biodiversity

insufficiently recognised, resulting in ongoing threats

primarily from: overexploitation; water pollution; flow

Freshwaters comprise less than 1% of the Earth’s surface

modification; habitat destruction or degradation; and

and yet hold almost 10% of all described species, including

invasive species, all coupled with global environmental

approximately a third of all vertebrates (Strayer & Dudgeon,

change (Dudgeon et al., 2006). As a result of these often

2010). Given the low coverage of Earth by freshwaters,

interacting threats, 65% of global river discharge and the

this species richness is relatively high compared with

associated aquatic habitats are now classed as under

both terrestrial and marine ecosystems (Gleick, 1996),

moderate to high levels of threat (Vorosmarty et al., 2010) and

with the most comprehensive assessment of freshwater

only 37% of rivers that are over 1,000 km long remain free-

fauna to date describing 125,530 species (Balian et al.,

flowing along their length (Gril et al., 2019). At the population

2008). However, even this value is likely to be a large

level, freshwater species have on average declined by 83%

underestimate as Balian et al. (2008) highlight the severe

since 1970, declines that are far greater than those seen in

lack of knowledge of freshwater biodiversity in some

the terrestrial or marine ecosystems (WWF, 2018).

geographic areas (particularly the tropics, which are

generally areas of high diversity) and taxonomic groups

At the level of species, the most commonly used tools for

(particularly invertebrates).

assessing status are the IUCN Red List Categories and

Criteria (IUCN, 2012), which provide a quantitative and

This diverse and species-rich realm is of great importance

consistent approach by which to assess relative extinction

to people’s livelihoods, both directly as a source of food,

risk that can be applied across different taxonomic groups.

medicine and income, for example, and indirectly through

The IUCN Red List of Threatened SpeciesTM (hereafter

services such as nutrient cycling, flood control and water

IUCN Red List) publishes the results of these assessments

filtration (Juffe-Bignoli & Darwall, 2012). Freshwaters

online at www.iucnredlist.org, along with information on the provide
ecosystem goods and services with a global value

taxonomy, distribution, population, habitats and ecology, use

of tril ions of United States Dol ars (USD) per year, although

and trade, threats, and conservation and research actions

estimating the true value is difficult and the resulting

relevant to individual species. IUCN are partway through

estimates vary (e.g. global value per year: USD 70 billion

their global freshwater biodiversity assessment (Figure 1.1),

(Schuyt & Brander, 2004); over USD 4 trillion (Costanza et

the aim of which is to complete Red List assessments of

al., 2014); and up to USD 15 trillion (Millennium Ecosystem

all described species of freshwater decapod crustaceans,

Assessment, 2005)).

fishes, molluscs and odonates, and of selected freshwater

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

2 South African Institute for Aquatic Biodiversity (SAIAB), Somerset

Street, Grahamstown, 6139, South Africa 1

plants. Al described freshwater decapod crustaceans were

by taxonomists. Importantly, the estimated proportion of

comprehensively assessed by Cumberlidge et al. (2009),

undescribed species is exceptionally high in freshwaters

De Grave et al. (2015) and Richman et al. (2015). Since

(Lundberg et al., 2000).

these studies were completed, however, more species

within these taxonomic groups have been described,

meaning these assessments, although near complete, are

1.2 Situation analysis for the Lake

not currently comprehensive. Global assessments of all

Malawi/Nyasa/Niassa Catchment

odonates and freshwater fishes are aimed for completion in

2020 and 2021, respectively. These five taxonomic groups

1.2.1 Location

have been prioritised for assessment as they represent a

range of trophic levels and ecological roles within freshwater

The Lake Malawi/Nyasa/Niassa Catchment (hereafter

systems (Smith et al., 2014). Adding these priority taxa to

LMNNC) is situated in East Africa and includes parts of

existing assessments of freshwater-dependent mammals,

Malawi, Mozambique and Tanzania (Figure 1.2), with the land

birds, crocodiles, turtles and amphibians will provide

area split approximately 68%, 7% and 25%, respectively,

a relatively comprehensive overview of the state of the

among the three riparian countries (Bootsma & Jorgensen,

planet’s freshwater biodiversity. Of the 30,510 freshwater or

2005). Lake Malawi/Nyasa/Niassa (hereafter LMNN) (Figure

freshwater-dependent species currently assessed for the

1.3) is the southernmost of the African Great Lakes, and the

IUCN Red List, close to 27% are globally threatened with

catchment is bordered by the Lake Tanganyika Basin to the

extinction (IUCN, 2019).

north and drains into the Zambezi via the Shire River to the

south. The LMNNC as defined for this study also includes

One caveat with the current process is that undescribed

Lake Chilwa. This lake is situated in the headwaters of the

species, which are species that have not yet been formal y

Rovuma system and is closely associated with the LMNNC

taxonomically described and assigned a scientific name,

in that the rifting that created LMNN truncated numerous

are not considered. This can be a large fraction of the

headwaters of the Rovuma system. All the rivers flowing

species in a clade or geographical area, and especially so

into LMNN were, therefore, formerly Rovuma tributaries

in species rich groups that have not yet been well studied

(Crossley & Crowe, 1980).

Legend

Comprehensively assessed regions

Regions requiring assessment

Figure 1.1 Progress in the global freshwater biodiversity assessment. In

‘comprehensively assessed regions’ (red) the extinction risk of all native
freshwater decapods, fishes, molluscs and odonates, and selected
freshwater plants, has been assessed using the IUCN Red List
Categories and Criteria. In ‘regions requiring assessment’ (grey)
assessment of all or some of these taxonomic groups have not been
completed.

2
¯

0 20 40

80

120

160Kilometers

Songwe

North

Rukuru

Ruhuhu

South

Rukuru
Kasitu

Dwangwa

Bua

Msinje

Lilongwe

Linthipe

Shire

Sources: Esri, HERE, Garmin,

Intermap, increment P Corp.,

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

GEBCO, USGS, FAO, NPS,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

NRCAN, GeoBase, IGN,

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 1.2 Location of the Lake Malawi/Nyasa/Niassa Catchment in

East Africa, with a close up of the catchment.

3
1.2.2 Physical characteristics

The catchment has an area of 126,500 km2 and there are nine

major rivers flowing into LMNN. Over half of the river inflow

LMNN is the fourth largest freshwater lake in the world by

is from the Ruhuhu River (in Tanzania) and Songwe River

volume, containing 8,400 km3, equivalent to c. 7% of the

(on the border of Tanzania and Malawi) and therefore, land

Earth’s available freshwater. It is the ninth largest by surface

use in these regions may have a disproportionate effect on

area, at 28,800–30,800 km2. The lake has a maximum length

the lake. There is only one outflowing river, the Shire River

of 603 km, but is generally narrow, with a mean width of

in Malawi (Figure 1.4), which flows into the Zambezi River in

50–60 km. The maximum depth is approximately 785 m, with

Mozambique (Bootsma & Jorgensen, 2005; Weyl, Ribbink &

a mean depth of 290–426 m. The southern half of the lake is

Tweddle, 2010) (Figure 1.2).

general y shal ower and more productive than the north. The

lake is permanently stratified and waters are anoxic below

The other water body considered in this study is Lake Chilwa

depths of 170–200 m (Weyl, Ribbink & Tweddle, 2010).

(Figure 1.5), one of two shallow headwater lakes of the

Figure 1.3 Satellite image of Lake Malawi/Nyasa/Niassa. © Richard Petry

(CC BY-NC-ND 2.0) 4
Rovuma River system. Lake Chilwa is an endorheic, saline
from the freshwater Lake Chiuta by a sandbar dated at

lake with highly variable lake extent and water quality,

approximately 9,000 years old (Lancaster, 1979; Tweddle,

which when full covers almost 2,000 km2. It is separated

2005).

Figure 1.4 The Shire River in Malawi is the only outflowing river of
Lake Malawi/Nyasa/Niassa. © David Davies (CC BY-SA 2.0) Figure 1.5
Lake Chilwa is an endorheic, saline lake in the south of the Lake
Malawi/Nyasa/Niassa Catchment. © Gio la Gamb (CC

BY-SA 3.0)

5
1.2.3 Climate

to human settlements and agricultural land. Droughts,

increasingly frequent in the LMNNC, are caused primarily by

The LMNNC experiences a tropical-continental climate

El Nino and Southern Oscil ation (ENSO) events (Chairuca,

with two seasons: a dry season between September and

2016; Chavula, 2016; Faraji, 2016).

November, and a wet season between late November and

April. The lake level rises during the wet season, resulting in

It is widely acknowledged that climate change is becoming

fluctuations of between 0.8–1.4 m (Weyl, Ribbink & Tweddle,

an increasingly important threat to both human populations

2010).

and biodiversity (IPCC, 2014). Climate change effects have

been suggested for LMNN, as in other lakes of the Great

The hydrological regime of LMNN is dominated by

African Rift, (Hampton et al., 2018) and recent climate data

precipitation over the lake (41 km3 per year) and evaporation

indicate that there has been a temperature increase of 0.9°C

(54 km3 per year), with river inflow and outflow only at
within the LMNNC between 1960 and 2006 (McSweeney,

29 km3 per year and 12 km3 per year, respectively (Weyl,

New & Lizcano, 2008). These recent warming trends have

Ribbink & Tweddle, 2010). This dominance means that the

resulted in historical lake-level fluctuations and seasonal

lake is susceptible to changes in climate. An increase in the

extremes in limnological parameters that are measurable,

precipitation to evaporation ratio can result in flooding, for

although they do not reach the extent of those during mega-

example as documented in the late 1970s (Neuland, 1984),

drought times that occurred some 135 thousand years

whereas a decrease can result in the lake becoming closed

and 75 thousand years ago (Lyons et al., 2015; Scholz et

with no outflow at the Shire River, for example as recorded

al., 2007). Climate change is likely to have an even greater

between 1915–1935 and in 1997 (Chavula, 2016; Kidd, 1983).

impact on Lake Chilwa as it is a very shal ow lake.

Flooding, following heavy rainfall, is a recurring and

The adverse effects of climate change on biodiversity have

increasing phenomenon in the Malawian and Tanzanian

already been observed at multiple biological levels, ranging

parts of the catchment (Figure 1.6), where the floodplains

from genes to biomes (Scheffers et al., 2016). Effects on

have a low gradient, whereas the risk in the Mozambican

freshwater systems wil also have human health and socio-

part is low. The severity of the impact of flood events is

economic impacts with increased demand for water and

exacerbated by the increasing conversion of the floodplains

deteriorating water quality.

Figure 1.6 Flooding in southern Malawi in 2015. © GovernmentZA

DIRCO (CC BY-ND 2.0)

6
1.2.4 Biodiversity

Genner et al. (2013) reported the first introductions of

two invasive tilapiines (Nile tilapia, Oreochromis niloticus

1.2.4.1 Native species

and blue-spotted tilapia, O. leucostictus) to the LMNNC,

LMNN is considered to be the most species-rich lake on

which occurred during initiatives to develop aquaculture

Earth, containing over 800 cichlid fishes of which 99% are

and capture fisheries. Nile tilapia has already been widely

endemic (Snoeks, 2000). The family Cichlidae is dominant

implicated in global biodiversity loss and is a competitor

in the lake in terms of species richness, diversity, and the

and predator of native species, as well as having potential

abundance of individuals, and can generally be divided

to hybridise with native Oreochromis species. Introduction

into two groups: haplochromine and tilapiine cichlids.

of a pelagic sardine ( Limnothriossa spp.), to fil a perceived

Haplochromine cichlids are either smaller, primarily

vacant niche for exploiting the lakefly ( Chaoborus edulis),

rock-dwelling mbuna (e.g. Figure 1.7), or larger primarily

has also been proposed historical y (Turner, 1982). However,

sand-dwelling species. Many of these species are not

it has since been shown that lakefly production is efficiently

only endemic to LMNN but locally endemic to islands or

utilised by the existing lake fish community, and any attempt

sections of rocky shoreline within the lake, and around 600

to improve fishery production through introduction of a

of the haplochromine cichlids are known only from relatively

non-native plantivorous fish species would have a negative

shallow water along the coast (Konings, 2016). Tilapiine

impact on the stability and productivity of the lake ecosystem

cichlids in the genus Oreochromis form a small, endemic

(Darwal et al., 2010; Eccles, 1985). Three other alien species

species flock, commonly referred to as chambo (e.g. Figure

occur in the LMNNC: rainbow trout ( Oncorhynchus mykiss),

1.8), which are valuable food fishes. There are an additional

found on the Nyika Plateau; lungfish ( Protopterus annectens

12 families of fishes native to the LMNNC and two introduced

brieni) found mainly in the Mpatsanjoka Dambo near Salima

non-native families (Weyl, Ribbink & Tweddle, 2010). Lake

but also spreading to other swamp margins of the lake (Weyl,

Chilwa has a less diverse fauna with 30 fish species in 10

2004); and guppies ( Poecilia reticulata), recorded in the

families (Tweddle, 1979, 2005).

Lilongwe River catchment in Lilongwe. There are currently no

known alien fish species in the Lake Chilwa catchment, with

1.2.4.2 Non-native species

the exception of rainbow trout on the Zomba Plateau.

Invasive alien species are considered to be the second

greatest threat to biodiversity globally (Bellard, Cassey

Another invasive species, water hyacinth ( Eichhornia

& Blackburn, 2016). In freshwater systems, introductions

crassipes) (Figure 1.9), was introduced to the LMNNC in

can lead to biodiversity loss both directly through biotic

the 1960s via the Zambezi River in Mozambique (Faraji,

interactions, such as predation, and indirectly by decreasing

2016). This species is now widespread in the Shire River,

the availability of resources, facilitating the spread of

where it affects the generation of hydroelectric power

pathogens and parasites, or hybridising with native taxa

with cost estimates of USD 27,000 per day for shut downs

(Vitule, Freire & Simberloff, 2009). They can have devastating

resulting from weeds (Chavula, 2016). The abundance of

consequences on native biodiversity, for example as

the plant is currently relatively low within LMNN itself, most

resulted from the introduction of the Nile perch ( Lates

likely due to the low nutrient concentrations in the lake.

niloticus) to Lake Victoria (Sayer, Máiz-Tomé & Darwall,

However, if nutrient inputs in the lake increase, there is a

2018).

potential for water hyacinth to become more problematic

Figure 1.7 A male Chindongo flavus at Chinyankwazi Island

Figure 1.8 A male Oreochromis squamipinnis, locally known as

in Malawi. Chindongo flavus is assessed as Near Threatened

chambo, in Chinuni, Mozambique. Oreochromis squamipinnis

(NT). © Ad Konings

is assessed as Critically Endangered (CR). © Ad Konings

and have negative consequences for freshwater biodiversity,

Communities with high dependency on freshwater

particularly given the richness of haplochromine cichlid

ecosystems in the LMNNC can be classified into four

species found near to the shore (Bootsma & Jorgensen,

types: i) river communities, who live along rivers and use

2005).

their water for small-scale irrigation of vegetable gardens;

ii) lake shore communities, who live along the shore of

Introductions are typically irreversible and their conse-

LMNN and are primarily dependent on fisheries for their

quences on native species are often unclear until many

livelihoods; i i) floodplain communities, who are dependent

years later. Given the reliance of communities within the

on floodwater for irrigation of their rice crops; and iv) gold

LMNNC on the lake, future introductions should be avoided

panning communities, who make their livelihoods from

and efforts should be made to strengthen measures to block

gold panning in the upper reaches of the Ruhuhu River

pathways for introduction and to eradicate species that do

(Kafakoma, 2019). The conservation of the freshwater

invade (Genner et al., 2013).

ecosystems of LMNNC is vital to support the livelihoods of

these communities.

1.2.5 Socio-economic characteristics

1.2.5.1 Agriculture
In Africa there is a clear spatial congruence between

Agricultural production (including forestry, hunting and

centres of rural poverty and of threatened freshwater

fishing, in addition to cultivation of crops and livestock

species, particularly in the Great Lakes region of eastern

production) is the mainstay of the economy of the LMNNC,

Africa (Darwal et al., 2011). It is wel known that declines in

accounting for 26%, 21% and 30% of gross domestic

freshwater ecosystems, species diversity and their services

product (GDP) in Malawi, Mozambique and Tanzania,

will have a strong impact on local communities, especially

respectively, in 2017 (World Bank, 2019a).

the rural poor who are often heavily dependent on aquatic

resources (Brooks et al., 2016; Phil ips, Reantaso & Bueno,

In Malawi, the main crops for smallholders are maize,

2002). Therefore, there is a great risk to rural livelihoods

cassava and sweet potatoes, whereas large-scale farming

unless freshwater biodiversity conservation efforts improve

is of higher value crops, such as tobacco, tea, sugar, coffee

markedly.
and macadamia, that are then exported (Chavula, 2016).

Figure 1.9 Invasive water hyacinth ( Eichhornia crassipes) at Senga Bay

in Malawi. © Amy Palmer-Newton 8

In Mozambique, agriculture is predominantly small-scale

issues (Bootsma & Jorgensen, 2005; Chavula, Brezonik &

and the main crops are rice, soya (Figure 1.10), maize and

Bauer, 2011; Kafakoma, 2019).

beans (Chairuca, 2016). In Tanzania, small-scale agriculture

is primarily cultivation of cassava, rice and groundnuts, and

Soil erosion is the primary manner by which agricultural

plantation agriculture is primarily of tea, coffee and cocoa

practices affect the rivers and lakes of the LMNNC. There is a

(Faraji, 2016).

need to identify areas of the catchment where erosion is the

greatest, and then to implement strategies to reduce erosion

The northern two-thirds of the LMNNC are predominantly a

in those areas. The impacts of herbicides, fertilisers and

mixture of miombo ( Brachystegia) woodland and agricultural

pesticides is thought to be comparatively small, although

land, whereas the southern third is miombo in Mozambique

there may be localised areas of high impact, for example

and almost al cultivated land in Malawi (Chavula, Brezonik

in regions where cotton is grown, or near large sugar

& Bauer, 2011). The LMNNC has a high population density

plantations (Bootsma & Jorgensen, 2005). Where these

(100 per km2) (Chavula, Brezonik & Bauer, 2011), with the

agricultural chemicals enter freshwaters, they can lead

greatest density in the southern part of the catchment

to eutrophication and damage to or mortality of aquatic

in Malawi (Kafakoma, 2019). The population is growing,

organisms.

with the annual population growth rate of all three riparian

countries at approximately 3% (World Bank, 2019b). Around

Irrigation for agriculture (Figure 1.11) is the most significant

80% of people living within the LMNNC rely on agriculture

consumer of water in the LMNNC (Chavula, 2016; Faraji,

for subsistence, and this rapid growth is sustained by smal -

2016). This competes with other sectors with water demands,

scale agriculture. However, as this is low-income work, many

including domestic consumption, livestock production,

farmers are unable to purchase resources to improve their

hydropower generation, industrial production and mining,

agronomic practices and are unable to manage their lands

alongside the maintenance of healthy ecosystems. There

in an optimal manner. This is resulting in increasing pressure

has been increasing investment into irrigation schemes in the

on the land and increasingly marginal lands (such as steeply

LMNNC with the aim of boosting agricultural productivity, for

sloping lands, those with poor soils, wetlands etc.) are

example through the Greenbelt Initiative in Malawi, and the

starting to be cultivated, which wil exacerbate environmental

Lower Songwe Irrigation Development Project on the border

Figure 1.10 A soya bean famer in Malawi. © Mitchel Maher, International

Food Policy Institute (CC BY-NC-ND 2.0) 9

of Malawi and Tanzania (Kafakoma, 2019). Irrigated rice

schemes are also found on the lakeshore plain of the Lake

Chilwa catchment in Malawi.

1.2.5.2 Fisheries

The fisheries of LMNN are characterised by diversity in

the species harvested, fishing techniques and end uses.

The majority of fishes are harvested for food, either by

industrial or artisanal fisheries (Weyl, Ribbink & Tweddle,

2010) (Figure 1.12). The fisheries provide a key source of

protein for communities within the LMNNC, with suggestions

that freshwater fishes make up around 70% of animal

protein consumed by Malawians (Bland & Donda, 1995).

Figure 1.12 Fish for sale at the Chia Lagoon fish market.

The fisheries also make significant contributions to the

© USAID Biodiversity & Forestry (CC BY-NC 2.0)

economies of the three riparian countries, with data for

Malawi alone in 2015 indicating the sector directly employed

The fisheries are largely of a small-scale, non-mechanised

60,600 people, with over a further 300,000 people directly

nature, although a mechanised trawl fishery operates in the

engaged in secondary activities (Chavula, 2016), and

southern part of the lake in Malawi. The non-mechanised

contributed between 2–4% of the country’s GDP (Kafakoma,

fishery mainly employs paddle-powered dugout canoes

2019). The yield of the artisanal fishery, which makes up

fishing gill and seine nets, handlines and longlines, and

around 80% of landings, is primarily composed of utaka

basket and fence traps. Small-scale programs of catch

( Copadichromis spp.), usipa ( Engraulicyrpis sardella)

assessment were initiated for a short period in 1998 in

and chisawasawa ( Lethrinops spp.). Catfish ( Bagrus and

Tanzania and Mozambique, but the only reasonable time

Bathyclarias spp.) and chambo ( Oreochromis spp.) comprise

series of data is for the Malawian fisheries. Total yield from

less than 20% of the total catch (Bootsma & Jorgensen,

the ‘traditional fishery’ fluctuated around 30,000 tonnes

2005).

throughout the 1990s (Bulirani et al., 1999). An industrial pair-

Figure 1.11 A sprinkler irrigation system in a maize field in Malawi. ©

Melissa Cooperman, IFPRI (CC BY-NC-ND 2.0) 10
trawl fishery was established in 1968 in Malawi to harvest

demersal cichlid stocks in the south-east arm of the lake that

were not being exploited by the artisanal fishery. In the early

to mid-1970s, stern trawlers were introduced to the fishery

for bottom and mid-water trawling (Weyl, Ribbink & Tweddle,

2010). The small-scale fisheries exploit an estimated 110

species, with 25 species making up 80% of the total catch by

weight (Darwal , 2003). The demersal trawl fisheries capture

an estimated 250 species of which 15–20 species make up

70–80% of the catch (Palsson, Banda & Bulirani, 1999).

However, this important resource is declining with data

indicating decreasing catch rates, depleting biomass,

including of high-value species, and declining diversity,

particularly where fishing effort is highest in the south of

Figure 1.13 Aquaculture on Lake Malawi/Nyasa/Niassa. ©

LMNN and in Lake Malombe (Weyl, Ribbink & Tweddle, 2010).

Jamie Oliver via WorldFish (CC BY-NC-ND 2.0)

These declines are despite formal fisheries management

activities in the lake since the 1930s and therefore, there is a

ments. However, repeated trawl surveys in the southern part

need to improve fisheries management strategies (Bootsma

of the lake have noted the depletion of larger, slow-growing

& Jorgensen, 2005).

and late-maturing species (Weyl, Ribbink & Tweddle, 2010).

The total yield of LMNN has remained stable at around

Smallholder and limited commercial aquaculture occurs in

80,000 tonnes per year (Weyl, Ribbink & Tweddle, 2010).

LMNN (Figure 1.13) and both sectors are based on native

However, this yield has only been maintained through

tilapi ne cichlids, with use of non-native species prohibited

increasing fishing effort (Weyl, Ribbink & Tweddle, 2010) and

by legislation (Weyl, Ribbink & Tweddle, 2010). At present,

a reduction in net mesh sizes (Bootsma & Jorgensen, 2005).

total fish production from aquaculture in Malawi is estimated

Declines in catch per unit effort (CPUE) indicate the stocks

at 2,600 MT per year (Kafakoma, 2019).

are overfished (Weyl, Ribbink & Tweddle, 2010).

The export of live nearshore rocky cichlids (mbuna) for

The decline in the endemic chambo species flock, of

ornamental use (e.g. Figure 1.14) is another key source of

which individuals are valued over two times as much as

employment within the catchment. There are limited data

comparatively sized haplochromine cichlids, is an example

available to determine whether there are any negative

of depletion of high-value species (Tweddle et al., 2015;

impacts of harvesting for this trade on the abundance and

Weyl, Ribbink & Tweddle, 2010). During the 1950s over 3,000
diversity of these species. However, one known impact of

tonnes per year of chambo were harvested from LMNN’s

the trade is the introduction of mbuna to parts of the lake

south-east arm alone. However, the total catch for chambo

where they do not occur natively as a result of escaping or

in this part of the lake has shown a steady decline since the

early 1990s. CPUE in the main harvesting fisheries has also

declined dramatical y due to overfishing. In Lake Malombe,

chambo catches were around 4,000 tonnes in the late 1970s,

increasing to over 6,000 tonnes in the early 1980s. In the late

1980s a drastic decline was observed with catches fal ing to

less than 600 tonnes per year by the early 1990s and to less

than 200 tonnes per year in the late 1990s. This decline in

total catch in Lake Malombe is directly matched by severe

declines in CPUE in the two main fisheries harvesting the

stock, namely gill nets and chambo seines. The chambo

stocks in Lake Malombe are considered to have been in

a state of collapse or near collapse since the early 1990s

(Kanyerere, Phiri & Shechonge, 2018; Konings, 2018; Phiri &

Kanyerere, 2018).
Figure 1.14 A male Labidochromis chisumulae in an aquarium.

It is more difficult to assess how species diversity has

Labidochromis chisumulae is assessed as Least Concern (LC).

changed over time due to a lack of species level assess-

© Ad Konings

11

dumping of individuals. Several introduced species can be

of close to 4% per annum, which declined to close to 2%

seen at Thumbi West and Otter Point. Introduced mbuna

by 1994 because little forested land remained (Chavula,

compete with native mbuna for resources, including food

Brezonik & Bauer, 2011). Savanna/shrub/woodland habitat

and space (Ribbink et al., 1983).

in Malawi has also been heavily impacted by deforestation,

showing overall declines from over 43,000 km2 of this habitat

Lake Chilwa, when holding water, supports a thriving

in 1982 to close to 27,000 km2 in 2005, with a low of only

fishery, notably for the small cyprinid species Enteromius

around 5,300 km2 in 1995 (Chavula, Brezonik & Bauer, 2011).

paludinosus. Catches of this species can exceed 9,000 t

per year if the lake holds water for up to three years or more

1.2.5.4 Industry

(Furse, Morgan & Kalk, 1979).

Industry (including mining, manufacturing, construction,

electricity, water and gas) accounted for 14%, 25% and 26%

1.2.5.3 Forestry

of GDP in Malawi, Mozambique and Tanzania, respectively,

Forestry resources are pivotal in supporting local livelihoods

in 2017 (World Bank, 2019c). However, it should be noted

and the functioning of wider ecosystems. Communities

that much of this industrial production occurs outside of the

rely on forests as sources of foods, construction materials,

LMNNC.

medicines and, especially within the LMNNC, fuels (Figure

1.15). The governments of the three riparian countries

In Malawi, industry (including mining) is the second highest

have not been successful in enforcing forest management

source of employment after agriculture. The main industries

regulations and, as a result, the public has open access to

include the processing of tea (Figure 1.16), tobacco, sugar,

the majority of forest resources. The reliance on forests and

coffee, cement and cotton. Coal, uranium and bauxite are

their relatively easy accessibility has resulted in widespread

also mined within the country but provide employment to

deforestation in the LMNNC, including within conservation

only a smal proportion of the population. There are four main

areas (Bootsma & Jorgensen, 2005).

industrial areas (Blantyre, Liwonde, Lilongwe and Mzuzu)

and the quality of water in rivers flowing through these cities

In Malawi, rapid expansion of agricultural land in the 1970s

is negatively affected by industrial discharges (Chavula,

and 1980s resulted in extensive deforestation, with rates

2016).

Figure 1.15 A cyclist carrying firewood on the road to Lilongwe in

Malawi. © Mitchel Maher International Food Policy Institute (CC BY-NC-
ND 2.0)

12

Both gold and coal are found in the Mozambican part of

Malawi’s Southern Region (Kafakoma, 2019). In 2016, 11%

the LMNNC, but neither are currently mined at an industrial

of the total population of Malawi had access to electricity

scale. However, artisanal mining of gold is a source of

(World Bank, 2019d), but only 4% in rural areas (World Bank,

pollution and soil erosion into the lake (Chairuca, 2016).

2019e) where over 80% of the population lives (World Bank,

2019f).

There is also relatively little industrial activity in the Tanzanian

part of the LMNNC, with the majority of the population

In contrast to Malawi, there is no current or potential for

instead involved in agricultural activities. However, mining

hydropower generation in the Mozambican part of the

for coal and iron ore, along with related industries, is planned

LMNNC. The majority of the population (65% in 2017) in

within the catchment (Faraji, 2016).

Mozambique, including within the LMNNC, lives in rural

areas (World Bank, 2019f) and in 2016 only 5% of the rural

1.2.5.5 Energy

population had access to electricity (World Bank, 2019e).

Over 90% of Malawi’s energy requirements are provided

for by fuelwood, which includes both charcoal and

In Tanzania, energy is primarily sourced from electricity,

firewood. This high dependence has resulted in large-

fuelwood, charcoal and petroleum products. At present,

scale deforestation and, as forested areas are decreasing,

there are no hydropower plants in the Tanzanian part of the

there has been increasing incidence of flash floods and

LMNNC that are connected to the national grid, although

levels of sedimentation, which has impacts on hydropower

a number of isolated, off-grid hydropower plants exist.

generation. Around 3% of Malawi’s energy requirements

However, there are plans to develop hydropower within the

are met by electricity, which is primarily generated through

catchment, for example the proposed Songwe Hydroelectric

hydropower at the Shire River outflow of LMNN. Hydropower

Power Station on the border between Tanzania and Malawi

generation is negatively affected by low water levels in

(Faraji, 2016). In 2017, 67% of the Tanzanian population

LMNN and there are frequent power black-outs (Chavula,

lived in rural areas (World Bank, 2019f) and 17% of this rural

2016). To mitigate this situation, the governments of Malawi

population had access to electricity (World Bank, 2019e),

and Mozambique have signed a power sharing agreement

which is higher than for the other riparian countries. In rural

to enable interconnection of electrical power from Matombo

areas, electricity is primarily used for lighting, instead of

Power Station in Mozambique’s Tete Province to the

heating and cooking (Faraji, 2016).

Figure 1.16 Mulanje tea estate in Malawi. © David Davies (CC BY-SA
2.0)

13

1.2.5.6 Tourism
Instead, each of the riparian countries has its own policies

Travel and tourism contributed 4% to the GDP of Malawi,

and regulations regarding management, although they are

Mozambique and Tanzania in 2018. Both Malawi and

bound to international treaties, which provide established

Mozambique have experienced a positive year-on-year

ways of managing threats to biodiversity (Kafakoma, 2019;

average growth rate since 1995 of 6% and 5%, respectively,

Miriti, 2019). Fisheries are the primary focus of research

whereas Tanzania has a slight negative growth rate (World

and management activities in LMNN, with the majority of

Bank, 2019g).

work on CPUE data to set fishing restrictions. This work is

carried out by the Department of Fisheries, the Institute for

When compared with Mozambique and Tanzania, the tourism

Fisheries Research (Instituto de Investigação Pesquiera;

industry within the Malawian part of LMNNC is relatively well

IIP), and the Tanzania Fisheries Research Institute (TAFIRI)

developed, with many tourists visiting the lake and taking

in Malawi, Mozambique and Tanzania, respectively. In

part in water activities, including snorkel ing and diving to see

Malawi, research and management activities are also

the colourful rock-dwelling cichlid fishes (Figure 1.17). Many

conducted by Malawi Wildlife Services and the Malawi

hotels and resorts can be found along the shores of the lake

Department of National Parks and Wildlife. Within the wider

in Malawi, including within the Lake Malawi National Park

LMNNC, management of natural resources is done on the

(Chavula, 2016). There are many opportunities for tourism in

basis of sector (Bootsma & Jorgensen, 2005). The lack of

Mozambique and Tanzania as wel , but the industry within the

integration among sectors, meaning in some cases there

LMNNC of both countries is relatively undeveloped at present

are incompatible sectoral policies, has resulted in conflicts

and would benefit from improved transportation networks

in resource use, particularly water use, in the LMNNC

(Chairuca, 2016; Faraji, 2016).

(Kafakoma, 2019).

1.2.6 Environmental policies

The majority of development and conservation funding for

the LMNNC is provided by national governments or external

In contrast to other African Great Lakes, there is no regional

development partners, and is limited to project terms.

institution to coordinate policies and regulations for

Efforts are, therefore, restricted by this funding, which is

management of resources within LMNN and its basin.

limited in both quantity and time. Establishment of a regional

Figure 1.17 Snorkelling off Cape Maclear in Malawi. © Sandra Mal inson
(CC BY-NC-ND 2.0) 14

institution to coordinate a management plan for the three

1. Species Red List assessments ( Chapters 3 –7,

riparian countries is a priority (Kafakoma, 2019; Miriti, 2019).

summarised in Chapter 8) – The previously published

This would allow an ecosystem approach to management

IUCN Red List assessments of the freshwater species

( Bootsma & Jorgensen, 2005 ) and help to improve

native to the LMNNC were updated (with the exception of

management of the resources of the basin by coordinating

the odonates) and newly described freshwater species of

policies and regulations, increasing institutional capacity,

the region were assessed.

sharing information and best practices, influencing

2. Red List Index (RLI) (Chapter 9) – Regional RLIs for

stakeholders, finding sustained funding and promoting

each comprehensively assessed freshwater taxonomic

the importance of the lake and associated resources

group within the LMNNC were calculated to investigate

(Kafakoma, 2019; Miriti, 2019). A draft convention for

trends in the status of biodiversity.

management of the LMNNC was written in 2003 by the

3. Key Biodiversity Areas (KBAs) (Chapter 10) – KBAs

governments of Malawi, Mozambique and Tanzania, with

for freshwater biodiversity were delineated within

assistance of FAO (WWF, 2003). However, this convention

the LMNNC, using data from the updated Red List

has not been finalised and negotiations are still underway

assessments described in this report.

(Chidammodzi & Muhandiki, 2015).

4. Dissemination (Chapter 11) – Results, including this

report and the summary policy brief, wil be disseminated.

