Plant Ecology and Evolution 144 (2): 126–137, 2011
doi:10.5091/plecevo.2011.539
REGULAR PAPER
Coastal dry forests in northern Mozambique
Jonathan Timberlake1,*, David Goyder1, Frances Crawford1, John Burrows2,
G. Philip Clarke3, Quentin Luke4,5, Hermenegildo Matimele6, Tom Müller7,
Olivier Pascal8, Camila de Sousa6 & Tereza Alves6
Herbarium, Royal Botanic Gardens Kew, Richmond, Surrey UK-TW9 3AB, United Kingdom
Buffelskloof Herbarium, P.O. Box 710, Lydenburg 1120, South Africa
3
The Hermitage, Crewkerne, Somerset UK-TA18 8HA, United Kingdom
4
East African Herbarium, National Museums of Kenya, P.O. Box 45166, 00100 Nairobi, Kenya
5
Centre for Tropical Plant Conservation, Fairchild Botanic Garden, 11935 Old Cutler Road, Coral Gables, FL 33156-4242, USA
6
Herbarium, Instituto de Investigação Agrária de Moçambique, C.P. 3658, Maputo, Mozambique
7
5 Vollendam, 13 J. Tongogara, Harare, Zimbabwe
8
Pro-Natura International, 15 Avenue de Ségur, Paris FR-75007, France
*Author for correspondence: j.timberlake@kew.org
1
2
Background and aims – The Coastal Forests of Eastern Africa, stretching along the Indian Ocean coastline
from Somalia to Mozambique, are considered by Conservation International to be a global biodiversity
hotspot – an area of high diversity and endemism under increasing threat. Although the largest remaining
extent of these forests is reported to be found in Mozambique, very little is known on their extent, condition
and composition here. In addition, the term ‘coastal forest’ has been used in different ways by different
authors. This paper deines and characterises coastal dry forests found in northern Mozambique and assesses
their present extent, botanical composition, conservation importance and the threats to these forests.
Methods – The study area of 18,150 km2 lies in Cabo Delgado Province in north-east Mozambique, adjacent
to Tanzania. Its limits are determined primarily by geological substrate and landform. Four smaller study
sites were chosen covering a range of landforms. Manual interpretation of satellite imagery dating from
1999–2002 was used to calculate possible previous and present extent of ‘dense vegetation’. Extensive
ield collecting was used in determining botanical composition and distribution patterns. IUCN Red List
assessments were carried out on selected species using distributional criteria.
Results – Dry forests similar to those in southern Tanzania are found widely scattered across coastal Cabo
Delgado, sitting in a matrix of miombo woodland and other vegetation types. However, forest cover is not
as extensive was believed. We calculate that the original extent of ‘dense vegetation cover’, which includes
coastal dry forest, was 6087 km2. Owing to clearance over the last 150 years this is now only 1182 km2, of
which perhaps only 400 km2 is moderately-intact dry forest.
In this southern part of their range such forests are essentially dry, not moist and mesic, and dominated by
a high proportion of deciduous or sclerophyllous evergreen trees. The plant species composition differs
signiicantly from that of the surrounding woodlands. There is a marked change in species composition
between forest patches along the coast, and they contain numerous species with restricted global distribution.
Since 2003, 68 species new to Mozambique have been recorded from Cabo Delgado in addition to 36
possible new species. Many new records are of species previously only known from south-eastern Tanzania.
Previously recorded patterns of restricted distribution and high species turnover between forest patches in
Kenya and Tanzania are conirmed. Seven coastal forest species were assessed as Endangered.
Regional context and conservation – Coastal dry forests are discussed in relation to the more widespread
‘sand forests’ of the continental interior of south-central Africa, and shown to have similarities in ecology,
species composition, soils and ecology. Very little of the present extent of coastal forests in Mozambique
lies within protected areas. The threats to their continued existence in the face of exploitation for timber,
agriculture and oil exploration are outlined.
Key words – Mozambique, coastal forest, endemics, hotspot, conservation.
All rights reserved. © 2011 National Botanic Garden of Belgium and Royal Botanical Society of Belgium – ISSN 2032-3921
Timberlake et al., Coastal dry forests in northern Mozambique
INTRODUCTION
The Coastal Forests of Eastern Africa are considered by Conservation International (CI), an international conservation
organisation, to be a global biodiversity hotspot – an area of
high species diversity and endemism that is under increasing
threat (Myers et al. 1999, Burgess et al. 2004b). The same
area along the eastern African coastline has also been recognised by the World Wide Fund for Nature (WWF) as a globally important ecoregion – the Eastern Africa Coastal Forest
Ecoregion (Burgess et al. 2004a) – with a speciic programme
directed towards its conservation (WWF-EARPO 2006).
This hotspot or ecoregion extends from southern Somalia
through Kenya and Tanzania to southern Mozambique (ig.
1), although most of our knowledge on the area comes from
studies in Kenya and Tanzania (e.g. Moomaw 1960, Greenway 1973, Brenan 1978, Robertson & Luke 1993, Hawthorne
1993, Vollesen 1994, Burgess & Clarke 2000, Burgess et al.
2004a).
It has often been assumed that the major extent of remaining intact vegetation in this hotspot lay in northern Mozambique in the provinces of Cabo Delgado and Nampula (Burgess et al. 2000, Burgess et al. 2004a, WWF-EARPO 2006),
but to date there has been little direct evidence of this in terms
of mapped forest extent. Historical plant collections from the
1940s to mid-1960s suggested that some restricted-range species, otherwise only known from coastal southern Tanzania,
were also found here (Brenan 1978, Clarke et al. 2000: 137)
along with some Mozambique endemics, but the extent of
forest remaining and the signiicant presence of range-restricted or endemic species was speculative.
Following the instability resulting from the independence
and civil wars, development is now occurring at a rapid rate
in the country, including exploration for onshore oil and gas,
and this is likely to accelerate the loss of what remains of
these coastal forests. The investigation of coastal northern
Mozambique is an obvious priority for botanists and conservationists.
