Finisterra, LVI(116), 2021, pp. 99‑114
ISSN: 0430-5027
doi: 10.18055/Finis20156
Artigo
PLANT COMMUNITIES OF NAMIBE SALTMARSHES
(SOUTHWEST OF ANGOLA)
João Francisco Cardoso1
José Carlos Costa2
Carlos da Silva Neto3
Maria Cristina Duarte4
Tiago Monteiro-Henriques5,6
ABSTRACT – This work constitutes the first phytosociological analysis of saltmarshes
on the Angolan coast. Sixty-five relevés were carried out resulting in the description of six
new plant associations. These saltmarshes are characterized by a lower floristic richness
when compared to the Holarctic saltmarshes. Eighteen taxa were identified, some of them
succulent. Saltmarshes occur from the mouth of the Cunene River to the Cuanza River,
although in this last part they are already very impoverished. In the Cuanza river, saltmarshes occupy only a narrow strip in the inner sector of the mangroves in contact with continental ecosystems and are often made up of just one taxon, Sarcocornia natalensis subsp.
affinis. Mangroves reach their southern limit in the city of Lobito, although they are almost
extinct there. The occurrence and distribution of saltmarshes are affected by the Cold Benguela Current, that influences the west coast of Africa between Cabo da Boa Esperança and
Benguela. The height of the saltmarsh’s platforms colonized by halophyte, sub-halophyte or
halotolerant plants determines the flooding period and thus the plant community’s floristic
composition. Soil granulometry also plays an important role in the spatial organization of
Recebido: maio 2020. Aceite: março 2021.
1
Universidade José Eduardo dos Santos, Huambo, Angola. E-mail: joaofca1974@gmail.com
2
Linking Landscape, Environment, Agriculture And Food (Leaf), Instituto Superior de Agronomia, Universidade de Lisboa,
Tapada da Ajuda, 1349-017, Lisboa, Portugal. E-mail: jccosta@isa.ulisboa.pt
3
Centro de Estudos Geográficos, Instituto de Geografia e Ordenamento do Território, Universidade de Lisboa, Lisboa, Portugal.
E-mail: cneto@campus.ul.pt
4
Centre for Ecology, Evolution and Environmental Changes (cE3c), Faculdade de Ciências, Universidade de Lisboa, Lisboa, Portugal. E-mail: mcduarte@fc.ul.pt
5
Centro de Investigação e Tecnologias Agroambientais e Biológicas (CITAB), Universidade de Trás-os-Montes e Alto Douro
(UTAD), Vila Real, Portugal. E-mail: tmh@isa.ulisboa.pt
6
Global Change and Conservation Lab (GCC), Faculty of Biological and Environmental Sciences, University of Helsinki,
Helsínquia, Finlândia.
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Cardoso, J. F., Costa, J. C., Neto, C. S., Duarte, M. C., Monteiro-Henriques, T. Finisterra, LVI(116), 2021, pp. 99-114
plant communities. One of the main originalities of Angolan saltmarshes is the predominance of fine sandy or sandy-loam soil texture as a consequence of the proximity of the
Namibe desert. The PCA segregated the different plant communities described.
Keywords: Saltmarshes; phytosociology; PCA; vegetation.
RESUMO – COMUNIDADES VEGETAIS DOS SALGADOS DO NAMIBE
(SUDOESTE DE ANGOLA). Este trabalho constitui a primeira análise fitossociológica de
sapais da costa angolana. Sessenta e cinco inventários foram realizados resultando na descrição de seis novas associações de plantas. Estes sapais são caracterizados por uma menor
riqueza florística quando comparadas aos sapais holárticos. Dezoito táxons foram identificados, alguns deles suculentos. Os sapais ocorrem desde a foz do rio Cunene até ao rio Cuanza,
embora nesta última parte já estejam muito empobrecidos. No rio Cuanza, os sapais ocupam
apenas uma faixa estreita no setor interno dos mangais em contato com os ecossistemas continentais e geralmente são constituídos por um único táxon, Sarcocornia natalensis subsp. affinis. Os mangais atingem seu limite sul na cidade de Lobito, embora estejam quase extintos. A
ocorrência e distribuição dos sapais são afetados pela Corrente Fria de Benguela, que influencia a costa oeste da África entre o Cabo da Boa Esperança e Benguela. A altura das plataformas
do sapal colonizadas por plantas halófitas, subhalófitas ou halotolerantes determina o período
de inundação e, portanto, a composição florística das comunidades vegetais. A granulometria
do solo também desempenha um papel importante na organização espacial das comunidades
de plantas. Uma das principais originalidades dos sapais angolanos é o predomínio de solos de
textura arenosa fina ou franco-arenosa, em consequência da proximidade com o deserto do
Namibe. A ACP segregou as diferentes comunidades de plantas descritas.
Palavras-chave: Sapais; fitossociologia; ACP; vegetação.