The importance of the biodiversity of LMNN is recognised

Red List assessments are already available through

but management actions to address its conservation have

the IUCN Red List website (www.iucnredlist.org) and

been limited so far (Bootsma & Jorgensen, 2005). Actions

species point records collated through the Red List

undertaken to address freshwater biodiversity conservation

assessment process are published on the Global

have included establishment of the Lake Malawi National

Biodiversity Information Facility (GBIF; www.gbif.org).

Park in Malawi in 1980, fol owed by its inscription as a World

KBA data are available through the World Database of

Heritage Site in 1984 (World Heritage Committee, 1984),

Key Biodiversity Areas (www.keybiodiversityareas.org).

and establishment of the Lake Niassa and its Coastal Zone

Red List assessments and KBA data will also be made Ramsar site in
Mozambique in 2011 (Ramsar Secretariat,

available through the Integrated Biodiversity Assessment

2019).

Tool (IBAT; www.ibat-al iance.org).

1.3 Objectives of this study

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‘East African megadroughts between 135 and 75

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Malawi: fishes, fisheries, biodiversity, health and habitat’,

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Aquatic Ecosystem Health & Management, 13(3), pp.

of Sciences, 104(42), pp. 16416–16421. https://doi.

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Schuyt, K. and Brander, L. (2004). Living Waters Conserving

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Smith, K.G., Barrios, V., Darwall, W.R.T. and Numa, C. (2014).

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The status and distribution of freshwater biodiversity in

[Accessed April 2019].

the Eastern Mediterranean. Cambridge, UK, Malaga,

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added (% of GDP). Available at: https://data.worldbank.

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https://doi.org/10.2305/IUCN.CH.2014.01.en

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Snoeks, J. (2000). ‘How well known is the ichthyodiversity

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2504(00)31005-4

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one of the great rivers of Africa’, Fisheries Management

18
Potamonautes lirrangensis. © Oliver-Mengedoht.de/Panzerwelten.de

Nothobranchius kirki. © B.R. Watters

Lanistes ovum. © Katharina C.M. Heiler

Drosera dielsiana. © David Midgley (CC BY-NC-ND 2.0)

19
Chapter 2
Species assessment methodology

Sayer, C.A.1 and Darwall, W.R.T.1

Contents

2.1 Selection of priority freshwater taxa

...........................................................................................................................
......................................20

2.1.1 Decapod crustaceans

...........................................................................................................................
...................................................21

2.1.2
Fishes.................................................................................................................
......................................................................................21

2.1.3 Mol uscs

...........................................................................................................................
........................................................................22

2.1.4 Odonates
...........................................................................................................................
......................................................................23

2.1.5 Plants
...........................................................................................................................
............................................................................24

2.2 Nomenclature
...........................................................................................................................
........................................................................25
2.3 Species mapping
...........................................................................................................................
..................................................................25

2.4 Assessment of species extinction risk

...........................................................................................................................
..................................25

2.5 Data col ection and quality control

...........................................................................................................................
........................................27

Bibliography
...........................................................................................................................
.................................................................................29

2.1 Selection of priority freshwater

information to conserve other components of the food web

taxa

that are essential to the maintenance of healthy functioning

wetland ecosystems, even if they are neither publically

In the majorit y of case s, large -scale biodiver sit y

charismatic nor often noticed, as is generally the case

assessments have focused on a limited range of taxonomic

for submerged species. As it is not practical to assess

groups, most often including those groups that provide

all species, a number of taxonomic groups have been

obvious benefits to humans through direct consumption, or

prioritised for comprehensive assessment at the global

the more charismatic groups, such as mammals and birds.

scale (i.e. assessment of all described species within the

In the case of aquatic systems, wetland birds, amphibians

taxonomic group on the global IUCN Red List of Threatened

and fishes have received most attention. However, it is

SpeciesTM, www.iucnredlist.org) as part of IUCN’s global important that

we take a more holistic approach by col ating

freshwater biodiversity assessment.

Figure 2 .1 The freshwater

biodiversity of Lake Malawi/

Nyasa/Niassaisvital

tothelivelihoodsofthe

l oc a l c o m m u n i t i e s of t h e

catchment,forexample

through provision of fishes for

food. © Benoit Rivard (CC BY-

NC-ND 2.0)

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

20
The taxonomic groups selected represent a range of trophic

Freshwater shrimps are also important as a human food

levels within the food webs that underlie and support

source (Holthuis, 1980), as wel as increasing in significance

wetland ecosystems, and are also groups for which there

in the aquarium trade. Relatively few freshwater groups

is thought to be a reasonable level of existing information.

have such a wide diversity of ecological traits, and occupy

These priority taxonomic groups are: freshwater decapod

such a wide range of freshwater habitats and environmental

crustaceans (crabs, crayfishes and shrimps), freshwater

conditions as do the shrimps. As such they also provide

fishes, freshwater molluscs, odonates (dragonflies and

a potentially useful indicator for the status of freshwater

damselflies), and freshwater plants1. Freshwater fishes, as

ecosystems (De Grave et al., 2015).

well as some freshwater decapods and molluscs, provide

clear benefits to the livelihoods of many people global y, either

There are 1,592 species of freshwater crabs, 751 species of

as a source of income or as a valuable food supply (Figure

freshwater shrimps and 657 species of freshwater crayfishes

2.1). Benefits provided by the other taxa may be indirect and

globally (N. Cumberlidge pers. comm. 2019). Cumberlidge

therefore poorly appreciated, but are nonetheless important.

et al. (2009), De Grave et al. (2015) and Richman et al. (2015)

Given the wide range of trophic levels and ecological roles

completed assessments of all freshwater crabs, shrimps

encompassed within these five taxonomic groups, it is

and crayfishes, respectively, for the global IUCN Red List.

proposed that information on their distributions and extinction

However, since these studies were completed more species

risk, when combined, will provide a useful indication for the

of crabs and crayfishes have been described. Therefore,

overal status of the associated ecosystems.

while al species of freshwater shrimps have been assessed

for the IUCN Red List, now only 82% (1,299 species) and

The work presented in this report follows on from earlier

87% (572 species) of freshwater crabs and crayfishes,

studies of the same freshwater taxonomic groups across

respectively, have been assessed (IUCN, 2019).

eastern Africa (Darwall et al., 2005) and mainland Africa

(Darwal et al., 2011). The same taxonomic groups have also

Five of the six freshwater crab species native to the LMNNC

been assessed for other parts of the world, beyond mainland

(e.g. Figure 2.2) were previously assessed for the IUCN Red

Africa (see www.iucn.org /theme /species /our-work /

List by Darwal et al. (2011). The sixth species was described

freshwater-biodiversity/freshwater-publications for other since the work by

Darwall et al. (2011) and was assessed

published regional freshwater biodiversity assessments).

through this study for the first time. The three freshwater

As such, the assessments presented here through this

shrimp species native to the LMNNC were previously

regional y-focussed study also contribute to building a global

assessed by De Grave et al. (2015). No freshwater crayfishes

coverage of these taxonomic groups, and to keeping their

are native to the LMNNC. The Red List assessment results

Red List assessments up to date.

presented here primarily reflect reassessments, building on

and updating the previous assessments.

2.1.1 Decapod crustaceans

2.1.2 Fishes

Freshwater decapod crustaceans include crabs, crayfishes

and shrimps. Of these three groups, only the freshwater

Fishes are arguably the most important products (in terms

crabs and shrimps have species native to the LMNNC.

of human use) of freshwater ecosystems at a global scale. In

2016 the total capture of fishes from inland waters global y

Freshwater crabs are one of the most ecological y important

was 11.6 mil ion tonnes and this represents a 11% increase

freshwater macro-invertebrate groups globally. They play

in comparison to the 2005–2014 average (FAO, 2018). Within

a key role in nutrient cycling due to the high importance

Africa, which accounts for 25% of global inland catches

of detritus in the diet of many species, coupled with their

(FAO, 2018), fishes provide an important food source for

abundance and high biomass (Cumberlidge et al., 2009).

over 400 million people and contribute essential proteins,

As freshwater crabs are found in a wide variety of aquatic

fats, minerals and vitamins to their diets (WorldFish Center,

habitats, and as they are normal y associated with relatively

2005). As well as essential nutrition, this capture provides

good quality water, they are excellent indicators of water

income for and supports the livelihoods of the poorest of

quality (Yeo et al., 2008). Additionally, they are a key

communities, through both consumption and non-food uses

component of tropical aquatic food webs, acting as prey

(Dugan et al., 2010).

items for a large number of predators, as wel as being widely

consumed by humans (Cumberlidge et al., 2009).

For the purposes of this assessment, freshwater fishes are

defined as those species that spend all or a critical part

of their life cycle in freshwaters. There are approximately

1 It should be noted that freshwater plants do not strictly represent a

17,800 freshwater fish species globally (R. van der Laan

taxonomic group. However, this terminology wil be used throughout

the report when discussing the high-level species groups assessed.

pers. comm. 2019) and at present, global extinction risk has

21
Figure 2.2 Potamonautes lirrangensis, commonly known as the Malawi
blue crab, is native to the Lake Malawi/Nyasa/Niassa Catchment. It is
assessed as Least Concern (LC). © Oliver-Mengedoht.de/Panzerwelten.de

been assessed for approximately 51% (9,138 species) of

assessments completed through this study were limited to

freshwater fishes using the IUCN Red List Categories and

taxonomically described species. There are currently 459

Criteria (IUCN, 2019). A global freshwater fish assessment is

taxonomically described freshwater fish species native

currently under way with the aim of assessing al species for

to the LMNNC. Darwall et al. (2011) previously assessed

the Red List by 2021.

all species (that were taxonomically described at the

time of assessment) native to the LMNNC and a number

Although LMNN is estimated as hosting up to 1,000

of those species widespread in eastern Africa were also

freshwater fish species (e.g. Figure 2.3), the Red List

recently assessed by Sayer et al. (2018). However, there

have been a significant number of taxonomic descriptions

in recent years resulting in 123 freshwater fish species

being assessed for the first time through this study. One

species, Anguil a bengalensis, was not reasssessed through

this study because assessments of anguillid eels are

completed by the IUCN Species Survival Commission (SSC)

Anguillid Eel Specialist Group. The remaining 335 species

were reassessed, building on and updating the previous

assessments.

2.1.3 Molluscs

Freshwater molluscs were found to be the group most at

risk of extinction and most poorly known in the continental

African assessment by Darwall et al. (2011), with 29% of

Figure 2.3 Aulonocara gertrudae is native to the Lake Malawi/

species assessed as threatened and 30% assessed as Data

Nyasa/Niassa Catchment. It is assessed as Least Concern

(LC). © Ad Konings

Deficient (DD). Freshwater mol uscs are mostly unobtrusive

22
2.1.4 Odonates

Larvae of almost al species of dragonflies and damselflies

(order Odonata) are dependent on freshwater habitats. The

habitat selection of adult odonates strongly depends on the

terrestrial vegetation type, and their larvae develop in water

where they play a critical role with regards to water quality,

nutrient cycling and aquatic habitat structure. The larvae

are voracious predators, often regarded as important in the

control of insect pest species. A wide array of ecological

niches is represented within the group and, as they are

susceptible to changes in water flow, turbidity or loss of

aquatic vegetation (Trueman & Rowe, 2009), they have been

widely used as an indicator of wetland quality.

Figure 2.4 Melanoides tuberculata is native to the Lake Malawi/

Nyasa/Niassa Catchment. It is assessed as Least Concern

There are currently 6,310 extant described species of

(LC). © Dennis L (CC BY 2.0)

odonate (Schorr & Paulson, 2019) but, even though the group

is well studied and relatively easily surveyed, it is believed

and are not normally considered to be charismatic, rarely

that the actual number is closer to 7,000 species (Kalkman

attracting the attention of the popular media, unless

et al., 2007). At present, the global risk of extinction has

in a negative light as some species are vectors in the

been assessed for approximately 61% (3,869 species) of

transmission of human and livestock parasites and

described odonates using the IUCN Red List Categories

diseases. This is unfortunate as freshwater molluscs play

and Criteria (IUCN, 2019). A global odonate assessment is

a vital role in the provision of ecosystem services and are

currently under way with the aim of assessing al species for

essential to the maintenance of wetlands, primarily due

the Red List by 2020.

to their contribution to water quality and nutrient cycling

through filter-feeding, algal-grazing and as a food source

There are 155 odonate species native to the LMNNC (e.g.

to other animals (Howard & Cuffey, 2006; Vaughn, Gido &

Figure 2.5). These species were previously assessed for

Spooner, 2004; Vaughn, Nichols & Spooner, 2008).

the IUCN Red List by Darwall et al. (2011) and a number of

species widespread in eastern Africa were also recently

There are approximately 6,500 freshwater mol usc species

reassessed by Sayer et al. (2018). Additional y, the IUCN SSC

described worldwide (M. Seddon pers. comm. 2019). At

Dragonfly Specialist Group has reassessed species native to

present, the global risk of extinction has been assessed for

the region in recent years and therefore, no reassessments

approximately 57% (3,683 species) of described freshwater

of odonates were conducted through this study. The

mollusc species using the IUCN Red List Categories and

results presented here relate to these previously published

Criteria (IUCN, 2019).

assessments.
There are 38 freshwater mollusc species native to the

LMNNC (e.g. Figure 2.4) and all were previously assessed

for the IUCN Red List by Darwal et al. (2011) and a number

of those species widespread in eastern Africa were also

recently assessed by Sayer et al. ( 2018 ). Therefore,

the Red List assessment results presented here reflect

reassessments, building on and updating the previous

assessments. For one species, Biomphalaria pfeifferi,

records from the LMNNC (Alharbi et al., 2019) were only

published after the assessment work for this study was

completed. This species was assessed as Least Concern

( LC ) (see 2.4 Assessment of species extinction risk

for terminology) in 2011 on the basis of its widespread

distribution across Africa, which excluded the LMNNC at

the time of assessment (Van Damme, 2015). It is included

Figure 2.5 Crocothemis erythraea is native to the Lake Malawi/

as a LC species in this study on the basis that this range

Nyasa/Niassa Catchment. It is assessed as Least Concern

extension would not change its Red List category.

(LC). © Bernard Dupont (CC BY-SA 2.0)

23

2.1.5 Plants

related to the proportion of aquatic species, availability

of information, stability of taxonomy, representation of

Freshwater plants are the building blocks of wetland

ecological niches, and representation geographically.

ecosystems, providing food, oxygen and habitats for

Additionally, Darwall et al. (2011) assessed 353 species

many other species. They are also a hugely important

from other plant families. These 718 species were used as

natural resource providing direct benefits to human

a base dataset for continental Africa from which species

communities. Numerous freshwater plants are highly valued

native to the LMNNC were extracted for this study. The

for their nutritious, medicinal, cultural, structural or biological

species distribution maps produced by Darwall et al. (2011)

properties. Some species also provide important wetland

were compared with the LMNNC (as defined in Figure 1.2)

ecosystem services, such as water filtration and nutrient

and the species occurring (with native presence) within the

recycling.

boundary were chosen for assessment. Additional species

for assessment (e.g. all freshwater-dependent Cyperaceae)

Following Cook (1996), freshwater plants are defined here

were added based on the expertise of specialists. This

as “vascular plants whose photosynthetically active parts

produced a list of 247 species of freshwater plants in 57

are permanently or, at least, for several months of the year,

families native to LMNNC for assessment (e.g. Figure 2.6).

submerged in water or float on the surface of the water”.

Fol owing this definition, it is estimated that freshwater plants

It is recognised that a significant number of additional

represent between 1–2% of all plant species, equivalent

plant species found in the LMNNC could be classified as

to approximately 2,900–5,800 of the approximate 300,000

freshwater-dependent but have not been included in this

species of vascular plants (Vié, Hilton-Taylor & Stuart,

assessment. Efforts to include these additional species will

2009). However, if considering non-vascular plants, such as

be made in the future. As discussed above, this work is part

bryophytes, the number of freshwater-dependent plants is

of a global effort and the original intention was to only assess

higher by at least an order of magnitude (R. Lansdown pers.

families for which a globally distributed set of freshwater

comm. 2019).

species could be identified. This approach was taken as it is

currently not feasible to assess all families of plant, given the

The species list considered for assessment in this study

high number of plant species. This approach is comparable

represents a subset of the species assessed by Darwal et

to that taken for animals, where the IUCN global freshwater

al. (2011), who assessed all known freshwater plant species

assessment is focussing on assessing selected taxonomic

(365 species) in 21 families. These families were selected

groups (decapods, fishes, molluscs and odonates).

from those identified by Cook (2004) based on criteria

Assessment of all species within the selected families al ows

Figure 2.6 Nymphaea nouchali, commonly known as blue lotus, is native

to the Lake Malawi/Nyasa/Niassa Catchment. It is assessed as Least
Concern (LC). © Bernard Dupont (CC BY-SA 2.0)

24

for comparative analysis of the status of global y assessed

The standard IUCN Red List attributes were used to

plant families with a significant component of freshwater

indicate the presence and origin of species at different

species. However, identifying plant families with such a

localities within their distribution ranges (IUCN Red List

freshwater component when working at the global scale has

Technical Working Group, 2018). Where data were available,

proved chal enging to achieve due to the need for additional

point localities (the latitude and longitude for a species

resources to enable expert judgement as to which species

collection record) were used to identify sites containing

are freshwater-dependent.

known occurrences of the species (coded as Presence 1:

Extant). These point data were supplemented by expert

knowledge of presence in sub-basins where no specific

2.2 Nomenclature

collection records were available. The preliminary species

distribution maps were digitised and then further edited at

Taxonomic schemes are constantly changing as results

the Red List review workshop (see 2.5 Data collection and

from ongoing studies, in particular with the introduction of

quality control) where errors were deleted from the maps

molecular techniques, are made available. As in many cases

and dubious records were recoded as Presence Uncertain

it is difficult to find a universal y agreed taxonomic hierarchy,

(Presence 6). Inferred distributions (coded as Presence 3:

the taxonomy followed here is that adopted by the IUCN

Possibly Extant), where a species is expected to occur but

Red List which, where possible, employs existing published

has not yet been confirmed, were determined through a

world checklists. For this study, fish classification general y

combination of expert knowledge, coarse scale distribution

follows Eschmeyer’s Catalog of Fishes (Fricke, Eschmeyer

records and unpublished information. Distributions where

& van der Laan, 2019) and odonate classification general y

the species were Possibly Extinct (Presence 4), Extinct

fol ows the World Odonata List maintained at the University

(Presence 5) and Introduced (Origin 3) were also captured

of Puget Sound (Schorr & Paulson, 2019). For plants, where

where known.

appropriate, we fol ow the World Checklist of Selected Plant

Families hosted by the Royal Botanic Gardens, Kew (WCSP,

Detailed in-lake distribution maps were produced for any

2019), but other more specialist lists are also followed.

decapods, fishes and molluscs native to LMNN. These

There is currently no widely accepted single taxonomy

in-lake maps were in polygon format and were based on

for molluscs and decapods, and we, therefore, follow

point localities (where available) and expert knowledge, in

the standards recommended by the IUCN SSC Mollusc

combination with bathymetry data.

Specialist Group and the IUCN SSC Freshwater Crustacean

Specialist Group, respectively.

All mapping was done using ArcGIS software (Environmental

Systems Research Institute (ESRI), 2018).

2.3 Species mapping

HydroBASIN distribution maps, with point data overlays and/

or detailed in-lake polygon overlays for selected species, are

Species distributions were mapped to river and lake sub-

published online on the IUCN Red List website (www.

basins as delineated by level 8 HydroBASINS (as il ustrated

iucnredlist.org) and are freely available to download for non-in Figure 2.7
for the LMNNC), a global standardised hydro-

commercial use.

logical framework that delineates catchments at 12

resolutions and includes information on hydrological

connectivity (Lehner & Gril , 2013). Where spatial data were

2.4 Assessment of species extinction

of sufficiently high detail, species were mapped to smaller

risk

sub-basins (level 12 HydroBASINS). River basins were

selected as the spatial unit for mapping and analysing

The Red List Categories and Criteria are widely accepted

species distributions because it is generally accepted that

as the most objective and authoritative system available

this is the most appropriate management unit for inland

for assessing the risk of a species becoming extinct (Mace

waters (Col ares-Pereira & Cowx, 2004).

et al., 2008; Rodrigues et al., 2006). The IUCN Red List of

Threatened Species™ is the world’s most comprehensive

The majority of species had published distribution maps

information source on the global conservation status of

from previous IUCN Red List assessments, for example as

plant, animal and fungi species, and is widely used to help

conducted by Darwal et al. (2011). These distribution maps

inform conservation priority setting. The risk of extinction

were used as a starting point and updated based on current

was assessed according to the IUCN Red List Categories

knowledge. The global (including beyond the LMNNC) native

and Criteria: Version 3.1 (IUCN, 2012) for all species in the

distribution of each species was mapped.

priority taxonomic groups native to the LMNNC.

25
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

HydroBASINS Level 8

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 2.7 Level 8 HydroBASINS used to map freshwater species

distributions for the IUCN Red List.

26

The nine Red List Categories at the global level are shown
must be met for a species to qualify under each category

in Figure 2.8. A species is assessed as Extinct (EX) when

please refer to The IUCN Red List Categories and Criteria:

there is no reasonable doubt that the last individual has

Version 3.1 (IUCN, 2012).

died. A species is assessed as Extinct in the Wild (EW)

when it is known only to survive in cultivation, captivity or

Red List assessments are published online on the IUCN Red

as a naturalised population well outside its native range.

List website (www.iucnredlist.org).

A species assessed as Critically Endangered (CR) is

considered to be facing an extremely high risk of extinction

in the wild. A species assessed as Endangered ( EN )

2.5 Data collection and quality control

is considered to be facing a very high risk of extinction

in the wild. A species assessed as Vulnerable (VU) is

The assessments of species extinction risk required

considered to be facing a high risk of extinction in the wild.

sourcing and collating the best information on all known,

All species listed as Critically Endangered, Endangered or

described species within the priority taxonomic groups. As

Vulnerable are termed threatened. A species is assessed

the primary source for this information, the best regional

as Near Threatened (NT) when it is close to qualifying for

and international experts for these taxa were first identified

a threatened category, or if it the focus of a specific and

through consultation with the relevant IUCN SSC Specialist

targeted conservation programme, the cessation of which

Groups.

would result in the species soon qualifying as threatened.

A species is assessed as Least Concern (LC) if it does not

These experts first collated the relevant information within

qualify (and is not close to qualifying) as threatened or Near

the IUCN Species Information Service (SIS) database (sis.

Threatened. Least Concern species are general y common

iucnsis.org) and applied the IUCN Red List Categories and

and widespread. A species is assessed as Data Deficient

Criteria (IUCN, 2012) to assess the risk of extinction of each

(DD) if there is insufficient information to make a direct or

species. Species range distributions were also mapped.

indirect assessment of its risk of extinction. DD is therefore

not a category of threat and instead indicates that further

All information related to the Red List assessments of

information on the species is required. Species assessed

freshwater fishes and plants was then peer reviewed at a

as DD are priorities for additional research and should be

Red List review workshop held in Nkopola, Malawi in May

acknowledged as potential y threatened.

2018 (Figure 2.10). During this workshop each Red List

assessment and distribution map was evaluated by at least

To determine whether a species should be assigned to one

one independent expert to ensure that the information

of the three threatened categories, there are five criteria with

presented was both complete and correct, and that the Red

quantitative thresholds (Figure 2.9), reflecting biological

List Category and Criteria assigned to each species were

indicators of populations threatened with extinction. For a

supported by the information provided. Data for freshwater

detailed explanation of the categories and of the criteria that

crabs and freshwater mol uscs were reviewed remotely.

Figure 2.10 Participants of the Red List review workshop held

in Nkopola, Malawi in May 2018 from left to right: T. Phiri,

R. Lansdown, A. Konings, D. Tweddle, J. Snoeks, G. Kanyerere,

M. Mwanyambo, E. Gobo, C. Sayer, W. Darwall, A. Shechonge

Figure 2.8 Global IUCN Red List Categories.

and A. Palmer-Newton. © Wil iam Darwal

27

SUMMARY OF THE FIVE CRITERIA (A-E) USED TO EVALUATE IF A

TAXON BELONGS IN AN IUCN RED LIST

THREATENED CATEGORY (CRITICALLY ENDANGERED,

ENDANGERED OR VULNERABLE).1

A. Population size reduction. Population reduction (measured over the

longer of 10 years or 3 generations) based on any of A1 to A4

Critically Endangered

Endangered

Vulnerable

A1

≥ 90%

≥ 70%

≥ 50%

A2, A3 & A4

≥ 80%
≥ 50%

≥ 30%

A1 Population reduction observed, estimated, inferred, or suspected in

(a) direct observation [except A3]

the past where the causes of the reduction are clearly reversible AND

(b) an index of abundance

understood AND have ceased.

appropriate to the taxon

A2 Population reduction observed, estimated, inferred, or suspected in the

(c) a decline in area of occupancy

past where the causes of reduction may not have ceased OR may not be

(AOO), extent of occurrence

understood OR may not be reversible.

based on

(EOO) and/or habitat quality

any of the

A3 Population reduction projected, inferred or suspected to be met in the

following:

(d) actual or potential levels of

future (up to a maximum of 100 years) [(a) cannot be used for A3].
exploitation

A4 An observed, estimated, inferred, projected or suspected population

(e) effects of introduced taxa,

reduction where the time period must include both the past and the future

hybridization,

pathogens,

(up to a max. of 100 years in future), and where the causes of reduction may

pollutants, competitors or

not have ceased OR may not be understood OR may not be reversible.

parasites.

B. Geographic range in the form of either B1 (extent of occurrence)

AND/OR B2 (area of occupancy) Critically Endangered

Endangered

Vulnerable

B1. Extent of occurrence (EOO)

< 100 km²

< 5,000 km²

< 20,000 km²

B2. Area of occupancy (AOO)

< 10 km²

< 500 km²

< 2,000 km²

AND at least 2 of the following 3 conditions:

(a) Severely fragmented OR Number of locations

=1

≤5

≤ 10

(b) Continuing decline observed, estimated, inferred or projected in any of:

(i) extent of occurrence; (ii) area of occupancy; (iii) area, extent and/or
quality of habitat; (iv) number of locations or subpopulations; (v) number of
mature individuals (c) Extreme fluctuations in any of: (i) extent of
occurrence; (ii) area of occupancy; (iii) number of locations or
subpopulations; (iv) number of mature individuals

C. Small population size and decline

Critically Endangered

Endangered

Vulnerable

Number of mature individuals

< 250

< 2,500

< 10,000

AND at least one of C1 or C2

C1. An observed, estimated or projected continuing decline

25% in 3 years or
20% in 5 years or

10% in 10 years or

of at least (up to a max. of 100 years in future):

1 generation

2 generations

3 generations

(whichever is longer)

(whichever is longer)

(whichever is longer)

C2. An observed, estimated, projected or inferred continuing

decline AND at least 1 of the following 3 conditions:

(a) (i) Number of mature individuals in each subpopulation

≤ 50

≤ 250

≤ 1,000

(ii) % of mature individuals in one subpopulation =

90–100%

95–100%

100%

(b) Extreme fluctuations in the number of mature individuals

D. Very small or restricted population

Critically Endangered

Endangered

Vulnerable

D. Number of mature individuals

< 50

< 250

D1. < 1,000

D2. Only applies to the VU category

Restricted area of occupancy or number of locations with

D2. typically:

a plausible future threat that could drive the taxon to CR

AOO < 20 km² or

or EX in a very short time.

number of locations ≤ 5

E. Quantitative Analysis

Critically Endangered

Endangered
Vulnerable

≥ 50% in 10 years or 3

≥ 20% in 20 years or 5

Indicating the probability of extinction in the wild to be:

generations, whichever generations, whichever

is longer (100 years

is longer (100 years

≥ 10% in 100 years

max.)

max.)

1 Use of this summary sheet requires full understanding of the IUCN Red
List Categories and Criteria and Guidelines for Using the IUCN Red List
Categories and Criteria.

Please refer to both documents for explanations of terms and concepts used
here.

Figure 2.9 Summary of the five criteria (A–E) used to evaluate if a

species belongs in an IUCN Red List threatened category: Critically
Endangered (CR), Endangered (EN) or Vulnerable (VU).

28

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30

Chapter 3

The status and distribution of freshwater

decapods in the Lake Malawi/Nyasa/

Niassa Catchment

N. Cumberlidge1

Contents

3.1 Introduction
...........................................................................................................................
...........................................................................31

3.2 Red List assessments

...........................................................................................................................
...........................................................32

3.3 Patterns of species richness

...........................................................................................................................
................................................ 34

3.3.1 Overal species richness

...........................................................................................................................
.............................................. 34

3.3.2 Endemic species richness

...........................................................................................................................
.......................................... 34

3.3.3 Data Deficient (DD) species richness

...........................................................................................................................
...........................37
3.4 Major threats
...........................................................................................................................
.........................................................................37

3.4.1 Agricultural and urban expansion

...........................................................................................................................
.................................37

3.4.2 Invasive species

...........................................................................................................................
.......................................................... 39

3.5 Recommended research and conservation actions

...........................................................................................................................
............. 39

Species in the spotlight

...........................................................................................................................
................................................................42

The hidden mountain crabs

...........................................................................................................................
..................................................42

Bibliography
...........................................................................................................................
................................................................................ 44

3.1 Introduction

elsewhere in continental Africa but is absent in North Africa

north of the Sahara (except for the Nile Basin) and in

The freshwater decapod fauna (freshwater crabs and

Madagascar (Bott, 1955; Cumberlidge, 1999; Daniels, Phiri &

freshwater shrimps) of the Lake Malawi/Nyasa/Niassa

Bayliss, 2014). The LMNNC’s freshwater crab fauna is

Catchment (hereafter LMNNC) comprises nine species in

relatively impoverished in comparison with similar-sized and

two families. Recent exploration and new taxonomic studies

better-studied areas of continental Africa, but it is likely that

have shown that the LMNNC has six species of freshwater

the number of known species will rise as exploration

crabs (Potamonautidae) and three species of freshwater

continues and taxonomic skil s are refined. The northern part

shrimps (Atyidae). All six species of freshwater crabs

of the LMNNC (south-western Tanzania) lies in the East

( Potamonautes lirrangensis, P. montivagus, P. choloensis,

African region, which supports 47 species and four genera

P. suprasulcatus, P. obesus and P. bellarussus) have a

spread across five countries. Three additional freshwater

wide distribution that extends outside of the LMNNC

decapod species ( Macrobrachium idella, Macrobrachium

(Cumberlidge, 2011; Daniels, Phiri & Bayliss, 2014; Reed &

rude and Potamonautes loveridgei) in this region may occur

Cumberlidge, 2004, 2006), meaning none are endemic to

in the LMNNC, although at present there is insufficient

the basin. Two of the freshwater shrimp species ( Caridina

information on their distributions to confirm this. Fol owing a

malawensis and C. kaombeflutilis) are endemic to the

recent taxonomic revision of the freshwater crabs, there is

LMNNC (De Grave et al., 2015; Richard & Clark, 2009, 2010).

likely to be at least one new species which is found in south-

Overal , rates of endemism in decapods in the LMNNC are

western Tanzania, and the Malawi blue crab should

low (Table 3.1).

potential y be recognised as a unique species and removed

from its current species assignment as P. lirrangensis

All six species of freshwater crabs found in the LMNNC

(unpublished data). The majority of the LMNNC is, however,

belong to the genus Potamonautes MacLeay, 1837, in the

within the Southern African region, which supports 46

e xc l u s i v e l y A f r o t r o p i c a l f r e s h w a t e r c r a b f a m i l y

species of freshwater crabs in two genera spread across ten

Potamonautidae. This genus has a wide distribution

countries (unpublished data).

1 Department of Biology, Northern Michigan University, Marquette,
Michigan, USA

31

The three species of freshwater shrimps found in the

Cumberlidge et al. (2009), and again through this study.

LMNNC are all of the genus Caridina in the family Atyidae.

The freshwater shrimps were assessed for the first time

Species of atyids found in African freshwaters feed by

in 2013, with the results summarised by De Grave et al.

scavenging, filtering suspended matter from the water or

(2015), and again through this study. Although there is a

sweeping up microbial films (Fryer, 1977). The feeding habits

need to collect more comprehensive information on these

of freshwater shrimps make them important components of

freshwater species, the data available were sufficient to

food webs in tropical streams and rivers because they not

make valid assessments of the extinction risk of the majority

only control invertebrate populations but are important food

of the freshwater decapod fauna. However, two species,

for carnivorous fishes (Browder, Gleason & Swift, 1994; De

Caridina kaombeflutilis (known only from three specimens

Resende et al., 1996; Fredrick & Spalding, 1994). All of the

in eastern Malawi; De Grave & Cumberlidge, 2018a) and

species of atyid shrimps found in LMNNC complete their

C. malawensis (known only from a single specimen in the

entire life cycle in freshwaters (De Grave et al., 2015).

littoral zone of Lake Malawi/Nyasa/Niassa (hereafter LMNN)

in Malawi; De Grave & Cumberlidge, 2018b) are assessed

With six species, the LMNNC’s freshwater crabs are slightly

as Data Deficient (DD) (Figure 3.1, Table 3.1) based on there

more speciose than the region’s freshwater shrimps (Table

being insufficient information available on their distributions,

3.1) but this may be deceptive because new species of both

populations and threats.

crabs and shrimps are still being described (unpublished

data; De Grave et al., 2015).

All six species of freshwater crabs and one of the three

species of freshwater shrimps found in the LMNNC are

Least Concern (LC), equivalent to 78% of the species

3.2 Red List assessments

assessed ( Figure 3.1, Table 3.1). The LC shrimp, C.

togoensis (Figure 3.2), is found along the west coast of

The extinction risk of the LMNNC’s freshwater crab species

Africa, throughout the Nile Basin and south to Malawi, and

was assessed in 2008, with the results summarised by

from Chad south to Botswana (De Grave & Cumberlidge,

Table 3.1 Numbers of freshwater decapod species native and endemic to

the Lake Malawi/Nyasa/Niassa Catchment in each Red List Category.
For a list of species native to the Lake Malawi/Nyasa/Niassa Catchment
and their Red List Categories please see Appendix 1.