Coastal Mozambique falls within what White (1983)
terms the Zanzibar–Inhambane regional mosaic, a phytochorion stretching along the East African coast from Somalia
down to South Africa. This area has a distinctive lora, different from that found further inland in the Zambezian regional
centre of endemism. Unfortunately, White did not map any
of the coastal dry forest patches within this rather broad regional mosaic, and his descriptions of dry forest vegetation
is skewed towards that found in Kenya. Within the mosaic
he describes ten vegetation types of which four (Zanzibar–
Inhambane undifferentiated forest, Zanzibar–Inhambane
transition woodland, Zanzibar–Inhambane woodland and
scrub woodland and Zanzibar–Inhambane secondary grassland and wooded grassland) seem to occur in north-east Mozambique. Much of the Cabo Delgado study area falls within
what White calls Zanzibar–Inhambane woodland and scrub
woodland. Surprisingly, he does not explicitly recognise dry
coastal forest as a separate type in Mozambique, unlike Wild
& Barbosa (1967) on which much of White’s work for the
country was based.
More recently, on the basis of species richness and composition, Clarke (1998) has split White’s Zanzibar–Inham-
Figure 1 – Area with coastal forests in Eastern Africa (adapted from
Burgess et al. 2004b).
bane regional mosaic into two separate phytochoria – the
Swahilian regional centre of endemism in the north and the
Swahilian/Maputaland regional transition zone in the south.
The only vegetation maps available for this part of northern Mozambique are those by Pedro & Barbosa (1955) and
the Flora Zambesiaca vegetation map (Wild & Barbosa 1967),
which was partly based on the former. Both maps show that
Cabo Delgado has a somewhat different vegetation from other parts of Mozambique, and that most of it is covered in a
type of miombo woodland. However, these maps show areas
of Dry Deciduous Lowland Forest on the Mueda plateau, a
vegetation type that appears to have largely disappeared, and,
of particular interest to us, Dry Deciduous Thicket with Guibourtia schliebenii (Harms) J.Léonard, the type that formed
part of the main focus of the present study.
In recent years there has been renewed interest in coastal
Mozambique. In 2003, Quentin Luke, who had previously
studied the moister coastal forests of Kenya and northern
Tanzania, visited Cabo Delgado on behalf of WWF and CI
and collected a number of new plant records (Luke 2006),
followed by John and Sandie Burrows in 2005–2008 in the
course of writing a tree ield guide for Mozambique. Results from these collections are incorporated here. In 2008,
the French NGO Pro-Natura International, together with the
Muséum National d’Histoire Naturelle in Paris, obtained
funding to carry out preliminary biological surveys of plants
and various vertebrate and invertebrate groups in this hotspot
in Cabo Delgado Province, resulting in two major expeditions in 2008 and 2009. Other institutional partners in the
study were the Instituto de Investigação Agrária de Moçambique, the herbarium of the Royal Botanic Gardens, Kew and
the Buffelskloof Herbarium in Lydenburg, South Africa.
This paper outlines initial indings on the extent and botanical composition of forests in Cabo Delgado Province arising from these recent expeditions and studies, and discusses
their links to similar dry forests elsewhere in Eastern and
Southern Africa.
127
Pl. Ecol. Evol. 144 (2), 2011
WHAT ARE COASTAL FORESTS?
The term ‘coastal forest’ has been used widely in recent years
(e.g. Hawthorne 1993, Robertson & Luke 1993, Myers et al.
1999, Burgess et al. 2004a, Burgess & Clarke 2000, Clarke
2000a, WWF-EARPO 2006) but there has been inconsistency in deinition. In some cases, most of the dense vegetation
formations found in the coastal area (e.g. within 100–150 km
of the coast) or within White’s (1983) Zanzibar–Inhambane
phytochorion are included, whilst others (e.g. WWF-EARPO
2006) have included various forest or woodland formations
(except mangroves) up to 300 km inland, possibly as they
were lowland vegetation types or contained some species
with a typically East African coastal distribution. But unless
such coastal species are dominant they can not realistically be
termed coastal forests.
Clarke (2000a) formally deines East African Coastal Forests as forests (i.e. a continuous stand of trees with crowns
overlapping or interdigitating, usually comprising several
layers and/or interlaced with lianas, often with a sparse or absent ground layer) dominated by Swahilian endemic or nearendemic tree species, and describes six different types. The
term is used collectively to encompass both typical Eastern
African coastal dry forests as well as variant and transitional
formations where they share features with forests of other
phytochoria. On the other hand, Hawthorne (1993) adopts a
more geological and geomorpohological deinition, deining
‘coastal’ as lying on sedimentary (or volcanic) sediments of
the coastal plains and plateaux, excluding any vegetation formations on Basement Complex substrates.
However, in practice areas termed ‘coastal forest’ shown
on some recent maps have covered a wide range of vegetation
from dense woodland through dry forest to true moist forest.
More importantly, in some cases there appears to be no common or linking feature between these ‘forest patches’ in terms
of species composition or ecology, which has led to confusion in determining the distribution of coastal dry forests in
Mozambique, their biodiversity attributes, ecology, possible
origins and conservation signiicance. The assumed commonality in terms of the origin and conservation importance of
this group of vegetation types has masked our understanding
of them, and perhaps inhibited selection of important or representative areas for conservation.
In this study we have attempted to deine ‘coastal forest’
in a more restricted way, at least as regards those in northern
Mozambique. It is hoped this deinition and understanding
will apply equally well to forests in Tanzania south of the
Ruiji River.
We deine coastal dry forest here as essentially dry forest or thicket formations that are found within 50–100 km
of the coast. These are generally vegetation formations with
a closed or almost-closed canopy (80% cover or more when
undisturbed) with a high proportion of deciduous woody species that lose most of their leaves during the long dry season.
The deinition does not include moist forest, i.e. forest dominated by species with non-drought adapted leaves, nor does it
include vegetation that is dominated or characterised by what
are primarily woodland species (such as Brachystegia or Julbernardia). Moist forests of the continental interior may contain some tree species that are found in coastal forests, but,
128
unless such species are dominant, this does not mean they
are coastal forests. These moist forests, miombo and similar
woodlands and mangroves are excluded, as is vegetation associated with watercourses such as rivers. Coastal dry forests
are characterised as much by their species composition as by
their physical structure (which can of course be modiied by
land uses such as logging or cultivation). Under our deinition
the main characteristics of coastal dry forests are:
• Dry, deciduous and semi-deciduous forest (80% canopy
cover or more), becoming thicket-like with disturbance.