RÉSUMÉ – COMMUNAUTÉS VÉGÉTALES DES MARAIS SALÉS DE NAMIBE
(SUD-OUEST D’ANGOLA). Ce travail constitue la première analyse phytosociologique des
marais salants de la côte angolaise. Soixante-cinq relevées ont été réalisés aboutissant à la
description de six nouvelles associations végétales. Ces marais salants se caractérisent par
une richesse floristique plus faible que les marais salants holarctiques. Dix-huit taxons ont
été identifiés, certains d’entre eux succulents. Les marais salants se trouvent dès l’embouchure de la rivière Cunene à la rivière Cuanza, bien que dans cette dernière partie ils soient
déjà très appauvris. Dans la rivière Cuanza, les marais salants n’occupent qu’une bande
étroite dans le secteur intérieur des mangroves en contact avec les écosystèmes continentaux
et sont souvent constitués d’un seul taxon, Sarcocornia natalensis subsp. affinis. Les mangroves atteignent leur limite sud dans la ville de Lobito bien qu’elles y soient presque éteintes.
La présence et la distribution des marais salants sont affectées par le courant froid de Benguela qui influence la côte ouest de l’Afrique entre le Cabo da Boa Esperança et Benguela. La
hauteur des plates-formes du marais salé colonisées par des plantes halophytes, soushalophytes ou halotolérantes détermine la période d’inondation et donc la composition floristique des communautés végétales. La granulométrie du sol joue également un rôle important dans l’organisation spatiale des communautés végétales. L’une des principales
originalités des marais salants angolais est la prédominance de sols de texture sableuse ou
sablo-limoneuse en raison de la proximité du désert de Namibe. L’ACP a séparé les différentes communautés végétales décrites.
Mot clés: Marais salés; phytosociologie; ACP; végétation.
Cardoso, J. F., Costa, J. C., Neto, C. S., Duarte, M. C., Monteiro-Henriques, T. Finisterra, LVI(116), 2021, pp. 99-114
101
RESUMEN – COMUNIDADES VEGETALES DE LAS MARISMAS DE NAMIBE
(SUROESTE DE ANGOLA). Este trabajo constituye el primer análisis fitosociológico de las
marismas del litoral angoleño. Se realizaron sesenta y cinco inventarios que dieron como
resultado la descripción de seis nuevas asociaciones de plantas. Estas marismas salinas se
caracterizan por una menor riqueza florística en comparación con las marismas salinas de
Holarctic. Se identificaron dieciocho taxones, algunos de ellos suculentos. Las marismas se
dan desde la desembocadura del río Cunene hasta el río Cuanza, aunque en este último
tramo ya están muy empobrecidas. En el río Cuanza, las marismas ocupan solo una franja
estrecha en el sector interior de los manglares en contacto con los ecosistemas continentales
y, a menudo, están formadas por un solo taxón, Sarcocornia natalensis subsp. affinis. Los
manglares alcanzan su límite Sur en la ciudad de Lobito, aunque allí están casi extintos. La
ocurrencia y distribución de las marismas se ven afectadas por la Corriente Fría de Benguela, que influye en la costa occidental de África entre Cabo da Boa Esperança y Benguela.
La altura de las plataformas de la marisma colonizada por plantas halófitas, subhalófitas o
halotolerantes determina el período de inundación y, por tanto, la composición florística de
las comunidades vegetales. La granulometría del suelo también juega un papel importante
en la organización espacial de las comunidades vegetales. Una de las principales originalidades de las marismas angoleñas es el predominio de la textura del suelo de arena fina o
franco-arenosa como consecuencia de la proximidad del desierto de Namibe. El ACP
segregó las diferentes comunidades vegetales descritas.
Palabras clave: Marismas; fitosociología; ACP; vegetación.
I.
INTRODUCTION
In the biosphere, amphibious and coastal tidal mudflat ecosystems are formed by
saltmarsh and mangrove habitats. The latter consist of a shrubby or arboreal, helo-halophytic vegetation formation, characteristic of the low-lying muddy areas, such as coastal
swamps or lagoon banks, within intertropical environments. Mangrove species are physiologically and morphologically adapted to the amphibian eu-haline environment,
withstanding high osmotic pressures and developing aerial root systems and pneumatophores (Moreira, 1984). Unlike mangroves, marshes don’t have arboreal species and colonise mainly temperate zones in the western part of the continents, under the influence of
cold sea currents. A marsh is defined as a “characteristic ecosystem of the highest part of
the coastal low-lying platforms, occupying the space between the low water neaps and the
high water springs. Morphologically it corresponds to the high slike and schorre; it is,
therefore, an amphibious ecosystem, with muddy or silty soils, salty, colonized by halo-helophytic herbaceous vegetation. It develops in estuaries, deltas, lagoons and marine
swamplands, especially in extratropical regions” (Moreira, 1984, p. 109).
In the intertropical zone of the African continent, mangrove ecosystems are found
mainly on the eastern coasts influenced by warm sea currents, as in Mozambique, for
instance (Beentje & Bandeira, 2007) where the coastline is influenced by the Mozambique Warm Current, which extends southwards by the Agulhas Current. Mangroves are
also common along the coasts of West Africa, from Senegal to Angola (Ward et al., 2016).
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Cardoso, J. F., Costa, J. C., Neto, C. S., Duarte, M. C., Monteiro-Henriques, T. Finisterra, LVI(116), 2021, pp. 99-114
Although at a similar latitude to Mozambique, Angola’s coastline is influenced by the
Benguela Cold Current and, therefore, the low-energy environments at the mouths of the
major watercourses and sheltered inlets are dominated by marshes, although mangroves
also occur. However, mangroves on the western coasts of the continents, apart from being
less frequent, have a much lower floristic richness than those on the eastern coasts
(Moreira, 1977). In Angola, mangroves have shown a very significant decrease due to
anthropic action (cutting trees for wood, systematic felling of all vegetation to eradicate
diseases, land conversion for agriculture and pollution, among others). In the past, they
extended south to Lobito where they are now extinct. Mangroves do not occur in the
south of Lobito and along the coast of southwest Angola and Namibia, and the amphibian
vegetation of muddy or sandy tidal platforms consists exclusively of saltmarshes.