Number of species native to the LMNNC

Number of species endemic to the LMNNC

IUCN Red List Category

Crabs

Shrimps

All decapods

Crabs
Shrimps

All decapods

Extinct (EX)

Extinct in the Wild (EW)

Critically Endangered (CR)

0
0

Endangered (EN)

Vulnerable (VU)

Near Threatened (NT)

0
0

Least Concern (LC)

Data Deficient (DD)

Total

6
3

Endemic species

Native species

0%

20%

40%

60%

80%

100%

EX

EW

CR

EN

VU

NT

LC
DD

Figure 3.1 Percentage (%) of freshwater decapod species

native and endemic to the Lake Malawi / Nyasa / Niassa

Catchment in each Red List Category. For a list of species

Figure 3.2 Caridina togoensis is native to the Lake Malawi/

native to the Lake Malawi/Nyasa/Niassa Catchment and their

Nyasa/Niassa Catchment. It is assessed as Least Concern

Red List Categories please see Appendix 1.

(LC). © Martin Grimm (CC BY-NC-SA 2.0)

32
2018c), and it lacks any major threats throughout its

Potamonautes bellarussus (LC) is found in south-western

range.

Tanzania near LMNN and in north-western Mozambique in

large boulder-strewn mountain streams in forested areas

The six species of freshwater crabs live in rivers, streams

on the slopes of Mounts Mecula, and in Yao close to the

and marshy lowlands, and in LMNN itself (Cumberlidge et

Mulanje Massif in southern Malawi (Daniels, Phiri & Bayliss,

al., 2009; Daniels, Phiri & Bayliss, 2014; Reed & Cumberlidge,

2014). The distributional range of this species includes one

2006). Potamonautes obesus (LC) is found in the coastal

protected area (the Niassa National Reserve in Mozambique)

regions of Somalia, Kenya and Tanzania, and in inland

and the majority of the localities from which the species has

localities in Malawi, central Mozambique and north-eastern

been collected are at high altitude (Cumberlidge, 2018c).

Zambia. This is a semi-terrestrial species that is capable

Potamonautes montivagus (LC) is a widespread species

of breathing air, living in marshy, swampy areas in coastal

that is found in rivers and lakes, and its distributional range

and inland localities. However, where it inhabits temporary

includes both low and high altitude localities. This species

wetlands in seasonal y arid regions it is vulnerable to changes

occurs in northern Malawi and south-western Tanzania, and

in rainfal patterns and the expansion of destructive farming

in southern Malawi and nearby localities in eastern Zambia,

practices driven by human population growth (Cumberlidge,

eastern Zimbabwe and western Mozambique (Cumberlidge,

2018a; Reed & Cumberlidge, 2004, 2006). Potamonautes

2018d). Potamonautes suprasulcatus (LC) has a distribution

choloensis (LC) is found in Malawi, south-western Tanzania

centred mainly in Tanzania but the southern part of its range

and western Mozambique, and most of the collection

extends as far as north-east Zambia and the LMNNC in

localities are at high altitudes. For example, in northern

Malawi (Reed & Cumberlidge, 2006). This is a large species

Malawi it is found on the Nyika Plateau (1,829–2,134 m asl)

that lives in large rivers and streams at both low altitude

(Figure 3.3) and near Rumphi (1,981 m asl), both of which

and high altitude localities (Cumberlidge, 2018e). Finally,

lie partly in the Nyika National Park. In southern Malawi this

Potamonautes lirrangensis (LC) (Figure 3.4) is a widespread

species is found on the Zomba Plateau (1,800 m asl) and

species found in Central Africa in the Middle Congo River

the Chambe Peak in the Mulanje Massif (2,500 m asl). In

basin in the cuvette centrale and at Kisangani, and in a

western Mozambique P. choloensis occurs on the forested

significant part of the African Rift Val ey from Lake Kivu south

slopes of Mount Mabo at 1,700 m asl (Cumberlidge, 2018b).

to LMNN. In the LMNNC, this species lives in LMNN itself but

Figure 3.3 The Nyika Plateau is home to the freshwater crab

Potamonautes choloensis. © Ludger Heide (CC BY-SA 2.0) 33
Figure 3.4 Potamonautes lirrangensis, commonly known as the Malawi
blue crab, is native to Lake Malawi/Nyasa/Niassa. It is assessed as Least
Concern (LC). © Oliver-Mengedoht.de/Panzerwelten.de

not in the tributaries of the lake or in any other rivers in Malawi

the Nyika Plateau, and also at the Kaombe River in Malawi

(Cumberlidge, 2018f). It has also been reported from Lake

(Figure 3.5). Parts of the region are relatively poorly sampled

Tanganyika from a locality in the southern part of the lake in

(e.g. Mozambique) and, therefore, these spatial patterns

Zambia (Marijnissen et al., 2006). Recent unpublished work

could be in part a reflection of sampling effort.

indicates that P. lirrangensis sensu lato might be a species

complex, and that the Malawi blue crab and the specimens

Three species of freshwater decapod are native to LMNN

from Lake Kivu (all currently recognised a P. lirrangensis)

itself (Figure 3.5): the shrimp C. malawensis, and the two

might belong to two different species. The low extinction

crabs, P. lirrangensis and P. montivagus. These species are

risk of these species is due to the general lack of threats

all restricted to a narrow band around the coasts of the lake

across the majority of their ranges, as fortunately large

(Figure 3.6), which reflects the distribution of littoral habitats

areas of their freshwater habitats are only lightly affected

(e.g. Figure 3.7).

by industrial development, aquatic habitat degradation and

pol ution events found near urban areas. However, even these

3.3.2 Endemic species richness

LC assessments are based on relatively poor quality data

and we stil need a great deal of survey work to learn more

Only two freshwater decapods are endemic to the LMNNC

about the biology, threats and distributional ranges of these

(Table 3.1): the shrimps C. kaombeflutilis and C. malawensis.

crustaceans.

Their known distributions are non-overlapping and are

depicted in Figure 3.8.

3.3 Patterns of species richness

Caridina kaombeflutilis was described in 2010 on the basis

of three specimens collected from eastern Malawi and no

3.3.1 Overall species richness

further specimens have been collected since (Richard &

Clark, 2009). The exact ecological requirements of this

The greatest species richness of freshwater decapods is in

species are not known, but one specimen was collected

the tributaries of LMNN, with the regions of highest species

from a pool in a dry river bed, and the other two specimens

richness on the northern side of the catchment near the

were collected from a large river. Part of the range of this

border between Malawi and Tanzania stretching south to

species lies in a protected area, the extensive Nkhotakota

34
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of freshwater decapod species

1
2

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 3.5 Richness of freshwater decapod species in the Lake

Malawi/Nyasa/Niassa Catchment based on spatial data coded as
Presence 1 (Extant). Richness data are classified using quantiles.
Mapping is to HydroBASINS and does not imply species occur across
the entire HydroBASIN (i.e. across the entirety of Lake
Malawi/Nyasa/Niassa). See Figure 3.6 for the distribution of species
within Lake Malawi/Nyasa/Niassa.

35
¯

20 40

80

120

160Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

Freshwater decapod distributions

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 3.6 Heat map showing relative richness of freshwater decapod

species in Lake Malawi/Nyasa/Niassa based on spatial data coded as
Presence 1 (Extant).

36
Wildlife Reserve (Figure 3.9), which includes wooded hills

also the two DD species. Please see section 3.3.2 Endemic

and mountains that are drained by numerous rivers that flow

species richness for a discussion of these species.

through miombo forests down into the lake at Nkhotakota.

This species is assessed as DD given the very limited

information on its distribution and ecology (De Grave &

3.4 Major threats

Cumberlidge, 2018a).
While none of the freshwater decapods native to the LMNNC

Caridina malawensis is assessed as DD because it is known

are assessed as threatened, close to half (44%) of the

only from a single specimen collected in 1974 from the

species had threats recorded in their Red List assessments.

rocky shores of LMNN in southern Malawi and no further

specimens have been collected since (Richard & Clark,

3.4.1 Agricultural and urban expansion

2009). This lake-living freshwater shrimp is associated with

the littoral zone and rocky shores of LMNN but otherwise

The human population in the LMNNC is growing rapidly.

very little is known of its habitat and ecological requirements.

Agricultural and urban expansion, required to support this

human population growth, are potential y major future threats

3.3.3 Data Deficient (DD) species richness

to freshwater biodiversity in this region (Darwal et al., 2005),

with agriculture (Figure 3.11) coded as a threat to 11% of the

The pattern of richness of freshwater decapods native to the

native decapods at present. Habitat loss and degradation

LMNNC and assessed as Data Deficient is shown in Figure

is of particular concern to species found outside protected

3.10. This figure reflects the distribution shown on Figure 3.8

areas. The conservation of freshwater decapods depends on

for endemic species, because the two endemic species are

preserving large enough areas of natural freshwater habitat

Figure 3.7 The littoral habitats of Lake Malawi/Nyasa/Niassa, such as at

Cape Maclear in Malawi, have the highest richness of freshwater
decapods in the lake. © Christian Albrecht

37
¯

0 25 50
100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of endemic freshwater

decapod species

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 3.8 Richness of endemic freshwater decapod species in the Lake

Malawi/Nyasa/Niassa Catchment based on spatial data coded as
Presence 1 (Extant). Richness data are classified using quantiles.
Mapping is to HydroBASINS and does not imply species occur across
the entire HydroBASIN (i.e. across the entirety of Lake
Malawi/Nyasa/Niassa). See Figure 3.6 for the distribution of species
within Lake Malawi/Nyasa/Niassa.

38
Figure 3.9 The freshwater shrimp Caridina kaombeflutilis is native to
Nkhotakota Wildlife Reserve in Malawi. © Catherine Sayer to maintain
good water quality in the LMNNC, given the

3.5 Recommended research and

species found there are sensitive to polluted or silted waters

conservation actions

and may not survive exposure. Pollution is coded as a threat

to 11% of the native decapods.

The limited nature of historical sampling means that there is a

lack of data available for freshwater decapods in the LMNNC.

3.4.2 Invasive species

Research was a recommended action for al of the freshwater

decapods native to the catchment. Our knowledge of

Although there are no reports of invasive crayfish species

species is based mainly on preliminary distribution data, with

in the LMNNC at present, it is nevertheless relevant to

information on specific threats and biological, ecological and

discuss their possible future impacts on freshwater decapod

population data stil lacking. There are stil many parts of the

habitats and populations should they arrive in the catchment.

LMNNC that have either never been studied for decapods,

Continental Africa has no native species of crayfishes, but

or require further surveying, and most of the region requires

there are two species of invasive crayfish that have become

more research attention. This no doubt reflects the chronic

established in southern Africa: the Louisiana red crayfish

lack of survey work in the freshwater ecosystems in this part

( Procambarus clarki ; Figure 3.12) and the Australian red claw

of Africa. This lack of basic information makes it difficult

crayfish ( Cherax quadricarinatus) (Foster & Harper, 2006a,

to make meaningful predictions about how species will

2006b, 2007). Both of these crayfishes are fast-growing

respond to changing freshwater environments and external

species that impact freshwater habitats because they feed

pressures, including those driven by changing climates, in

on aquatic plants, molluscs, small fishes and crustaceans

the future.

(Smart et al., 2002). In addition, these species, which can

travel over land, can easily expand their ranges, and have

Although the majority of freshwater decapods found in the

the potential to reach the LMNNC from nearby drainages,

LMNNC are LC, the two DD species of freshwater shrimps

with the Australian red claw crayfish already present in the

indicate that there are stil substantial gaps in our knowledge

Zambezi system (Nunes et al., 2016). Conservationists are

that need to be addressed. There is a need to intensify

concerned that if these two destructive species of crayfishes

ecological fieldwork, biotic inventories and conservation

reach LMNN they wil put the hundreds of endemic species of

activities to help gather data and establish the extinction

fishes and invertebrates at high risk of extinction (Kaufman,

risk of these species, and to define their true distributions,

1992) and could also have negative impacts on fisheries, and

abundance and threats. Making informed decisions about

therefore on the livelihoods of communities in the LMNNC,

the conservation and monitoring of poorly documented

as observed fol owing the spread of the Australian red claw

species, and about the management of ecosystems requires

crayfish to the Kafue River in Zambia (Weyl et al., 2017).

targeted surveys of the above nature be undertaken. The

39
¯
0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of Data Deficient freshwater

decapod species

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 3.10 Richness of Data Deficient (DD) freshwater decapod species

in the Lake Malawi/Nyasa/Niassa Catchment based on spatial data
coded as Presence 1 (Extant). Richness data are classified using
quantiles. Mapping is to HydroBASINS and does not imply species
occur across the entire HydroBASIN (i.e. across the entirety of Lake
Malawi/Nyasa/Niassa). See Figure 3.6 for the distribution of species
within Lake Malawi/Nyasa/Niassa.

40
Figure 3.11 A maize field near Lilongwe in Malawi. Agriculture is one of
the primary threats to freshwater decapods in the Lake
Malawi/Nyasa/Niassa Catchment. © Lars Plougmann (CC BY-SA 2.0)

Figure 3.12 The Louisiana red crayfish ( Procambarus clarkii) is now

established in southern Africa and is a potential future threat to
freshwater decapods in the Lake Malawi/Nyasa/Niassa Catchment. ©
fra928 (CC BY-NC-ND 2.0) 41

impacts of threats, such as habitat disturbance and pollution,

Final y, measures aimed at stopping the spread of invasive

need to be quantified.

species of crayfishes need to be considered before their

destructive impact reaches the wetland ecosystems

While it is encouraging that most of the LMNNC’s freshwater

associated with the African Rift Valley lakes. Management

decapods are LC, the long-term security of this fauna means

strategies for control ing the invasive Louisiana red crayfish

that it is important to actively protect the freshwater habitats,

include trapping and removing individuals, creating barriers to

primarily streams and rivers, where they live. Given the high

prevent its spread, prohibiting the transport of live crayfishes,

sensitivity of freshwater ecosystems to alterations, there is

and improving public education about its negative impacts

good reason to consider establishment of new protected

on aquatic ecosystems (Global Invasive Species Database,

areas better representing freshwater ecosystems as part of

2015). All of these strategies should be considered for the

conservation actions for these habitats and their crustacean

LMNNC. However, the main priority is to ensure these alien

faunas.

invasive species are prevented from entering the LMNNC as

their removal once present wil be highly chal enging.

Species in the spotlight

The hidden mountain crabs

Palmer-Newton, A.F.1 and Cumberlidge, N.2

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

2 Department of Biology, Northern Michigan University, Marquette,

Michigan, USA

Recent exploration and new taxonomic studies have shown that there are six
species of freshwater crabs within the LMNNC. However, there are stil
many areas that have either never been studied for decapods, or that require
further surveying. This is especially the case for remote, high-altitude
freshwater habitats (Daniels, Phiri & Bayliss, 2014).

Recent surveys of the freshwater crab fauna of the mountainous areas in

southern Africa have led to the description of a number of new species, and
it is likely that there are more undiscovered species living in the remote
highlands of the LMNNC.
One such recent discovery is the red river crab ( Potamonautes bellarussus
Daniels, Phiri & Bayliss, 2014) (Figure 3.13).

This blood-red coloured freshwater crab (LC) has been recorded from only
six highland localities living in boulder strewn mountain streams in the
Niassa Province of Mozambique (Figure 3.14) and in south-western
Tanzania, and its range includes the north-eastern part of the LMNNC
(Cumberlidge, 2018c; Daniels, Phiri & Bayliss, 2014).

In addition to this, four new species of freshwater crabs have recently been
described from high-altitude localities just south of the LMNNC in Malawi,
Mozambique and Zimbabwe. The first of these species is the Mount Mulanje
crab ( Potamonautes mulanjeensis) (DD) that is endemic to pools and
streams on Mount Mulanje in southern Malawi (Cumberlidge, 2018g;
Daniels & Bayliss, 2012). The second is the small-bodied Numali river crab
( Potamonautes namuliensis), which is endemic to boulder-strewn mountain
streams on Mount Namuli in central Mozambique (Cumberlidge, 2018h;
Daniels & Bayliss, 2012). The third new species is the Mount Mutare crab,
Potamonautes mutareensis, from highland habitat in the Nyanga mountains
in Mutare, Zimbabwe, in the Eastern Zimbabwe Highlands (Phiri & Daniels,
2013). Final y, the fourth is Potamonautes gorongosa, which is found in fast-
flowing mountain streams and rivers on Mount Gorongosa (1,863 m asl) in
Gorongosa National Park, northern Mozambique, close to the border with
Zimbabwe (Cumberlidge, Naskrecki & Daniels, 2016). The extinction risk
of two of these new species has not been evaluated, while the other two
species are currently assessed as DD, due insufficient information on their
threats to assess their extinction risk (Cumberlidge, 2018h, 2018g).

Freshwater crabs that live in high-altitude, mountainous habitats tend to have

a general y oval smooth body that lacks teeth on the sides, and the movable
finger of the largest claw is typically highly arched. Phylogenetic results
suggest that this mountain specialism has evolved independently on several
occasions, because these mountain-living species are not monophyletic
(Daniels et al., 2015; Daniels & Bayliss, 2012).

These new discoveries show that continued exploration of these remote

inland freshwater systems in the LMNNC will likely result in the discovery
of even more unique freshwater crab species (Daniels, Phiri & Bayliss,
2014).

42
Figure 3.13 Potamonautes bellarussus, commonly known as the red river
crab, has been recorded from only six highland localities, including in
the Lake Malawi/Nyasa/Niassa Catchment, where it lives in boulder
strewn mountain streams. It is assessed as Least Concern (LC). © Prof.
Julian Bayliss

Figure 3.14 The habitat of Potamonautes bellarussus at Mt Mecula in

Niassa Province, northern Mozambique. © Prof. Julian Bayliss

43

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iucngisd.org/gisd/species.php?sc=608 o n 28-09-2017.

Kaufman, L. (1992). ‘Catastrophic Change in Species-Rich

Freshwater Ecosystems’, BioScience, 42(11), pp. 846–

858. https://doi.org/10.2307/1312084

45

Chapter 4

The status and distribution of freshwater

fishes in the Lake Malawi/Nyasa/Niassa

Catchment

Kanyerere, G.Z.1, Phiri, T.B.2, Sayer, C.A.3, Shechonge, A.4, Snoeks, J.5
and Tweddle, D.6

Contents

4.1 Introduction
...........................................................................................................................
.......................................................................... 46

4.1.1 Freshwater fishes of Lake Malawi/Nyasa/Niassa, Lake Malombe and

the Upper Shire River .................................................................47

4.1.2 Freshwater fishes of Lake Chilwa

...........................................................................................................................
..................................51

4.2 Red List assessments

...........................................................................................................................
...........................................................52
4.3 Patterns of species richness
...........................................................................................................................
................................................ 53

4.3.1 Overal species richness

...........................................................................................................................
.............................................. 53

4.3.2 Endemic species richness

...........................................................................................................................
...........................................57

4.3.3 Threatened species richness

...........................................................................................................................
.......................................57

4.3.4 Data Deficient (DD) species richness

...........................................................................................................................
...........................57

4.4 Major threats

...........................................................................................................................
.........................................................................57

4.4.1
Fisheries.............................................................................................................
......................................................................................57

4.4.2 Agriculture
...........................................................................................................................
....................................................................61

4.4.3 Invasive species

...........................................................................................................................
...........................................................62

4.4.4 Climate
change................................................................................................................
....................................................................... 63
4.5 Recommended research and conservation actions
...........................................................................................................................
............. 63

Species in the spotlight

...........................................................................................................................
............................................................... 65

The one who sleeps – kalingono, Nimbochromis livingstoni

...........................................................................................................................
65

Volcanic evolution: a unique species in a crater lake – Chungruru tilapia,

Oreochromis chungruruensis ....................................................... 66

Liwonde National Park – a fish sanctuary for the ntchila, Labeo mesops
........................................................................................................67

Bibliography
...........................................................................................................................
................................................................................ 69

4.1 Introduction

differences and, so for the species summaries discussed in

this section, LMNN and Lake Chilwa are treated separately.

The Lake Malawi/Nyasa/Niassa Catchment (hereafter

LMNNC) as defined in this study primarily reflects the

LMNN is known for its high richness in freshwater fish

catchment of Lake Malawi /Nyasa /Niassa (hereafter

species, currently estimated at 800–1,000 native species

LMNN) and the outflowing Shire River, but also includes

including undescribed species (Konings, 2016; Snoeks,

the catchment of Lake Chilwa, an endorheic lake at the

2000, 2004; Turner et al., 2001). The majority of these species

headwaters of the Rovuma River, which is a major east

are from a single family, the Cichlidae, with approximately 60

coast river system that drains southern Tanzania and

species belonging to 12 other families (excluding non-native

northern Mozambique (Figure 1.2). The two systems have

species). Over 99% of the cichlid fish species are endemic

many freshwater fish species in common because the rift

to LMNN (Ribbink, 1991; Turner, 1996), meaning they are not

valley in which LMNN lies cuts across former east flowing

native to any other freshwaters globally. Moreover, there is

tributaries of the Rovuma system (Crossley & Crowe, 1980;

also a high degree of intra-lacustrine endemicity, with many

Weyl, Ribbink & Tweddle, 2010). There are, however, notable

species found only at particular islands or stretches of shore

1 Malawi Department of Fisheries, Ministry of Agriculture, Irrigation and

Water Development, Box 30134, Lilongwe 3, Malawi 2 Senga Bay Fisheries
Research Centre, P.O. Box 316, Salima, Malawi

3 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

4 Tanzania Fisheries Research Institute, P.O.BOX 98 Kyela, Mbeya,

Tanzania

5 Laboratory for Biodiversity and Evolutionary Genomics, KU Leuven

(Leuven University) and Department of Biology, Section Vertebrates, Royal
Museum for Central Africa, Leuvensesteenweg 13, B-3080 Tervuren,
Belgium

6 South African Institute for Aquatic Biodiversity (SAIAB), Somerset

Street, Grahamstown, 6139, South Africa 46

within the lake (Eccles & Trewavas, 1989; Owen et al., 1990;

and lakes, particularly in sheltered and well-vegetated

Ribbink, 1991; Snoeks & Hanssens, 2004).

areas. Their diet includes algae, water plants and aquatic

invertebrates (Ribbink, 2001).

Lakes Chilwa, which is included in the LMNNC for this study,

contains 30 species, of which 22 species are shared with

Members of the mouthbrooding Oreochromis are relatively

the rest of the LMNNC, although this is likely to change with

large and deep-bodied. Within the genus three endemic

ongoing taxonomic investigation. One family, Schilbeidae

species, Oreochromis karongae, O. lidole (Figure 4.2) and

(the species Pareutropius longifilis) is found in the Lake

O. squamipinnis (Eccles, 1992; Turner, 1995), local y known

Chilwa catchment but is absent from the rest of the LMNNC.

as chambo, were one of the most important fisheries in

LMNN, supporting both commercial and artisanal fisheries

4.1.1 Freshwater fishes of Lake Malawi/

but catches have severely declined in the last two decades

Nyasa/Niassa, Lake Malombe and the

(Weyl, Ribbink & Tweddle, 2010). A fourth endemic species,

Upper Shire River

O. chungruruensis, is restricted to a single crater lake in

the catchment in Tanzania (see Species in the spotlight:

4.1.1.1 Cichlids (Cichlidae)

Volcanic evolution: a unique species in a crater lake –

Cichlids are a large family of fresh and brackish water

Chungruru tilapia, Oreochromis chungruruensis p. 66).

fishes found in Africa and Madagascar, the Middle East,

The non-endemic species, O. shiranus, which also occurs

in South, Central and the southern part of North America,

in the Middle Shire River south of the LMNNC, is more

and in South-east Asia. They have many unusual biological

abundant in smaller water bodies, lagoons and swamps

features, which include a highly evolved parental care system

where it supports major artisanal fisheries .

and the potential to evolve rapidly into an array of species

capable of colonising almost all habitats from shallow to

4.1.1.1.2 Haplochromine Cichlids

deep waters whether vegetated, rocky, sandy or muddy

The LMNNC is home to many haplochromine genera, the

(Lewis, Reinthal & Trendall, 1986; Skelton, 2001). Many

great majority of which are endemic to the lake. Only three

cichlids are important food fishes, while others are popular

haplochromine cichlids within the catchment ( Astatotilapia

in the aquarium trade because of their attractive colours

cal iptera, Pseudocrenilabrus philander and Serranochromis

(e.g. Figure 4.1) (Eccles, 1992; Konings, 2016; Ribbink et al.,

robustus) are non-endemic to the LMNNC (Eccles, 1992;

1983; Turner, 1995). Within the LMNNC, there are two main

Tweddle, 1996). They occur in rivers, swamps, lagoons

lineages: tilapi nes and haplochromines.

and inshore swampy areas of lakes (Turner, 1995) and are

important for the artisanal fisheries, being harvested using

4.1.1.1.1 Tilapi ne Cichlids

hand lines, fish traps and seine nets. Pseudocrenilabrus

Three tilapiine cichlid genera are found in the LMNNC:

philander is a small species restricted to inflowing rivers

Oreochromis, Tilapia and Coptodon. Tilapia sparrmanii and

and swamps. It is widespread throughout the Zambezi

Coptodon rendalli are substrate spawners, while the five

system and in other river systems to the south. Astatotilapia

Oreochromis species are mouthbrooders (Trewavas, 1983).

calliptera (Figure 4.3) is a similar sized species that is most

They are valuable food fishes, supporting both commercial

abundant in rivers but is also found in shallow sheltered

and artisanal fisheries, besides being cultured. They are

areas of LMNN. This species is also found in the Lower

widely distributed, occurring in rivers, swamps, lagoons

Zambezi, but not further upriver, and it is also found in rivers

Figure 4.2 Oreochromis lidole, locally known as chambo, at

Figure 4.1 Aulonocara stuartgranti is popular in the aquarium

Domwe Island in Malawi. This species is assessed as Critically

trade. It is assessed as Least Concern (LC). © Ad Konings

Endangered (CR). © Ad Konings

47

further south, although taxonomic study may separate

metres from the lake (Tweddle & Willoughby, 1978). Most

some of these populations. Serranochromis robustus is a

species are habitat specific and many are endemic to a

much larger predatory fish preferring deep mainstream river

very smal area of the lake, some, particularly in the mbuna

channels, lagoons and inshore areas of lakes (Genner et al.,

group of rocky shore species (see below), being restricted

2007). The identity of specimens in the Luangwa River in

to a single rocky outcrop, while others are wider ranging

Zambia needs taxonomic review but they are believed to be

(Ribbink, 2001).

this species rather than the related S. jal ae of the Upper and

Middle Zambezi and Kavango Rivers.

The demer sal species are stratified by depth and

species composition of trawl catches is strikingly

In contrast to the non-endemics, the endemic haplochromine

different at different depths (FAO, 1976; Ribbink, 2001).

cichlids comprise what is commonly referred to as a “species

The demersal cichlid genera targeted by fishers include

flock”, representing unparalleled evolution of species not

Buccochromis, Maravichromis, Lethrinops, Aulonocara,

seen to the same degree outside of the African Rift Valley

Alticorpus, Placidochromis, Otopharynx, Protomelas and

lakes (Brawand et al., 2014). Approximately 800–1,000

Nyassachromis. In demersal trawl catches, Turner (1977b)

species are currently recognised in LMNN (Konings, 2016;

and Tómasson & Banda (1996) reported up to 140 species

Snoeks, 2000, 2004; Turner et al., 2001) and the species are

caught in the trawl in the south-west and south-east arms

the focus of numerous books (e.g. Eccles & Trewavas, 1989;

of the lake, while Ribbink (2001) reported 145–170 species

Konings, 2016; Turner, 1996). The cichlids dominate the lake

caught in trawling transects in the south-west arm in

in both species richness and numerical abundance (Ribbink,

1998–1999. The greatest species richness is in the shal ows

2001).

(Ribbink, 2001).

These cichlids are widely studied in the development

Pelagic and semi-pelagic genera include Diplotaxodon,

of models of rapid speciation and adaptive radiation by

Rhamphochromis, Copadichromis and Dimidiochromis

evolutionary biologists (e.g. Genner & Turner, 2005; Owen

with about 50–80 species, many of which are undescribed

et al., 1990; Salzburger & Meyer, 2004; Seehausen, 2006;

(Ribbink, 2001). By far the most important pelagic fishery

Sturmbauer, 1998).

species are the utaka, genus Copadichromis, which are

targeted by chirimila nets (Jackson et al., 1963). The potential

The cichlids are important species for both food (Banda,

for further development of offshore fisheries in the lake was

Tomasson & Tweddle, 1996; FAO, 1976, 1993; Turner, 1995,

explored by Thompson and Al ison (1997).

1977a; Tweddle & Turner, 1977; Weyl, Ribbink & Tweddle,

2010) and the ornamental fish trade (Konings, 2016; Lewis,

The ornamental fishery is based largely on the mbuna

Reinthal & Trendal , 1986; Ribbink et al., 1983). They occupy

group of rocky shore dwelling cichlids in the genera

a wide range of littoral, sublittoral, demersal, pelagic

Pseudotropheus, Labeotropheus, Maylandia (= Metrioclima),

and semi-pelagic habitats in the lake, extending down

Melanochromis, Labidochromis, Cynotilapia, Petrotilapia,

to depths where deoxygenation limits their metabolism

Ge nyochromis , Cyathochromis , G e phyrochromis ,

(Ribbink, 2001). A small number of species penetrate into

Iodotropheus and Tropheops, but also includes most of

the lowermost reaches of inflowing streams and associated

the other non-mbuna genera, most notably Aulonocara

fringing wetlands but seldom more than a few hundred

(Konings, 2016). The rocky shore cichlid fauna comprises

about 400 species (Ribbink, 2001).

4.1.1.2 Non-cichlids

Other equally important fish groups in the LMNNC are the

non-cichlid families. In LMNN three families of non-cichlid

fishes (Cyprinidae, Clariidae and Mochokidae) include

pelagic, open water species, but the majority are demersal,

or bottom dwel ing.

4.1.1.2.1 Cyprinids (Cyprinidae)

Fishes of the family Cyprinidae include a wide range of

sizes and shapes, with diverse life-history strategies and

habitat preferences (Eccles, 1992; Skelton, 2001). In the

LMNNC there are five native cyprinid genera ( Engraulicypris,

Figure 4.3 A male Astatotilapia calliptera at Thumbi East in

Opsaridium, Labeo, Labeobarbus and Enteromius) with

Malawi in Malawi. This species is assessed as Least Concern

(LC). © Ad Konings

26 described native species (Eccles, 1992; Snoeks, 2004;

48

Tweddle, 1996). The riverine cyprinids form artisanal fisheries

currents and overcoming obstacles such as rapids. Their

in central and northern LMNNC in rivers flowing from the

main diet includes algae and detritus from the substratum.

Nyika and Viphya plateaus.

There are more than 80 species in Africa and most of them

support major artisanal river fisheries (Eccles, 1992; Skelton,

Engraulicypris sardella (usipa; Figure 4.4) is a small lake

2001) employing fish traps and gil nets. In the LMNNC, there

species endemic to and widely distributed throughout

are two extant species: Labeo mesops and L. cylindricus.

LMNN. It is pelagic, feeding exclusively on zooplankton.

Labeo mesops migrates into flooded grassland, including

It is currently (2019) the most important fishery species,

permanent and temporary streams, to breed. It used to be

supporting small-scale fisheries and, to a smaller extent,

caught in large quantities by artisanal fishing gears, such as

large-scale lift net fisheries. It is mainly caught in open

gil nets, weirs and baskets, but has now become very rare

water seines, such as chilimira where light attraction is

except in streams and rivers with relatively low or negligible

used, and usipa beach seine nets. The species is also a

impact, draining relatively undisturbed catchments, such as

major prey item for predatory pelagic species in the genera

in Liwonde National Park (Tweddle, 2018a) (see Species in

Rhamphochromis, Diplotaxodon and Opsaridium.

the spotlight: Liwonde National Park – a fish sanctuary

for the ntchila, Labeo mesops p. 67). On the other

Members of the genus Opsaridium, which have a salmonid-

hand, L. cylindricus, which prefers rocky habitats, is still

like appearance (Tweddle & Lewis, 1983), in general

quite abundant in LMNN and, to a lesser extent, Lake

favour the clear, flowing waters of larger perennial rivers,

Malombe. It is very common in the inflowing rivers, where

frequenting pools below rocky rapids, but two of the three

there are permanent riverine populations in addition to

species present in the LMNNC, Opsaridium microcephalus

potamodromous fish ascending from the lake to breed. It

and O. microlepis, have evolved to also favour lacustrine

is also found commonly throughout the Shire River. This

conditions (Tweddle & Lewis, 1983; Tweddle & Turner, 2014).

species is targeted by the gil net and trap fisheries. A third

The third species, O. tweddleorum, lives in the rivers where

species, L worthingtoni, is only know from the holotype and

it feeds on a variety of food items, including aquatic insects,

two paratypes and is considered extinct (Tweddle, 2018b).

invertebrates and other small organisms. The two larger

species, O. microcephalum and O. microlepis, are both

African species in the LMNNC formerly included in the

pelagic piscivores but they migrate into major rivers to breed

Eurasian genus Barbus are now placed in the genera

(Tweddle, 1983), although O. microcephalum also breeds

Enteromius (small barbs) and Labeobarbus (larger barbs).

on rocky exposed shores in the lake (Tweddle & Turner,

There are 15 known Enteromius species in the LMNNC, and

2014). Both species contribute greatly to riverine fisheries

three species of Labeobarbus, one of which, L. nthuwa, is

especially when they migrate upstream to breed. In LMNN

restricted to the South Rukuru River. They inhabit a wide

they are also caught in gil nets, while juveniles are a bycatch

range of habitats from smal streams to large rivers and lake

of the fishery for usipa employing mosquito netting.

margins. They are of minor importance to artisanal fisheries

in the LMNNC.

Labeo is a widely distributed genus in Africa and South-

east Asia. Labeo species migrate upstream to breed and,

4.1.1.2.2 Catfishes (Clari dae, Bagridae, Amphili dae,

as such, they are wel adapted to swimming against strong

Schilbeidae and Mochokidae)

Catfishes comprise several families of scaleless fishes found

in a wide range of habitats from mountain streams, swamps

and lagoons to large lakes. The catfish families found in the

LMNNC are Clariidae, Bagridae, Amphiliidae, Schilbeidae

and Mochokidae, many of which include important food

fishes.