The main species are not moisture-demanding or even
mesic. They occur in areas subject to a lengthy dry season
(in excess of 6 months), with the majority of tree species
responding by losing leaves.
• Contain a signiicant number of sclerophyllous evergreen
tree and shrub species in the understorey.
• Have a species composition signiicantly different from
that of the surrounding woodland (mostly miombo). The
overlap in species composition between the two types is
often less than 30%.
• Have a very patchy distribution, and are often apparently
restricted to particular soils, such as unconsolidated sands,
and geomorphological positions, such as in the upper part
of the catena.
• Show a marked change in species composition between
patches with very few species found regularly or widely.
There are a high number of species of restricted distribution, often from particular families and genera (e.g. Annonaceae, Leguminosae: Caesalpinioideae, Rubiaceae).
Typical sclerophyllous species commonly found in
coastal forests across Cabo Delgado include Manilkara sansibarensis (Engl.) Dubard, Warneckea sansibarica (Taub.)
Jacq.-Fél. and Baphia macrocalyx Harms, while Pteleopsis
myrtifolia (M.A.Lawson) Engl. & Diels and M. sansibarensis
are among the very few that are found in most forest patches.
STUDY AREA
The study area runs the length of Cabo Delgado Province in
north-eastern Mozambique from the Rovuma river, the border
with Tanzania, south to Pemba, with an extent of 18,150 km2
(ig. 2). Forming a long triangle, about 280 km at its longest and 100 km at its widest, between Mueda and Mocímboa
do Rovuma in the west, Quionga and the Rovuma estuary
in the northeast, and Pemba in the south, its limits are based
on geology and landform, encompassing only Cretaceous and
more recent deposits. Full details are given in Timberlake et
al. (2010).
Four smaller study sites were chosen (ig. 2) on the basis of their apparent good condition and size, uniqueness and
accessibility, and as they were different from each other in
terms of landscape and substrate.
a) Pundanhar–Nangade in Palma and Nangade Districts,
west of the Nhica area along the W-E higher ground associated with the Rio Rovuma, including a hunting concession (approx. 750 km2).
b) Palma–Nhica do Rovuma area in Palma District, along
the W-E higher ground associated with the lower reaches
of the Rio Rovuma and along the main road south from
Timberlake et al., Coastal dry forests in northern Mozambique
Palma (approx. 1400 km2).
c) Quiterajo in Macomia District, situated on the coast 45
km south of Mocímboa da Praia, just south of the Rio
Messalo (approx. 750 km2).
d) Lupangua in Quissanga District inside the Quirimbas National Park, 15 km south of Quissanga and opposite Ilha
Mefunvo (approx. 25 km2).
Our principal focus was on coastal dry forest formations,
so associated vegetation types such as Brachystegia-dominated miombo and similar woodlands, pan grassland and
riverine formations, were only looked at in order to obtain a
broader context.
Geomorphology, geology and soils
The area comprises a gently tilting interior plateau, rising
from about 100 m above sea-level along the Palma–Mocímboa road to over 600 m in the west along the Mueda escarpment. However, most of our study sites lie between 80 and
180 m, although patches of dry forest were found down to an
altitude of 40 m. To the east of the Palma–Mocímboa road the
land drops down to a narrow coastal plain. Much of the interior plateau acts as a ‘sponge’ with pans and edaphic grasslands
resulting from seasonally-poor drainage. There are numerous
drainage lines lowing to the south east or, in the northernmost section, to the north east, and some are quite deeply
incised closer to the coast. On the northern margin, the Rio
Rovuma has cut through these plateau sediments to create a
wide valley (c. 10 km wide), forming the border with Tanzania. Similar land forms are found in south-east Tanzania.
North-eastern Cabo Delgado is seen to have a different
geological origin from much of the rest of Mozambique (national geological map, ING 1987), comprising younger formations dating from the Lower Cretaceous period up to the
Neogene lying adjacent to the much older continental block of
Precambrian granites and other rocks. There is also a narrow
coastal strip comprising uplifted recent Quaternary deposits.
As strata in these apparently marine deposits are relatively
level, the landform is primarily determined by differential resistance to erosion by the different strata and retreating scarp
erosion, resulting in numerous lat-topped plateaux.
Nearly all the coastal dry forest patches seen were located
on iron-rich sandstone and conglomerates of the Mikindani
Formation (mid-Neogene, ± 15–10 mya), while associated
miombo and similar woodlands were found on more recent
Quaternary formations (Pleistocene, ± 1.6–0.01 mya). This
is clearly seen in the Quiterajo area with its forest-capped
plateau, but is less obvious along the Rovuma rim (Nhica do
Rovuma to Nangade).
It would appear that the extent of soils derived from the
Mikindani Formation, comprising coarse, unstructured, welldrained sands, red-brown in colour, and possibly quite acidic,
are a major determinant of the distribution of coastal forests
here. However, this link has not yet been tested and causation
is not fully established.
Climate
Summary monthly data have been taken from Kassam et al.
(1981), which covers various nearby localities. Rainfall is
around 1000 mm/year falling in a single rainy season, while
potential evapo-transpiration signiicantly exceeds rainfall
for the period May to November–December, giving a growing season of around 4–5 months. There is a long hot dry season before a single clearly-deined rainy season from December to April, while atmospheric humidity during November
and early December is high.
Although there is a coastal inluence and some effects
from the Indian Ocean monsoon, the climate of the study
area generally follows that more typical of the continental
interior. This part of northern Mozambique lies partly in the
rain-shadow of Madagascar (Clarke 2000b), so has a somewhat lower rainfall than areas of the country further south or
central Tanzania.
SURVEY RESULTS
Past and present extent of forest
Figure 2 – Cabo Delgado study area showing the four detailed study
sites (white lines), and present extent of ‘dense vegetation’ cover
(green blocks).