The occurrence of saltmarshes in the west African coast north of the Cape of Boa
Esperança is associated with the Benguela Cold Current, which constitutes a wide flow
towards northwest forming the eastern branch of the South Atlantic circulation cell. Typically, cold water from the South Atlantic circulation flows up the west coast of Africa to
the latitude of Benguela, about 16° South. The upwelling system known as the Benguela
Upwelling System consists of a rise in cold, nutrient-rich water from depths of around
200-300m, which is responsible for the high productivity of Angola’s southwest ecosystems (Kirkman & Nsingi, 2019). However, its influence is irregular as to water characteristics, temperature, northern latitude reach, current intensity, etc. These oscillations are
related to two phenomena: on the one hand, frequently warm and nutrient-poor waters
from the South Indian Ocean Current, called the Agulhas Current, mix with the Benguela Current, making the thermal (and other) characteristics of the latter chaotic (Veitch
& Peven, 2017); on the other hand, about once every 10 years (Mann & Lazier, 2006),
during the process known as the Benguela Niño Current (Imbol Koungue et al., 2019),
the Angolan Warm Current extends abnormally southwards from the usual 15° S up to
25° S, extending also offshore up to 150km and with a depth of 50m (Richter et al., 2010).
Due to the double influence of the Angolan Hot Current and the Benguela Cold Current on the Angolan coast, we find that all mangroves are located north of Lobito, and, in
this entire sector, marshlands are floristically reduced to few succulent species (in most
cases only Sarcocornia natalensis subsp. affinis (Moss) S.Steffen, Mucina & G.Kadereit),
which are characteristic of marshlands of the inland sector of the mangrove forest, in
contact with continental ecosystems. These very impoverished versions of marshes had
also been identified for the coast of Mozambique in the inland mangrove sector (Moreira,
1977). In these versions, which Moreira (1977) calls “salty swamps”, are present species as
Sesuvium portulacastrum (L.) L. and Sporobolus virginicus (L.) Kunth, among others,
which also occur in Angolan marshlands. No mangroves occur south of Lobito, a reality
that extends through all the Namibian territory, as in both the southwest of Angola and
in the Namibian coast, the tide-influenced muddy or sandy-muddy platforms, where in
low energy environments, are colonised by marsh vegetation. Globally, in terms of floristic richness, these marshlands are frankly poorer than the Western Europe ones (and
poorer than all the Holarctic Kingdom saltmarshes) of both Mediterranean and temperate oceanic climates (Cardoso, 2014; Costa et al., 2009). This area is characterized by a
Cardoso, J. F., Costa, J. C., Neto, C. S., Duarte, M. C., Monteiro-Henriques, T. Finisterra, LVI(116), 2021, pp. 99-114
103
desert to semi-desert climate, where fresh water supplies are extremely scarce, unlike in
the European marshes, especially those in the Eurosiberian Region. In Angola the saltmarshes have their northernmost position in the Cuanza River estuary (near Luanda),
where a very impoverished version of these communities occupies a narrow strip in the
most inland sector of the mangrove forest, which occures extensively along this river.
The study we are presenting is the first phytosociological analysis of the saltmarshes
and brackish marshes of the southwest of Angola and, as such, the communities identified are unprecedented for science. We also make a description of the environmental
characteristics in which each community is included, as well as their floristic separation
mainly through indicator species.
II. MATERIAL AND METHODS
The vegetation relevés were collected according to the sigmatist and dynamic-catenal
phytosociology approach (Braun-Blanquet, 1979; Géhu & Rivas-Martínez, 1981; Rivas-Martínez, 1976, 2005; Theurillat et al., 2020). A floristic-statistical group analysis (Müller-Dombois & Ellenberg, 1974) was used for community definitions and synthetic table arrangements. The bioclimatic typology was based on the work of Rivas-Martínez et al. (2011). For
the identification of collected specimens, the most relevant bibliography for the area was used,
namely Flora Zambesiaca, Flora of Tropical Africa, Flora of Southern Africa, Flora of Tropical
East Africa, Conspectus Florae Angolensis (Brummitt, 1992; Exell & Fernandes, 1961; Exell
et al., 1970; Exell & Mendonça, 1937, 1951, 1954; Klaassen & Craven, 2003; Ornold & De
Wet, 1993; Retief & Herman, 1997; Vollesen, 2000) and their comparison with duly identified
herbarium specimens (LISC herbarium, University of Lisbon). The nomenclature used follows the Plants of the World Online database (Plants of the World Online [POWO], 2019).
The Braun-Blanquet scale values were converted to numerical values based on the
Müller-Dombois and Ellenberg methodology (1974), and a Principal Component Analysis (PCA) was applied to the matrix containing all the relevés using the Canoco 5.5 software (Ter Braak & Šmilauer, 2012).