The two clariid catfish genera, Clarias and Bathyclarias,

include 13 species native to the LMNNC . Members of the

genus Clarias occur in almost any habitat but most species

prefer floodplains, large slow flowing rivers, lakes and dam

impoundments. They are omnivorous feeding on a wide

range of materials such as frogs, birds, small mammals,

snails, other invertebrates and plant matter, including seeds

Figure 4.4 Engraulicypris sardella, commonly known as

and fruits. Breeding occurs in summer early in the rains,

usipa, is currently the most important fishery species in Lake

when large numbers of mature fishes migrate to flooded

Malawi/Nyasa/Niassa. It is assessed as Least Concern (LC).

© Ad Konings

shallow grassy fringes of rivers and lakes (Skelton, 2001).

49

They are mostly caught in fish traps and on baited hooks.

nets and seine nets. It also occurs in small numbers in the

Five species, C. gariepinus, C. ngamensis, C. theodorae,

lower reaches of the larger rivers. On one occasion, an

C. stappersii and C. liocephalus, occur in the LMNNC. Of

upstream mass breeding migration was observed during

these, two have restricted distributions within the LMNNC,

a flood in the Bwanje/Livulezi delta adjacent to the south-

with C. stappersi only known from the Limphasa Dambo and

west arm of the lake (D. Tweddle pers. obs.). Fishers had

C. liocephalus in the South Rukuru River, although they are

placed fish traps in the road culverts to catch fish migrating

more widely distributed outside of the catchment.

upstream through them. These traps were packed with

Synodontis such that they were difficult to extract from

Members of the endemic genus Bathyclarias inhabit both

the traps. Undescribed mochokid species in the genus

shallow and deep waters of LMNN (Lewis, Reinthal &

Chiloglanis are common in the rapids and riffles of inflowing

Trendall, 1986). Those species which frequent surface

rivers, while another Chiloglanis species has localised

waters feed on zooplankton. Others are piscivorous, while

distributions in intermediate zones in the lake itself (Fryer,

those living in deep waters feed on a variety of invertebrates

1959; R. Bil s pers. comm.).

(Lewis, Reinthal & Trendall, 1986). Locally called bombe

or sapuwa, these fishes are the largest in LMNN and can

4.1.1.2.3 Spiny eels (Mastacembelidae)

reach 1.5 m in length and weigh over 30 kg (Lewis, Reinthal

Spiny eels (family Mastacembelidae), with an estimated 43

& Trendal , 1986). It is, therefore, not surprising that they are

species in Africa, are eel-like fishes (Skelton, 2001). In the

caught in both large and small-scale fisheries using trawl

LMNNC there is just one described species, Mastacembelus

nets, gil nets, long lines and seine nets.

shiranus, which commonly occurs in the vegetated fringes

of rivers, lagoons and lakes but is not frequently caught in

In the LMNNC the family Bagridae is represented by a single

fisheries.

species, Bagrus meridionalis, locally known as kampango

or kampoyo. It occurs in both commercial and small-scale

4.1.1.2.4 Anguil id eels (Anguil idae)

fisheries but has declined sharply in the commercial fishery.

There are four species of anguillid eels in eastern and

southern Africa, but only one species, A. bengalensis labiata,

Members of the family Amphili dae, smal African catfishes,

is recorded in the LMNNC having migrated to the lake from

are found mainly in clear flowing streams and rivers. In the

the Indian Ocean via the Zambezi and Shire Rivers (Jackson,

LMNNC, the family includes two genera ( Amphilius and

1961; Tweddle, 1996). Although it is a popular food fish

Zaireichthys) and at least five species. Members of the

elsewhere, in the LMNNC it is only caught occasional y, using

genus Amphilius, known as mountain catfishes, inhabit fast

baited hooks at night, and is seldom eaten.

flowing, mainly rocky rivers. Individuals of Amphilius in the

LMNNC are referred to as A. uranoscopus, but ongoing

4.1.1.2.5 Elephant fishes (Mormyridae)

taxonomic research will undoubtedly lead to changes to

This African family has 21 genera and some 228 valid species

this nomenclature. The genus Zaireichthys, known as sand

distributed throughout tropical Africa (Froese & Pauly, 2019).

catlets, includes at least three species located in rivers

There are seven described species native to the LMNNC

flowing into LMNN (Eccles, Tweddle & Skelton, 2011) where

(Kadye et al., 2008; Tweddle & Wil oughby, 1982). Mormyrids

they prefer sandbanks in a gentle to moderate water flow.

are usual y active at night and can generate and receive weak

Zaireichthys lacustris is a dwarf species maturing at around

electric currents to communicate with each other, and to

17 mm. This endemic species lives and guards its young

detect prey and predators. Their preferred habitats include

in empty shells of the mollusc Lanistes nyassanus (Eccles,

rivers, floodplains, swamps and vegetated, deep and quiet

Tweddle & Skelton, 2011).

areas of lakes. They are caught in smal numbers in artisanal

fisheries, especial y those of shal ow water bodies and large

One species of the family Schilbeidae, Pareutropius longifilis,

rivers. The fishing gears used include gill nets, seine nets,

occurs in the wider LMNNC but is native only to Lake Chilwa

fish traps and hand lines.

within the catchment, and discussed in section 4.1.2

Freshwater fishes of Lake Chilwa.

4.1.1.2.6 Alestidae

Two alestid species, Brycinus imberi and Hemigrammo-

Squeakers or suckermouth catlets belong to the family

petersius barnardi, occur in the LMMNC (Eccles, 1992).

Mochokidae, the largest African catfish family (Skelton,

Brycinus imberi is a shoaling species found in sheltered

2001). A single Synodontis species, S. njassae, is native to

inshore habitats in lakes, lagoons and in the lower reaches

the LMNNC. It is primarily a lacustrine species that occurs

of rivers, while H. barnardi, one of the smal est fish species

throughout LMNN in both demersal and pelagic habitats,

in Malawi at under 4 cm total length, is common in sheltered

including surface and deep waters (Eccles, 1992). It is

weedy areas on the lakeshore plain. They are of negligible

abundant in the lake and is harvested using trawl nets, gill

importance in the LMNNC fisheries.

50

4.1.1.2.7 Nothobranchi dae

the LMNNC. For the latter, individuals in Lake Chilwa are

Nothobranchius species are small and colourful killifishes

considered to be a subspecies, O. shiranus chilwae. This is

that live in relatively small ephemeral bodies of water and

one of the three most important fishery species in the lake,

have extreme life-history adaptations that allow them to

the others being Clarias gariepinus and E. paludinosus, with

survive the periodic drying up of their natural habitats.

the latter being by far the most important (Furse, Morgan &

Nothobranchius wattersi (Figure 4.5) occurs in temporary

Kalk, 1979).

pans on the southern lakeshore plains of the LMNNC.

4.1.2.1.2 Haplochromine Cichlids

4.1.1.2.8 Poecili dae

Unlike LMNN, Lake Chilwa does not support an endemic

Poeciliids are a large family of about 30 genera and 300

cichlid species flock. There are three known small haplo-

species found in North, Central and South America, the

chromines, two of which, Astatotilapia calliptera and

Caribbean and Africa. The family includes well-known

Pseudocrenilabrus philander, are shared with the rest of the

live-bearing aquarium fish such as the guppy. Due to the

LMNNC. The third, Haplochromis (= Astatotilapia) tweddlei,

release of aquarium specimens and the widespread use of

is restricted to the inflowing streams of Lake Chilwa.

species for mosquito control, poeciliids can now be found

non-natively in all tropical and subtropical countries. Only

4.1.2.2 Non-cichlids

one described species of Poecili dae is native to the LMNNC,

4.1.2.2.1 Cyprinids (Cyprinidae)

Micropanchax johnstoni. This species prefers inshore well

There are 12 cyprinid species recorded in Lake Chilwa.

vegetated shallow habitats in the lake and inflowing rivers.

Eleven species of Enteromius are recorded from the lake,

A second undescribed species has been recorded in an

three of which are undescribed and four of which are not

inflowing stream in Mozambique (M. Genner pers. comm.).

found elsewhere in the LMNNC. Labeo cylindricus (Figure

4.6) is shared with the rest of the LMNNC.

4.1.2 Freshwater fishes of Lake Chilwa

The fish fauna of Lake Chilwa is less diverse than in LMNN.

4.1.2.2.2 Catfishes (Clari dae, Amphili dae, Schilbeidae

Lake Chilwa is an ephemeral, shal ow, saline, endorheic lake.

and Mochokidae)
Due to its high salinity the fish fauna is depauperate, with only

Two clariids occur in Lake Chilwa, both of which are also

three species commonly found in the lake itself, Enteromius

found elsewhere in the LMNNC. These are the ubiquitous

paludinosus, Clarias gariepinus and Oreochromis shiranus

Clarias gariepinus (Figure 4.7) and C. theodorae.

chilwae. These species do, however, support a fishery

yielding over 10,000 tonnes per year when the lake is full

Two amphili d species are known from the catchment of Lake

(Furse, Morgan & Kalk, 1979). The inflowing streams have

Chilwa, the mountain catfish Amphilius cf. uranoscopus,

greater species diversity (Tweddle, 1979, 2005).

which occurs in the fast-flowing rocky sections of streams

flowing into the lake, and an undescribed sand catlet,

4.1.2.1 Cichlids (Cichlidae)

Zaireichthys sp., found over sand banks in flowing streams.

4.1.2.1.1 Tilapi ne Cichlids

The two tilapiines in Lake Chilwa, Coptodon rendalli and

The small East African schilbeid Pareutropius longifilis is

Oreochromis shiranus, are shared with the main part of

fairly common in Lake Chilwa streams, though unimportant

Figure 4.5 A male Nothobranchius wattersi caught near the

village of Hoba, upper Shire River floodplain, just south of

Figure 4.6 Labeo cylindricus occurs throughout the Lake

Lake Malombe in Malawi. This species is assessed as Near

Malawi/Nyasa/Niassa Catchment. It is assessed as Least

Threatened (NT). © B.R. Watters

Concern (LC). © Ad Konings

51

in the fishery. This species is not found elsewhere in the

Turner et al., 2001), the Red List assessments completed

LMNNC.

through this study were limited to taxonomical y described

species. As a result, Red List assessments of 459 freshwater

The family Mochokidae is represented by an undescribed

fish species native to the LMNNC were completed through

species of Chiloglanis in the faster flowing rocky streams.

this study with a high proportion (407 species, 89%) being

considered endemic to the catchment (Table 4.1). The

4.1.2.2.3 Elephant fishes (Mormyridae)

undescribed species native to the LMNNC are primarily

There are three mormyrid species in Lake Chilwa, Petro-

cichlids endemic to the lake, such that the true number of

cephalus catostoma, Marcusenius livingstonii and M.

endemic species will be significantly higher than recorded

macrolepidotus. At present, they are considered to be

here. Given the high degree of endemicity, patterns in Red

shared with the rest of the LMNNC, although taxonomic

List categories are similar when displayed as either native or

study is ongoing.

endemic species groupings (Figure 4.8, Table 4.1).

4.1.2.2.4 Alestidae

Thirty-eight native species (9% of extant species excluding

The same two species, Brycinus imberi and Hemigrammo-

species assessed as Data Deficient, DD) and 34 endemic

petersius barnardi, that are found in main body of the

species (9% excluding those assessed as DD) are assessed

LMNNC also occur in Lake Chilwa streams.

in a threatened category (Figure 4.8, Table 4.1). This level

of threat is significantly lower than recorded for continental

4.1.2.2.5 Nothobranchi dae

Africa, where 27% of native species (excluding DD species)

The species found adjacent to Lake Chilwa, Nothobranchius

were assessed as threatened (Darwall et al., 2011), and

kirki, is a close relative of N. wattersi found elsewhere in the

for the Lake Victoria Basin, where 56% of native species

LMNNC. Nothobranchius kirki is found in temporary habitats

(excluding DD species) were assessed as threatened

on both the Malawian and Mozambican sides of Lake Chilwa,

(Sayer, Máiz-Tomé & Darwal , 2018). However, although this

including in the extensive rice plantations in Malawi.

message is currently positive, note should be taken of other

large lakes and lake systems, such as Lake Victoria in East

4.1.2.2.6 Poecili dae

Africa (Sayer, Máiz-Tomé & Darwal , 2018) or the Malili Lakes

Micropanchax johnstoni is the only poeciliid species in

in Sulawesi, Indonesia (Herder et al., 2012), where threats,

streams in the Lake Chilwa catchment.

such as the introduction of invasive alien species, have

contributed to dramatic and fast declines in the status of

freshwater biodiversity and endemic species flocks.

4.2 Red List assessments

Only one species is listed as Extinct (EX) (Table 4.1): the

For the remainder of this chapter, the species of the LMNNC

endemic Labeo worthingtoni. This is a poorly known species

as defined for this study (Figure 1.2), including those of Lake

that has not be recorded since its description from several

Chilwa, are discussed together.

sites around LMNN in the early 1930s (Tweddle, 2018b). This

is, however, potential y an underestimate of the true number

Although LMNN is estimated as hosting 800 –1,000

of species extinctions in the catchment. Four species, al of

freshwater fish species (Konings, 2016; Snoeks, 2000, 2004;

which are cichlids, are assessed as Critically Endangered

(CR) and flagged as Possibly Extinct (PE) (e.g. Figure 4.9),

representing 20% of al CR species. Surveys are required to

determine whether these species are stil extant.

Forty-one native species (9%) and 38 endemic species

(9%) are listed as DD (Figure 4.8, Table 4.1), indicating

that further information is required to assess their relative

extinction risk. This is significantly lower than the results for

continental Africa and the Lake Victoria Basin, where 18%

and 33% were assessed as DD, respectively (Darwal et al.,

2011; Sayer, Máiz-Tomé & Darwall, 2018), indicating that

the freshwater fishes of the LMNNC are, in general, better

known than for the continent as a whole and for some other

Great Lakes. However, species level data are stil lacking for

Figure 4.7 Clarias gariepinus occurs throughout the Lake

Malawi/Nyasa/Niassa Catchment. It is assessed as Least

many of the freshwater fishes native to the LMNNC meaning

Concern (LC). © Ad Konings

that their Red List statuses were assigned based on inferred

52
information. Systematic surveys of the lake and catchment

(Figure 4.8, Table 4.1). Although the majority of species within

that identify individuals to species level need to be resumed

the LMNNC are restricted to the lake, many have widespread

and extended to all areas of the lake in order to effectively

distributions within LMNN and are at low relative risk of

track changes in the lake fauna and to inform conservation

extinction. There are, however, many species restricted to

and development planning.

small islands or short sections of the shoreline. For these

species, with sometimes highly restricted distributions,

The majority of species (75% of native and 74% of endemic

the risk of extinction is still currently considered as low if

species) are Least Concern (LC), and 8% and 9% of native

they face no threats or only minor threats (e.g. Figure 4.10).

and endemic species, respectively, are Near Threatened (NT)

However, it is important that these species are monitored

closely and threats are contained because, given their

Table 4.1 Number of freshwater fish species native and endemic

restricted distributions, a single new threatening event could

to the Lake Malawi/Nyasa/Niassa Catchment in each Red List

put them at risk of global extinction.

Category. For a list of species native to the Lake Malawi/Nyasa/

Niassa Catchment and their Red List Categories please see

Appendix 1.

Number of

Number of

4.3 Patterns of species richness

species native to species endemic

IUCN Red List Category

the LMNNC

to the LMNNC

4.3.1 Overall species richness

Extinct (EX)

Extinct in the Wild (EW)

The greatest richness of freshwater fishes in the LMNNC

Critically Endangered (CR)

20

18

is found in LMNN itself, where 427 described species were

Endangered (EN)

mapped to be present. The next richest areas are Lake

Vulnerable (VU)
11

Near Threatened (NT)

37

35

Malombe and the catchments surrounding the south-

Least Concern (LC)

342

299

east arm of the lake, including the Shire River, which host

Data Deficient (DD)

41

38

79–90 species. The rivers and tributaries of LMNN on the

Total

459

407

western side of the lake and Middle Shire River south to

Endemic species

Native species
0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

EX

EW

CR

EN

VU

NT

LC

DD

Figure 4.8 Percentage (%) of freshwater fish species native and endemic
to the Lake Malawi/Nyasa/Niassa Catchment in each Red List Category.
For a list of species native to the Lake Malawi/Nyasa/Niassa Catchment
and their Red List Categories please see Appendix 1.

Figure 4.9 Nyassachromis breviceps is assessed as Critically

Figure 4.10 A territorial male Aulonocara hueseri at Likoma

Endangered (Possibly Extinct) (CR(PE)). It has not been

Island in Malawi. This species is restricted to a single island but

collected since 1997 and is thought to have been extirpated by

there are no significant threats to the population. Therefore, it

artisanal fishing. © Ad Konings

is assessed as Least Concern (LC). © Ad Konings

53
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of freshwater fish species

6 - 13

14 - 24

25 - 40

41 - 90

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

91 - 427

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 4.11 Richness of freshwater fish species in the Lake
Malawi/Nyasa/Niassa Catchment based on spatial data coded as
Presence 1 (Extant). Richness data are classified using quantiles.
Mapping is to HydroBASINS and does not imply species occur across
the entire HydroBASIN (i.e. across the entirety of Lake
Malawi/Nyasa/Niassa). See Figure 4.12 for the distribution of species
within Lake Malawi/Nyasa/Niassa.

54
¯

20 40

80

120
160Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

Freshwater fish distributions

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 4.12 Heat map showing relative richness of freshwater fish

species in Lake Malawi/Nyasa/Niassa based on spatial data coded as
Presence 1 (Extant).

55
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of endemic freshwater fish

species

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

4 - 401

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 4.13 Richness of endemic freshwater fish species in the Lake
Malawi/Nyasa/Niassa Catchment based on spatial data coded as
Presence 1 (Extant). Richness data are classified using quantiles.
Mapping is to HydroBASINS and does not imply species occur across
the entire HydroBASIN (i.e. across the entirety of Lake
Malawi/Nyasa/Niassa). See Figure 4.12 for the distribution of species
within Lake Malawi/Nyasa/Niassa.

56

Lake Chilwa have moderate species richness, with between

with the next highest levels of data deficiency, with three

25–40 species mapped. Lowest species richness is found

DD species mapped to occur in each of these catchments

in the extreme north-west and north-east of the LMNNC

(Figure 4.16).

in Tanzania (Figure 4.11) but this is likely a product of low

sampling effort in these regions.

There are two main reasons that species within the LMNNC

are assessed as DD. First, a number of species are known

Richness of freshwater fish species is not uniformly

from only a few specimens, meaning there is insufficient

distributed across LMNN, with the majority of species found

information on their distribution, population status and

within the peripheral shal ow waters at less than 100 m depth

ecology to assess their relative extinction risk. Other DD

in sand and rock lakeshore habitats. Regions of the lake

species are primarily harvested species for which there

with vast expanses of shallow waters, such as the south-

is insufficient population information, for example from

east and south-west arms, the western shoreline within

fisheries catch statistics, to assess long-term population

the Malawian districts of Salima, Nkhotakota and Karonga,

trends and hence, the status of populations (e.g. Figure 4.17).

and small sections on the eastern shoreline, for example

Systematic surveys of the lake and catchment that identify

around Chizumulu and Likoma islands, have highest relative

individuals to species level would help to provide further

richness of fishes. Pelagic species are found throughout the

information on these species, which could be used to inform

lake and there are demersal species living at depths down

their Red List assessments.

to an estimated 130 m close to the anoxic zone (Figure 4.12).

There are two regions with no DD species: the catchments

4.3.2 Endemic species richness

around the Kasungu National Park in Malawi, and the region

stretching from the southern Nyika Plateau in the north to the

LMNN supports the highest richness of endemic species

headwaters of the South Rukuru River in the south (Figure

in the catchment, with 401 endemic species mapped to

4.16).

occur in the lake. Lake Malombe and the Upper Shire

River represent the next richest areas with 52–60 endemic

species mapped. Other areas of relatively high endemic

4.4 Major threats

species richness are the Kiwira River in southern Tanzania

and the South Rukuru River in central Malawi, each with four

Threats affecting the integrity of an ecosystem can be

to five species endemic to the catchment. Three areas of

categorised into two groups: i) those that occur naturally,

the LMNNC have no endemic species: the extreme north-

such as floods and natural fires; and ii) anthropogenic

west and the majority of the north-east of the catchment in

activities. Threats induced by human activities are currently

Tanzania, and a significant portion of the LMNNC north and

recorded as the most significant in the LMNNC due to the

west of Lilongwe in Malawi (Figure 4.13).

agro-based economy. Such economies rely heavily on

utilisation of natural resources, which often manifests in

4.3.3 Threatened species richness

the form of agriculture, fisheries, mining and associated

activities. The rapidly growing human population in the

LMNN also supports the highest richness of threatened

LMNNC makes the demand for farmland increase further

freshwater fishes with 35 threatened species mapped to

and also puts additional pressure on other resources, such

occur in the lake (Figure 4.14). Again, the Upper Shire River

as fuelwood and fisheries.

(Figure 4.15) and Lake Malombe also have relatively high

numbers of threatened species, with nine to ten threatened

4.4.1 Fisheries

species mapped to occur in these catchments. The Kiwira

River in Tanzania and the Shire River below Lake Malombe

The freshwater fishes of LMNN support a fishery that is of

in southern Malawi also have relatively high numbers (four)

great importance to the livelihoods of communities and

of threatened species. Mirroring total and endemic species

the economies of countries in the LMNNC, but overfishing

richness (Figure 4.11, Figure 4.13), the extreme north-west

and use of destructive fishing gears pose a threat to these

and the north-east sections of the LMNNC in Tanzania have

species with 75% of freshwater fish species native to the

no threatened species (Figure 4.14).

catchment recorded as threatened by fishing and harvesting

of aquatic resources. LMNN supports a highly diverse

4.3.4 Data Deficient (DD) species richness

capture fishery that can be categorised as large-scale

commercial, small-scale commercial and subsistence

LMNN is also richest in DD species, supporting 35 species

(Banda et al., 2001). The large-scale commercial fishery is

assessed in this category. Catchments on the northern,

a mechanised fishery that operates trawls, purse seines

western and southern sides of LMNN represent the areas

or lift nets. The smal -scale commercial fishery includes all

57
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of threatened freshwater fish

species
0

3-4

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

5 - 35

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 4.14 Richness of threatened freshwater fish species in the Lake

Malawi/Nyasa/Niassa Catchment based on spatial data coded as
Presence 1 (Extant). Richness data are classified using quantiles.
Mapping is to HydroBASINS and does not imply species occur across
the entire HydroBASIN (i.e. across the entirety of Lake
Malawi/Nyasa/Niassa). See Figure 4.12 for the distribution of species
within Lake Malawi/Nyasa/Niassa.

58

fishers that use engines of less than 20 horsepower or no

catch and CPUE for the main gears in the artisanal fishery.

engine (Figure 4.18) to fish with the catch intended primarily

Most dramatic has been the increase in CPUE in the chilimira

for sale. Gears used in this sector include beach seines,

net fishery, which has increased from 60 kg/trip in 2006 to

open water seines, gil nets, fish traps, long lines and hand

more than 100 kg/trip after 2009. The increased catch rates

lines. By contrast, most of the catch from subsistence

in the chilimira net fishery may be due to increases in

fishing is consumed at the household level and, only if there

technology, the increased use of light attraction and the

is a surplus, is it sold (Banda et al., 2001). Species are under

inclusion of usipa in the fishery. At a regional-level, catch

threat from both smal and large-scale fisheries. The large-

rates of beach seine, gill net and/or longline and handline

scale fishery has a much greater impact where it takes

fisheries appear to be declining, while catch rates in the

place, being implicated in some possible extinctions (e.g.

chilimira net fishery have generally increased. On a broad

Oreochromis lidole; Konings, 2018), but it is more limited in

scale, this suggests that inshore and demersal fisheries,

spatial scope. Therefore, overall more species are recorded

which target more resident species, are in decline. During the

on the Red List as under threat from the smal -scale fishery

surveys, it was also observed that mbuna, the diverse

(67%) than the large-scale fishery (13%).

community of rock dwel ing cichlids that is synonymous with

the lake, appear to be directly targeted in the fishery. As this

In the last decade, annual catch landings in the LMNNC have

practise may not be sustainable and as these inshore mbuna

declined significantly, particularly in the Malawian section of

communities are highly restricted in their distribution, their

LMNN where fishing effort is concentrated in the southern

harvest may conflict with the biodiversity mandate of the

end (Weyl, Ribbink & Tweddle, 2010). An assessment based

Lake Niassa Reserve and needs to be incorporated in the

on 1976–2002 data from the Malawian section of the lake

monitoring system (Weyl et al., 2018).

demonstrated that, although total catch was relatively

stable, increasing fishing effort had decreased catch rates,

Fish stocks in the pelagic zone of LMNN were previously

depleted larger, more valuable species and led to species

reported to be unexploited or marginal y exploited because

changes, most notably in the large, and valuable, endemic

of the shortage of stable and powered boats and modern

Oreochromis “Nyasalapia” (chambo) species flock (Weyl,

fishing equipment (Menz, 1995). Weyl et al. (2010), however,

Ribbink & Tweddle, 2010). More recently, this decline has

stated that pelagic fishes may be exploited at higher levels

continued with the fishery in southern LMNN now being

than previously thought given Banda et al. (2002) reported

based largely on E. sardella, with catches of cichlids, and

that truly pelagic species contributed 36% to the total

particularly chambo, continuing to decline (Irvine, Etiegni &

catch. Weyl et al. (2010), therefore, stated that with a pelagic

Weyl, 2018).

yield already in excess of 28,000 tonnes per year the stock

cannot be considered unexploited. Thus, although there is

In the Lake Niassa Reserve in Mozambique, with its steep

a concentration of fishing effort in inshore shal ower waters,

shoreline and deep-water close inshore, the fishery is

which has resulted in extensive overfishing, the offshore

dominated by pelagic species, usipa and pelagic

pelagic stocks are also effectively exploited as the fish move

haplochromine cichlids, which together contribute in excess

extensively throughout the lake frequently placing them

of 80% to the total landings in the fishery (Weyl et al., 2018).

close to shore and, therefore, vulnerable to the heavy inshore

Inspection of annualised catch, effort and catch per unit

fishing pressure.

effort (CPUE) data between 2006 and 2015 indicated

relatively stable effort, and stable or increasing trends in

The use of fishing weirs in rivers flowing into LMNN can

be detrimental to fisheries, although there is scope for

their use in some circumstances. Weirs with basket traps

inserted in gaps to catch fish passing through have been

used for millennia with no ill effects on fish stocks, even

when they appear to be complete barriers to fish migration.

The situation is, however, changing with rising human

populations leading to more intense fishing pressure. In

addition, the use of mosquito netting for traps, instead of

branches or reeds, prevents juvenile fishes from passing

through and returning to the lake. The change in character

of the rivers from permanent flow to flash flood regimes, as a

result of deforestation, also means the weirs can be used for

longer periods as the floods recede earlier. In the Bua River,

Figure 4.15 The Shire River in Malawi is rich in freshwater fish

species, including threatened and endemic species. © Denis

the permanent concrete weir, currently in use to divert the

Tweddle

flow into irrigation canals, prevents fish fry and fingerlings

59
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of Data Deficient freshwater

fish species

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

4 - 35

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 4.16 Richness of Data Deficient (DD) freshwater fish species in
the Lake Malawi/Nyasa/Niassa Catchment based on spatial data coded
as Presence 1 (Extant). Richness data are classified using quantiles.
Mapping is to HydroBASINS and does not imply species occur across
the entire HydroBASIN (i.e. across the entirety of Lake
Malawi/Nyasa/Niassa). See Figure 4.12 for the distribution of species
within Lake Malawi/Nyasa/Niassa.

60

them extremely destructive. On the other hand, if used for

open water, such as chirimila nets using light attraction at

night, they are an appropriate and effective fishing gear for

usipa, E. sardella. A blanket ban on the use of mosquito

netting in LMNN would, therefore, be inadvisable. Any

prohibition should be targeted at the use of mosquito netting

used in shore seines.

4.4.2 Agriculture

The increasing demand for more farmland has caused

Figure 4.17 Otopharynx selenurus is assessed as Data Deficient

(DD) because a decline in the population is suspected based

deforestation of fragile environments, burning of vegetation,

on its capture for use as food but there are insufficient data

degradation of river catchments, destruction of wetlands

available to quantify this decline. © Ad Konings

on river banks for agricultural purposes and the cultivation

of marginal areas, which were previously untouched. All

from returning to the lake, becoming trapped in the canals

of these activities weaken the soil structure by removing

instead. The weir also restricts the river into channels that

the vegetation cover, and soil and its nutrients are washed
enable the use of fish traps to completely block the river and

directly into rivers by rains, ultimately arriving in the lake.

prevent any adults from ascending above the weir much

The introduction of heavy sediment loads into rivers and

earlier in the year than would otherwise be possible (see

other large water bodies causes increased water turbidity

Sites in the spotlight: Mpasa and the Lower Bua and

and direct smothering of river beds and lakeshore habitats,

Nkhotakota Wildlife Reserve Key Biodiversity Areas

which ultimately eliminates food organisms (algae in

p. 160 in Chapter 10). Steps, therefore, need to be taken to

particular for many mbuna species), and smothers breeding

ensure gaps are left in weirs to al ow adult spawning fish to

sites, eggs and larvae (Skelton, 2001). This leads to declining

ascend, and juveniles to descend to the lake.

fish populations and impacts human communities within the

basin through reduced catches leading to loss of income

Other fishing gears can also be used destructively. For

and malnutrition. Pollution from agricultural and forestry

instance, most seine nets in use in the lake are now made of,
effluents is recorded as a threat to 29% of freshwater fish

or lined with, mosquito netting (Figure 4.19) and, therefore,

species native to the LMNNC. Fish species that are likely

catch everything from the size of fish eggs upwards, making

to be disadvantaged by these practices include those that

Figure 4.18 Fishing canoes and nets at Nkhata Bay in Malawi. ©

joepyrek (CC BY-SA 2.0)

61
migrate up rivers to breed in habitats now degraded through

can completely cover large expanses of water surfaces

sedimentation, pol ution and loss of riparian shading, such

and in the process smother under water habitats through

as the cyprinids including species belonging to the genera

reduction of both light and oxygen (Skelton, 2001). Both

Labeo, Labeobarbus and Opsaridium. Labeo mesops has

these species are present in the LMNNC and their spread

completely disappeared in most rivers in the catchment

needs to be monitored and controlled. Elsewhere within

because of degraded rivers and streams (Skelton et al., 1991)

southern Africa, invasive predatory fishes like bass

(see Species in the spotlight: Liwonde National Park – a

( Micropterus spp.) and rainbow trout ( Oncorhynchus mykiss)

fish sanctuary for the ntchila, Labeo mesops p. 67). For

have wiped out entire populations of certain Enteromius

species with narrow distribution ranges, such as some of the

species and mountain catlets (Ellender & Weyl, 2014).

rock-dwel ing cichlids (mbuna), an entire rock habitat can be

Similarly, in East Africa, introduction of the Nile perch ( Lates

lost to siltation (Konings, 2016). Siltation of mountain streams

niloticus) has contributed to severe declines in the endemic

may also represent a serious threat to some species, as

haplochromine cichlid flock of Lake Victoria (Sayer, Máiz-

recorded in some parts of southern Africa where it has had

deleterious effects on distributions of endemic species of

cyprinids and catfishes, especial y for those that prefer clear

flowing streams (Skelton, 2001; Skelton et al., 1991).

4.4.3 Invasive species

Invasive alien plant and animal species represent a serious

threat to freshwater species in the LMNNC. Invasive

species are, however, currently recorded as a threat to

only 2% of native freshwater fish species. Despite this low

level of current threat, it is extremely important to prevent

introduction of alien invasive species and to control those

already present in the catchment as once established

these species are extremely difficult to eradicate and

Figure 4.20 Non-native Nile tilapia ( Oreochromis niloticus) is

being farmed in Tanzanian rivers in the Lake Malawi/Nyasa/

their impacts can be extremely rapid and widespread. For

Niassa Catchment and presents a future threat to native

example, invasive plant species, such as water hyacinth

freshwater fish species. © Samuel Stacey via WorldFish (CC BY-

( Eichhornia crassipes) and Kariba weed ( Salvinia molesta),

NC-ND 2.0)

Figure 4.19 A fishing net at Lake Malawi/Nyasa/Niassa. © Benoît Rivard

(CC BY-NC-ND 2.0) 62

Tomé & Darwal , 2018). At present, there is no evidence that

these surveys have not been carried out in some parts of the
any LMNNC species have been negatively impacted by alien

lake, especial y in Nkhata Bay, due to insufficient human and

invasive species, but this situation is likely to change. Nile

financial resources. As such, it is strongly recommended that

tilapia ( Oreochromis niloticus; Figure 4.20) is being farmed

these data collection initiatives should be strengthened, in

in Tanzanian rivers in the catchment and will, therefore,

particular to provide information for the 52% of freshwater

inevitably escape and become established in the lake,

fishes native to the catchment for which increased monitoring

with potential adverse impacts on the native Oreochromis

was recommended.

species (Genner et al., 2013).