Estimates of the remaining extent of coastal dry forest and
similar formations in the study area were made using photographic copies of 1999–2002 false-colour Landsat ETM imagery at 1 : 250,000 scale. An approximation was also made
of the possible original extent of dense vegetation cover some
129
Pl. Ecol. Evol. 144 (2), 2011
Table 1 – Original and remaining extent of dense woodland and dry forest vegetation types, based on interpretation of 2002 Landsat
imagery.
area
original extent
(km2)
NW, Mueda plateau & E slopes
NE, Nangade–Pundanhar–Nhica–Quionga–Palma–Mocímboa
EC, Rio Messalo–Quiterajo
S, Chai–Mucojo–Pemba, S of Rio Messalo
total
2332
2173
576
1006
6087
100–150 years ago, before signiicant changes in land use
occurred. Delineation was by manual interpretation using a
transparent mylar sheet overlay; a dot planimeter was used to
determine area. Results are presented by geographical block.
Original extent – The possible original extent of dry forest was determined using a combination of (a) suitable upland landform, (b) underlying geology, and (c) relectance
(smooth, reddish texture on false-colour imagery). A reduced
image at 1: 800,000 scale was used to obtain a better synoptic
view. Based on ield observations of current species composition and distribution, as dry forest patches appear to be restricted to certain soils and landscape units, it is believed that
only 10% of the area delineated as ‘dense vegetation cover’
would have been dry forest, with the majority being miombo
or similar woodland types. However, these igures may be
signiicantly in error.
The suggested original extent of ‘dense vegetation cover’
was 6080 km2 (table 1), and within that the extent of dry forest is estimated at around 615 km2, or 3% of the entire study
area. Even with a higher proportion of dry forest vs. woodland, the original extent is unlikely to have been more than
1000 km2. Although the assumptions are not substantiated,
given the lack of historical data this is probably the best igure that can be obtained. The main densely vegetated areas
were thought to be in the north along the Rovuma margin and
along the eastern slopes of the Mueda plateau, but there were
also signiicant blocks around the lower reaches of the Rio
Messalo and in the Macomia area.
Present extent – A more detailed assessment using 1 : 250,000
scale images was made of the present extent of dense vegetation cover. Various key assumptions were made, including (a) that relatively smooth-textured non-mottled reddish
areas on false-colour imagery indicate well vegetated sandy
soils from the Mikindani Formation, most of which lie up on
the plateau, while blue areas indicate woodland or grassland;
(b) that deeper red-hued, ± homogeneous blocks are likely to
be dry forest or dense woodland; (c) mottled areas include
signiicant cultivation and were excluded; (d) around a third
(rounded igures) of suitable substrate is likely to support dry
forest patches rather than dense woodland, transitional areas
or areas with just a sprinkling of dry forest species; and (e)
rugged terrain along the Rovuma valley rim was included as
ield experience showed these areas support some good dry
forest patches.
Based on these assumptions, the total extent of ‘dense
vegetative cover’ was calculated to be 1182 km2 in 2002, with
130
present extent (km2)
dense veg.
dry forest
cover
89
30
769
260
166
55
158
53
1182
398
loss of dense
veg. cover (%)
96.2
64.6
71.2
84.3
80.6
398 km2 of this being dry forest (table 1), or 2% of the total
study area. The overall loss of dense vegetation cover from
the entire study area over the last 100–150 years appears to
be around 80%, with losses ranging from 96% on the Mueda
plateau to around 65% in the Nhica area.
The main dry forest areas are now found in the north-east
part of the study area associated with the southern margin
of the Rovuma valley from Pundanhar to Nhica do Rovuma,
and in the Quionga area associated with the Rio Luvumba/
Macanga that low into the Rovuma estuary (ig. 2). There are
also sizeable areas along the Palma–Mocímboa da Praia road.
Other signiicant areas of dry forest include Nahavara forest
near Quiterajo (31 km2) and others south of this, and the patch
at Lupangua (20 km2) in the Quirimbas National Park.
It is clear that large parts of the area have been cleared
for agriculture over the last 100 years, yet our ield work suggests signiicant additional expanses have been cleared close
to population centres and main roads since 2002. It is also
thought that forest quality on the ground is often low owing
to previous logging, old clearance for ields (5–50 years ago),
and frequent ire, none of which are readily detectable on the
imagery. Hence the igure of almost 400 km2 of remaining
dry forest extent given here is likely to be an overestimate
from a conservation viewpoint.
Botanical composition
The main focus of the botanical study was on plant collection, particularly from the dry forest patches. More open vegetation types, such as around pans, on the coastal margins and
in the Rovuma valley, were not well-collected. The expeditions took place just before the rains, so grasses and herbs are
poorly represented. Representative sets are held at RBG Kew
(K), IIAM in Maputo (LMA), and at the Muséum National
d`Histoire Naturelle, Paris (P), with any additional duplicates
in Maputo (LMU), Nairobi (EA) and Buffelskloof (BRNH).
Earlier identiications were done by Quentin Luke (EA) and
John Burrows (BRNH), with all others and conirmation done
at Kew, primarily by David Goyder, Frances Crawford and
Iain Darbyshire.
Over 3000 numbered collections were made during the
various earlier trips and two expeditions in 2008 and 2009.
A total of 738 plant taxa were recorded from 105 families,
of which 36 taxa are either entirely new to science or were
known previously from material too fragmentary to describe
formally (table 2; see Burrows 2009, Burrows & Burrows
2010). The largest family recorded was Rubiaceae with 83
Timberlake et al., Coastal dry forests in northern Mozambique
Table 2 – New and undescribed species recorded from Cabo
Delgado study area, 2003–2009.
Monocotyledons
Asparagaceae
Asparagus ?sp. nov.
Araceae
Stylochaeton sp., uncertain status
Dicotyledons
Annonaceae
Xylopia sp. nov.
Xylopia sp. A of FTEA
Asteraceae
Vernonia ?sp. nov. aff. inhacensis G.V.Pope
Vernonia ?sp. nov. 2
Celastraceae
Pleurostylia ?sp. nov. aff. serrulata Loes.
Convolvulaceae
Ipomoea ?sp. nov.
Euphorbiaceae
Euphorbia ?sp. nov. aff. ambroseae L.C.Leach
Flacourtiaceae
Casearia ?sp. nov.
Lamiaceae
Vitex ?sp. nov. aff. buchananii
Vitex cf. mossambicensis Gürke
Leguminosae: Mimosoideae
Acacia latispina J.M.& S.M.Burrows
Leguminosae: Papilionoideae
Baphia ?sp. nov.