III. CHARACTERIZATION OF THE REGION
This work was carried out in the coastal saltmarshes and brackish areas of the Namibe
Province. This territory comprises the coastal strip of the Namib Desert and is crossed by
the Bero and the Pindo rivers with water almost all year round. The Bero estuary is next
to the city of Namibe (formerly known as Moçâmedes) and the Pindo estuary near Tombua (formerly known as Porto Alexandre). Occurring in a desert area (mostly a sandy
desert), the Angolan saltmarshes in the studied region belong to the silt-loam textural
class or even the sandy-loam class (Cardoso et al., 2019).
This area is characterized by a tropical hyperdesert, upper thermotropical, lower
hyperarid, attenuated hyper-oceanic bioclimate (fig. 1) (Cardoso, 2014; Rivas-Martínez
et al., 2011).
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Cardoso, J. F., Costa, J. C., Neto, C. S., Duarte, M. C., Monteiro-Henriques, T. Finisterra, LVI(116), 2021, pp. 99-114
Fig. 1 – Thermopluviometric diagram for Namibe city (Namibe Weather Station – ID 857 331,
Lat. 15 159; Long. 12 178; Alt. 11m; Hyperdesertic tropical, superior thermotropical,
inferior hyperarid, attenuated hyperoceanic), 2014-2020.
Fig. 1 – Diagrama termopluviométrico da cidade do Namibe (Estação Meteorológica do Namibe – ID
857331, Lat. 15 159; Long. 12 178; Alt. 11m; Tropical hiperdesértico, termotropical superior, hiperárido
inferior, hiperoceânico atenuado), 2014-2020.
Source: Southern African Science Centre for Climate Change and Adaptative Land Management [SASSCAL] WeatherNet (2020)
IV. RESULTS AND DISCUSSION
1. New associations
The occurrence of 18 species, belonging to eight families (Appendix 1) and six new
associations were reported for the saltmarshes and brackish marshes along the coast of
the southwest of Angola, which we now describe:
1. Sarcocornietum affinis J.F. Cardoso, J.C. Costa, Neto, Maria C. Duarte & Monteiro-Henriques ass. nov.
(table I, holotypus relevé no. 7)
Halophilous chamaephytic association of Sarcocornia natalensis subsp. affinis, often
monospecific, although sometimes Sporobolus virginicus and Sesuvium portulacastrum,
may also occur in its floristic composition.
The bioclimate of the areas where it occurs varies from tropical xeric to tropical
hyperdesert. We place this community in the Arthrocnemetea franzii class (coastal saltwater marshes flooded by sea tides or occasionally interior saline waters: succulent shrub,
forbs and grass communities, growing in infra-thermotropical desertic and xeric bioclimate of tropical west coast of Africa), order Sarcocornio-Salicornietalia meyerianae (halophytic vegetation of small succulent plants); Sarcocornion decumbentis alliance (flooded
communities; Boucher & Jarman, 1977; Hanekom et al., 2009; Mucina et al., 2003; Rivas-Martínez et al., 2017).
Cardoso, J. F., Costa, J. C., Neto, C. S., Duarte, M. C., Monteiro-Henriques, T. Finisterra, LVI(116), 2021, pp. 99-114
105
Sarcocornia natalensis subsp. affinis was first reported for Angola by Cardoso (2014)
as part of the fieldwork conducted during this study. This species was listed in Plantas de
Angola (Figueiredo & Smith, 2008) as Arthrocnemum macrostachyum (currently Arthrocaulon macrostachyum). However, the genus Arthrocnemum has one undivided cavity in
the axis after the detachment of the flowers while the specimens we observed, in the
Namibe saltmarshes, have three cavities which correspond to the genus Sarcocornia
(S. natalensis subsp. affinis). It is a prostrate, rhizomatous chamaephyte, woody at the
base, succulent, with segments of 5-15mm long and 2-3mm in diameter, corresponding
to the description made by Le Roux and Wahl (2005).
Table I – Phytosociological table of the association Sarcocornietum affinis.
Quadro I – Quadro fitossociológico da associação Sarcocornietum affinis.
Area (m2)
1
1
1
1
4
2
2
2
2
4
2
4
4
1
2
4
No. of taxa
1
1
1
Relevé no.
6
7
8
1
2
2
2
2
2
3
2
4
3
2
2
2
9
10 11 12 13 14 15 16 17 18 19 20 21
Sarcocornia natalensis subsp. affinis
5
5
Sporobolus virginicus
.
.
5
4
3
4
4
3
5
4
3
2
3
5
5
4
V
.
.
.
.
.
.
.
2
1
2
1
+
1
3
Sesuvium portulacastrum
.
II
.
.
.
.
3
1
2
1
2
.
.
.
.
.
.
Cyperus laevigatus
II
.
.
.
.
.
.
.
.
.
.
.
.
3
.
.
.
Suaeda merxmuelleri
+
.
.
.
.
.
.
.
.
.
.
.
1
.
.
.
.
+
Chloris flabellata
.
.
.
.
1
.
.
.
.
.
.
.
.
.
.
.
+
Eragrostis prolifera
.
.
.
.
.
.
.
.
.
.
.
+
.
.
.
.
+
Mesembryanthemum dimorphum
.
.
.
.
.
.
.
.
.
.
.
+
.
.
.
.
+
Characteristics
Companions
Places: 6, 15, 16 Bero river (Namibe); 7, 11, 12, 13, 17 Pindo river (Tombua); 8, 14, 20, 21 Lobito; 9, 10 Cubai
(Sumbe); 18, 19 Salinas Sol (Namibe).