Where resources permit, the Malawi Department of Fisheries

4.4.4 Climate change

also carries out detailed biological studies of fishes landed

by fishers in various parts of the country. Information is

With the advent of climate change, occurrences of

gathered on gear selectivity, breeding biology, species

destructive weather phenomena like floods (Figure 4.21),

diversity and composition. These surveys have mostly been

drought, abnormal thunderstorms and strong winds

conducted in southern LMNN and Lake Malombe and more

(hurricanes or typhoons) are likely to increase. All these

work is needed in northern areas of the lake.

pose future threats to the freshwater biodiversity of the

LMNNC but are currently coded as threats to only 1%

The Malawi Department of Fisheries also occasionally

of freshwater fishes. For example, when lake levels drop

carries out fish biomass monitoring programmes using the

beyond their usual range shoreline rocky habitats may

research vessel R.V. Ndunduma in areas where the bottom

become unavailable to some of the restricted range mbuna

topography permits trawling. In some areas, particularly in

species. Stronger winds may also lead to more frequent

the northern parts of the lake, uneven and rocky bottoms

upwelling of deep waters in the lake, potentially leading to

make bottom trawling impractical, and as a result, the

fish kills as deoxygenated waters rise to the shallow water

demersal assemblage of fishes in this part of the lake is not

habitats of many fish species. Stratification of the lake

wel known. Both pelagic and demersal surveys need to be

may also be weakened through deep-water warming by

undertaken periodically to quantify and ascertain status

sub-littoral water supply during increasing milder winters

of fish stocks in the bottom and pelagic environments of

(Vol mer et al., 2005).

the lake. Where fishing methods are impractical, such as in

rocky areas, the use of video surveys to assess the state of

the stocks is a valuable technique that is becoming more

4.5 Recommended research and

frequently adopted, as currently in Mozambique where

conservation actions

baited remote underwater video was recently used to

provide baseline data on species richness and abundance in

In Malawi, the Department of Fisheries, through its monthly

key monitoring sites of the Lake Niassa Reserve (Weyl et al.,

catch assessment surveys (CAS), collects information on

2018; van Wyk et al., 2018).

catch, effort and beach price of fresh fish from all capture
fisheries in major rivers and al lakes. This is a form of fishery

In Mozambique, the National Institute of Fisheries Research

monitoring programme, the main aim of which is to provide

( Instituto Nacional de Investigação Pesqueira ; IIP)

insight into the performance of the fisheries sector. However,

implements a functional catch assessment system which

has col ected catch and effort data on the fisheries of LMNN

since 2006. Data are stored in a Microsoft Access database

and are retrievable on a habitat (rocky shore, beach and river

mouth) basis. Improvements to the system were suggested

by Weyl et al. (2018) and the continuation of the time series

is essential.

In Tanzania, a weekly CAS is also implemented in LMNN by

the Tanzania Fisheries Research Institute (TAFIRI) with the

primary aim of collecting data on fish composition, effort,

landing catch and price. Other studies are conducted on a

monthly to yearly basis, for example those on fish

parasitology or cichlids in the crater lakes. The use of native

species for cage aquaculture in the LMNNC is now

emphasised in Tanzania given the dangers of using non-

Figure 4.21 Flooding in Malawi in 2015. © George Ntonya UNDP

(CC BY-NC-ND 2.0)

native species.

63

Improved coordination between the three countries sharing

highlight the urgent need for conservation strategies that

LMNN fisheries resources is needed, particularly with regard

include effort limitation and elimination of environmentally

to catch assessment, and fishing gear and vessel counts.

destructive fishing methods in these lakes.

This will allow for more comprehensive assessment of the

status of the fisheries and for the publication of annual

Another conservation strategy is the identification and

reports on the overal status of the lake fisheries as a whole.

establishment of further protected areas in addition to the

Lake Malawi National Park, as suggested by Chafota et

In addition to increased monitoring of fish stocks, it is

al. (2005) and developed through the identification of Key

essential to develop new approaches to regulate fisheries

Biodiversity Areas (KBAs) for freshwater biodiversity in this

and enforce those regulations. The monitoring that has been

study (see Chapter 10).

conducted over the last decades has shown a clear decline

in the fisheries with a possible loss of some species due to

It is recommended that a system for monitoring the col ection

excessive fishing pressure and destructive fishing gears.

and export of fishes for the ornamental trade be devised to

ensure there is no over-col ection, in particular for many of

The severe impact of excessive fishing pressure in the lake

the highly restricted range species that are highly popular in

needs to be addressed. Various attempts have been made

the aquarium trade (e.g. Figure 4.22).

in the past to control rising effort but without success. For

instance, the Malawi National Fisheries and Aquaculture

Restoration of forest habitat in al catchments of the LMNNC

Policy of 2001 provided strategies and guidelines for

is also required if the impacts from sedimentation, habitat

participatory fisheries management. This included a chambo

loss and pollution mentioned above are to be reduced. In

restoration policy, which aimed, through participatory

particular, the stream catchments of Lake Chilwa have been

management between the Fisheries Department and District

severely degraded, possibly leading to more frequent drying

Assemblies, to change the fishery from “open access” to

up of the lake. Catchment restoration and reforestation are

restricted and limited access. The aims were to protect

urgently needed for both lakes.

chambo breeding areas and promote enhancement through

community culture based fisheries. Alternative livelihood

Finally, there remains a critical need for further taxonomic

strategies were to be engaged for fishers to move away from

studies on al of the freshwater fish species in the LMNNC,

fishing. In the two decades since the strategy was approved

with as many as half of the known species still to be

by government, however, the chambo fishery has continued

described. Until they are formally described many species

to deteriorate to the point that chambo species are now

cannot be considered for the IUCN Red List and so no

considered to be CR. Declines in al species in the fisheries,

assessments of their relative extinction risk are available. It

apart from usipa, are evident in both LMNN and Lake

is likely that there are many more species in the ecosystem

Malombe (Tweddle et al., 2015; Weyl, Ribbink & Tweddle,

whose status is not wel known due to paucity of information

2010). Although no authoritative published data are available

and thus, further research is essential, as recommended

since the early 2000s, unpublished data suggest that the

for 27% of freshwater fishes native to the LMNNC. In Lake

declines continue. The continued declines and assessment

Chilwa the fishes are wel -known but some species are stil to

of important fisheries species in threatened categories

be formal y described.

Figure 4.22 A male Aulonocara baenschi in breeding colouration at

Nkhomo Reef in Malawi. This species is listed as Critically Endangered
(CR) based on overcollection for the ornamental fish trade. © Ad
Konings
64

Species in the spotlight

The one who sleeps – kalingono, Nimbochromis livingstonii

Konings, A.1

1 Cichlid Press, El Paso, Texas, USA

Nimbochromis livingstoni is common in the shal ow muddy

habitat of LMNN. Its characteristic behaviour is responsible

for its Chichewa name kaligono, meaning “the one who

sleeps”. Nimbochromis livingstonii is a piscivore that has

developed a remarkable ambush-hunting technique.

When not breeding, both males and females exhibit several

characteristic liver-coloured patches on a whitish body,

a colour pattern that is unique among LMNN cichlids.

The kaligono feeds on small, inexperienced cichlids but

never pursues its prey. It has instead ‘invented’ another

Figure 4.23 Nimbochromis livingstonii lying in the sediment

technique to obtain its daily meal, wherein its unique

and waiting for its prey at Gome in Malawi. © Ad Konings

colouration plays the key role. White-coloured objects,

resembling decaying fish, are very attractive to any cichlid and msima
(white, boiled corn flour) is widely used as bait by young fishers angling for
fish from the shore. However, although the white colour may attract smal
fishes, they would never approach within striking distance of the kaligono. It
has, therefore, developed several ‘procedures’ to prevent its prey from
recognising it as a piscivore. One of these is employed when the bottom is
covered with a few centimetres-thick layer of mud: the predator lies down on
its side and wriggles itself into the sediment, remaining in that position
without moving a fin.

If the bottom is sandy it may stir up some sand as it lies down on its side, but
most of the time it just drops on the substrate.

The result is that the outline of the fish is partial y camouflaged and while it
lies on one side, it is not directly recognised as a threat by smal cichlids
(Figure 4.23). N. livingstoni may lie motionless for several minutes before it
moves to another site. The sand and debris that it occasional y stirs up (as it
lies down) may attract all kinds of smal fishes, but often the predator just
waits until a smal cichlid inspects this very interesting white coloured ‘thing’
that is lying on the bottom.
The death-shamming predator does not appear to be recognised by them.
When the smal fishes come within striking distance, the kaligono pounces
upon them with a sideways stroke. Larger fishes are also sometimes
attracted, but in this case the sleeper avoids contact by ‘waking up’. Every
adult N. livingstoni has its own feeding territory of about 40 metres of
shoreline (McKaye, 1981). Neighbouring individuals contest territorial
boundaries with a short display, and then return to their own feeding
grounds. However, at some places around the lake up to 30 sub-adult
specimens have been observed within an area about 100 metres in diameter.
Adults, however, seem to have large private feeding areas.

Breeding males of N. livingstonii are a dark sky-blue, which completely

obscures the blotched pattern. Neighbouring individuals probably recognise
each other, and females in adjacent territories notice the sexual ripeness of
the male. It is not uncommon to find a few breeding males forming a

lek but such groups are not usually more than a handful

of males. Spawning usually takes place at the edge of

the rocky habitat, where the male wil have dug a shal ow

saucer-shaped spawning site beside a large rock. Both

male and female circle around each other and at some

point the female deposits a small batch of eggs. She then

moves forward to make room for the male, who fertilises

them while they are still on the spawning substrate. On

the next pass the female picks up the fertilised eggs in her

mouth and then deposits a new batch. Mouthbrooding

females normally remain solitary and guard their free-

swimming offspring for several weeks after they have

Figure 4.24 A female Nimbochromis livingstonii guarding fry

been released for the first time (Figure 4.24).

at Cobwé in Mozambique. © Ad Konings

Observations from A. Konings and documented in Konings (2016)

65

Volcanic evolution: a unique species in a crater lake – Chungruru

tilapia, Oreochromis

chungruruensis

Shechonge, A.1 and Palmer-Newton, A.F.2

1

Tanzania Fisheries Research Institute, P.O.BOX 98 Kyela, Mbeya, Tanzania

Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

The Chungruru tilapia ( Oreochromis (Nyasalapia) chungruruensis) (CR)

(Figure 4.25, Figure 4.26) is endemic to Tanzania and found only in the very
small oligotrophic crater Lake Kyungululu (Figure 4.27) in the Rungwe
district in the north of the LMNNC. This lake is a Key Biodiversity Area
(KBA) and Al iance for Zero Extinction (AZE) site recognised for its global
importance to freshwater fishes, including the Chungruru tilapia (see
Chapter 10 and the KBA Datasheets in Supplementary Material). This
isolated lake was formed through volcanic activity and fil ed with
groundwater. It is just over 400 m in diameter, 30 m deep and has very steep
sides with no shal ow productive area where plant life can grow. The waters
appear dark and turbid, and the bottom has fresh volcanic dirt and gravel. It
is, therefore, likely that plankton can survive only near the surface and when
they die they sink to the bottom, such that nutrients are not recycled to
maintain the fertility of the lake.

The Chungruru tilapia is smal bodied with a maximum size of 19 cm TL and

it is clearly distinct from al other species in the chambo group, with most
found to have large heads and thin bodies. It is thought that the low nutrient
levels in the lake have led to stunted body growth. Adults are observed to
feed in the middle of the lake, while juveniles are likely dependent on
benthic production in the shal ows (Turner, Shechonge & Tweddle, 2018).

The Chungruru tilapia has declined significantly, with only one adult found
in a survey in 2017 (Turner, Shechonge & Tweddle, 2018). The main threat
is from two introduced tilapia species, Tilapia sparrmani and Coptodon
rendal i, which were stocked in the lake. As fish populations are extremely
low and there are extremely limited resources, this unique native tilapia
species is likely to suffer from competition, especial y in the limited area of
shal ow water on which juveniles are heavily dependent. Nearby crater lakes
also hold other invasive species which, if introduced to Lake Kyungululu,
could also compete and hybridise, leading to even further declines (Turner,
Shechonge & Tweddle, 2018). Fishing is also a threat in the smal , enclosed
lake as, although it is unlikely to support a fishery due to its low population
size, there is no regulation. Activities at the lake margins present additional
threats as ongoing tree fel ing and cultivation is likely causing Figure 4. 25
A male Oreo-

chromis chungruruensis,

commonlyknownas

Chungurutilapia.This

speciesisassessedas

Critically Endangered (CR).

© Martin Genner

Figure 4.26 A female Oreo-

chromis chungruruensis,

commonlyknownas

Chungurutilapia.This

speciesisassessedas

Critically Endangered (CR).

© Martin Genner

66
siltation, especial y in the shal ow water habitat (Turner, Shechonge &
Tweddle, 2018). To protect this species, population monitoring is required
alongside invasive species control to prevent any more introductions and to
reduce populations of those already in the lake. Fishing should be prohibited
or restricted to subsistence levels without the use of nets, and the creation of
a forest reserve covering the lake catchment would ensure that the habitat is
not degraded (Turner, Shechonge

& Tweddle, 2018).

In addition to the Lake Kyungululu endemic, a number of phenotypical y

distinct taxa have been observed in other nearby crater lakes. Six of these
populations have been grouped as the O. “crater lake chambo”, and they bear
pigmentation patterns resembling species from the LMNN chambo group.
Additional research is needed to establish their taxonomic status, and it is
possible that more intensive sampling of these crater lakes wil find more
unique and threatened species (Shechonge et al., 2019).

Figure 4.27 Lake Kyungululu

is a small oligotrophic crater

lake in Tanzania home to

the endemic Oreochromis

chungruruensis. © Martin

Genner

Liwonde National Park – a fish sanctuary for the ntchila, Labeo mesops

Tweddle, D.1

1 South African Institute for Aquatic Biodiversity (SAIAB), Somerset

Street, Grahamstown, 6139, South Africa The ntchila, Labeo mesops (Figure
4.28), is endemic to the LMNNC and is listed as CR (Tweddle & Gobo,
2018). As such, conservation initiatives for this species are essential. Ntchila
was formerly a species of major importance in the LMNN and Lake
Malombe fisheries, particularly in the south-east arm of the lake. As a result,
it was the subject of fisheries research in the early 1960s (Anon, 1964, 1965;
Iles, 1962) when it was suggested that catches of ntchila could be increased
tenfold without detriment to the stocks (Iles, 1962). This prediction was
based on its known high fecundity and the assumption that fishing mortality
was lower than natural mortality over the first breeding period. In the GFTC
annual report for 1960, Figure 4.28 Labeo mesops,

commonlyknownas

ntchila, is endemic to the

Lake Malawi/Nyasa/Niassa

Catchment and assessed as

Critically Endangered (CR).

© Denis Tweddle

67

however, declining catches of ntchila were already being

noted in spite of increased fishing effort (Anon, 1961;

Skelton et al., 1991). By the mid-1960s the fishery had

effectively collapsed. The species continued to exist in

smal numbers in the Malawian waters of the lake but has

not been seen in recent decades. In the Mozambican

waters of the lake with, until recently, a much smaller

human population, ntchila continued to exist in good

numbers but catches here are now also in steep decline as

the lakeshore becomes more heavily populated (Tweddle &

Gobo, 2018).

The assumptions of Iles (1962) appeared reasonable. The

fish is highly fecund and, even with high fishing pressure,

it should have continued to thrive and contribute to the

Figure 4.29 Liwonde National Park in Malawi plays a major

fishery. Thus, other factors are in play. Unlike most other

role in the conservation of Labeo mesops. © Denis Tweddle

migratory cyprinids in the LMNNC that are fractional

spawners (i.e. ripening and releasing batches of eggs over a period of

weeks), ntchila is a total spawner, releasing all its eggs in one annual
spawning event (Msiska, 1990). It runs up temporary streams when in spate
to breed in the flooded grassy margins. The character of these streams
changed dramatical y with increased human populations and deforestation.
The risks of changes in stream character were noted as early as the 1940s by
Lowe (1952) who stated: “It seems certain that the extensive clearing on the
watershed of these rivers in the last fifty years has changed their character
from more or less permanent streams to streams of very sudden rise and fal
”. The streams are now flash flood streams, rather than running for several
weeks at a time as previously. Ntchila eggs are consequently either
smothered in silt or left high and dry, and with only one annual spawning
event, the likelihood of spawning failure is very high. The loss of suitable
spawning habitat is considered to be a major reason for col apse of the
fishery (Skelton et al., 1991) and this is compounded by overexploitation.
Basket traps, fish weirs, seine nets and gil nets were al used to catch this
fish. The factors that caused this decline appear insurmountable in Malawi
because of the high and stil increasing human population that has caused
apparently irreversible degradation of the spawning streams along the
lakeshore.
There is, however, a beacon of hope for this species. While the species has
effectively vanished from the Malawian waters of LMNN and from Lake
Malombe, a population continues to survive, indeed thrive, in the Upper
Shire River, extending from the outlet from Lake Malombe down to the
Liwonde Barrage (Tweddle, 2018a), with smaller numbers found
downstream in the Shire River as far as Matope.

In gil net surveys in the Liwonde National Park (Figure 4.29) from 2016 to
2018, ntchila was the most important species by weight, comprising 27% of
the catch in the latest 2018 survey. While the protection provided against
fishing is a large contributory factor to its abundance, the protection of
spawning grounds within the park is thought to be the main benefit. In
contrast to the rest of the ntchila’s range, Liwonde National Park protects the
riparian zone and particularly the catchments of numerous smal streams and
swamps where ntchila can successful y breed. As a result, the river supports
a healthy stock. Ntchila breed at three years old (Anon, 1964), and in the
2018 survey al year-classes were wel -represented, showing that they were
able to breed successful y every year.

Liwonde National Park thus plays a major role in the conservation of this
CR species and it is recommended that ntchila is made a flagship species for
the park in order to emphasise the critical importance of protecting the Shire
River between the outlet of Lake Malombe and the southern tip of the park
near Liwonde. Liwonde National Park is also now recognised as a Key
Biodiversity Area (KBA) for this species (see Chapter 10 and the KBA
Datasheets in Supplementary Material).

This story of the ntchila demonstrates the great potential benefit that
protected areas, such as Liwonde National Park, can provide to the less
iconic freshwater species when rivers, lakes and their catchments are ful y
incorporated within the park boundary. Rivers are more than just useful
boundary markers for protected areas and need to become the focus of
targeted protection.

68

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72

Chapter 5

The status and distribution of freshwater

mol uscs in the Lake Malawi/Nyasa/

Niassa Catchment

Van Damme, D.1 and Albrecht, C.2

Contents

5.1 Introduction
...........................................................................................................................
...........................................................................73

5.2 Red List assessments

...........................................................................................................................
...........................................................75

5.3 Patterns of species richness

...........................................................................................................................
.................................................77

5.3.1 Overal species richness

...........................................................................................................................
...............................................77

5.3.2 Endemic species richness

...........................................................................................................................
...........................................81

5.3.3 Threatened species richness

...........................................................................................................................
...................................... 84

5.4 Major threats

...........................................................................................................................
........................................................................ 84

5.4.1 Pol ution

...........................................................................................................................
....................................................................... 84
5.4.2 Fisheries
...........................................................................................................................
...................................................................... 84

5.4.3 Invasive species

...........................................................................................................................
.......................................................... 84

5.4.4 Climate change

...........................................................................................................................
........................................................... 85

5.5 Recommended research and conservation actions

...........................................................................................................................
............. 85

5.5.1 Conservation actions recommended

...........................................................................................................................
.......................... 85

5.5.2 Research actions recommended

...........................................................................................................................
................................ 85

Species in the spotlight

...........................................................................................................................
............................................................... 86

Melanoides
species...............................................................................................................
.......................................................................... 86

Bibliography
...........................................................................................................................
.................................................................................87

5.1 Introduction

and, more than any other aquatic invertebrates, were

considered valuable expedition material.

On October 15th 1859, Dr David Livingstone wrote to the Earl

of Malmesbury: “My Lord, I have the honour to convey the

The original agreement was that the molluscs collected

information that we have traced the river Shire up to its point

from LMNN during the second Zambesi Expedition would

of departure from the hitherto undiscovered Lake Nyienyesi

be handled by the German malacologist Heinrich Dohrn.

or Nyassa”. Having thrown an explorator’s look at the lake,

However, John Kirk sent his najad bivalve material instead

Livingstone promptly returned to his prime mission,

to Richard Owen, superintendent of the British Museum

navigating the Zambesi River. He returned a few years later,

of Natural History, who passed them on to the American

appointed head of the second Zambesi Expedition,

malacologist Isaac Lea in Philadelphia (Lea, 1864). Kirk’s

accompanied by a group of scientists, and traversed Lake

other molluscs were, after an appropriate delay, sent to

Malawi/Nyasa/Niassa (hereafter LMNN; Figure 5.1) from

Dohrn in Berlin (Dohrn, 1865) (Figure 5.2).

south to north on a small sailboat.

Dohrn was far from happy with this unexpected arrange-

John Kirk, the expedition’s official physician/botanist, was

ment, as is evident from his comment: “I regret very much

the first to col ect mol uscs from the lake in 1861. The precise

that there are no Unionidae in the collection which I got for

origin of these shells is unknown, but they were collected

examination. All I can state from the above list [of species]

along the western shore somewhere between Cape Maclear

is, that the conchological fauna of Lake Nyassa seems

and Livingstonia in Malawi. Mol uscs, being non-perishable,

to belong to the same region with Natal; but most of

were beloved col ection objects for 19th century naturalists

the freshwater species from the lake having turned out

1 Laboratory of Palaeontology, Department of Geology, University of Ghent,

Ghent, Belgium

2 Aquatic Biodiversity and Biogeography, Department of Animal Ecology,

Justus Liebig University Giessen, Giessen, Germany 73
Figure 5.1 Sunrise over Lake

Malawi/Nyasa/Niassa.

© David Barton (CC BY-NC 2.0)

to be hitherto unknown, and some of the other species

LMNN’s malacofauna virtual y ground to a halt between 1890

having been found by Captain Speke and others far more

and 1960.
northwards, it is rather diffcult to come to any conclusion

from the present collection” (Dritsas, 2005).

It is only during the second half of the 20th century that

the interest in LMNN’s malacofauna and in the connection

When a few years later, descriptions of some of the spec-

between Lake Tanganyika and LMNN was revived by the

tacular sea-like shells of nearby Lake Tanganyika became

discovery, in the deeper waters, of an exceptional y large and

public (Woodward, 1859), speculations about a former faunal

strange viviparid. This species, described as Neothauma

connection between the two similar looking deep lakes and

ecclesi by Crowley & Pain (Crowley, Pain & Woodward,

the possible existence of equally deviating shells in LMNN

1964), was considered to belong to a genus that was only

ran rife. This theory appeared confirmed when a few decades

known from fossils and from Lake Tanganyika.

later the French malacologist Jules René Bourguignat

published a paper on four new marine-like thiarid genera (30

Subsequently, scientists realised that their knowledge of the

species) discovered in the lake (Bourguignat, 1889) (see

LMNN malacofauna was appallingly poor, being limited to a

Species in the spotlight: Melanoides species p. 86).

few handfuls of empty shel s col ected along the lakeshore.

The World Health Organisation (WHO) Snail Identification

A belief that exploration of the vast depths of LMNN would

Centre in Denmark, therefore, set up an exploration with the

bring to light additional strange shells, as predicted by

purpose of collecting live molluscs and dredging for

Bourguignat, became widespread. However, it remained

specimens in deeper waters. It was C.C. Cridland, well

wishful thinking for nearly a century, as further exploration of

known for his studies on Schistosoma, who set off to

Monkey Bay in 1969 and with the aid of J.C. Eccles, Head of

the Fisheries Research Station there, succeeded in dredging

the deeper areas in the southern part of the lake, a feat not

since repeated.

Cridland not only succeeded in recovering new specimens

of Neothauma ecclesi (subsequently placed within the

genus Bellamya) but also found a second deep water form,

Lanistes nasutus. The material, constituting the first

systematic collection of molluscs in LMNN, including the

soft parts, was studied by Georg Mandahl-Barth (1972).

Since then, no species new to science have been discovered

and it is considered unlikely that this will happen. This does

not mean that research on the molluscs stopped. On the

Figure 5.2 Shells collected from the shores of Lake Malawi/

contrary, the molluscs of LMNN were the first African

Nyasa/Niassa north of Mangochi and described as Lanistes

nyassanus by Dohrn in 1865. © Naturalis Biodiversity Center

molluscs to be ecologically studied using scuba (Louda et

Wikimedia Commons (CC0)

al., 1983, 1984).

74

Major new thinking on the LMNN endemic molluscs came a

B. jeffreysi and B. rober tsoni ( Figure 5.4), and one

few decades later when, fol owing molecular investigations

ampullariid, the endemic Lanistes nasutus. The two deep-

on the endemic species flock of Melanoides (Eldblom

water species B. ecclesi and L. nasutus were the last

& Kristensen, 2003; Genner et al., 2007; Von Gersdorff

endemic species discovered (in the early 1970s) and are only

Sørensen, Sørensen & Kristensen, 2005), our understanding

known from a few sites in the relatively shal ow southern end

of taxonomy and species of the Great Lakes took a dramatic

of the lake. The lack of additional records, despite renewed

turn. The 19th century notion that these molluscs had

survey efforts, suggests that the deeper water populations

evolved into species flocks appeared to be erroneous.

were rare and localised to start with.

Investigation of the endemic Melanoides clade of LMNN

showed that, while they had quite differently shaped and

The rarefication of the viviparids B. robertsoni and B. jeffreysi

ornamented shells, genetically they had not divided and

seems to be relatively recent starting during the last decades

hence still remained a single species (see Species in the

of the 20th century. While B. robertsoni was never abundant,

spotlight: Melanoides species p. 86).

it was still quite widespread in the southern part in the late

1990s, while B. jeffreysi was also relatively common. Since

This low genetic diversity is not surprising because, although

then, the depth ranges of these species, as wel as their areas

modern day LMNN is not as young as Lake Victoria, it is not

of extent, appear to have contracted significantly.

especially old having originated around 60,000 years ago,

but with probable significant lake level fluctuations over that

Endangered (EN) species

time. Paleontological evidence for a freshwater malacofauna

Only one species, the endemic pulmonate Bulinus

is less than 10,000 years old, and the whole southern half of

succinoides, is Endangered (EN) (Table 5.1). This small,

the present lake was probably not yet flooded at that time

fragile species is restricted to patches of aquatic vegetation

(Van Damme & Gautier, 2013).

along the littoral zone where the habitat is heavily impacted

by pollution and weed removal. A fur ther decline in

In summary, the modern scientific view on the taxonomy

abundance is expected.

of the LMNN mol uscs is one cause for the apparent recent

decrease in species richness of this fauna. The second

Vulnerable (VU) species

cause is the significant negative impact of human activities,

Two species, the large endemic Lanistes nyassanus and the

which primarily affects molluscs in the shallow sublittoral

endemic Gabbiella stanleyi, are Vulnerable (VU) (Table 5.1).

and deep-water zones. This chapter is the first to attempt

The large heavy-shelled Lanistes nyassanus was formerly

to evaluate the status of the LMNN mollusc fauna, taking

a common sight in shallow sandy sublittoral areas and

into account recent views on taxonomy, as well as human

became popular amongst aquarists keeping LMNN cichlids.

impacts.

Around the turn of the century, however, large piles of

dying snails were seen on the southern beaches, a bycatch

discarded by fishers. Since then, the populations of Lanistes

5.2 Red List assessments

nyassanus have not recovered and are declining further due

to increased seine net fishing and pollution. In the 1990s the

There are 38 species of freshwater molluscs native to the

small Gabbiella stanleyi was still common, in particular in

Lake Malawi/Nyasa/Niassa Catchment (hereafter LMNNC),

areas with aquatic vegetation. The severe reduction in this

of which 12 are endemic to the catchment (Table 5.1). These

endemic species face a high relative risk of extinction

Table 5.1 Number of freshwater mollusc species native and

with 58% assessed as threatened (Figure 5.3, Table 5.1).

endemic to the Lake Malawi/Nyasa/Niassa Catchment in each

Red List Category. For a list of species native to the Lake Malawi/

The remaining approximately 70% of native molluscs are

Nyasa/Niassa Catchment and their Red List Categories please see

widespread African species or species characteristic of the

Appendix 1.

Zambezi Drainage. Overall, 19% (excluding Data Deficient

Number of

Number of

(DD) species) of freshwater mollusc species native to the

species native

species endemic

IUCN Red List Category

to the LMNNC

to the LMNNC
LMNNC are assessed as threatened (Figure 5.3, Table 5.1).

Extinct (EX)

This is slightly lower than for the Lake Victoria Basin and

Extinct in the Wild (EW)

continental Africa, where 26% and 29% of species (excluding

Critically Endangered (CR)

DD species) are considered threatened, respectively (Darwall

Endangered (EN)

et al., 2011; Sayer, Máiz-Tomé & Darwal , 2018).

Vulnerable (VU)

2
Near Threatened (NT)

Critically Endangered (CR) species

Least Concern (LC)

29

Four species are Critically Endangered (CR) (Table 5.1),

Data Deficient (DD)

namely the three endemic viviparids, Bellamya ecclesi,

Total

38

12

75
vegetated habitat, combined with pollution, is the most likely

species was assessed as LC in 2011 on the basis of its

causes of its subsequent population decline.

widespread distribution across Africa, which excluded the

LMNNC at the time of assessment (Van Damme, 2015). It

Near Threatened (NT) species

is included as a LC species in this chapter on the basis that

The fourth endemic Lanistes species, L. solidus, is the only

this range extension would not change its Red List category.

species assessed as Near Threatened (NT) (Table 5.1). Since

LC species also include all morphotypes of Melanoides,

this species is most commonly observed in weed beds,

formerly considered as distinct endemic species but

the decline of this vegetated habitat and the species are

currently as strains of the Melanoides polymorpha complex

probably correlated.

(see Species in the spotlight: Melanoides species p. 86),

some of which, for example M. magnifica, were previously

Least Concern (LC) species

considered rare and localised. The endemic but widespread

The majority of species (76%) in the LMNNC are assessed

unionid bivalve Nyassunio nyassaensis (Figure 5.5) and the

as Least Concern (LC) (Figure 5.3, Table 5.1), a higher figure

iridinid bivalves Chambardia nyassensis and Mutela alata,

than for the Lake Victoria Basin (60%) (Sayer, Máiz-Tomé

both formerly considered as endemic but which now appear

& Darwall, 2018) and much higher than for continental

to have a wider distribution (Graf & Cummings, 2007, 2018),

Africa (34%) (Darwall et al., 2011). It should be noted that

are also LC. Finally, the LC species include all remaining

one species native to the LMNNC previously assessed

molluscs with ranges either covering much of the Zambezi

as LC, Biomphalaria pfeifferi, was not reassessed here as

drainage or which are widespread throughout continental

new records from the catchment (Alharbi et al., 2019) were

Africa.

published after the assessment work was completed. This

Data Deficient (DD) species

There is only one DD species (Table 5.1), the widespread

Endemic species

Sphaerium bequaerti. The reported distribution of this

species includes Zambia, Tanzania, Burundi, Central

African Republic and Ghana, in addition to the LMNNC.

Native species

Due to incomplete information it was not, however, possible

to map its distribution fully so it is not included in Section

5.3 Patterns of species richness below. The percentage

0%

20%

40%

60%

80%

100%

of DD species in the LMNNC (3%; Figure 5.3, Table 5.1) is

significantly lower than previously recorded for continental

EX

EW

CR

EN

VU

NT

LC

DD

Africa (30%) (Darwall et al., 2011) but is comparable to the

Lake Victoria Basin (6%) (Sayer, Máiz-Tomé & Darwall,

Figure 5.3 Percentage (%) of freshwater mollusc species native

and endemic to the Lake Malawi/Nyasa/Niassa Catchment

2018). This suggests levels of knowledge on freshwater

in each Red List Category. For a list of species native to the

molluscs are higher in these Great Lakes than across the

Lake Malawi/Nyasa/Niassa Catchment and their Red List

continent.

Categories please see Appendix 1.

Figure 5.5 Shells of Nyassunio nyassaensis collected from

Figure 5.4 Bellamya robertsoni is endemic to Lake Malawi/

Domwe Island in Malawi. This species is assessed as Least

Nyasa/Niassa and is assessed as Critically Endangered (CR).

Concern (LC). © Naturalis Biodiversity Center Wikimedia Commons

© Thies Geertz

(CC0)

76
5.3 Patterns of species richness

of Mutela alata, Chambardia nyassaensis and Coelatura

hypsiprymna.

5.3.1 Overall species richness

According to the sparse available information on the current

Areas high in species richness of freshwater molluscs

malacofauna in LMNN, there are currently probably no

include: i) LMNN itself; ii) the near-shore swampy region

more than 20 species present. Individual molluscs are still

at the southern end of the lake, including the Upper Shire,

present in reasonable numbers but only a few opportunistic

the Lilongwe River with the adjacent swamps and Lake

species are abundant, namely Lanistes ovum, Melanoides

Malombe; i i) Chia Lagoon (Figure 5.6), a satel ite lake at the

tuberculata (although possibly the invasive Asian form;

central western edge of LMNN; and iv) the upper part of the

Genner et al., 2004), Bel amya capil ata, Bulinus tropicus and

Lufira and northern Rukuru river basins in the north-western

Corbicula africana (Kamtambe et al., 2019).

parts of the LMNNC (Karonga District) (Figure 5.7).

In the marshlands and swamps in the Lilongwe area and

The highest species richness, 34 species, is in LMNN

in Chia Lagoon a similarly impoverished malacofauna is

and includes a number of lake endemics. The maximum

expected. In the latter, Lanistes ellipticus and Coelatura

species richness outside of LMNN was formerly thought

mossambicensis were formerly present, but the situation

to be 22 species but this is now considered an optimistic

in this satellite lake has recently deteriorated and so it is

estimate. Regions with the highest species richness were

suspected these species may no longer be present.

the Upper Shire between LMNN and Lake Malombe, and

Lake Malombe. On the basis of our knowledge of the late

Additional species recorded from the marshes around Lake

20th century, the Upper Shire contained a malacofaunal

Malombe and LMNN include Radix natalensis, Biomphalaria

extension of LMNN and Lake Malombe contained a swamp-

angulosa, Bulinus forskalii, Pisidium pirothi, P. reticulatum

adapted form of the LMNN fauna, including Melanoides

and Sphaerium bequaerti. There are also records of

tuberculata, the type morph of the Melanoides polymorpha

Aspatharia subreniformis and Chambardia wahlbergi, but

complex (see Species in the Spotlight: Melanoides p. 86),

these date from the middle of the 20th century or even earlier

Lanistes ellipticus, L. ovum (Figure 5.8), a low-winged form

(Mandahl-Barth, 1972).