Erythrina ?sp. nov.
Melastomataceae
Warneckea sp. nov.
Meliaceae
Trichilia ?sp. nov.
Ochnaceae
Ochna ?sp. nov.
Rubiaceae
?Chassalia cf. umbraticola Vatke
Didymosalpinx callianthus J.E.& S.M.Burrows
Oxyanthus sp. A of FZ
Oxyanthus bilorus J.E.& S.M.Burrows
Polysphaeria ?sp. nov.
Psilanthus sp. nov., cf. sp. A of FTEA
Pyrostria sp. B of FZ
Pyrostria sp. D of FTEA
Pyrostria ?sp. nov. = Luke 9724
Rytigynia cf. umbellulata (Hiern) Robyns
= de Koning et al. 9759 of FZ
Tarenna sp. 53 of Degreef 2006
Tricalysia sp. A of FZ
Tricalysia sp. B of FZ
Rutaceae
Vepris sp. nov.
Zanthoxylum lepreurii Guill.& Perr., subsp. nov.?
Sapindaceae
Deinbollia ?sp. nov.
Sterculiaceae
Cola sp. nov. 1 aff. clavata Mast.
Cola ?sp. nov. 2 aff. clavata Mast.
taxa, followed by Leguminosae: Papilionoideae with 43 (table 3). The family with the largest number of new species was
Rubiaceae (thirteen), followed by Annonaceae, Asteraceae,
Lamiaceae, Papilionoideae, Rutaceae and Sterculiaceae with
two each. A full list of species identiied is given in Annex
2 of Timberlake et al. (2010). Excluding new species, 68
new records are reported for Mozambique (table 4) from the
study area and immediate surrounds. We believe that this is
an exceptionally high number of new records to discover anywhere in southern and eastern Africa, and indicates not just
the marked lack of previous collecting, but also the richness
of the area and the high number of range-restricted species.
Distribution patterns
One of the study’s objectives was to see if the patterns of local endemism recorded from the Tanzania coast, especially in
the Lindi region (Bidgood & Vollesen 1992, Vollesen 1994,
Clarke 1998, 2001), were also seen in coastal northern Mozambique. This would require the mapping of distributions of
a large number of species using records both from this study
and from herbaria elsewhere, and has not yet been done. Instead, some preliminary observations are given on broad species distribution patterns for seven selected species and on
differences between vegetation types.
All available records for these seven species, known to be
restricted to coastal areas and showing differing distribution
patterns, were collated and mapped (ig. 3). Two species are
endemic to coastal northern Mozambique [Micklethwaitia
carvalhoi (Harms) G.P.Lewis & Schrire and Thespesia mosTable 3 – Number of taxa from the main plant families found in
the Cabo Delgado study.
family
Aracaeae
Orchidaceae
Acanthaceae
Annonaceae
Apocynaceae
Asteraceae
Capparaceae
Celastraceae
Combretaceae
Ebenaceae
Euphorbiaceae
Lamiaceae
Leg.: Caesalpinioideae
Leg.: Mimosoideae
Leg.: Papilionoideae
Rubiaceae
Rutaceae
Sapindaceae
Sterculiaceae
Tiliaceae
Vitaceae
no. taxa
12
11
18
23
28
11
19
14
17
12
41
28
24
29
43
83
9
12
11
12
6
no. new
species
1
2
2
1
1
2
1
2
13
2
1
2
-
no. new Moz
records
4
5
2
5
2
1
2
1
3
3
3
2
1
3
8
2
1
1
2
131
Pl. Ecol. Evol. 144 (2), 2011
Table 4 – New records for Mozambique from Cabo Delgado study area, 2003–2009.
monocotyledons
Amaryllidaceae
Crinum aurantiacum Lehmiller
Anthericaceae
Chlorophytum amplexicaule Baker
Araceae
Amorphophallus maximus (Engl.) N.E.Br. subsp. ischeri
(Engl.) Govaerts & Frodin
Anchomanes abbreviatus Engl.
Culcasia orientalis Mayo
Stylochaeton euryphyllus Mildbr.
Arecaceae
Hyphaene petersiana Mart.
Dracaenaceae
Sansevieria cf. metallica Gérôme & Labroy
Orchidaceae
Eulophia acutilabra Summerh.
Eulophia guineensis Lindl.
Microcoelia megalorrhiza (Rchb.f.) Summerh.
Microcoelia physophora (Rchb.f.) Summerh.
Nervilia bicarinata (Blume) Schltr.
dicotyledons
Acanthaceae
Lepidagathis plantaginea Mildbr.
Whitieldia orientalis Vollesen
Anacardiaceae
Lannea schweinfurthii (Engl.) Engl. var. acutifolia
(Engl.) Kokwaro
Annonaceae
Artabotrys modestus Diels subsp. macranthus Verdc.
Lettowianthus stellatus Diels
Monanthotaxis faulknerae Verdc.
Monanthotaxis trichantha (Diels) Verdc.
Monodora minor Engl.& Diels
Apocynaceae
Baissea myrtifolia (Benth.) Pichon
Cryptolepis hypoglauca K.Schum.
Asteraceae
Vernonia zanzibarensis Less.
Balanitaceae
Balanites maughamii Sprague subsp. acuta Sands
Burseraceae
Commiphora fulvotomentosa Engl.
Commiphora pteleifolia Engl.
Capparaceae
Maerua bussei (Gilg & Gilg-Ben.) Wilczek
Ritchiea capparoides (Andr.) Britton var. capparoides
Celastraceae
Elaeodendron buchananii (Loes.) Loes.
Connaraceae
Vismianthus punctatus Mildbr.
Cucurbitaceae
Peponium leucanthum (Gilg) Cogn.
Ebenaceae
Diospyros kabuyeana F.White
132
dicotyledons
Ebenaceae
Diospyros magogoana F.White
Diospyros shimbaensis F.White
Euphorbiaceae
Croton polytrichus Pax subsp. polytrichus
Drypetes sclerophylla Mildbr.
Omphalea mansieldiana Mildbr.
Lamiaceae
Orthosiphon scedastophyllus A.J.Paton
Premna gracillima Verdc.
Premna hans-joachimii Verdc.