2. Suaedetum merxmuelleri J.F. Cardoso, J.C. Costa, Neto, Maria C. Duarte & Monteiro-Henriques ass. nov.
(table II, holotypus relevé no. 52)
This plant community, with a distribution in tropical desertic and hyperdesertic,
upper thermotropical, ultrahyperarid to hyperarid, attenuated hyperoceanic (semi-hyperoceanic) bioclimate, is characterized, almost exclusively, by Suaeda merxmuelleri, an endemic nanophanerophyte (bush) of the southwest of Angola, sometimes
accompanied by Sporobolus virginicus, Sesuvium portulacastrum and Juncus rigidus. It
is found in the highest areas of the saltmarsh, with a brackish water table, and it is only
occasionally flooded, sometimes by brackish waters after heavy rains. Being a desert
with tropical characteristics, the soil is subjected to strong salinization for most of the
year. Salinity oscillations can be high although low values can be recorded very sporadically during the scarce rainy periods. It is a frequent association in nitrified sites
(with anthropic disturbance) along roads and paths. Very common in the studied area,
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also occurring further in the north, in Lobito marshes. We have placed this community
in Arthrocnemetea franzii, Sarcocornio-Juncetalia kraussii (halophilous nanophanerophytic vegetation), Spergulario mediae-Puccinellion angustae (dense communities rich
in Amaranthaceae and halotolerant grasses, adapted to brackish water flooding)
(Boucher & Jarman, 1977; Hanekom et al., 2009; Mucina et al., 2003; Rivas-Martínez
et al., 2017).
Table II – Phytosociological table of the association Suaedetum merxmuelleri.
Quadro II – Quadro fitossociológico da associação Suaedetum merxmuelleri.
Area (m )
1
1
1
1
2
2
4
4
8
4
4
8
8
4
No. of taxa
1
1
1
1
1
2
2
2
5
2
2
3
3
2
Relevé no.
52 53 54 55 56 57 58 59 60 61 62 63 64 65
2
Characteristics
Suaeda merxmuelleri
5
5
5
4
4
3
4
3
3
5
5
4
3
4
V
Sporobolus virginicus
.
.
.
.
.
.
+
1
3
+
2
2
2
.
III
Sesuvium portulacastrum
.
.
.
.
.
.
.
.
.
.
.
.
.
2
+
Juncus rigidus
.
.
.
.
.
.
.
.
1
.
.
.
.
.
+
Cressa salina
.
.
.
.
.
.
.
.
.
.
.
+
.
.
+
Phragmites mauritianus
.
.
.
.
.
.
.
.
.
.
.
.
4
.
+
Mesembryanthemum pseudoschlichtianum
.
.
.
.
.
1
.
.
.
.
.
.
.
.
+
Chloris flabellata
.
.
.
.
.
.
.
.
.
.
.
1
.
.
+
Felicia mossamedensis
.
.
.
.
.
.
.
.
+
.
.
.
.
.
+
Mesembryanthemum dimorphum
.
.
.
.
.
.
.
.
+
.
.
.
.
.
+
Companions
Places: 52, 55, 65 Bero river (Namibe); 53, 63 Namibe; 54, 61 Lobito; 56, 57, 58, 59, 60, Pindo river (Tombua);
62, 64 Salinas Sol (Namibe).
3. Cypero laevigati-Cressetum salinae J.F. Cardoso, J.C. Costa, Neto, Maria C. Duarte
& Monteiro-Henriques ass. nov.
(table III, holotypus relevé no. 3)
This community is dominated by halophilous annual (therophytes) and stoloniferous
(hemicryptophytes) species, with fugacious hydromorphy (only when heavy rainfall
occurs), inserted in a tropical hyperdesert, upper thermotropical, lower ultrahyperarid,
attenuated hyperoceanic (semi-hyperoceanic) bioclimate. It is characterized by Cressa
salina, Sporobolus virginicus and Cyperus laevigatus. This community is infrequent,
having been observed only in moderately nitrified saline habitats with short flood periods,
near the city of Namibe. We consider it to be a geovicarious of the Cressetum salinae association, present in Cabo Verde, where it also occurs in temporarily flooded sandy soils
(Rivas-Martínez et al., 2017). This association has been positioned in the Arthrocnemetea
franzii class, Sarcocornio-Juncetalia kraussii order, Spergulario mediae-Puccinellion angustae alliance.
Cardoso, J. F., Costa, J. C., Neto, C. S., Duarte, M. C., Monteiro-Henriques, T. Finisterra, LVI(116), 2021, pp. 99-114
107
Table III – Phytosociological table of the association Cypero laevigati-Cressetum salinae.
Quadro III – Quadro fitossociológico da associação Cypero laevigati-Cressetum salinae.
Area (m2)
1
1
4
2
4
No. of taxa
1
1
2
2
2
Relevé no.
1
2
3
4
5
Cressa salina
4
5
5
4
3
Sporobolus virginicus
.
.
.
1
2
Cyperus laevigatus
.
.
+
.
.
Characteristics
Places: 1, 3 Namibe; 2, 4, 5 Salinas Sol (Namibe)
4. Junco rigidi-Sporoboletum virginici J.F. Cardoso, J.C. Costa, Neto, Maria C. Duarte
& Monteiro-Henriques ass. nov.