Figure 5.6 Chia Lagoon, a satellite lake in the Lake

Malawi/Nyasa/Niassa Catchment, is rich in freshwater molluscs. ©
Catherine Sayer

77
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of freshwater mollusc species

12 - 16
17 - 19

20

21

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

22 - 34

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 5.7 Richness of freshwater mollusc species in the Lake

Malawi/Nyasa/Niassa Catchment based on spatial data coded as
Presence 1 (Extant). Richness data are classified using quantiles.
Mapping is to HydroBASINS and does not imply species occur across
the entire HydroBASIN (i.e. across the entirety of Lake
Malawi/Nyasa/Niassa). See Figure 5.9 for the distribution of species
within Lake Malawi/Nyasa/Niassa.

78
Species richness in other parts of the catchment is largely

of the underwater slopes towards anoxic waters. The vertical

determined by current hydrology. The LMNNC is a very

distribution of the species in the lake is clearly limited by the

narrow basin with high and steep rift shoulders, situated in a

dissolved oxygen content of the water column with most

zone where seasonal rainfal strongly fluctuates. As a result,

species occurring in the oxygen-rich shal ow zone (0–15 m).

the upper parts of small, steeply descending rivers can

Only three gastropod species, of which two ( Lanistes

alternate between moist and arid conditions. The present

nasutus and Bellamya ecclesi) show adaptations (inflated,

dominance of dry shrublands and semi-desert habitats,

light shel s with large mouth apertures) to life on fluid muds

in particular in the highlands of the western rift shoulder,

in oxygen-poor depths, have been col ected in deep waters

suggests that development of water-rich environments is

(at depths of 50–90 m). They have, however, not been

not particularly favoured. Decreasing rainfal since the 1980s

recorded since the 1970s and may have become extinct due

has probably led to the dominance of such habitats. Aridity

to increasing anoxia at depths greater than 50 m, as based

and the alternation of long periods of drought with periods of

on circumstantial evidence (Van Bocxlaer et al., 2012). Only

intense floods have been further exacerbated by destruction

the bivalve Coelatura hypsiprymna has been found in deeper

of woodlands and intensifying use of water. Against this

waters (80 m) with all other bivalves apparently restricted to

backdrop of a changing hydrology, richness of species in

above 30 m depth (Figure 5.10).

the upper catchments of small river basins in north-western

Malawi fluctuates according to the severity and length of

Given the above, species richness distributions (Figure 5.9)

these dry periods. It is therefore possible that during formerly

fol ow the oxygenated depth zone above the 100 m contour.

wet decades species richness could have increased to

This is extremely narrow or absent in the northern deep half

include c. 20 of the more widespread common species. In

of the lake, where the underwater bottom quickly plunges

most other parts of the LMNNC, due to the steep relief and

to depths below 100 m along the coast. In the geological y

erratic rainfall, the diversity of freshwater molluscs is not

much younger southern basin of the lake, the south-east

expected to exceed 17 species (Figure 5.7), and is probably

and south-west arms are relatively shallow and it is at the

much lower, taking into account increasing human impacts

edges of this region that the specialised deep-water species

(see Section 5.4 Major threats below) .

Lanistes nasutus and Bellamya ecclesi have been found.

The littoral zone in this part of the lake harbours the most

In LMNN itself, species are primarily concentrated in a

diversified mollusc fauna, but it is also the area most heavily

shal ow coastal band (Figure 5.9) due to the steep descent

affected by human activities.

Figure 5.8 Lanistes ovum is a common and widespread species, and is

assessed as Least Concern (LC). © Katharina C.M. Heiler 79
¯

20 40

80

120
160Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

Freshwater mol usc distributions

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 5.9 Heat map showing relative richness of freshwater mollusc

species in Lake Malawi/Nyasa/Niassa based on spatial data coded as
Presence 1 (Extant).

80
LMNNC has only 10 endemic gastropods and two endemic

bivalves (Table 5.1), the majority of which occur in LMNN

(Figure 5.11). As already noted, the apparent recent decline

in endemic species richness is due to the molecular

investigation that compacted the eight or nine endemic

Melanoides species into a single M. polymorpha complex

(see Species in the spotlight: Melanoides species p. 86).

This reduced the total number of endemic gastropods in the

LMNNC from 17–18 to 10 species. Two of these remaining

endemics, the deep-water species Bellamya ecclesi

(recorded from Chipoka and from east of Monkey Bay only)

and Lanistes nasutus (recorded from north-east of Monkey

Bay and from north of Boadzula Island only; Figure 5.12) are

here assessed as CR and in decline due to increasing

Figure 5.10 A Coelatura species in its natural habitat in Lake

pol ution and climate change.

Malawi/Nyasa/Niassa. © Thies Geertz

While the majority of endemic gastropods are threatened,

the two endemic unionid bivalves are not. This situation is

5.3.2 Endemic species richness

similar in Lake Victoria where, in its present state of nutrient

enrichment, bivalves seem to be less affected when

In the LMNNC, endemic species richness is relatively low

compared with gastropods. In addition, bivalves live buried

when compared with that in other African rift val ey lakes of a

in the sediment and so may be less susceptible to being

similar size, such as Lakes Tanganyika and Victoria. The

swept up by fishing nets.

Figure 5.12 The view of Boadzulu Island from Nkopola in Malawi.

Lanistes nasutus is recorded from north of Boadzulu Island.
© Catherine Sayer

81
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of endemic freshwater mollusc

species

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

3 - 11

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 5.11 Richness of endemic freshwater mollusc species in the Lake

Malawi/Nyasa/Niassa Catchment based on spatial data coded as
Presence 1 (Extant). Richness data are classified using quantiles.
Mapping is to HydroBASINS and does not imply species occur across
the entire HydroBASIN (i.e. across the entirety of Lake
Malawi/Nyasa/Niassa). See Figure 5.9 for the distribution of species
within Lake Malawi/Nyasa/Niassa.

82
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of threatened freshwater

mollusc species

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

1-7
Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 5.13 Richness of threatened freshwater mollusc species in the

Lake Malawi/Nyasa/Niassa Catchment based on spatial data coded as
Presence 1 (Extant). Richness data are classified using quantiles.
Mapping is to HydroBASINS and does not imply species occur across
the entire HydroBASIN (i.e. across the entirety of Lake
Malawi/Nyasa/Niassa). See Figure 5.9 for the distribution of species
within Lake Malawi/Nyasa/Niassa.

83

5.3.3 Threatened species richness

where beach tourism is high and continues to be developed

(Ngochera et al., 2018) (Figure 5.14). Nearshore levels of E.

All threatened freshwater mollusc species of the LMNNC

coli contamination, for example, are higher than offshore and

are restricted to LMNN itself (Figure 5.13). For a detailed

related to settlement densities (Tyner et al., 2018). Given that it

discussion of these species, see Section 5.2 Red List

is the southern two arms of LMNN which harbour the highest

assessments above.

share of endemic and overall mollusc diversity (Figure 5.9)

this rising source of domestic pol ution is of concern. Metal

contaminants have also been identified in LMNN stemming

5.4 Major threats

from mining activities in Malawi (Kinnaird & Nex, 2016).

A number of threats to molluscs within the LMNNC have

5.4.2 Fisheries

been identified with 42% of species having specific coded

threats.

Fishing and harvesting of aquatic resources is the second

most frequently recorded threat to freshwater molluscs,

5.4.1 Pollution

affecting 24% of species. Unsustainable fisheries and

severe overfishing have been reported for LMNN (Bootsma

The most commonly recorded threat to native freshwater

& Hecky, 1993; Weyl, Ribbink & Tweddle, 2010) and Lakes

molluscs is pollution, specifically from agriculture and

Malombe and Chilwa (Jamu et al., 2011). This is proposed to

forestry effluents (34% of species) and domestic and urban

have caused significant changes in community compositions

waste water (24%). Pol ution is particularly severe in certain

with trophic cascades leading to general ecosystem

hotspot areas of the comparatively small basin (Otu et al.,

changes (Van Bocxlaer & Albrecht, 2015). For example, it

2011). Nutrient loads from atmospheric sources and surface

has been suggested that overfishing of some molluscivorous

runoff, particularly of nitrogen and phosphorous, have a

fish species has led to an increase in population sizes of

significant impact (Hampton et al., 2018). Logging and

intermediate host snails ( Bulinus) for schistosomiasis. It

intensified agriculture extending to the lakeshore are further

has been reported that due to this “prey release effect” and

exacerbating the situation through increasing sediment

the resulting increase in snail populations there has been a

loads and nutrient influx to the lake. Increased aquaculture

corresponding increase in the incidence of schistosomiasis

of tilapia fishes has also resulted in local increases in carbon,

around LMNN (Madsen & Stauffer, 2011).

nitrogen and phosphorous levels (Gondwe, Guildford &

Hecky, 2011).

5.4.3 Invasive species

Residential and commercial development also contribute to

Increased prevalence in schistosomiasis over the past few

increasing pol ution, especial y in the southern half of LMNN,

decades has also been associated with the arrival of a non-

Figure 5.14 Use of the beach

at Cape Maclear with result-

ing pollut ion. © Christian

Albrecht

84
native form of the gastropod Melanoides tuberculata from

Asia, which has significantly altered the native Melanoides

communities and also affected Bulinus abundances (Genner

et al., 2004; Van Bocxlaer, Albrecht & Stauffer, 2014; Van

Bocxlaer & Albrecht, 2015). There are other invasive species

in the vicinity of the lake, such as Physa acuta (Figure 5.15)

and another physid species, which has been provisionally

identified as Stenophysa marmorata (Clewing & Albrecht,

unpublished data). Invasive species are currently reported as

a threat to 3% of mollusc species.

5.4.4 Climate change

Shifts in the oxycline and depletion of the profundal as a

result of climate induced changes have been associated

with the observed decline in deep-water mollusc species

in the lake (Van Bocxlaer et al., 2012). Such decreases

in productivity and declines in biomass have also been

demonstrated as a result of increased water temperatures

in Lake Tanganyika (Cohen et al., 2016) and so are thought

Figure 5.15 Physa acuta is an invasive species already in the

likely to occur in LMNN too.

vicinity of Lake Malawi/Nyasa/Niassa that poses a potential

future threat to freshwater molluscs native to the lake. © Robert

Aguilar, Smithsonian Environmental Research Center (CC BY 2.0)

5.5 Recommended research and

conservation actions

the ecosystem services provided by aquatic habitats and

resources in the LMNNC. Poverty related issues should

5.5.1 Conservation actions recommended

also be addressed as part of a sustainable development

framework that includes radical changes in water usage

No specific conservation actions were noted as part of

and sanitisation practices if the incidence of bilharzia is to

the species Red List assessments. However, the following

be reduced and the balance in the mollusc fauna restored

general conservation measures are suggested as being

(Ngochera et al., 2018), although it is acknowledged that

relevant.

these wil need to be local and low maintenance systems for

this recommendation to be realised.

In order to reduce water pollution, nutrient loads in

rivers and in LMNN could be reduced by introduction of

5.5.2 Research actions recommended

waste-water treatment systems. More broadly, effective

water management plans should be developed on a

This study has revealed a significant lack of information on

transboundary basis, which might require formation of a

al aspects of freshwater molluscs in the LMNNC. In the Red

regional institution to coordinate policies and regulations

List assessments 76% of native species have recommended

for management of resources within the LMNNC. Efforts

research actions. The most common of these is for research

must also be stepped up to avoid introduction of non-

into the population sizes, distributions and trends of species

native species given their potential for significant impacts

(71% of species), fol owed by research (Figure 5.16) to better

to mol usc biodiversity and associated human health within

understand their life histories and ecology (66%) and threats

the LMNNC. Finally, sustainable forestry and agricultural

(47%). At present we lack sufficient information on the

practices, as well as land use management systems are

distribution of species to even confirm levels of endemism

needed in order to further reduce run-off of nutrients and

in the LMNNC. Taxonomic work, including genetic and

sediments. Forest restoration, particularly in the upper

genomic approaches, is needed to disentangle emerging,

catchments, is a priority if the hydrology is to be stabilised

evolutionary young, species, which seem to dominate the

such that sediment erosion and influx to the river and lake

recent fauna. We also need to better understand species

systems are reduced.

level differentiation of lacustrine and basin forms that have

been overlooked for a long time or could not be detected

Conservation actions are more likely to be successful if

formerly due to ecophenotypic plasticity, as shown for the

local communities are involved. This primarily requires

genera Lanistes and Bel amya (Schultheiß et al., 2009, 2011).

awareness of the value of the lake for human wel -being and

Satellite water bodies in the LMNNC and island habitats

85
Figure 5.16 Detailed malacological and parasitological studies

Figure 5.17 Mollusc surveys taking place on Lake Malawi/

taking place in the Lake Malawi/Nyasa/Niassa Catchment.

Nyasa/Niassa. © Stefan Schmid

© Stefan Schmid

within LMNN require much more malacological attention.

Population-level differences studies are needed that would

There also remain several cryptic species or even species

address seasonal dynamics. Approaches like that of Genner

complexes that require in-depth taxonomic analysis (see

& Michel (2003) should be used in various regions of the lake

Species in the spotlight: Melanoides species p. 86).

covering al major habitat types. They should be set up in a

comparative framework.

The apparent “creeping biodiversity crisis” facing endemic

molluscs of the LMNN over the last decades requires

Finally, we are far from understanding the impact to the

extensive large-scale biodiversity surveys (Figure 5.17)

native freshwater mollusc fauna of introduced species, such

in both horizontal and bathymetrical dimensions as our

as tilapia fishes or invasive gastropod species. Future studies

knowledge on species distributions and population trends is

should employ a holistic ecosystem approach to evaluate the

stil fragmentary in large portions of the LMNNC, specifical y

importance of such species and to model their potential

on the Tanzanian and Mozambican sides. The lack of recent

future impacts.

population information is particularly severe for deep water

species (Van Bocxlaer et al., 2012).

Species in the spotlight

Melanoides species

Van Damme, D.1

1 Laboratory of Palaeontology, Department of Geology, University of Ghent,

Ghent, Belgium

Study of the LMNN gastropods, more precisely of the flock of

endemic Melanoides species, considerably altered our views on the

taxonomy of freshwater molluscs and hence, had a significant impact

on the Red List assessment of species and on species conservation

general y.

The story began in the late 19th century when Jules René Bourguignat

published a paper on the Thiaridae (= Mélanidées) of LMNN (Figure

5.18), which he received from Victor Giraud. Giraud was an intrepid

French traveller and after months of being held captive near Lake

Bangweulu, succeeded in escaping, reached Lake Tanganyika, and

Figure 5.18 Plate I from Bourguignat’s 1889 paper

from there, after surviving a mutiny of his bearers, travel ed to LMNN

on the Thiaridae (= Mélanidées) of Lake Malawi/

and collected the thiarid shel s at Karonga beach (Bourguignat, 1889).

Nyasa/Niassa.

86

Bourguignat considered the thiarids of LMNN so strikingly different from

other African ones that he placed them in four genera new to science:
Nyassia, Nyassella, Nyassomelania and Micronyassia. The approximately 30
new species that he described were, according to him, less marine-like than
the shel s from Lake Tanganyika but they nevertheless had marine-like
characters and he believed that more ‘thalassoid’ shel s would be found
when LMNN’s depths were explored.

After Bourguignat’s death, however, changes in our understanding on the

importance of intraspecific variability in taxonomy, obliterated the old static
species concept and many former ‘species’ and even genera were
downgraded to ecophenotypic variations or ‘formas’ of a single, variable
species.

Two groups of biota more or less escaped this mid-20th century taxonomic
judgement, namely the species flocks on oceanic islands and those of the
large African Rift lakes, in particular of LMNN, Lake Tanganyika and Lake
Victoria with their cichlid fishes and freshwater mol uscs. These lakes were
said to form “a unique comparative series of natural laboratories for
evolutionary studies” (Coulter et al., 1986).

In his study on the LMNN mol uscs, Mandahl-Barth (1972), a fervent

“lumper” (favouring the combining of species into single taxa), rejected, as
many before him, the endemic thiarid genera created by Bourguignat but
retained the existence of an endemic Melanoides species flock, though
reduced to eight species. This still remained the largest group of endemic
freshwater molluscs in Africa. Considering the differences of the radulas
(the teeth structure of molluscs) of these endemic ‘species’ as being
diagnostical y important, Mandahl-Barth consolidated the arguments for the
existence of this endemic Melanoides flock. His view, including the
suggestion that there could be more endemic Melanoides species, was taken
forward by Brown (1994), on whose magistral work the previous IUCN Red
List assessments (Darwall et al., 2011) were based.

However, this taxonomic structure and the conviction that the large Rift
lakes were so uniquely important collapsed with the results of genetic
studies on the Melanoides endemics. These were very clear: all the
distinctive forms, then considered as discrete endemic species, contained the
same molecular signature (Eldblom & Kristensen, 2003; Genner et al., 2007;
Von Gersdorff Sørensen, Sørensen & Kristensen, 2005). The marked
differences between shell morphologies, hence, do not imply that a species
flock radiated in LMNN. Melanoides being a parthenogenetic taxon, it
simply meant that somewhere in the past, some individuals with a somewhat
different shell morphology, produced exact morphological copies of
themselves, in which these differences were maintained. However,
individuals still remained one and the same species, namely Melanoides
polymorpha, or the Melanoides polymorpha complex, i.e. a complex of
different morphotypes.

About 270 years after Carolus Linnaeus wrote his Systema Naturae,
demonstrating how animals could be discerned and classified by external
characteristics, the LMNN Melanoides proved that this was erroneous as, in
many instances, they cannot. Nature is much more complex than we thought.

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89
Chapter 6
The status and distribution of odonates

(dragonflies and damselflies) in the

Lake Malawi/Nyasa/Niassa Catchment

Palmer-Newton, A.F.1, Clausnitzer, V.2 and Dijkstra, K.-D.B.3

Contents

6.1 Introduction
...........................................................................................................................
.......................................................................... 90

6.2 Red List assessments

...........................................................................................................................
...........................................................91

6.3 Patterns of species richness

...........................................................................................................................
.................................................92

6.3.1 Overal species richness

...........................................................................................................................
...............................................92

6.3.2 Endemic species richness

...........................................................................................................................
.......................................... 96

6.3.3 Threatened species richness

...........................................................................................................................
...................................... 96
6.4 Major threats
...........................................................................................................................
........................................................................ 96

6.4.1 Habitat loss and degradation

...........................................................................................................................
....................................... 99

6.4.2 Pol ution

...........................................................................................................................
......................................................................100

6.4.3 Water management

...........................................................................................................................
....................................................100

6.5 Recommended research and conservation actions

...........................................................................................................................
............101

6.5.1 Research actions recommended

...........................................................................................................................
..............................101

6.5.2 Conservation actions recommended

...........................................................................................................................
.........................102

Species in the spotlight

...........................................................................................................................
.............................................................102

The threatened forest yel owwings

...........................................................................................................................
......................................102

The (near) endemics of the lake

...........................................................................................................................
..........................................104
Bibliography
...........................................................................................................................
...............................................................................106

6.1 Introduction

of removing sperm from a female’s previous mate, which is

why they often guard females during egg laying and fight

Odonates include the conspicuous, colourful and diurnal

furiously with other males.

“damselflies” (Zygoptera; e.g. Figure 6.1) and “dragonflies”

( A nisoptera ; e.g. Figure 6. 2 ) . Commonly the term

“dragonflies” is used to refer to al odonates, as here in this

report. They are often referred to as “flying jewels” because

of their beautiful colouration and agility in the air. Dragonflies

are the dinosaurs of the insect kingdom, pre-dating them

with ancestors emerging over 300 mil ion years ago. Giants

from the Carboniferous period had wingspans of 70 cm or

more, although 5–10 cm is typical of modern dragonflies.

Adult dragonflies are usually found along or close to water

bodies where females lay their eggs (Figure 6.3) and their

larvae hatch and develop, and on sunny days dragonflies

can easily be observed patrolling water sites or perching on

exposed sticks. Males may hold territories for several days or

Figure 6.1 Ischnura senegalensis, commonly known as tropical

bluetail, is a species of damselfly native to the Lake Malawi/

even weeks and display courtship behaviour such as flashing

Nyasa/Niassa Catchment. It is assessed as Least Concern

their bright colours when females arrive. Males are capable

(LC). © Jean-Pierre Boudot

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

2 Senckenberg Museum of Natural History Görlitz, Görlitz, Germany

3 Naturalis Biodiversity Center, Leiden, Netherlands

90
Figure 6.2 Anax ephippiger, commonly known as vagrant

Figure 6.3 Oviposition by a female Anax imperator, commonly

emperor, is a species of dragonfly native to the Lake Malawi/

known as blue emperor. This species is assessed as Least

Nyasa/Niassa Catchment. It is assessed as Least Concern

Concern (LC). © Christian Fischer (CC BY-SA 3.0)

(LC). © Jean-Pierre Boudot

Dragonflies are a good taxon to use in biodiversity studies

against the IUCN Red List Habitats Classification Scheme,

because they are easy to collect, sensitive to changing

over 70% of these native species occur along permanent

environmental conditions, and are comparatively well

rivers, streams and creeks. The most common terrestrial

studied taxonomically, ecologically and ethologically

habitat is moist shrubland (58%), closely fol owed by moist

(Clausnitzer, 2001). Due to their presence both below water

lowland forest (55%). Approximately 27% of species are

as larvae and above it as adults they are great indicators of

recorded in permanent freshwater lakes such as Lake

overall wetland health, acting as environmental sentinels

Malawi/Nyasa/Niassa (hereafter LMNN). Lake specialists

and as “whistle-blowers” for declining habitats. Studies of

are rare but include the Malawi hooktail ( Paragomphus

dragonfly biodiversity can be used to minimise or mitigate

nyasicus), a shal ow lakeshore breeder (see Species in the

impacts of future development in site conservation planning

spotlight: The (near) endemics of the lake p. 104). Only

across freshwater systems, while species level-assessments

8% of species are recorded in artificial habitats such as

can be used to monitor the possible impacts of growing

irrigated land, water storage areas and drainage channels.

threats such as climate change. Their study can help us

Savannahs support high numbers of dragonflies in East

to understand the past and future of rapidly changing

Africa, but these are mainly widespread and eurytopic

environments. Their attractive appearance makes them key

species (Clausnitzer, 2001). There are a large number of

candidates as flagships for wetland conservation.

these wide ranging species in the LMNNC, such as the blue

emperor ( Anax imperator), which occurs from South Africa to

Of the 773 dragonfly species native to continental Africa

most of Europe, the Arabian Peninsula and Asia in a very

that have been assessed for the IUCN Red List (updated

wide variety of natural and man-made aquatic habitats

from Clausnitzer et al., 2012) the majority are considered

(Mitra, 2016). The most threatened species are Afromontane

Least Concern (LC; 79%), while 71 are threatened: 2% are

forest specialists, such as the CR Ntchisi yellowwing

Critically Endangered (CR), 4% are Endangered (EN), and

( Al ocnemis maccleeryi) (see Species in the spotlight: The

3% are Vulnerable (VU) (IUCN, 2019). Taking into account

threatened forest yellowwings p. 102).

Data Deficient (DD) species, 10% of dragonflies assessed

within continental Africa are considered threatened.

6.2 Red List assessments

For the Lake Malawi/Nyasa/Niassa Catchment (hereafter the

LMNNC), 156 dragonfly species belonging to 10 families

The majority of the 156 dragonfly species recorded in

have been recorded. Of these, there are 55 damselfly

the LMNNC are assessed as LC (97%). Four species are

(Zygoptera) species within five families (Calopterygidae,

threatened (3%), and none are assessed as DD (Table

Chlorocyphidae,Coenagrionidae,Lestidae,

6.1, Figure 6.4). These Red List assessments indicate that

Platycnemididae) and 101 dragonflies (Anisoptera) from

dragonflies native to the LMNNC are, in general, at a lower

another set of five families (Aeshnidae, Corduliidae,

level of threat than the group across continental Africa

Gomphidae, Libellulidae, Macromi dae). According to coding

(updated from Clausnitzer et al., 2012) (Figure 6.4).

91

Table 6.1 Number of odonate (referred to throughout the chapter as

dragonflies) species native and endemic to the Lake Malawi/Nyasa/
Niassa Catchment and native to continental Africa in each Red List
Category. For a list of species native to the Lake Malawi/Nyasa/

Niassa Catchment and their Red List Categories please see Appendix 1.

IUCN Red List Category

Number of species native to

Number of species native to Number of species endemic to

continental Africa

the LMNNC

the LMNNC

Extinct (EX)

Extinct in the Wild (EW)

Critically Endangered (CR)

17

1
Endangered (EN)

28

Vulnerable (VU)

26

Near Threatened (NT)

33

Least Concern (LC)

608

151

Data Deficient (DD)

61

0
Total

773

156

levels of species richness can be found around the Songwe

LMNNC

River on the border of Malawi and Tanzania in the north of

the catchment. As wel as being rich in odonates, this area

Continental Africa

is the site of the Lower Songwe River Key Biodiversity Area

(KBA) as it is an important site for freshwater biodiversity at a

global level (see Chapter 10). This river enters north-western

LMNN through a deltaic system with extensive swampy

0%

10%

20%

30%

40%

50%

60%
70%

80%

90% 100%

areas covered in reeds and other plants. However, there are

also extensive areas of rice cultivation on both Malawian

EX

EW

CR

EN

VU

NT

LC

DD

and Tanzanian sides of the border. Upstream from the delta

Figure 6.4 Percentage (%) of odonate (referred to throughout

the river is rocky and fast flowing (See Songwe River KBA

the chapter as dragonflies) species native to the Lake Malawi/

Datasheet in Supplementary Material). Another area of

Nyasa/Niassa Catchment and to continental Africa in each Red

List Category. For a list of species native to the Lake Malawi/

high richness is the Northern Region of Malawi in the area

Nyasa/Niassa Catchment and their Red List Categories please

extending from the Nyika Plateau (Figure 6.6) in the north

see Appendix 1.

to the South Rukuru River in the south, including the city of

Mzuzu and Nkhata Bay in the east. This includes part of the

A large number of species found in the LMNNC are wide-

protected Nyika National Park and Vwaza Marsh Wildlife

spread open-land species found across sub-Saharan Africa

Reserve, as wel as a number of smal er forest reserves. The

that are tolerant to some degree of habitat degradation

Vwaza Marsh is an extensive wetland of reedbeds, patches

(Clausnitzer et al., 2012). Of the four threatened species,

of papyrus and seasonally flooded grassland, while Nyika

all but the Mulanje damsel (Oreocnemis phoenix) are

National Park includes grasslands, forest, dambos and

Afromontane forest stream specialists that can only be

miombo woodland. Nyika National Park is an important

found in remnant forest patches. This includes the only

catchment area that contains the source of four large rivers

CR and endemic species in the catchment, the Ntchisi

draining into LMNN, including the Rukuru, Chilinda, Rumphu

yel owwing ( A. maccleeryi) (see Species in the spotlight:

and Rynyina. A large number of species are also associated

The threatened forest yellowwings p. 102).

with the lake itself, mostly along the shoreline.

Some of the lowest levels of species richness are in

6.3 Patterns of species richness

the north-eastern corner of the catchment in southern

Tanzania (including the Ruhuhu and Kitwaka rivers), and the

6.3.1 Overall species richness

southern tip of the catchment in Malawi and Mozambique

encompassing all of Lake Chilwa (Figure 6.7). Lake Chilwa

The spatial pattern of species richness of dragonflies in the

is the second largest lake in Malawi with poorly studied

LMNNC is shown in Figure 6.5.

seasonally flooded grasslands that are declining from the

expansion of rice cultivation and increasingly extreme

The diversity of dragonflies in Africa is strongly correlated

droughts. The point locality data for dragonflies collected

with lotic forest habitats in a heterogenic landscape

within the LMNNC recorded in the Odonata Database of

(Clausnitzer et al., 2012). In the LMNNC some of the highest

Africa (ODA) (Figure 6.8) (Clausnitzer et al., 2012; Dijkstra,

92
¯

0 25 50

100

150

200Kilometers

Legend
Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of odonate species

22 - 116

117 - 121

122 - 125

126 - 129

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

130 - 135

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 6.5 Richness of odonate species (referred to throughout the

chapter as dragonflies) in the Lake Malawi/Nyasa/Niassa Catchment
based on spatial data coded as Presence 1 (Extant). Richness data are
classified using quantiles. Mapping is to HydroBASINS and does not
imply species occur across the entire HydroBASIN (i.e. across the
entirety of Lake Malawi/Nyasa/Niassa).

93
Figure 6.6 The Nyika Plateau is rich in odonate species (referred to
throughout the chapter as dragonflies). © Joachim Huber (CC

BY-SA 2.0)

Figure 6.7 Lake Chilwa has relatively low recorded richness of odonates
(referred to throughout the chapter as dragonflies).

However, this could be a result of poor sampling effort in its seasonally

flooded grasslands. © Gio la Gamb (CC BY-SA 3.0) 94
¯

0 25 50

100

150
!

Lake Malawi/Nyasa/Niassa Catchment

Odonate records from the Odonata

Database of Africa (ODA)

Records from 1990 onwards

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

Records from pre-1990

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 6.8 Distribution of odonate species (referred to throughout the

chapter as dragonflies) records in the Lake Malawi/Nyasa/

Niassa Catchment based on data from the Odonata Database of Africa

(ODA). Records dated from before 1990 are coloured purple and those
from 1990 onwards are coloured green. A single point on the figure may
contain multiple records from that location.

95

2016; addo.adu.org.za) show that these areas of low east of the basin in
Tanzania in the Mbinga region. Like other

diversity are at least in part due to col ection gaps as large

Allocnemis species, it is dependent on clear forest streams

areas of the catchment have not been extensively or recently

(Clausnitzer, 2010a).

sampled (Clausnitzer, 2001).

The VU eastern horntail ( Nepogomphoides stuhlmanni;

6.3.2 Endemic species richness

Figure 6.11) is confined to clear forest streams in the Eastern

Arc Mountains in Tanzania and adjacent southern mountains

There is only one endemic species recorded within the

in Malawi and Mozambique, mostly in headwaters but

LMNNC: the CR Ntchisi yellowwing ( A. maccleeryi). This

also larger shaded streams (Clausnitzer, 2010b; Dijkstra,

species is restricted to Mount Ntchisi in central Malawi

2019a). This species has a highly disjunct distribution in part

(Figure 6.9), a forest reserve that includes a small remnant

because of the island like situation of forested mountains in

patch of Afromontane forest surrounded by farmland. A

an otherwise dry matrix of Miombo and savannah vegetation.

detailed account of this species is given in Species in the

spotlight: The threatened forest yellowwings p. 102.

The clear, shaded streams of these forest specialists are

all under intense pressure from expanding agricultural

Aside from the Ntchisi yel owwing, a form of the wide ranging

activity. Much of this habitat has already been lost and

dancing jewel ( Platycypha caligata) cal ed the Malawi jewel

encroachment is ongoing (Clausnitzer, 2001). Their habitats

has only ever been recorded from the shores of LMNN. Its

are fragmented, isolated and most are unprotected from

status as a separate species is not confirmed. A detailed

further degradation and conversion. These specialists have

account of this form is given in Species in the spotlight:

a low dispersal capacity and are outcompeted by more

The (near) endemics of the lake p. 104.

generalist dragonflies when forests are opened up. To ensure

their continued survival their unique forest habitats need

This low level of endemism is in contrast to the ‘explosive’

protection and restoration (Dijkstra & Clausnitzer, 2006).

levels in aquatic groups, such as fishes, from the East

African Great Lakes. This paucity can be explained by

The EN Mulanje damsel ( Oreocnemis phoenix; Figure

adult dragonflies, whose capacity for flight inhibits genetic

6.12), known for its bright red colouration, is endemic to

isolation of lakeshore populations from surrounding habitats

Mount Mulanje in southern Malawi, but its distribution lies

(Dijkstra et al., 2011).

mostly outside of the LMNNC on the southern boundary

(Clausnitzer, 2018). Unlike other threatened LMNNC

6.3.3 Threatened species richness

dragonflies it can be found in both montane forest and

grassland streams, so is not as restricted in habitat (Dijkstra,

There are four threatened and one Near Threatened (NT)

2019). However, its population is just as isolated, having

dragonfly species within the LMNNC. These are the Ntchisi

only been recorded on the Mulanje plateau in a very small

yellowwing (CR), the blue-lipped yellowwing ( Allocnemis

area with no other suitable habitats found within several

montana; EN), the Mulanje damsel ( Oreocnemis phoenix;

hundred kilometres (Parr, 1983). Although Mount Mulanje is

EN), the eastern horntail ( Nepogomphoides stuhlmanni; VU)

a forest reserve, there have been cases of encroachment of

and the eastern yel owwing ( Allocnemis abbotti; NT). All but

agriculture and logging that directly threatens the Mulanje

the Mulanje damsel are restricted to shaded, clear

damsel’s habitat. Bauxite mining is also a threat and this

Afromontane forest streams scattered throughout the

would have devastating impacts on its entire population

LMNNC (Figure 6.10). These ecosystems are biologically

(Dijkstra, 2004). Although subpopulations are apparently

highly valuable, poorly studied and unique (Jocque,

healthy at present (Clausnitzer, 2018), without ongoing

Geeraert & Jones, 2018), and have been described as one of

protection this unique species could disappear in a very

the most important habitats in terms of diversity for

short period of time.

dragonflies in East Africa.

More details on the Ntchisi and blue-lipped yellowwings

6.4 Major threats

can be can be found in Species in the spotlight: The

threatened forest yellowwings p. 102. The NT eastern

The main threat to African and tropical dragonflies worldwide

yellowwing is known from the Taita Hills, Mt. Kilimanjaro,

is habitat loss and destruction through deforestation,

and the Eastern Arc and Pare Mountains in Tanzania and

urbanisation and agricultural encroachment, and the

Kenya. In the LMNNC, this species occurs in the Central

subsequent alteration of water bodies by erosion,

Region of Malawi including the southern tip of Nkhotakota

eutrophication and siltation (Dijkstra et al., 2011). Globally,

Wildlife Reserve and the Ntchisi Forest reserve (where the

all wetlands are economically vital but poorly protected

CR Ntchisi yel owwing is also found), as well as in the north-

and so are vulnerable to clearance and overuse (Darwal et

96
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of endemic odonate species

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 6.9 Richness of endemic odonate species (referred to throughout

the chapter as dragonflies) in the Lake Malawi/Nyasa/

Niassa Catchment based on spatial data coded as Presence 1 (Extant).