Leguminosae: Caesalpiniodeae
Scorodophloeus ischeri (Taub.) J.Léonard
Senna auriculata (L.) Roxb.
Leguminosae: Mimosoideae
Newtonia paucijuga (Harms) Brenan
Leguminosae: Papilionoideae
Dalbergia lactea Vatke
Erythrina haerdii Verdc.
Erythrina sacleuxii Hua
Loganiaceae
Strychnos xylophylla Gilg
Myrtaceae
Eugenia capensis (Eckl.& Zeyh.) Sond. subsp. multilora Verdc.
Ochnaceae
Ochna ovata F.Hoffm.
Passiloraceae
Adenia kirkii (Mast.) Engl.
Rubiaceae
Coffea schliebenii Bridson (Coffea sp. D of FTEA)
Gardenia transvenulosa Verdc.
Kraussia kirkii (Hook.f.) Bullock
Leptactina papyrophloea Verdc.
Pavetta lindina Bremek.
Rhodopentas parvifolia (Hiern) Kårehed & B.Bremer
Rothmannia macrosiphon (Engl.) Bridson
Vangueria cf. randii S.Moore subsp. vollesenii Verdc.
Rutaceae
Vepris sansibarensis (Engl.) Mziray
Zanthoxylum lindense (Engl.) Kokwaro
Sapindaceae
Haplocoelum inoploeum Radlk.
Thymelaeaceae
Synaptolepis kirkii Oliv. sensu stricto
Tiliaceae
Grewia stuhlmannii K.Schum.
Violaceae
Rinorea welwitschii (Oliv.) Kuntze subsp. tanzanica GreyWilson
Viscaceae
Viscum gracile Polhill & Wiens
Vitaceae
Cissus phymatocarpa Masinde & L.E.Newton
Cissus sylvicola Masinde & L.E.Newton
Timberlake et al., Coastal dry forests in northern Mozambique
A
B
C
D
E
F
G
Figure 3 – Distribution records of selected coastal species.
A, Guibourtia schliebenii; B, Hexalobus mossambicensis;
C, Micklethwaitia carvalhoi; D, Monodora minor; E,
Scorodophloeus ischeri; F, Thespesia mossambicensis; G,
Vismianthus punctatus. Black line = extent of occurrence
(EOO); red squares = area of occupancy (AOO) using 4 km2
grid; blue polygon = extent of subpopulations using Rapoport’s
mean propinquity technique.
133
Pl. Ecol. Evol. 144 (2), 2011
Table 5 – Global IUCN Red Data status for seven selected Eastern African dry forest species and 2010 assessments from the East
African Plant Red List Authority (EAPRLA).
family/species
Annonaceae
Hexalobus mossambicensis N.Robson
Monodora minor Engl. & Diels
Connaraceae
Vismianthus punctatus Mildbr.
Leguminosae: Caesalpinioideae
Guibourtia schliebenii (Harms) J.Léonard
Micklethwaitia carvalhoi (Harms) G.P.Lewis &
Schrire
Scorodophloeus ischeri (Taub.) J.Léonard
Malvaceae
Thespesia mossambicensis (Exell & Hillc.) Fryxell
No.
records
EOO
(km2)
AOO
(4 km2 cell)
status
EAPRLA assessment
11
23
45,043
54,062
28
68
EN
EN
NT
11
22,864
40
EN
VU B1ab(iii)+2ab(iii)
18
164,778
64
EN
VU B2ab(ii,iii,iv,v)
8
9,502
28
EN
-
72
137,578
212
EN
LC
5
16,416
20
EN
-
sambicensis (Exell & Hillc.) Fryxell], one shows a Cabo
Delgado–Lindi distribution (Vismianthus punctatus Mildbr.), and three show a somewhat broader Tanzania–Mozambique coastal distribution [Monodora minor Engl. & Diels,
Guibourtia schliebenii and Scorodophloeus ischeri (Taub.)
J.Léonard], although Guibourtia is also found inland on the
Eastern Arc mountains and Scorodophloeus is also found in
coastal Kenya.
During the identiication process it was noted that twelve
taxa were endemic to coastal northern Mozambique (area
Moz N: of Flora Zambesiaca) plus around nineteen of the
new species. There are 53 taxa known primarily from northern Mozambique and adjacent areas of south-east Tanzania
(area T8 of Flora of Tropical East Africa), showing the strong
links between them. Most of these appear to be local endemics. A further 46 taxa appear to be at the southern end of their
East African (Swahilian) distribution in Cabo Delgado.
It is also apparent that species in miombo and similar
woodland types, fallows, grassland and wetlands, are far
more widespread than those from dry forests. Many of them
are found across the Miombo Ecoregion of south central Africa (Timberlake & Chidumayo 2001) or even more widely, e.g. Afzelia quanzensis Welw., Brachystegia spiciformis
Benth., Parinari curatellifolia Benth., Pseudolachnostylis
maprouneifolia Pax and Uapaca nitida Müll.Arg., in marked
contrast to the local distribution of so many dry forest species. An interesting exception is Berlinia orientalis Brenan
that commonly occurs in fallows and miombo woodland (and
sometimes in dry forest), but which is known only from a
limited area of Cabo Delgado across to south-east Tanzania
(Luke 2004).
Future work will probably conirm these distribution patterns and show the very marked East African coastal element
in the overall distribution of the dry forests of the study area,
with particularly strong links to that found with the Lindi–
Mtwara region of south-east Tanzania, while also showing
that woodland and grassland lora species have a much wider
distribution.
134
Red Data assessments
Preliminary conservation assessments to determine IUCN
Red Data List categories were carried out on the seven woody
species used to map distribution patterns. All were chosen
primarily for their known restricted coastal distribution patterns or ecological importance and because there were adequate data points.
By using the Kew GIS Unit’s extension tool (www.kew.
org/gis/projects/cats/catsdoc.pdf), rapid conservation assessments based on IUCN categories and criteria were produced
(IUCN 2008). Data points were derived from collections
made during this study and from historic specimens held
at Kew, those available from the African Plants Initiative
(http://plants.jstor.org) and from Missouri Botanic Garden’s
Tropicos database (http://www.tropicos.org), which includes
specimens from the East African Herbarium (EA). The assessments calculate the Extent of Occurrence (EOO), Area of
Occurrence (AOO), number of sub-populations and number
of locations (table 5). The cell size used for AOO was 4 km2
(2 × 2 km), recommended by IUCN (2008) for restricted distribution species.
