ ϯϲ Ϭ͘ϬϬϭ ŽŵŵƵŶŝƚLJϳ͗ƌĂĐŚLJƐƚĞŐŝĂƐƉŝĐŝĨŽƌŵŝƐͲWƚĞůĞŽƉƐŝƐĂŶŝƐŽƉƚĞƌĂǁŽŽĚůĂŶĚƐ ƌĂĐŚLJƐƚĞŐŝĂƐƉŝĐŝĨŽƌŵŝƐ ϭϬϬ Ϭ͘ϬϬϭ WƚĞůĞŽƉƐŝƐĂŶŝƐŽƉƚĞƌĂ ϱϴ Ϭ͘Ϭϭ :ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂ ϯϭ Ϭ͘ϬϬϭ

436 C A sĞŐĞƚĂƚŝŽŶŽŵŵƵŶŝƚŝĞƐ ϭ Ϯ ϯ ϰ ϱ ϲ ϳ ϴ ϵ ϭϬ ϭϭ ϭϮ ϭϯ ϭнϮ ϭнϰ Ϯнϯ Ϯнϱ Ϯнϲнϳ ϭϮнϭϯ ƉͲǀĂůƵĞ ŽŵŵƵŶŝƚLJϴ͗:ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂͲƵƌŬĞĂĂĨƌŝĐĂŶĂǁŽŽĚůĂŶĚƐ ƵƌŬĞĂĂĨƌŝĐĂŶĂ ϱϬ Ϭ͘ϬϬϭ :ƵůďĞƌŶĂƌĚŝĂƉĂŶŝĐƵůĂƚĂ ϯϬ Ϭ͘ϬϬϭ ŽŵŵƵŶŝƚLJϵ͗,ĞdžĂůŽďƵƐŵŽŶŽƉĞƚĂůƵƐͲWƚĞůĞŽƉƐŝƐĂŶŝƐŽƉƚĞƌĂǁŽŽĚůĂŶĚƐ ,ĞdžĂůŽďƵƐŵŽŶŽƉĞƚĂůƵƐ ϰϮ Ϭ͘ϬϬϭ WƚĞůĞŽƉƐŝƐĂŶŝƐŽƉƚĞƌĂ ϯϯ Ϭ͘Ϭϭ ŽŵŵŝƉŚŽƌĂŵŽůůŝƐ ϯϭ Ϭ͘ϬϬϭ ŽŵŵƵŶŝƚLJϭϬ͗ĂŝŬŝĂĞĂƉůƵƌŝũƵŐĂͲĂƉŚŝĂŵĂƐƐĂŝĞŶƐŝƐǁŽŽĚůĂŶĚƐ ĂŝŬŝĂĞĂƉůƵƌŝũƵŐĂ ϭϬϬ Ϭ͘ϬϬϭ ĂƉŚŝĂŵĂƐƐĂŝĞŶƐŝƐƐƵďƐƉ͘ŽďŽǀĂƚĂ ϲϯ Ϭ͘ϬϬϭ ŽŵŵƵŶŝƚLJϭϭ͗ĂƉŚŝĂŵĂƐƐĂŝĞŶƐŝƐͲdĞƌŵŝŶĂůŝĂƐĞƌŝĐĞĂǁŽŽĚůĂŶĚƐ ĂƉŚŝĂŵĂƐƐĂŝĞŶƐŝƐƐƵďƐƉ͘ŽďŽǀĂƚĂϴϬϬ͘ϬϬϭ dĞƌŵŝŶĂůŝĂƐĞƌŝĐĞĂ ϳϬ Ϭ͘ϬϬϭ ŽŵŵƵŶŝƚLJϭϮ͗ŽůŽƉŚŽƐƉĞƌŵƵŵŵŽƉĂŶĞͲ^ƉŝƌŽƐƚĂĐŚLJƐĂĨƌŝĐĂŶĂǁŽŽĚůĂŶĚƐ ŽůŽƉŚŽƐƉĞƌŵƵŵŵŽƉĂŶĞ ϵϬ ϴϰ Ϭ͘ϬϬϭ ^ƉŝƌŽƐƚĂĐŚLJƐĂĨƌŝĐĂŶĂ ϲϮ Ϭ͘ϬϬϭ Biodiversity dĞƌŵŝŶĂůŝĂƉƌƵŶŝŽŝĚĞƐ ϲϮ ϰϮ Ϭ͘ϬϬϭ ĐĂĐŝĂŶŝůŽƚŝĐĂ ϰϱ Ϭ͘ϬϬϭ ŽŵŵƵŶŝƚLJϭϯ͗ŽůŽƉŚŽƐƉĞƌŵƵŵŵŽƉĂŶĞͲWƚĞƌŽĐĂƌƉƵƐůƵĐĞŶƐƐƵďƐƉ͘ĂŶƚƵŶĞƐŝŝǁŽŽĚůĂŶĚƐ ŽůŽƉŚŽƐƉĞƌŵƵŵŵŽƉĂŶĞ ϴϰ Ϭ͘ϬϬϭ ĐĂĐŝĂŶŝůŽƚŝĐĂ ϱϴ Ϭ͘ϬϬϭ ŽŵŵŝƉŚŽƌĂŵŽůůŝƐ ϱϯ Ϭ͘ϬϬϭ <ŝƌŬŝĂĂĐƵŵŝŶĂƚĂ ϱϯ Ϭ͘ϬϬϭ WƚĞƌŽĐĂƌƉƵƐůƵĐĞŶƐƐƵďƐƉ͘ĂŶƚƵŶĞƐŝŝ ϰϮ Ϭ͘ϬϬϭ ŽŵŵŝƉŚŽƌĂĂŶŐŽůĞŶƐŝƐ ϯϳ Ϭ͘ϬϬϭ ƌŽƚŽŶŵƵďĂŶŐŽ             ϯϮ       Ϭ͘Ϭϭ

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