Richness data are classified using quantiles. Mapping is to
HydroBASINS and does not imply species occur across the entire
HydroBASIN (i.e. across the entirety of Lake Malawi/Nyasa/

Niassa). The single species endemic to the Lake Malawi/Nyasa/Niassa

Catchment is the Critically Endangered (CR) Ntchisi yellowwing (
Allocnemis maccleeryi).

97
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of threatened odonate species

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 6.10 Richness of threatened odonate species (referred to
throughout the chapter as dragonflies) in the Lake Malawi/

Nyasa/Niassa Catchment based on spatial data coded as Presence 1

(Extant). Richness data are classified using quantiles.

Mapping is to HydroBASINS and does not imply species occur across

the entire HydroBASIN (i.e. across the entirety of Lake
Malawi/Nyasa/Niassa).

98
Figure 6.11 The eastern horntail ( Nepogomphoides stuhlmanni)

Figure 6.12 The Mulanje damsel ( Oreocnemis phoenix) is

is assessed as Vulnerable (VU). © Viola Clausnitzer

assessed as Endangered (EN). © K.-D.B. Dijkstra

al., 2018). Despite having only four threatened species, the

Africa (Clausnitzer et al., 2012, 2018; Dijkstra et al., 2011;

majority (76%) of LMNNC dragonfly species have recorded

Dijkstra, Tchibozo & Ogbogu, 2009). Annual and perennial

threats in their Red List assessments.

non-timber crops are coded as a threat to 67% of species

native to the catchment. Overall, 37% and 64% species

6.4.1 Habitat loss and degradation

are threatened by small holder and agro-industry farming,

respectively, while logging and wood harvesting threatens

The greatest threat to dragonflies within the LMNNC is

22% of native species. Dragonflies are subject to impacts

habitat loss and degradation primarily from agricultural

primarily through direct loss of essential habitat and

expansion (Figure 6.13) as human populations continue

also through increased siltation and floods. This not only

to grow. This is a similar story for dragonflies throughout

decreases food sources, but also disrupts natural life cycles

Figure 6.13 Agriculture in Malawi. Agriculture is the primary threat to

odonates (referred to throughout the chapter as dragonflies) in the Lake
Malawi/Nyasa/Niassa Catchment. © Sailing Nomad (CC BY-NC-ND 2.0)

99
through the destruction of breeding, ovipositon and larval

LMNNC, impacting 48% of native species. Urban hubs, such

development sites.

as the Malawian capital of Lilongwe (Figure 6.14), are major

contributors but levels of pollution will continue to rise as

The threatened forest specialists, including the blue-lipped

urban areas expand. Even widespread and relatively resilient

yellowwing ( A. montana), are dependent on clear forest

species, such as the dancing jewel ( Platycypha caligata), are

streams, and so the rapid loss and degradation of these

threatened by pollution in these highly populated areas

habitats caused by smal holder and agro-industry farming

(Clausnitzer, Suhling & Dijkstra, 2016). The second largest

is creating highly isolated and vulnerable subpopulations

source of pollution is from agricultural and forestry effluents,

(Clausnitzer, 2010c). The overall species richness does

threatening 47% of species. In addition to sedimentation

not usually change with habitat loss but the composition

from forest clearance, this includes the input of nutrients

can shift from localised and threatened forest species to

from fertilisers, as well as the run off of pesticides and

more common, widespread and robust open-land species

herbicides. Industrial and military effluents are coded as

(Dijkstra & Clausnitzer, 2006). However, the loss of these

threats to 35% of native species. There is often little control

forests does not just impact specialists, as shaded refuges

of industrial output and these harmful effluents can cause

along rivers are important for many savannah and woodland

enormous damage to al freshwater life, in particular through

dragonflies (Dijkstra, Tchibozo & Ogbogu, 2009).

their rapid and sometimes long distance dispersal to

downstream connected habitats. This will especially impact

6.4.2 Pollution

sensitive dragonflies and their larvae.

Pollution is a major and growing threat to freshwater

6.4.3 Water management

biodiversity throughout Africa and is coded as a threat to

half (50%) of native species in the LMNNC. Agricultural,

The third largest threat to dragonflies within the LMNNC

urban and industrial expansion is often not accompanied

is through natural system modifications, which is coded

by appropriate water treatment measures, meaning harmful

as a threat to 44% of native species. This is mainly from

pol utants can be expel ed directly into freshwater habitats.

the abstraction of surface (26%) and ground water (34%)

This impacts dragonflies specifical y through the destruction

for agricultural use. Abstraction results in large scale loss

of larval habitats and by altering wel -established patterns of

of wetland habitat that destroys larval habitats and alters

competition between species (Dijkstra et al., 2011).

species interactions. The issue of water abstraction for

human use wil continue to grow in the light of global climate

By a small margin, domestic and urban waste water is the

change, which wil put increased pressure on al freshwater

most common source of pollution affecting dragonflies in the

ecosystems (Darwal et al., 2011).

Figure 6.14 Lilongwe and other urban areas are a major source of
pollution in the Lake Malawi/Nyasa/Niassa Catchment.

© katymartin (CC BY-NC-ND 2.0)

100

Figure 6.15 The effects of localised threats on the black-kneed

duskhawker ( Gynacantha bullata), which is assessed as Least Concern
(LC), need to be monitored. © Charles J Sharp (CC BY-SA 4.0)

6.5 Recommended research and

51% of species, closely fol owed by research into life history

conservation actions

and ecology (46%). Knowledge on population ecology on all

Afrotropical dragonflies is especially deficient and equally

6.5.1 Research actions recommended

vital. As evidence for this lack of information, the IUCN

Red List Criteria A, C and D (IUCN, 2012) could be applied

Although in general dragonflies are well known ecologically,

in only a few cases for the African Odonata assessments.

knowledge of Afrotropical species is often poor (Dijkstra et

To enable more comprehensive assessments of species

al., 2011). The need for more knowledge on the dragonflies

extinction risk in the future, population trends and detailed

within the LMNNC is reflected in the fact that 87% of native

habitat requirements of (at least) selected species should be

dragonfly species have recommendations for further

investigated (Clausnitzer et al., 2018).

research.

For this to occur the capacity for research needs to be built

The monitoring of population trends is recommended for

on within the LMNNC and researchers need to be equipped

the majority (78%) of native species. While many species

with the tools to identify and monitor this biodiversity

are widespread with stable population trends, there are

themselves (Clausnitzer et al., 2018). A major step forward

often little data on the impacts of localised threats. The

in the identification of dragonflies in eastern Africa has been

black-kneed duskhawker ( Gynacantha bullata; Figure 6.15),

made with the publication of an illustrated field guide (Dijkstra

a native of the LMNNC, is an example of a widespread LC

& Clausnitzer, 2014), incorporating many photographs, and

species that is found in central, eastern and western Africa

with an online tool providing identification details, photos

(Clausnitzer & Dijkstra, 2016). As it is threatened by forest

and distribution maps: African Dragonflies and Damselflies

destruction the impacts of this need to be monitored to

Online (ADDO)(Dijkstra, 2016) (see addo.adu.org.za).

prevent local declines.

Accurate and accessible information on dragonfly distribu-

To effectively monitor changes in biodiversity, it is first

tions, population dynamics and ecology is vital for their

necessary to know current distributions. This information

effective conservation (Clausnitzer, 2001). Many areas have

is lacking for LMNNC species, with research into the

yet to be comprehensively sampled, and sampling of these

population size, distribution and trends recommended for

areas often leads to new country records such as in recent

101

surveys in the highlands of Mozambique (Jocque, Geeraert

Considering the impact of the major threats of agricultural

& Jones, 2018). Areas with little or no recent records (Figure

and urban expansion, the fol owing conservation actions are

6.8) could be the focus of future surveys, including the north-

suggested:

east of the basin in southern Tanzania which has some of

the lowest levels of dragonfly diversity recorded within the

■ Protect existing aquatic habitats, especial y remaining

LMNNC (Figure 6.5). This knowledge will help to further the

montane forest fragments;

role of dragonflies in freshwater conservation within East

■ Reforest hil -top and riparian zones of streams and rivers

Africa as bio-indicators of declining wetland health (Dijkstra

with indigenous trees;

et al., 2011).

■ Eliminate direct waste water influent into streams, rivers

and large lakes from agricultural, domestic and industrial

6.5.2 Conservation actions recommended

sources, including into LMNN itself.

Even though the majority of species within the LMNNC

For the threatened forest specialists, including the yellow-

are considered LC, 37% have recommended conservation

wings ( Allocnemis) and the eastern horntail ( Nepogom-

actions. The most common recommendation is land and

phoides stuhlmanni), the protection of remaining habitat

water management (33%), which includes habitat and

fragments is essential for their continued survival and the

natural process restoration (17%) and site/area management

restoration of degraded forest habitat is needed to help them

(33%).

to recover.
Species in the spotlight

The threatened forest yellowwings

Palmer-Newton, A.F.1, Dijkstra, K.-D.B.2 and Clausnitzer, V.3

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

2 Naturalis Biodiversity Center, Leiden, Netherlands

3 Senckenberg Museum of Natural History Görlitz, Görlitz, Germany

The Afromontane forest, shaded stream specialists are the most threatened
group of dragonflies within the LMNNC

because of their low adaptability, combined with the multitude of threats to

their habitat. Two of the most threatened species within the LMNNC are
detailed here.

The Ntchisi yellowwing ( Al ocnemis maccleeryi) is a Critically Endangered

(CR) damselfly found only within the 9,712 ha Ntchisi forest reserve in
central Malawi (BirdLife International, 2019; Clausnitzer, 2010d). This
reserve has an interesting cultural history, protected from habitat degradation
as it provided vital shelter for the local Chewa tribe against attacks by the
warring Ngonis in the 19th century. It is the only peak in the Dowa Hil s that
stil has forest within this densely cultivated, populated and eroded region
(BirdLife International, 2019). It is said to contain some of the last
remaining indigenous rainforest in Malawi and is also the most isolated of
these forests in the county (Dowsett-Lemaire, 1989). While some natural
vegetation has been cleared and replaced with exotic plantations to help al
eviate logging pressure, the forest is still in good condition and the woodland
to the north-east is relatively untouched (BirdLife International, 2019). The
blue-lipped yel owwing ( Al ocnemis montana) is an Endangered (EN)
damselfly that has a slightly wider distribution than its relative. It has been
recorded in the Mughese Forest in Misuku Hil s (north Malawi) and the
Matengo Highlands (southern Tanzania) (Clausnitzer, 2010c). As this
species is restricted to clear, shaded forest streams, its presence makes it a
good indicator of healthy forest ecosystems (Jocque, Geeraert & Jones,
2018).

These Afromontane ecosystems have high levels of biodiversity but are

poorly studied and under intense pressure from agricultural expansion and
wood extraction (Jocque, Geeraert & Jones, 2018). The specific habitat
preferences of forest specialists make them especially vulnerable as they are
easily outcompeted by open-land species when forests are broken up.
Ongoing loss of forest habitat mean that subpopulations are highly isolated
and unlikely to recolonise after loss, meaning when habitat is cleared
subpopulations have little chance of survival (Clausnitzer, 2010c). Without
protection and restoration of their remaining habitats the status of these
unique forest specialists wil only continue to worsen. With the ongoing
decline of unstudied Afromontane habitats throughout the basin, forest
specialists may disappear before they can even be recorded.

102
Figure 6.16 Afromontane forest, shaded streams, such as this stream in
the Mulanje Massif in Malawi, are home to the threatened forest
yellowings. © DJ Cockburn (CC BY-NC 2.0)

103

Figure 6.17 The eastern yellowwing ( Allocnemis abbotti) is a close

relative of the threatened Ntchisi yellowwing ( Allocnemis

maccleeryi) and blue-lipped yellowwing ( Allocnemis montana). © K.-

D.B. Dijkstra The (near) endemics of the lake

Palmer-Newton, A.F.1, Dijkstra, K.-D.B. 2 and Clausnitzer, V.3

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

2 Naturalis Biodiversity Center, Leiden, Netherlands

3 Senckenberg Museum of Natural History Görlitz, Görlitz, Germany

While there is only one confirmed endemic to the LMNCC, the dragonfly
populations outlined below just miss out on that title. These species
exemplify the need for ongoing research into the taxonomy and distributions
of dragonflies within the basin, especial y in understudied areas, which may
reveal more unique and potential y threatened dragonflies.

The first is the ‘lacus’ form of the dancing jewel ( Platycypha caligata). This
damselfly is widespread in eastern and southern Africa, from Ethiopia to
Angola and South Africa (Clausnitzer, Suhling & Dijkstra, 2016). Male
dancing jewels have a bril iant, multi-coloured appearance that is used to
court the cryptic coloured females by waggling their abdomens and legs.
Males in territorial displays can rise more than six metres above waterbodies
in an upward ‘dance’, ending only when one male leaves (Jennions, 1998).
The ‘lacus’ form, also known as the Malawi jewel, has only been recorded
on rocky shores of LMNN in areas such as Mbenji Island and Senga Bay. It
prefers large lakes in open landscapes, often in splash zones with rocks and
submerged roots (Dijkstra, 2019c). Also unlike the typical dancing jewel, it
is smal er and its darker colouration has been shown to darken with age with
unknown consequences on breeding behaviour. Genetic sampling of riverine
populations from Malawi is needed to substantiate the Malawi jewel as a
distinct species and not just a dwarf form, like those found in Lake Chala on
the Kenya-Tanzania border (Dijkstra, 2007).

The second near endemic is the Malawi hooktail ( Paragomphus nyasicus),

whose range extends just beyond the boundaries of the LMNNC. It is
another open lake species that occurs on the beaches of Malawi. It has also
been found on streams nearby and mysteriously at Victoria Fal s in
Zimbabwe, but it is uncertain whether it reproduces there or is vagrant
(Dijkstra, 2019d). Larvae have been seen emerging from the lake after
midnight at Chembe vil age, leaving beaches littered 104
with empty larval skins. This late emergence time is unusual for tropical
gomphids, and may be a unique characteristic or a response to the intense
human disturbance by local people and tourists for washing, swimming,
canoeing and fishing that continues wel after dusk. In the day, females dip
their abdomens into sand just touched by tiny lake waves to lay eggs, while
males patrol and perch on sand very close to the waterline (Reinhardt, 2006).
Both of these species are threatened by domestic and agricultural pollution
flowing directly into LMNN. Although currently assessed as Least Concern
(LC) any changes in the conditions of LMNN wil be a risk and may qualify
these near endemic populations for a higher threat category in the future
(Clausnitzer, 2010e; Clausnitzer, Suhling & Dijkstra, 2016).

Figure 6.18 The dancing jewel ( Platycypha caligata) is widespread in

eastern and southern Africa, and assessed as Least Concern (LC). The
‘lacus’ form (not pictured) is endemic to the Lake Malawi/Nyasa/Niassa
Catchment. © Hans-Joachim Clausnitzer

Figure 6.19 The Malawi jewel (‘lacus’ form of Platycypha caligata), has
only been recorded on rocky shores of Lake Malawi/

Nyasa/Niassa, such as at Senga Bay. © Amy Palmer-Newton

105

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107

Chapter 7

The status and distribution of freshwater

plants in the Lake Malawi/Nyasa/Niassa

Catchment

Palmer-Newton, A.F.1, Mwanyambo, M.2 and Lansdown, R.V.3

Contents
7.1 Introduction
...........................................................................................................................
.........................................................................108

7.2 Red List assessments

...........................................................................................................................
.........................................................109

7.3 Major threats

...........................................................................................................................
....................................................................... 111

7.3.1 Habitat degradation and destruction

...........................................................................................................................
.......................... 112

7.3.2 Biological resource use

...........................................................................................................................
.............................................. 112

7.3.3 Natural system modifications

...........................................................................................................................
..................................... 112

7.4 Recommended research and conservation actions

...........................................................................................................................
............ 114

7.4.1 Research recommendations

...........................................................................................................................
....................................... 114

7.4.2 Conservation
recommendations...............................................................................................
............................................................. 115

Species in the spotlight

...........................................................................................................................
.............................................................. 116
Freshwater plants of the Nyika Plateau
...........................................................................................................................
.............................. 116

The socio-economic value of freshwater plants in the Lake

Malawi/Nyasa/Niassa Catchment ....................................................................
117

Bibliography
...........................................................................................................................
............................................................................... 118

7.1 Introduction

The Lake Malawi/Nyasa/Niassa Catchment (hereafter

LMNNC) is home to a great diversity of freshwater habitats

The diverse climate and geology of Africa has led to

and associated plants. In the context of this current study

the development of an enormous range of freshwater

it was, therefore, not possible to assess the status of all

habitats, from the vast lakes of the Rift Valley and the

freshwater plants within the region, such that only a subset

extensive wetland complexes of the Okavango Delta to

of these species was assessed, total ing 247 native species

small seasonally inundated wetlands found throughout

in 57 families. See Chapter 2 for an explanation of how the

the continent. These wetlands support a correspondingly

species list for assessment was generated.

diverse flora, ranging from tall stands of papyrus ( Cyperus

papyrus; Figure 7.1) to the diminutive and poorly-known

The freshwater-dependent plant species assessed through

Crassula and Isoetes species. Wetland-dependant plants

this study are heavily dominated by those which occur in the

play a vital role in the creation and maintenance of these

shallower areas of Lake Malawi/Nyasa/Niassa (hereafter

habitats by providing nutrients and structural support

LMNN) and in marginal habitats such as marshes, ponds and

to species in higher trophic levels and, together with

backwaters associated with inflowing streams (Figure 7.2).

phytoplankton, supply the primary production upon which

The main wetland types in addition to those associated with

this life depends. They also help sustain local livelihoods

the LMNNC are the high altitude seasonal pools in the Mount

through direct use as food, medicines and structural

Mulanje range at the southern tip of the catchment and on the

materials, and provide a wide range of indirect ecosystem

Nyika Plateau in northern Malawi. Fol owing species coding

services including flood control, nutrient recycling,

according to the IUCN Red List Habitats Classification

bioaccumulation of pollutants and sediment trapping

Scheme (Figure 7.2), the majority of native species (62%)

(Mil ennium Ecosystem Assessment, 2005; IPBES, 2019).

occur along permanent rivers, streams and creeks, fol owed

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

2 National Herbarium and Botanic Gardens of Malawi, P.O. Box 528,

Zomba, Malawi

3 IUCN SSC Freshwater Plant Specialist Group, 45 The Bridle, Stroud,

Glos. GL5 4SQ, UK

108

7.2 Red List assessments

The majority of the 247 plant species native to the LMNNC

and assessed through this study are Least Concern (LC;

94%). Only two species are assessed as threatened, both

of which are Endangered (EN, 1%), and 14 are assessed

as Data Deficient (DD, 6%) (Table 7.1, Figure 7.3). These

Red List assessments suggest that the freshwater plants

native to the LMNNC are, in general, at a lower level of

threat than across continental Africa as a whole, for which

24% (excluding DD species) were considered threatened

(Darwall et al., 2011). However, due to limitations in both

Figure 7.1 Tall stands of papyrus ( Cyperus papyrus). This

available data and resources, the species assessed in this

species is assessed as Least Concern (LC). © Rod Waddington

study may not represent a comprehensive list of wetland-

(CC BY-SA 2.0)

dependent plant species native to the LMNNC and so these

closely by swamps and marshes (60%) and then permanent

results may not be representative of the situation overall.

freshwater pools (44%). A large proportion (42%) of species

The LMNNC is a region for which data on freshwater-

can be found in artificial aquatic environments, such as

dependent plants, excluding “true” aquatic plants (i.e. those

canals and drainage ditches (25%) and seasonal y flooded

that can only grow with their roots and photosynthetically

agricultural land (19%). Seasonal habitats also support

active parts submerged permanently or for extended

a high proportion of freshwater plants, and this includes

periods), are extremely sparse. This lack of data has

seasonal y flooded grasslands (31%), seasonal marshes and

resulted in a simplified understanding of species status

pools (35%) and seasonal flowing waters (26%). Permanent

in which widespread species are considered to be of low

lakes, such as LMNN, support relatively few species (29%),

extinction risk (in the absence of evidence to the contrary),

and these are mostly confined to shallow areas along the

while there is insufficient information available to assess the

shore, although it is recognised that the abundance and

extinction risk of the majority of other species and these are,

distribution of macrophytes around the lake itself is poorly

therefore, assessed as DD. In the future, it is recommended

understood (Kafakoma, 2019). While a large number of

that additional resources are made available to enable

wide ranging, generalist species occur within the LMNNC,

production of a more comprehensive and representative

there are also a number of species restricted to areas such

species list. This would be likely to include a higher

as the Nyika National Park (see Species in the spotlight:

proportion of restricted range and potentially threatened

Freshwater plants of the Nyika Plateau p. 116).

wetland-dependent plants. Northern Africa represents a

Permanent Rivers/Streams/Creeks (includes waterfal s)

61.5%

Bogs, Marshes, Swamps, Fens, Peatlands

60.3%

Permanent Freshwater Marshes/Pools (under 8 ha)

44.1%

Seasonal/Intermittent Freshwater Marshes/Pools (under 8 ha)

34.8%

ype

Subtropical/Tropical Seasonal y Wet/Flooded Grassland

31.2%

Permanent Freshwater Lakes (over 8 ha)

29.1%

Habitat t

Seasonal/Intermittent/Irregular Rivers/Streams/Creeks

25.5%

Canals and Drainage Channels, Ditches

25.1%

Seasonal y Flooded Agricultural Land

18.6%

Seasonal/Intermittent Freshwater Lakes (over 8 ha)

18.2%

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%
100%

Proportion (%) of species recorded in each habitat type

Figure 7.2 Proportions (%) of the 247 species assessed within the Lake
Malawi/Nyasa/Niassa Catchment found in each habitat type as defined
by the IUCN Red List Habitats Classification Scheme. Only the top 10
habitat types in terms of proportion of species are shown. Dark blue
bars represents permanent wetland habitats, light blue bars represent
seasonal/intermittent wetland habitats, and purple bars represent
artificial habitats.

109

region for which good quality data are available, not only

have restricted ranges and it is these groups that include the

to define which species are wetland-dependent but also

threatened and DD taxa.

to assess their extinction risk. Assessment of selected

freshwater plants of this region (Rhazi & Grillas, 2010)

The two species assessed as threatened are both sedges

yielded results that are probably more representative of the

(Cyperaceae). Carex brassii (EN) is a sedge that can be

true situation for freshwater plants, with a higher proportion

found along streams and rivers and on riverbeds in forests,

of threatened species.

bogs and on the forest floor. It is endemic to Malawi and is

only known from Nyika National Park in the west of the

In general, “true” aquatic plant species are likely to be

catchment and the Mount Mulanje Forest Reserve, which

widespread and abundant and less threatened. Of the

mostly lies outside of the LMNNC on its southern boundary.

species assessed in the LMNNC, 22% have an almost

Although there is little information on its population trends

global distribution, including greater duckweed ( Spirodela

and use, it is threatened by uncontrol ed fires and introduced

polyrhiza; Figure 7.4) and spiked water-milfoil ( Myriophyllum

plant species on the Nyika Plateau (see Species in the

spicatum) (Lansdown, 2017; Máiz-Tomé & Beentje, 2017).

spotlight: Freshwater plants of the Nyika Plateau p. 116),

Many others are distributed across a high proportion of Sub-

as well as fire, fuelwood collection (Figure 7.5), illegal

Saharan Africa (40%), such as papyrus ( Cyperus papyrus)

logging and invasive species in the Mount Mulanje area

(Beentje, 2017). While some of the widespread species may

(Bayliss et al., 2007; Larridon & Lansdown, 2018a). Its small

be exposed to local threats, their large distribution ranges

and disjunct distribution range means that it is vulnerable to

mean that the majority are considered LC with a low current

ongoing declines.

risk of global extinction. Only a smal proportion of species

Scleria richardsiae (EN), a nutrush sedge, occurs in peren-

nially wet places in thick grassy vegetation, often along

Table 7.1 Number of assessed freshwater plant species native and

endemic to the Lake Malawi/Nyasa/Niassa Catchment in each

mountain streams and in lush streamside bogs from 1,700 m

Red List Category. For a list of species native to the Lake Malawi/

upwards. It is only known from eight col ections on the Nyika

Nyasa/Niassa Catchment and their Red List Categories please

Plateau in northern Malawi, north-eastern Zambia and

see Appendix 1.

south-western Tanzania. There is a lack of information on

Number of

Number of

its exact range, population trends and life history, but like C.

assessed species assessed species

native to the
endemic to the

brassii it is threatened by uncontrolled fire and introduced

IUCN Red List Category

LMNNC

LMNNC

plant species within Nyika National Park (see Species in the

Extinct (EX)

spotlight: Freshwater plants of the Nyika Plateau p. 116)

Extinct in the Wild (EW)

and agricultural and urban expansion outside of protected

Critically Endangered (CR)

areas (Larridon & Lansdown, 2018b). Without continued

Endangered (EN)

2
0

protection the status of both of the threatened sedge

species will be likely to decline.

Vulnerable (VU)

Near Threatened (NT)

Only one of the species assessed is endemic to the LMNNC:

Least Concern (LC)

231

Helichrysum tithonioides, which is assessed as DD (Table

Data Deficient (DD)

14

7.1, Figure 7.3). This is a member of the daisy family

Total

247
1

(Asteraceae) and occurs on the Nyika Plateau in northern

Malawi (see Species in the spotlight: Freshwater plants

of the Nyika Plateau p. 116) where it grows in marshy areas.

Endemic species

It is thought to be fairly abundant within this smal range, but

it may be impacted by tourists and visitors within the park

and information on its population, ecology and threats is

Native species

scarce (Lansdown, 2018).

There are 14 species assessed as DD within the LMNNC,

0%

20%

40%

60%

80%

100%

including the endemic H. tithonioides. These are species for

which there is insufficient information available to evaluate

EX
EW

CR

EN

VU

NT

LC

DD

their extinction risk. Six of these occur on Mount Mulanje

Figure 7.3 Percentage (%) of freshwater plant species

(Figure 7.6), of which two are endemic to the mountain

assessed as native and endemic to the Lake Malawi/Nyasa/

range (Strugnel , 2002). Other DD species, such as Vernonia

Niassa Catchment in each Red List Category. For a list of

species native to the Lake Malawi/Nyasa/Niassa Catchment

tolypophora which occurs from Kenya south to Mozambique

and their Red List Categories please see Appendix 1.

(Mwanyambo, 2018), have a wider distribution but are equal y

110
deficient in information on their population trends and threats.
These DD species are a key focus for future research efforts

within the LMNNC as they may be threatened and in need of

conservation actions. This is especial y the case for Mount

Mulanje endemics where uncontrol ed fire, invasive species,

and il egal logging are serious concerns (Bayliss et al., 2007).

7.3 Major threats

Freshwater plants within the LMNNC face multiple threats,

the majority of which relate to changing land use practices.

Although only 1% of the freshwater plant species assessed

Figure 7.4 Greater duckweed ( Spirodela polyrhiza) is native to

the Lake Malawi/Nyasa/Niassa Catchment but has an almost

(excluding DD species) within the LMNNC meet the

global distribution. It is assessed as Least Concern (LC).

thresholds to be assessed as threatened (Figure 7.3), 9%

© Christian Fischer (CC BY-SA 3.0)

Figure 7.5 Fuelwood collection at the edge of Mount Mulanje Forest

Reserve. © David Davies (CC BY-SA 2.0) Figure 7.6 Mount Mulanje is
home to a number of Data Deficient (DD) freshwater plant species. ©
David Davies (CC BY-SA 2.0) 111
have recorded threats within seven major threat categories

farming is one of the major drivers for habitat destruction,

based on the standard IUCN Red List Threats Classification

particularly where new gardens are opened in forested

Scheme. There is also a lack of knowledge on localised

areas. This expansion of agriculture includes encroachment

threats within the catchment and so the number of species

into protected areas such as at the Dwambazi Forest

facing threats is likely higher. The major recorded threats

Reserve in Malawi (Figure 7.9), which lies just to the north of

to freshwater plants within the LMNNC are expanding

the Nkhotakota Wildlife Reserve on the border between the

agriculture, biological resource use and changing fire

Central and Northern regions of Malawi.

patterns, while the impacts of climate change, mining,

pol ution, and residential and commercial development are

Wetland habitats are also threatened by residential and

emerging threats.

commercial development, which is coded as a threat to 2%

of species. This is particularly the case in urban areas where

7.3.1 Habitat degradation and destruction

these threats are concomitant with small-scale garden

farming. An example of this is the Lunyangwa River wetland

The greatest threat to freshwater plants within the LMNNC

that runs through Mzuzu City in northern Malawi, where

is the degradation and destruction of wetland areas,

residential expansion and increasing levels of garden farming

predominately through agriculture. This is coded as a threat

have now encroached onto its fringes and have significantly

to 7% of species and is mainly due to the expansion of smal -

altered this wetland habitat.

holder farms (4%), but also agro-industry farming (2%) and

livestock farming and ranching (2%).

7.3.2 Biological resource use

Rapid expansion of agricultural land (Figure 7.7) within the

The second greatest threat to freshwater plant species

catchment can be seen clearly in land cover and land use

within the LMNNC is biological resource use. This is coded

patterns in Malawi from 1990 to 2010 and in 2016 (Figure

as a threat to 2% of species and includes the gathering of

7.8), with the central and southern areas of Malawi seeing the

terrestrial plants (2%), and logging and wood harvesting

highest rates of conversion. There is also increasing growth

(1%). The human population within the catchment is pre-

of agriculture along the lake shore in both Mozambique

dominantly rural based and is heavily dependent on natural

and Tanzania (Kafakoma, 2019). An increasing demand

resources for materials, medicinal plants and fuel. This

for agricultural land in the most densely populated areas

extraction is largely destructive and, although mostly small

in the south of the basin is causing accelerating declines

scale, is putting increasing pressure on freshwater plant

in soil fertility. Consequently, agricultural land is now

populations.
rapidly expanding into more mountainous central and

northern regions within the catchment, and increasing

The extraction of food plants such as edible orchid tubers is

population densities and land tenure reforms have forced

especial y destructive. The raw tubers of Satyrium species,

rural populations to cultivate in marginal areas such as

for example, are harvested for both domestic consumption

wetlands (Kafakoma, 2019). Demographic factors are largely

and trade across international borders. More details on the

attributable as the majority of people in the LMNNC derive

harvest of the montane grassland S. shirense can be found in

their livelihoods from agriculture or fishing, and smal -holder

Species in the spotlight: The socio-economic value of

freshwater plants in the Lake Malawi/Nyasa/Niassa

Catchment p. 117. Many Satyrium species threatened by

overharvesting are documented from seasonally wet

grasslands known as dambos or from wet open Brachystegia

woodland. These species include kajibatike ( S. carsonii),

mbuyeuye ( S. buchanani ), and mbuyeuye wa mudambo ( S.

ambylosaccos) (Kasulo, Mwabumba & Munthali, 2009;

Mwanyambo & Kananji, 2003), none of which have yet been

assessed for the IUCN Red List. These dambo habitats are

also are prime areas for small-scale farming, which adds

additional pressure on the wetland plant species and people

that rely on their harvest.

7.3.3 Natural system modifications

Figure 7.7 Birds eye view of agricultural land in Malawi. © lucianf

Modifications to natural systems are coded as a threat to

(CC BY 2.0)

2% of freshwater plant species within the LMNNC. This is

112
Figure 7.8 Land cover land use (LCLU) patterns in Malawi within the
Lake Malawi/Nyasa/Niassa Catchment between 1990–2010

and in 2016, based on public domain Serviresa data from the RCMRD
(Regional Centre For Mapping Resource For Development) Geoportal.
The 1990–2010 maps were produced from LandSat Imagery (30 m by 30
m) resolution using supervised classification.

The 2016 map was clipped from Sentinel-2 global land cover data. Both
accessed 22/05/2019 at http://geoportal.rcmrd.org.

113

Figure 7.9 An illegal small-

holderfarmwithinthe

Dwambazi Forest Reserve

in Malawi, 2018. Agricultural

expansionisamajor

threat to freshwater plants

w i t h i n t h e L a ke M a l aw i /

Nyasa/Niassa Catchment.

© Montfort Mwanyambo

mainly due to late dry season fires, which have become a

towards the creation of a more comprehensive species list

common feature in some areas. These fires burn intensely

for the catchment, which would be likely to include a larger

and devastate wetland habitats, and in the long term can

proportion of restricted range and potentially threatened

alter the species composition to favour more fire resilient

species which could be targeted for conservation actions.

species, such as bracken ferns (Burrows & Willis, 2005).