From the individual species assessments, distribution
maps were compiled (ig. 3) and the species’ IUCN status
(IUCN 2001) determined (table 5). By using the default 4
km2 cell size all species were assessed as being Endangered
(EN). If a larger cell sizes were used (e.g. 35–85 km sides),
the threat status decreased in most cases to Least Concern.
Of the seven species, the most narrowly distributed was the
Mozambique endemic Thespesia mosambicensis, occurring
in only twenty cells, while the other Mozambique endemics
Hexalobus mossambicensis and Micklethwaitia carvalhoi
were recorded from just 28 cells. Scorodophloeus ischeri,
found from northern Mozambique to southern Kenya, was
found in 212 cells. As the process is automated, this can lead
to some odd maps, such as in ig. 3F. Here two obviously
separate populations are artiicially linked by the EOO, which
is more an indication of the spread of risk. In this case the
restricted AOO should be regarded as the more deinitive indication of distribution and risk.
Timberlake et al., Coastal dry forests in northern Mozambique
Conservation assessments are already available on the
IUCN Red List website (http://www.iucnredlist.org) for three
of these species – Guibourtia schliebenii (VU B1+2b), Hexalobus mossambicensis (DD) and Monodora minor (Near
Threatened) – the irst two dating from 1998 before much
information was available for the Mozambique populations.
Recently, four of the seven species above were assessed by
the East African Plant Red List Authority (EAPRDA, Q.
Luke, pers. comm.) with results shown in table 5. In three
cases the assessments were lower, probably as more local information was available on their status in Tanzania and Kenya. Of particular note is Scorodophloeus, which the EAPRDA
assessed as Least Concern rather than Endangered.
Although the preliminary assessments are based solely on
recorded past and present distributions derived from herbarium specimens, we know the habitat is under marked threat,
hence the Endangered status may be appropriate for some
species. It is likely that a number of other dry forest species,
once assessed, will have a similar threat status. In contrast,
the great majority of species associated with woodland across
the study area are known to have a much wider sub-continental distribution and will likely be assessed as Least Concern.
CONTEXT AND CONSERVATION
Regional context
Across south-central Africa numerous, generally small (less
than 20 km2) patches of dry forest can be found within the
matrix of miombo or similar Caesalpinoid-dominant woodland types (Timberlake & Chidumayo 2001, Frost et al.
2002). Examples are the extensive Cryptosepalum exfoliatum De Wild. forests of north-west Zambia (Fanshawe 1973,
Burgess et al. 2004a, type 32); the Itigi thickets in central
Tanzania dominated by Baphia burttii Baker f., B. massaiensis Taub., Combretum celastroides M.A.Lawson and Bussea massaiensis (Taub.) Harms. (Burtt 1942, Burgess et al.
2004a, type 48); similar forests around Lake Mweru Wantipa in north-east Zambia; forest patches dominated by Xylia
torreana Brenan in the middle Zambezi valley and northern
Zimbabwe (Timberlake et al. 1993, type C2); and patches
characterized by Guibourtia conjugata (Bolle) J.Léonard on
Cretaceous sands in south-eastern Zimbabwe and the northern Kruger National Park in South Africa. Sometimes termed
‘sand forests’ these vegetation types are primarily found on
unconsolidated medium to coarse-textured sandy soils (often
acidic) in the higher part of the catena.
Such dry forests show much similarity to many of the
coastal forest patches seen in Cabo Delgado, particularly as
regards the substrate, geomorphological position and in the
families and even genera of characteristic plant species. As
with coastal forests, there is a characteristic change in species
composition across an area, and always a marked difference
in species composition compared to that of the surrounding
woodlands. In addition, many canopy species of dry forests
of the continental interior are also early-deciduous with scattered sclerophyllous evergreen species in the understorey.
With disturbance dry forest tends towards thicket, as can be
clearly seen with the ‘jesse bush’ of the mid-Zambezi valley
and in the Itigi thickets.
This raises an interesting question of whether all these dry
forests – coastal and those on the old African plateau of the
continental interior – are remnants of an earlier drier period,
or whether it is a case of convergent evolution of vegetation
type. However, coastal dry forests do show much higher
levels of endemism and contain more species of restricted
coastal distribution compared to those of the interior. For the
coastal forests it is not clear if the species are relictual or,
alternatively, what the driver might have been for their evolution given the relatively recent age of these patches in the
landscape. The soils on which they are found are generally
not old and infertile, as is the case with other areas of high
endemism such as the Chimanimani mountains, Mt. Mulanje
or the sandstones of the Cape mountains.
Conservation
The coastal strip of this part of Cabo Delgado has been settled for hundreds of years. Slave trading was signiicant during the 19th century and there was much trade at that time
in ivory, gum copal (from Hymenaea verrucosa Gaertn.) and
‘wild rubber’ (Landolphia spp.). During the 20th century, in
particular after the Second World War, there was extensive
exploitation of timber by the Portuguese colonial authorities,
focussing primarily on Pterocarpus angolensis Harms (Umbila), Afzelia quanzensis (Chanfuta), Dalbergia melanoxylon
Guill. & Perr. (Pau-preto), Milicia excelsa (Welw.) C.C.Berg
(Tule, Mvule), Millettia stuhlmannii Taub. (Jambiri, panga
panga) and Swartzia madagascariensis Desv. (Pau-ferro), the
effects of which can still be seen today (Timberlake et al.
2010).
Although many people moved out of rural areas during the independence struggle and civil war years (1960s to
1991), they are now starting to reoccupy many parts of the
interior. This has been helped by better roads and transportation, including the construction of cut-lines during recent
(2008) geophysical prospecting for oil, the sinking of wells
and boreholes, and an inlux of settlers from adjacent parts of
Tanzania. The biggest threats to coastal forests at present are
the rapid and uncontrolled clearance for subsistence agriculture, which is increasingly moving away from the main roads,
logging (much of it illegal), and uncontrolled ires associated
with settlement.