(table IV, holotypus relevé no. 48)
This community was observed in occasionally flooded depressions, in solonchak arenic
soils with very occasional hydromorphy (characterized by a strong concentration of soluble
salts due to strong evaporation, in desert and semi-desert climates). It is composed almost
exclusively by Sporobolus virginicus, sometimes accompanied by Juncus rigidus, and, in the
higher areas, by Suaeda merxmuelleri. In areas where the flooding period is longer, the
lower salinity allows the occurrence of reed (Phragmites mauritianus). Besides the study
area, we also observed this community further north, at Lobito. This community occurs in
tropical hyperdesert, upper thermotropical, ultrahyperarid to hyperarid, hyperoceanic attenuated (semi-hyperoceanic) bioclimate, although it can occur in tropical desert and xeric
bioclimate too. It has been placed in the Arthrocnemetea franzii class, Sarcocornio-Juncetalia
kraussii order and Spergulario mediae-Puccinellion angustae alliance.
Table IV – Phytosociological table of the association Junco rigidi-Sporoboletum virginici.
Quadro IV – Quadro fitossociológico da associação Junco rigidi-Sporoboletum virginici.
Area (m )
1 1 1 1 1 1 4 2 4 4 4 8 4
No. of taxa
1 1 1 1 1 2 2 2 2 2 3 3 2
Relevé no.
39 40 41 42 43 44 45 46 47 48 49 50 51
Characteristic
Sporobolus virginicus
5 5 5 5 4 4 2 3 4 4 4 4 4 V
Juncus rigidus
.
.
.
.
.
.
.
.
. + 1 .
. I
Suaeda merxmuelleri
.
.
.
.
.
.
.
.
.
. + 1 . I
Cressa salina
.
.
.
.
.
.
.
.
.
.
.
. 2 +
Companions
Phragmites mauritianus
.
.
.
.
.
.
.
. 3 .
. 2 . I
Brachiaria psammophila
.
.
.
.
.
. 3 .
.
.
.
.
. +
Stipagrostis prodigiosa
.
.
.
.
. + .
.
.
.
.
.
. +
Mesembryanthemum pseudoschlichtianum
.
.
.
.
.
.
. + .
.
.
.
. +
Places: 39 Namibe; 40, 50, 51 Salinas Sol (Namibe); 41, 43, 44, 45, 46, 47, 48, 49 Pindo (Tombua); 42 Lobito.
2
108
Cardoso, J. F., Costa, J. C., Neto, C. S., Duarte, M. C., Monteiro-Henriques, T. Finisterra, LVI(116), 2021, pp. 99-114
5. Sesuvietum crithmoidis J.F. Cardoso, J.C. Costa, Neto, Maria C. Duarte & Monteiro-Henriques ass. nov.
(table V, holotypus relevé no. 23)
Halonitrophilous, psammophilous and monospecific community of the perennial
Sesuvium crithmoides (= S. mesembryanthemoides, = S. crystallinum) sometimes accompanied by Zygophyllum simplex (table V). It occurs on intertidal sands, flooded by oceanic waters usually at maximum tidal level, where the sea deposits organic (responsible
for the nitrification) and inorganic debris. The relevés were collected in areas characterized by tropical hyperdesertic and desertic, thermotropical, ultrahyperarid to arid,
hyperoceanic attenuated (semi-hyperoceanic) bioclimate. According to our observations, this community occurs along the maritime dunes at least up to Luanda (tropical
xeric bioclimate).
The taxon Sesuvium crithmoides, previously considered endemic in Angola, was
recently (Sukhorukov et al., 2018) observed on the sandy coast of the Democratic Republic of Congo.
We have placed this association in the Arthrocnemetea franzii class, Sesuvietalia persoonii order (communities of succulent stolonate, hydro-halophytic and aero-halophytic
species from the western coasts of Africa, from Morocco to South Africa, in infra-thermotropical, ultrahyperarid to arid bioclimate) and Sesuvion persoonii alliance (see Costa,
2018; Rivas-Martínez et al., 2017).
Table V – Phytosociological table of the association Sesuvietum crithmoidis.
Quadro V – Quadro fitossociológico da associação Sesuvietum crithmoidis.
Area (m2)
10
10
10
10
10
10
10
No. of taxa
1
1
1
1
2
2
2
10
2
Relevé no.
22
23
24
25
26
27
28
29
4
4
2
3
3
3
2
2
V
Zygophyllum simplex
.
.
.
.
+
1
1
.
II
Brachiaria psammophila
.
.
.
.
.
.
.
1
I
Characteristic
Sesuvium crithmoides
Companions
Places: 22 Namibe foz do rio Bero; 23, 26, 27, 28 Praia Sacomar (Namibe); 24, 25, 29 Praia Amélia (Namibe)
6. Sarcocornio affinis-Sesuvietum portulacastri J.F. Cardoso, J.C. Costa, Neto, Maria
C. Duarte & Monteiro-Henriques ass. nov.
(table VI, holotypus relevé no. 37)
Halophilous community dominated by Sesuvium portulacastrum, a perennial, succulent, pioneer species, of pantropical distribution, accompanied by Sarcocornia natalensis
subsp. affinis, typical of saltmarshes and sandy depressions flooded and submerged by
brackish waters for more or less long periods. It occurs in tropical hyperdesertic and
desertic, thermotropical, ultrahyperarid to hyperarid, hyperoceanic attenuated (semi-
Cardoso, J. F., Costa, J. C., Neto, C. S., Duarte, M. C., Monteiro-Henriques, T. Finisterra, LVI(116), 2021, pp. 99-114
109
-hyperoceanic) bioclimate. According to our observations it occurs in Lobito marshes
and southward until the river Cunene estuary, almost always in monospecific formations.