There is no formal protection of any coastal forest patches
in Cabo Delgado except for a very small extent (less than 25
km2) inside the newly-proclaimed Quirimbas National Park.
Given their characteristic features – the high turnover in species composition between patches, which often makes each
patch unique, and the number of species with very restricted
distributions – it is dificult to identify a ‘typical’ area for
conservation protection or to cover the full range of important
species in just a few areas.
Timberlake et al. (2010) have described fourteen areas of
importance for conservation of coastal forests and associated
vegetation types from across the study area. These cover the
more diverse and intact areas identiied so far and also various sites containing species of interest. Four sites (Pundanhar–Nangade, Rio Macanga–Nhica do Rovuma, Quiterajo,
Lupangua) are considered high priority. What is apparent
is that in the northern parts along the Rovuma escarpment,
135
Pl. Ecol. Evol. 144 (2), 2011
landscape-level conservation would be the most appropriate,
covering dry forest, dense woodland, wooded grassland and
the large seasonal pans – an ecosystem approach. This area is
also a major source of water for coastal towns. Further south,
where settlement is much denser, a site approach would be
more appropriate, for example the raised plateau with Guibourtia schliebenii forest in the Quiterajo concession south
of the Messalo River.
Current research at RBG Kew and IIAM, Maputo is attempting to identify Important Plant Areas across the broad
study area. Once zoological indings from recent surveys are
available it is planned to incorporate these in order to identify
Key Biodiversity Areas. Such identiied and documented areas will be brought to the attention of government and provincial authorities in Mozambique, who have already indicated
interest in the study.
CONCLUSIONS AND FUTURE STUDIES
Although the area studied in detail was limited in extent,
compounded by dificulties in access over much of it, some
conclusions on the extent, distribution and biodiversity of
coastal forests in northern Mozambique can be drawn.
1. Coastal forests similar to those in south-eastern Tanzania
are present, but are far less extensive than had previously
been suggested. The present extent of dry forest is around
400 km2, although this may well be an over-estimate.
Parts of this are now being rapidly cleared for agriculture.
2. Most of the forest patches seen are relatively small and
conined to deeper, well-drained sandy soils, most apparently derived from the Mikindani sandstone formation.
They are surrounded by woodland vegetation dominated
by typical miombo species such as Brachystegia, other
caesalpinoids and some Euphorbiaceae trees, species that
(with the exception of the coastal Berlinia orientalis) are
mostly very widely distributed across the sub-continent.
3. The forests generally comprise an upper canopy of deciduous species, many from the Caesalpinioideae (e.g. Guibourtia schliebenii, Hymenaea verrucosa, Micklethwaitia
carvalhoi), with sclerophyllous evergreen species in the
sub-canopy (e.g. Manilkara spp., Warneckea sansibarica,
Baphia macrocalyx). The shrub layer is particularly rich
in Rubiaceae. A surprisingly high proportion of these species have quite restricted ranges or are local endemics,
being conined to parts of the coastal regions of southern
Kenya, Tanzania and northern Mozambique. Many are
conined to the Lindi–Mtwara–Cabo Delgado area, the
so-called Lindi centre of endemism in the Swahilian regional centre of endemism (Clarke 1998, 2001). It is this
attribute – the high proportion of range-restricted species
– that gave rise to the discovery of so many new species
and Mozambique records.
4. Despite the name, such coastal forests do not contain
mesic evergreen species and show little similarity in composition or ecology to moist forests in the region, whether
montane or lowland.
5. Another interesting feature is the high turn-over in species
composition between forest patches. Most patches appear
to be unique, and there is no species that is characteristic or found in most of them, the closest being Pteleopsis
136
myrtifolia (Combretaceae). Each patch appears to have a
different dominant and suite of associates. This feature is
either related to subtleties in the mineralogical or moisture status of the substrate, or could be solely an artefact
of serendipity – the irst species to arrive taking over.
6. The coastal forests are essentially ‘sand forests’ and, in
many cases, show marked similarity in their structure,
ecology and family (even generic) composition to areas
of dry forest or thicket found on well-drained sandy soils
across much of south-central Africa.
7. The high proportion of range-restricted species, the limited extent of the forest patches, and the increased threat,
show that the coastal forests of northern Mozambique
should be of international conservation concern. The possibilities of landscape or ecosystem-level conservation
are now very limited, so attention also needs to be given
to the selection of a range of sites across Cabo Delgado
in order to conserve the full range of forest types and species.
The origins and afinities of these coastal dry forests,
as well as determination of what the drivers for the marked
levels of speciation and endemism might have been, remain
important areas for research. But perhaps more urgent at this
stage is continued documentation of the extent of coastal forests in other parts of Mozambique and the development of
practical conservation measures needed to protect a rapidly
disappearing vegetation type.
ACKNOWLEDGEMENTS
We would like to thank Pro Natura International and the
Muséum National d’Histoire Naturelle, Paris, for organising
the expeditions under their Our Plant Reviewed programme,
and the Prince Albert II of Monaco Foundation, the Stavros
Niarchos Foundation and the Total Foundation for funding.
Much help in the ield was given by Salim Annebi, Aurelio
Banze, Sandie Burrows, Mervyn Lotter, Patricia Luke, Alice
Massingue, Frédéric Mathias and Barbara Turpin. Field support and logistics during the expedition were ably provided
by Mike Scott and his team from Khangela Safaris, Bulawayo, while Charlie Mackie provided aerial reconnaissance.
We are particularly grateful to Iain Darbyshire and Tim Harris of the Drylands Africa team at RBG Kew for assistance
in plant identiication; Aaron Davis, Diane Bridson and the
Rubiaceae team, Gill Challen, Martin Cheek, Anna Haigh,
Ana-Claudia Araujo and Paul Wilkin also made valuable contributions. Erik Prins produced the satellite images and Justin
Moat of the Kew GIS Unit helped prepare the maps. Neil
Burgess and an anonymous reviewer made valuable comments on an earlier version of the paper.
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Paper based on results presented during the XIXth AETFAT Congress (Madagascar 2010). Manuscript received 30 Sep. 2010; accepted in revised version 15 Feb. 2011. This paper will be reprinted
in the Proceedings of the XIXth AETFAT Congress.
Communicating Guest Editor: Mats Thulin.
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