It is a geovicarious of the Cape Verde association Sesuvietum portulacastri, also dominated by Sesuvium portulacastrum (Rivas-Martínez et al., 2017), however, in SW Angola
this species is accompanied by Sarcocornia natalensis subsp. affinis and Sporobolus virginicus, which do not occur in Cape Verde. We also include this community in the Sesuvietalia persoonii order and Sesuvion persoonii alliance.
Table VI – Phytosociological table of the association Sarcocornio affinis-Sesuvietum portulacastri.
Quadro VI – Quadro fitossociológico da associação Sarcocornio affinis-Sesuvietum portulacastri.
Area (m2)
1
2
1
1
1
1
1
1
No. of taxa
1
1
1
1
1
2
2
2
4
3
Relevé no.
30
31
32
33
34
35
36
37
38
Sesuvium portulacastrum
5
5
5
4
4
4
4
5
3
V
Sarcocornia natalensis subsp. affinis
.
.
.
.
2
2
3
1
2
III
Sporobolus virginicus
.
.
.
.
.
.
.
.
1
+
Characteristic
Places: 30, 31, 35, 35 rio Bero (Namibe); 32, 36, 37 rio Pindo (Tombua); 33, 38 Lobito.
2. Multivariate analysis (ordination) of the relevés
The relevés were collected in a table and the Braun-Blanquet scale values were converted into numerical values to be used in a Principal Component Analysis (PCA). The
purpose of this analysis was primarily to determine the dispersion patterns of the relevés
as a result of their floristic composition. The closest points correspond to similar relevés
in terms of floristic composition and the farthest points correspond to more dissimilar
relevés. In this way we can test the manual organisation of the relevés that has been carried out with the previously presented tables and the respective plant associations. The
observed structure, although being the result of the floristic composition of the relevés, is
also the result of the implicit effect of the environmental factors, which were discussed in
the description of each of the plant communities.
Figures 2 and 3, resulting from the PCA, show a good segregation of the plant communities described (fig. 2) and clearly characterized by the bioindicator species presented
(fig. 3). Suaeda merxmuelleri is the diagnostic species of the association Suaedetum merxmuelleri; Sporobolus virginicus is the diagnostic species of the association Junco rigidiSporoboletum virginici; Sarcocornia affinis is the diagnostic species of the association Sarcocornietum affinis; Sesuvium portulacastrum is the diagnostic species of the association
Sarcocornio affinis-Sesuvietum portulacastri; Cressa salina is the diagnostic species of the
association Cypero laevigati-Cressetum salinae; and Sesuvium crithmoides is the diagnostic species of the association Sesuvietum crithmoidis.
110
Cardoso, J. F., Costa, J. C., Neto, C. S., Duarte, M. C., Monteiro-Henriques, T. Finisterra, LVI(116), 2021, pp. 99-114
Fig. 2 – Principal Component Analysis (PCA) – plot showing the ordination of vegetation samples.
Fig. 2 – Análise de Componentes Principais (ACP) – plot com a ordenação de amostras de vegetação.
A. Cypero laevigati-Cressetum salinae (samples 1-5); B. Sarcocornietum affinis (samples 6-21); C. Sesuvietum crithmoidis
(samples 22-29); D. Sarcocornio affinis-Sesuvietum portulacastri (samples 30-38); E. Junco rigidi-Sporoboletum virginici
(samples 39-51); F. Suaedetum merxmuelleri (samples 52-65).
Fig. 3 – Principal Component Analysis – plot showing the ordination of species.
Fig. 3 – Análise de Componentes Principais – plot com a ordenação das espécies.
Chl (Chloris flabellata); Cr (Cressa salina); Cyp (Cyperus laevigatus); Era. (Eragrostis prolifera); Fel (Felicia mossamedensis);
Jun (Juncus rigidus); Bra (Brachiaria psammophila); Phr (Phragmites mauritianus); Med (Mesembryanthemum dimorphum);
Mesp (Mesembryanthemum pseudoschlichtianum); Sar (Sarcocornia natalensis subsp. affinis); Scr (Sesuvium crithmoides);
Spo (Sesuvium portulacastrum); Svir (Sporobolus virginicus); Sti (Stipagrostis prodigiosa); Sua (Suaeda merxmuelleri);
Zy (Zygophyllum simplex).
Cardoso, J. F., Costa, J. C., Neto, C. S., Duarte, M. C., Monteiro-Henriques, T. Finisterra, LVI(116), 2021, pp. 99-114
111
Syntaxonomical scheme (in bold, new syntaxa described in the context of this work):
Class: Arthrocnemetea franzii Rivas-Martínez, Lousã, J.C. Costa & Maria C. Duarte 2017
Order: Sesuvietalia persoonii Rivas-Martínez, Lousã, J.C. Costa & Maria C. Duarte 2017 corr.
Rivas-Martínez, Lousã, J.C. Costa & Maria C. Duarte in J.C. Costa 2018
Alliance: Sesuvion persoonii Lousã, J.C. Costa & Maria C. Duarte 2017 corr. Rivas-Martínez, Lousã,
J.C. Costa & Maria C. Duarte in J.C. Costa 2018
Sesuvietum crithmoidis J.F. Cardoso, J.C. Costa, Neto, Maria C. Duarte & Monteiro-Henriques (new)
Sarcocornio affinis-Sesuvietum portulacastri J.F. Cardoso, J.C. Costa, Neto, Maria C. Duarte &
Monteiro-Henriques (new)
Order: Sarcocornio-Salicornietalia meyerianae Boucher ined.
Alliance: Sarcocornion decumbentis Mucina, Jansen & O’ Callaghaen 2003
Sarcocornietum affinis J.F. Cardoso, J.C. Costa, Neto, Maria C. Duarte & Monteiro-Henriques (new)
Order: Sarcocornio–Juncetalia kraussii Boucher ined.
Alliance: Spergulario mediae-Puccinellion angustae Mucina, Jansen & O’Callaghan 2003
Suaedetum merxmuelleri J.F. Cardoso, J.C. Costa, Neto, Maria C. Duarte & Monteiro-Henriques (new)
Cypero laevigati-Cressetum salinae J.F. Cardoso, J.C. Costa, Neto, Maria C. Duarte & Monteiro-Henriques (new)
Junco rigidi-Sporoboletum virginici J.F. Cardoso, J.C. Costa, Neto, Maria C. Duarte & Monteiro-Henriques (new)
V. CONCLUSIONS
The coastal region of the southwest of Angola (south of Lobito), despite being under
a tropical macrobioclimate, presents no mangroves in its coastal swamps. Instead, saltmarshes are present, much probably owing to the influence of the cold oceanic waters of
the Benguela Cold Current. These marshes are floristically very poor and often monospecific. This is a substantially different situation from the South African and Holarctic marshes characterized by a much greater floristic richness. Angolan marshlands are characterized by a dominance of fine sandy soil texture which is associated to the presence of
the extensive field of dunes typical of the Namib Desert. The research carried out during
2010-2015 and 2018 allowed to analyse the floristic composition of the Namibe province’s
marshlands, as well as the originality of the floristic combinations associated to the more
or less differentiated environments.
In this way, a total of six new plant associations were described. Their spatial distribution is related to the ecological gradients associated to water table salinity, time of
submersion by sea waters, soil texture, as well as nitrogen levels. These communities,
clearly segregated by the PCA, do not present a seral logic and, therefore, as a whole,
constitute an original geopermasigmetum, presented for the first time in this work.
112
Cardoso, J. F., Costa, J. C., Neto, C. S., Duarte, M. C., Monteiro-Henriques, T. Finisterra, LVI(116), 2021, pp. 99-114
ACKNOWLEDGEMENTS
T. M. H. was funded by the European Social Fund (POCH) and by National Funds (MCTES),
through a FCT – Fundação para a Ciência e a Tecnologia postdoctoral fellowship (SFRH/
BPD/115057/2016), as well as by National Funds, through the same foundation, under the project
UIDB/04033/2020.
ORCID iD
João Francisco Cardoso https://orcid.org/0000-0003-1326-1368
José Carlos Costa https://orcid.org/0000-0002-7619-840X
Carlos da Silva Neto https://orcid.org/0000-0003-0912-0255
Maria Cristina Duarte https://orcid.org/0000-0002-3823-4369
Tiago Monteiro-Henriques https://orcid.org/0000-0002-4206-0699
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APPENDIX 1. LIST OF PRESENTED TAXA
Arthrocnemum macrostachyum (Moric.) K.Koch = Arthrocaulon macrostachyum (Moric.) Piirainen & G.Kadereit
(Amaranthaceae)
Brachiaria psammophila (Welw. ex Rendle) Launert (Poaceae)
Chloris flabellata (Hack.) Launert (Poaceae)
Cressa salina (J.A. Schmidt) Rivas Mart., Lousã, J.C.Costa & Maria C. Duarte (Convolvulaceae)
Cyperus laevigatus L. (Cyperaceae)
Eragrostis prolifera (Sw.) Steud. (Poaceae)
Felicia mossamedensis (Hiern) Mendonça (Asteraceae)
Juncus rigidus Desf. (Juncaceae)
Mesembryanthemum dimorphum Welw. ex Oliv. [=Psilocaulon dimorphum (Welw. ex Oliv.) N.E.Br.] (Aizoaceae)
Mesembryanthemum pseudoschlichtianum (S.M.Pierce & Gerbaulet) Klak (=Brownanthus pseudoschlichtianus
S.M.Pierce & Gerbaulet) (Aizoaceae)
Phragmites mauritianus Kunth (Poaceae)
Sarcocornia natalensis subsp. affinis (Moss) S.Steffen, Mucina & G.Kadereit
Sesuvium crithmoides Welw. (= S. mesembryanthemoides Wawra = S. crystallinum Welw. ex Oliv.) (Aizoaceae)
Sesuvium portulacastrum (L.) L. (Aizoaceae)
Sporobolus virginicus (L.) Kunth (Poaceae)
Stipagrostis prodigiosa (Welw.) De Winter (Poaceae)
Suaeda merxmuelleri Aellen (Amaranthaceae)
Zygophyllum simplex L. (Zygophyllaceae)