The two EN species, Carex brassii and Scleria richardsiae,

Species classed as LC are also often poorly known. For

are subject to such fire regimes. Both are found within

example, the river pumpkin ( Gunnera perpensa), which is

Nyika National Park, which is discussed in Species in the

found throughout sub-Saharan Africa, is a species thought

spotlight: Freshwater plants of the Nyika Plateau p. 116.

to be declining due to destruction of its wetland habitat and

unsustainable harvesting for food and medicine, although

no information on its trade is available outside southern

7.4 Recommended research and

Africa. In this case, to ensure the species is prevented from

conservation actions

becoming threatened in the future and to help prevent local

extinctions, its population and harvest trends need to be

7.4.1 Research recommendations

monitored throughout its wide range, especial y where it is

collected (Palmer-Newton, 2018). This example reinforces

There is a notable lack of information on wetland-dependent

the need to monitor species, even those at apparently low

plants in the LMNNC. There have been almost no targeted

relative current risk of extinction. This will be especially

surveys to document wetlands and their vegetation, while

important in areas where localised threats exist.

most of the restricted range species are known only from

notes taken as part of general botanical surveys (e.g. Burrows

Lack of information on the distribution of wetland-

& Wil is, 2005; Strugnell, 2002; Wil is et al., 2001). Although

dependant plants has meant that no species richness maps

only 14 (6%) of the species assessed were classed as DD,

could be produced for these plants within the LMNNC, with

17% were considered to require research, particularly into

many species only having distribution data at the country

population size and trends (15%), the threats that they face

presence level or none at al . This is especial y the case for

(11%) and their ecology (10%). The most important action

DD species. More precise and accessible distribution

to benefit conservation of freshwater wetland-dependent

information would allow centres of overall and threatened

plants in the LMNNC is, therefore, to collect and compile

freshwater plant species richness to be identified as a

data to improve our knowledge on these species and the

critical input to effective planning for future conservation

habitats upon which they depend. This would also help

actions.
114

As an example for the type of survey required, botanists

Finally, as freshwater plants are an integral part of wetland

from the Royal Botanic Gardens at Kew are currently

ecosystems and provide vital resources to people in the

undertaking surveys in Mozambique toward identification

LMNNC, it is recommended that vulnerable habitats are

of Important Plant Areas (IPAs). These include recording

protected and restored where needed, especially where

information on a range of poorly-known wetland-dependent

threatened by agricultural expansion and overharvesting.

plant species, particularly members of the Lythraceae family

This may require a programme of awareness raising and

(I. Darbyshire pers. comm. 2019). Similar projects throughout

development of an advisory programme for farmers. As

the LMNNC would likely generate new data for poorly known

many threats to aquatic plants will spread rapidly through

taxa and previously undescribed species.

wetland systems, it is important that management actions

are employed at the catchment scale using methods

7.4.2 Conservation recommendations

such as Environmental Flows (E-Flows) and Integrated

River Basin Management ( IRBM ). A strong focus on

Specific conservation actions were proposed for only two

conservation of wetlands habitats and their biodiversity

species, both of which are exploited by people. Satyrium

should also be included in future iterations of the National

shirense (DD) is considered to be threatened by over-

Biodiversity Strategies and Action Plans as, without this

col ecting for food (see Species in the spotlight: The socio-
increased attention, it is likely that these currently poorly

economic value of freshwater plants in the Lake Malawi/

valued habitats and their associated plants will continue

Nyasa/Niassa Catchment p. 117), and measures to protect

to be lost and degraded. In particular, as currently noted

this species from over-collection include development of

in Malawi’s National Biodiversity Strategy and Action

alternative food sources and ex situ propagation, combined

plan (Environmental Affairs Department, 2006) there is

with enforcement of trade legislation (Henry, Barker &

no wetlands policy framework (outside of the Ramsar

Hargreaves, 2018). Hygrophila auriculata (Figure 7.10) (LC)

convention) to guide management and conservation

is considered to be important for human medicine and,

of wetland resources. Such a policy is needed if these

although it is not currently considered to be threatened, it

wetlands and their associated plants are to be retained and

would be valuable to establish material in a genome resource

continue to benefit people.

bank as a precaution against future declines (Gupta, 2018).

Figure 7.10 Hygrophila auriculata (Least Concern, LC) is considered to
be important for human medicine and it would be valuable to establish
material in a genome resource bank as a precaution against future
declines. © Dinesh Valke (CC BY-SA 2.0) 115

Species in the spotlight

Freshwater plants of the Nyika Plateau

Palmer-Newton, A.F.1, Mwanyambo, M.2 and Lansdown, R.V.3

1 Freshwater Biodiversity Unit, Global Species Programme,

IUCN (International Union for Conservation of Nature), David

Attenborough Building, Pembroke Street, Cambridge, CB2 3QZ,

UK

2 National Herbarium and Botanic Gardens of Malawi, P.O. Box

528, Zomba, Malawi

3 IUCN SSC Freshwater Plant Specialist Group, 45 The Bridle,

Stroud, Glos. GL5 4SQ, UK

The Nyika Plateau (Figure 7.11) is a large montane complex

covering 3,134 km² in the Nyika National Park in northern

Malawi and a further 70 km² in a park of the same name

across the border in Zambia. It is one of Africa’s centres

of plant diversity and is a Key Biodiversity Area (KBA)

recognised for its global importance to freshwater plants

and fishes (see Chapter 10 and the KBA Datasheets in

Figure 7.11 The Nyika Plateau is a Key Biodiversity Area (KBA)

Supplementary Material), as well as birds. The plateau

recognised for its global importance to freshwater plants.

supports miombo woodland, rol ing montane grasslands,

© Dr Thomas Wagner (CC BY-SA 3.0)

wetland dambos and evergreen forests (Burrows & Wil is,

2005; Wil is et al., 2001), and it is an important catchment

area that includes the source of four large rivers that drain
into LMNN (see the KBA Datasheets in Supplementary

Material). It is likely that the Nyika Plateau supports more

than 2,000 plant taxa but, given that vast areas remain

under collected, this number is likely to be much larger

(Burrows & Willis, 2005; Willis et al., 2001). Its montane

grasslands are dominated by sedges (Cyperaceae)

including Carex brassii (EN) and Scleria richardsiae (EN),

as well as numerous orchid species (Burrows & Willis,

2005). It is also the only known location of the endemic

freshwater plant Helichrysum tithonioides, in addition to

another 33 recorded Nyika plant endemics (Lansdown,

Figure 7.12 Fire resilient bracken is spreading on the Nyika

2018; Larridon & Lansdown, 2018a, 2018b). Of these

Plateau as a result of uncontrolled and intense late season

endemic species, six occur in freshwater habitats but are

fires. © firesika (CC BY-NC-ND 2.0)

yet to be assessed.

The major threat to floral diversity in Nyika is uncontrol ed and intense late
season fires that have a great impact across the plateau. In the long term this
alters the species composition to favour more fire resilient bracken (
Pteridium aquilinum; e.g.
Figure 7.12) (Burrows & Wil is, 2005; Nyika-Vwaza (UK) Trust, 2016). An
additional threat to native freshwater plants is the impact of invasive pine (
Pinus) species that were introduced to form eventual y unsuccessful
plantations in the mid-20th century, and other invasive species such as the
Himalayan raspberry ( Rubus el ipticus) (Nyika-Vwaza (UK) Trust, 2016).

Increasing human populations on the border of the park are also creating
growing political pressure to release land, which would have serious impacts
on al of its freshwater biodiversity (see the KBA Datasheets in
Supplementary Material).

Through the Nyika-Vwaza Trust, work is being done to protect this unique
park, including early burning and debris clearing to prevent late-season fires
and improving access to facilities, al while employing local people (Nyika-
Vwaza (UK) Trust, 2016). In addition, a number of environmental
educational programmes are being operated in northern Malawi by the
Lilongwe Wildlife Trust (see the KBA Datasheets in Supplementary
Material). This continued protection against agricultural encroachment and
uncontrolled fire is needed to safeguard this understudied but biologically
diverse habitat for all freshwater taxa.

116
The socio-economic value of freshwater plants in the Lake
Malawi/Nyasa/Niassa Catchment

Palmer-Newton, A.F.1, Mwanyambo, M.2 and Sayer, C.A.1

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

2 National Herbarium and Botanic Gardens of Malawi, P.O. Box 528,

Zomba, Malawi

Freshwater plants provide a vital alternative resource for the rural poor who
lack access to, or funds to purchase, market goods and modern
pharmaceuticals, as wel as providing vital food supplies in times of drought
for both livestock and people (Sayer et al., 2018). In the LMNNC 34% of the
247 freshwater plant species assessed are recorded as being used.

Medicinal use is the most common (22%) fol owed by use as food (human –
12%; animal – 9%) and then for horticulture (9%) (Figure 7.13).

25% % of fresh

20% ater plant spe

15% cies assessed

10% with eac

5%

h end use
ial

ri

es

Figure 7.13 The proportion (%) of freshwater plant species from a total
of 247 assessed within the Lake Malawi/Nyasa/Niassa catchment, within
each end use category defined in the IUCN End Use Classification
Scheme. Source of Icons from left to right from the Noun Project
(www.thenounproject.com): carrot by ibrandify; Dog food by arif fajar
yulianto; Medicine by designvector; Perfume by Smalllike; Firewood by
Pixelicatom; Rope by Eucalyp; Construction by Arafat Uddin; Clothing
by GreenHill; Chair by Bartama Graphic; Necklace by Vectors Point;
Flower by Made by Made; growing business by SBTS; and Compost by
Juraj Sedlák. Icon for poisons from Microsoft Office.

One example of a freshwater plant with a wide variety of uses is Piper

capense (Figure 7.14), known as long black pepper or timiz. This is an
aromatic evergreen shrub that occurs in shaded wet areas of evergreen and
montane forest and swamps throughout sub-Saharan Africa (Palmer-
Newton, 2019). It is used as a cultivated food and spice, as wel as a
medicinal plant to treat a variety of conditions from stomach troubles and
sore throats to epilepsy and paralysis (Burkill, 1985).

Multiple experimental studies have supported its use in infection related

conditions (Woguem et al., 2013; Zimudzi, 2008), and it has shown potential
in cancer treatment (Kuete et al., 2013; Woguem et al., 2013) and as an
antimalarial (Bobasa et al., 2018). This plant is, therefore, important both in
helping support local communities who lack access to commercial medicine
and on the global scale in medical research.

The second most common use of freshwater plants after medicine is for
human consumption. An example of such a plant is Satyrium shirense, an
orchid that is found across Malawi and in southern Tanzania, north-eastern
Zambia and the central-eastern area of Mozambique. Its tubers are harvested
to produce chikande/chinaka, a traditional meat-loaf like dish, which can be
used as a relish or snack (Kasulo, Mwabumba & Munthali, 2009). It has
been rapidly growing in 117
popularity, especially in urban areas (Veldman et al.,

2017). This increasing demand has depleted Zambia’s

orchid resources, so cross-border trade is growing

and harvesters are going further afield, principally to

the southern highlands of Tanzania where Satyrium

shirense also occurs. Poaching to fulfil this demand

also occurs in protected areas and has been a serious

issue in Kitulo National Park (Figure 7.15), Tanzania, in

the north of the LMNNC (Lalika et al., 2013). Satyrium

shirense is assessed as DD because this harvest is

driving unknown but potential y rapid population declines

(Henry, Barker & Hargreaves, 2018), with estimates of

up to 3.5 million Tanzanian orchid tubers (Figure 7.16)

Figure 7.14 Leaves of Piper capense, commonly known

from a number of species exported to Zambia each year

as long black pepper or timiz, has a variety of uses. It is

assessed as Least Concern (LC). © Scamperdale (CC BY-NC

(Veldman et al., 2014). Research into population trends,

2.0)

increased protection from poaching, development of

ex situ cultivation, and ongoing public awareness campaigns are urgently

needed to protect this species, as wel as the people who rely on its harvest
(Henry, Barker & Hargreaves, 2018; Kasulo, Mwabumba & Munthali,
2009).

In conclusion, a better understanding of the use of freshwater plants within

the LMNCC is needed to more effectively protect both the plants and the
people that rely on them. Given that for 20% of species there is currently no
available information on their use or trade, research in this area could
provide many additional benefits to people both within and outside of the
LMNNC.

Figure 7.15 Collection of Satyrium shirense is a serious issue

Figure 7.16 Orchid tubers for sale in Mbeya, Tanzania. ©

in Kitulo National Park. © Jojona (CC BY-SA 3.0)

Jojona (CC BY-SA 3.0)

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120

Chapter 8

Synthesis for al taxonomic groups

Sayer, C.A.1

Contents

8.1 Introduction
...........................................................................................................................
.........................................................................121

8.2 Red List assessments

...........................................................................................................................
.........................................................121
8.3 Patterns of species richness
...........................................................................................................................
...............................................123

8.3.1 Overal species richness

...........................................................................................................................
.............................................123

8.3.2 Endemic species richness

...........................................................................................................................
.........................................123

8.3.3 Threatened species richness

...........................................................................................................................
.....................................126

8.3.4 Data Deficient (DD) species richness

...........................................................................................................................
.........................126

8.4 Major threats

...........................................................................................................................
.......................................................................126

8.4.1 Biological resource use

...........................................................................................................................
..............................................131

8.4.2 Pol ution

...........................................................................................................................
......................................................................131

8.4.3 Land use change for agriculture

...........................................................................................................................
.................................132

8.4.4 Water management

...........................................................................................................................
....................................................132
8.5 Recommended research and conservation actions
...........................................................................................................................
............132

8.5.1 Research actions recommended

...........................................................................................................................
...............................132

8.5.2 Conservation actions recommended

...........................................................................................................................
.........................133

Bibliography
...........................................................................................................................
...............................................................................134

8.1 Introduction

taxonomically described species were found to be native to

the catchment, of which 422 (64%) are endemic. Adding the

In this synthesis chapter, we combine information

subset of freshwater plant species assessed gives a total

presented in the individual taxonomic chapters (Chapters

of 909 native species considered in this study, of which 423

3-7) in order to discuss the status and distribution of

(47%) are endemic. It should be noted that these values do

freshwater biodiversity across the Lake Malawi/Nyasa/

not include species that have not been formal y described,

Niassa Catchment (hereafter LMNNC). We present a

which if included could add at least an additional 400

combined analysis of our results for all described species

hundred endemic but undescribed cichlid fish species based

of freshwater decapods, fishes, molluscs and odonates

on current estimates (Konings, 2016; Snoeks, 2000).

native to the catchment, and for selected freshwater plants,

considering their relative extinction risk (Red List status)

Only one freshwater species in the LMNNC is known

and spatial distributions, and highlighting major threats

to be Extinct (EX), the endemic freshwater fish Labeo

that are impacting many species. We then recommend

worthingtoni, and no species are Extinct in the Wild (EW)

research and conservation actions that could help to

(Figure 8.1, Table 8.1). Labeo worthingtoni is a poorly known

improve the conservation status of many of these species

species that has not be recorded since its description from

and subsequently of freshwater ecosystems, habitats and

several sites around Lake Malawi/Nyasa/Niassa (hereafter

species communities as a whole within the LMNNC. We feel

LMNN) in the early 1930s (Tweddle, 2018a). However, these

that the combined information on these taxonomic groups

results potential y underestimate the true number of species

provides a reasonable representation of the status and

extinctions in the catchment. Four species (16% of all

distribution of freshwater biodiversity in the LMNNC.

Critical y Endangered (CR) species), al of which are cichlid

fishes endemic to the lake, are assessed as CR and flagged

as Possibly Extinct. Surveys are required to determine

8.2 Red List assessments

whether these species are stil extant.

Of the freshwater taxonomic groups considered in this study

Looking at species at high relative risk of extinction, 51 fresh-

that were comprehensively assessed within the LMNNC

water species (6% of assessed extant species excluding

(freshwater decapods, fishes, mol uscs and odonates), 662

those assessed as Data Deficient, DD) native to the basin

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

121

are considered threatened (Figure 8.1, Table 8.1). The true

Levels of data deficiency are also higher in endemic species,

percentage of threatened species native to the catchment

with 10% (41 species) of endemic species assessed as DD,

lies between 6% and 12% depending on the actual patterns

compared with 6% (58 species) of native species (Figure

of extinction risk within the DD species. Al of these estimates

8.1, Table 8.1). These DD species are primarily endemic

are significantly lower than for both the Lake Victoria Basin

fishes, together with some non-endemic plants, highlighting

and for continental Africa where 20% and 21% of freshwater

the need for surveys and monitoring of freshwater species

species assessed (excluding DD species) were placed into

within the lake, catchment and beyond.

a threatened category, respectively (Darwall et al., 2011;

Sayer, Máiz-Tomé & Darwal , 2018). When considering only

The majority of freshwater species in the catchment are

endemics, 41 freshwater species (11% of assessed extant

assessed as Least Concern (LC), with 84% (760 species) of

species excluding those assessed as DD) are threatened

native species and 72% (303 species) of endemic species

(Figure 8.1, Table 8.1), with the true value fal ing between 10%

placed in this category (Figure 8.1, Table 8.1). This pattern is

and 20% depending on the actual patterns of extinction risk

reflective of all of the taxonomic groups considered.

within the DD species. Conservation actions are needed

urgently to combat the threats acting within the LMNNC that

These Red List assessments, with a number of notable

have resulted in the deterioration of these species.

exceptions highlighted in the preceding chapters, paint a

relatively positive picture of the current status of freshwater

biodiversity within the LMNNC when compared with other

Table 8.1 Number of freshwater species (including all decapods,

regions, such as continental Africa (Darwall et al., 2011).

fishes, molluscs and odonates, and selected plants) native and

However, a number of threats have been identified and

endemic to the Lake Malawi/Nyasa/Niassa Catchment in each

it is still vital that actions are taken now to conserve the

Red List Category. For a list of species native to the Lake

Malawi/Nyasa/Niassa Catchment and their Red List Categories

freshwater ecosystems and species of the LMNNC into

please see Appendix 1.

the future, particularly given the high richness of endemic

Number

Number

species, many of which are undescribed and so could not

of species

of species

be considered in this study. Key plausible future threats,

native to the

endemic to the

IUCN Red List Category

LMNNC

LMNNC

such as the introduction of an alien invasive species or an

oil spill, could have significant and rapid negative effects

Extinct (EX)

on much of the freshwater species within the LMNNC,

Extinct in the Wild (EW)

0

the consequences of which could be irreversible. The

Critically Endangered (CR)

25

23

vulnerability of lake systems with endemic species flocks,

Endangered (EN)

12

such as LMNN, has been demonstrated in the Lake Victoria

Vulnerable (VU)

14

11

Basin in East Africa where introduction of the non-native

Near Threatened (NT)

39

36

Nile perch ( Lates niloticus) contributed to severe declines

in the endemic haplochromine cichlid flock of the lake

Least Concern (LC)

760

303

(Sayer, Máiz-Tomé & Darwal , 2018). The introduction of the

Data Deficient (DD)

58

41

flowerhorn cichlid to the Malili Lakes system in Sulawesi,

Total

909

423

Indonesia has had similarly severe effects on the native,

endemic freshwater fauna (Herder et al., 2012).

Final y, it should be noted that we still lack basic distribution

Endemic species

and population information for most taxonomic groups

considered in this study, as standardised lake or catchment-

wide sur veys have not been conducted to monitor

Native species

freshwater biodiversity at the species level, either at all or in

recent years. There is much evidence to support declines in

water quality and loss of natural habitats through conversion

0%

20%

40%

60%

80%

100%

to other land uses, but there are few data available to link

these environmental changes to those of the freshwater

EX

EW

CR

EN

VU

NT

LC

DD

biodiversity of the LMNNC. As a result, many of the Red

Figure 8.1 Percentage (%) of freshwater species (including

List assessments summarised in this report are based on

all decapods, fishes, molluscs and odonates, and selected

inferred declines in species populations or distributions,

plants) native and endemic to the Lake Malawi/Nyasa/Niassa

rather than those estimated on the basis of scientific data.

Catchment in each Red List Category. For a list of species

This lack of monitoring also means that real-time changes in

native to the Lake Malawi/Nyasa/Niassa Catchment and their

Red List Categories please see Appendix 1.

populations of species are not necessarily being detected.

122

There is, therefore, an urgent need for standardised surveys

and the Limpimbi River basin, the majority of which are in

of freshwater species in the LMNNC. The results of such

Malawi (Figure 8.2). The Upper Shire River and Lake

surveys could be used to update and better inform Red List

Malombe are regions of high species richness for freshwater

assessments, which in turn could be used to track trends

decapods, fishes and molluscs, but with relatively few

in the status of freshwater biodiversity in the catchment

species of odonates. High numbers of species are found for

through tools such as the Red List Index (RLI; see Chapter

all of the taxonomic groups in areas associated with specific

9), and this information can be used to inform conservation

rivers and tributaries flowing into the lake but these patterns

and development planning.

are not consistent between groups, although it should be

noted that the number of species in each taxonomic group

varies considerably and so richness is relative.

8.3 Patterns of species richness

Regions of low overall species richness are found at the

Patterns of species richness discussed in this section

edges of the LMNNC, including the north-east of the basin

consider only the distributions of assessed species of

and the Poroto Mountains in Tanzania, Kisungu National

freshwater decapods, fishes, molluscs and odonates.

Park and the headwaters of the Bua River in Malawi, and

Distribution data, beyond country presence level records,

south of Lake Chilwa in Malawi and Mozambique. Despite

are largely lacking for freshwater plant species native to the

being of relatively low richness these areas still host

LMNNC and, as a result, it was not possible to map these

between 69–156 freshwater species each (Figure 8.2).

species to the same level of detail as for the other taxonomic

The north-east of the basin in Tanzania is the only area of

groups.

the LMNNC that is consistently poor in species richness

for all of the taxonomic groups considered. However, as

8.3.1 Overall species richness

some regions of the LMNNC have been poorly sampled for

freshwater species, Mozambique in particular, this pattern

Unsurprisingly, LMNN represents the region of the catch-

may be in part a reflection of sampling effort.

ment with the highest overall richness of freshwater species,

with 590 described species mapped to occur within the

8.3.2 Endemic species richness

lake (Figure 8.2). All taxonomic groups considered included

species occurring in the lake, but LMNN was the highest

LMNN is the area of highest endemic species richness in

richness in terms of species richness only for the freshwater

the catchment, with 413 freshwater species endemic to the

fishes and molluscs, supporting 427 and 34 species,

catchment occurring in the lake (Figure 8.5). For freshwater

respectively.

decapods, fishes and mol uscs, the lake represents the area

with highest richness of endemic species, but no species

Species richness within LMNN is not uniform throughout the

of odonate endemic to the catchment are thought to be

lake, with highest richness observed within the narrow band

present in the lake. The only species of odonate endemic

of shallow, oxygenated waters along the lakeshore and

to the LMNNC ( Allocnemis maccleeryi) occurs at Ntchisi

around islands (Figure 8.3), such as Chizumulu, Likoma,

Mountain in central Malawi (Clausnitzer, 2010).

Mbenji and Namalenje (Figure 8.4). Of the species with

distributions mapped within the lake, decapods have the

The next richest areas in terms of endemic species are the

narrowest depth range (to 80 m), followed by molluscs (to

Upper Shire River and Lake Malombe, which host 54 and 62

100 m) and finally fishes, of which there are some pelagic

endemic species, respectively, primarily fishes but including

species distributed through the lake at a range of depths,

two mollusc species. Numbers of endemic species then

and other demersal species living at depths down to an

drop with the next richest areas containing only five or six

estimated 130 m. Species richness of fishes is highest in

endemic species, again primarily fishes. These areas are the

less than 100 m depth, in particular within the shallower

Lufira River basin in Tanzania, and the Kaombe and Rumphi

southern arms of the lake and within the sand and rock

River basins in Malawi.

lakeshore habitats.

Areas of the LMNNC with no endemic species according to

Outside of LMNN, catchments with the greatest richness of

the Red List data include the extreme north-west and the

freshwater species are located around the Upper Shire River

north-east of the catchment in Tanzania (with the exception

(the only outflowing river of LMNN), Lake Malombe and

of the Rutukira River, which is home to the endemic but

many of the rivers and tributaries flowing into LMNN, notably

poorly known fish Zaireichthys compactus; Tweddle, 2018)

the Songwe River basin, Nyika Plateau, Rumphi River basin,

and Kasungu National Park in Malawi (Figure 8.5). These are

basins along the western central coast of LMNN, the

also areas of general y low overal species richness (Figure

Kaombe River basin, Chia Lagoon and surrounding areas,

8.2).

123
¯
0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of freshwater species

69 - 156

157 - 163

164 - 177

178 - 183

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

184 - 590

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 8.2 Richness of freshwater species (decapods, fishes, molluscs

and odonates) in the Lake Malawi/Nyasa/Niassa Catchment based on
spatial data coded as Presence 1 (Extant). Richness data are classified
using quantiles. Mapping is to HydroBASINS and does not imply
species occur across the entire HydroBASIN (i.e. across the entirety of
Lake Malawi/Nyasa/Niassa). See Figure 8.3
for the distribution of species within Lake Malawi/Nyasa/Niassa.

124
¯

20 40

80

120

160Kilometers

Legend
Lake Malawi/Nyasa/Niassa

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

Freshwater species distributions

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 8.3 Heat map showing relative richness of freshwater species

(decapods, fishes, molluscs and odonates) in Lake Malawi/

Nyasa/Niassa based on spatial data coded as Presence 1 (Extant).

125
8.3.3 Threatened species richness

wel as the Shire River in the south of the LMNNC (Figure 8.7).

These regions all contain three or four DD species and are

Consistent with the patterns of endemic species richness,

also regions of relatively high overal species richness (Figure

the greatest richness of threatened freshwater species is

8.2).

also found in LMNN, with 42 threatened species occurring

in the lake, followed by the Upper Shire River and Lake

There are two regions in the LMNNC with no DD species

Malombe, with 10 and 11 threatened species, respectively

(Figure 8.7). The first is the Kisungu National Park in Malawi,

(Figure 8.6). LMNN includes a number of threatened fishes

which is also an area of relatively low overal species richness

and mol uscs, while only threatened fishes are found in the

(Figure 8.2). In contrast, the second region with no DD

Upper Shire River and in Lake Malombe.

species stretches along the western boundary of the LMNNC

from the southern Nyika Plateau (Figure 8.8) in the north

Other regions with relatively high numbers of threatened

to the headwaters of the South Rukuru River in the south,

species are the Lufira River basin in Tanzania, the Shire River

an area with moderate species richness (164-177 species

as it passes through Liwonde and the Phalombe River, the

overal ; Figure 8.2).

latter two both in Malawi, al of which are home to four or five

threatened species.

8.4 Major threats

8.3.4 Data Deficient (DD) species richness

Documenting threats to species is an important starting

LMNN is the area with the most DD species, with 36 DD

point for guiding conservation actions. In this section, the

species occurring in the lake (Figure 8.7), all but one of

following major threats negatively impacting freshwater

which (a decapod) are fishes. Regions with the next highest

species in the LMNNC are discussed: biological resource

richness of DD species include the rivers and tributaries

use, pollution, land use change for agriculture and poor water

flowing into the lake on its northern and western coasts, as

management. It should be noted that three of these threats

Figure 8.4 Namalenje Island in Malawi is an area of high freshwater
species richness. © Amy Palmer-Newton 126
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of endemic freshwater species

1-2

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

5 - 413

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 8.5 Richness of endemic freshwater species (decapods, fishes,

molluscs and odonates) in the Lake Malawi/Nyasa/Niassa Catchment
based on spatial data coded as Presence 1 (Extant). Richness data are
classified using quantiles. Mapping is to HydroBASINS and does not
imply species occur across the entire HydroBASIN (i.e. across the
entirety of Lake Malawi/Nyasa/

Niassa). See Figure 8.3 for the distribution of species within Lake
Malawi/Nyasa/Niassa.

127
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of threatened freshwater

species

3
Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

4 - 42

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 8.6 Richness of threatened freshwater species (decapods, fishes,

molluscs and odonates) in the Lake Malawi/Nyasa/

Niassa Catchment based on spatial data coded as Presence 1 (Extant).

Richness data are classified using quantiles. Mapping is to
HydroBASINS and does not imply species occur across the entire
HydroBASIN (i.e. across the entirety of Lake Malawi/Nyasa/

Niassa). See Figure 8.3 for the distribution of species within Lake
Malawi/Nyasa/Niassa.

128
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

Number of Data Deficient freshwater

species

3-4

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

5 - 36

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 8.7 Richness of Data Deficient (DD) freshwater species
(decapods, fishes, molluscs and odonates) in the Lake Malawi/

Nyasa/Niassa Catchment based on spatial data coded as Presence 1

(Extant). Richness data are classified using quantiles.

Mapping is to HydroBASINS and does not imply species occur across

the entire HydroBASIN (i.e. across the entirety of Lake
Malawi/Nyasa/Niassa). See Figure 8.3 for the distribution of species
within Lake Malawi/Nyasa/Niassa.

129

Figure 8.8 The southern Nyika Plateau is an area with no freshwater

species mapped and assessed as Data Deficient (DD). © Dr Thomas
Wagner (CC BY-SA 3.0)

(clearance of land, pollution and water use) are linked, being

could benefit the most species. The introduction of invasive

jointly driven by agricultural expansion to support human

alien species is a key plausible future threat to species

population growth. Other threats are recorded as impacting

within the LMNNC and could have devastating effects on

freshwater species in the catchment (Figure 8.9) but here

the freshwater community. However, it is not included in the

we focus on those that are most prevalent to species at

discussion here because at present it is only considered a

present, to highlight the threatening activities that if reduced

threat to a relatively low proportion of species – 2% of all

or managed with consideration of freshwater biodiversity

species and 4% of threatened species (Figure 8.9).

Agriculture & aquaculture

Biological resource use

Climate change & severe weather

Energy production & mining

Human intrusions & disturbance

Invasive and other problematic species, genes & diseases

Natural system modifications

Pol ution
Residential & commercial development

Transportation & service corridors

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

Threatened species

All species

Figure 8.9 Percentage (%) of freshwater species (decapods, fishes,

molluscs, odonates and plants) in the Lake Malawi/Nyasa/

Niassa Catchment with each threat, coded to high level threats of the
IUCN Red List Classification Scheme. Results are displayed separately
for threatened species (red) and all species (blue).

130
8.4.1 Biological resource use

through the direct removal of individuals leading to species

mortality (for example as plant species are intentionally

Biological resource use, including harvesting of aquatic

gathered for use) or indirectly due to the consequent

resources, wood and plants, is the most frequently recorded

degradation and loss of species habitats. The former is

threat to freshwater species in the LMNNC, affecting 44% of

primarily a threat to freshwater plants in the catchment, while

all species. However, it is much more frequently recorded as

the latter threatens the odonates.

a threat to threatened species, affecting 86% of threatened

species, which identifies biological resource use as a key

8.4.2 Pollution

driver towards species extinctions in the catchment (Figure

8.9). It is predominantly fishing and harvesting of aquatic

The next most frequently reported threat to freshwater

resources that represent the major drivers of this threat in

species in the LMNNC is pollution, which affects 25% of

the LMNNC.

all species and 35% of all threatened species (Figure 8.9).

Pollution enters the riverine and lacustrine systems of the

Harvesting takes place at both small (artisanal) and large

LMNNC in many forms: as agricultural and forestry effluents,

(commercial /industrial) scales, with species caught

including nutrient loads, herbicides and pesticides, and

intentional y or accidental y as bycatch. Smal -scale targeted

sediments; as domestic sewage and waste water from

harvesting is the primary threat and mainly affects freshwater

urban areas; and as industrial effluents, including those from

fishes. The freshwater fishes of LMNN support a fishery

mining. The primary reported sources of pol ution threatening

(Figure 8.10) that is of great importance to the livelihoods of

freshwater species in the LMNNC are agricultural and forestry

communities, nutritional wellbeing, and the economies of

effluents and domestic and urban waste water.

countries in the LMNNC, with fishes harvested primarily for

use as food (19% of al species) or for the ornamental trade

Increasing use of marginal land for agriculture, to sustain

(39% of all species). The threat that this fishery poses is

the growing human population, is leading to higher levels

discussed in detail in Chapter 4.

of soil erosion in the LMNNC (Bootsma & Jorgensen,

2005). This in turn leads to increased sedimentation in

On the other hand, freshwater molluscs in LMNN are

water bodies, threatening fishes and mol uscs in particular

primarily threatened as an unintended consequence of

through degradation of habitat. For example, the algae

destructive fishing methods or bycatch, rather than by direct

grazing cichlid fish Metriaclima usisyae (Figure 8.12) (CR)

targeting. For example, submerged Vallisneria aethiopica

is negatively affected by sedimentation, which reduces

plants, which are the habitat of Bulinus succinoides, are

light penetration in the water column, resulting in less

damaged and uprooted by use of seine nets by fishers

algal growth, and directly smothers algae growing on the

(Albrecht et al., 2018), and dredging by fishers can disturb

rocks (Konings, 2018). Many other cichlid fishes within

the soft substrates where Lanistes nasutus occurs or result

this group of algae grazing species, commonly known as

in accidental col ection of molluscs from the sediment, which

mbuna, may similarly suffer impact. Other agricultural and

are then dumped on beaches where they die (Albrecht &

forestry effluents, such as herbicides and pesticides, were

Clewing, 2018).

not commonly recorded as threats, most likely due to a

lack of water quality monitoring. However, as deforestation

Logging of wood (Figure 8.11) and gathering of plants also

continues and landscapes change, the ease of these

threaten freshwater species. Impacts are realised either

pol utants reaching water bodies will increase.

Figure 8.10 A fisherman on a dugout canoe on Lake Malawi/

Figure 8.11 Firewood being sold at a roadside in Malawi. © Thies

Nyasa/Niassa. © Denis Tweddle

Geertz

131

Figure 8.12 A male Metriaclima usisyae at Mara Rocks. This

Figure 8.13 A maize field being worked in Lilongwe, Malawi. ©

species is negatively affected by sedimentation and is

Lars Plougmann (CC BY-SA 2.0)

assessed as Critically Endangered (CR). © Ad Konings

Pollution from domestic sources is a key threat to mol uscs

in rivers, and can also block passage of migratory species.

and odonates in the LMNNC. Levels of domestic pol ution,

This is a potential future threat to the migratory mpasa

particularly sewage and solid waste, are highest in the lake

( Opsaridium microlepis) (VU) (Tweddle, 2018c).

margins and in the most heavily populated areas, such as the

southern part of the lake (Kafakoma, 2019), corresponding

to the areas of greatest species richness (Figure 8.3). These

8.5 Recommended research and

pollutants both degrade habitats and lead to damage or

conservation actions

mortality in freshwater species if ingested.

Recommended research and conservation actions are

8.4.3 Land use change for agriculture

documented as part of Red List assessments, representing a

good starting point for guiding relevant conservation

Agriculture and aquaculture are recorded as threats to 16%

strategies. The research and conser vation actions

of all species and 18% of threatened species (Figure 8.9).

recommended for freshwater biodiversity native to the

Around 80% of people living within the LMNNC rely on

LMNNC are shown in Figure 8.14.

agriculture for subsistence (Bootsma & Jorgensen, 2005),

with a focus on cultivation of non-timber crops (Figure

8.5.1 Research actions recommended

8.13), both at small and agro-industry scales (Chairuca,

2016; Chavula, 2016; Faraji, 2016). Forest clearance for

Monitoring and research are the two most frequently

agriculture, in addition to the impacts of sedimentation

recommended actions, with monitoring recommended for

mentioned above, can also impact the functioning of

41% of all species and 67% of threatened species (Figure

freshwater systems, including through biogeochemical,

8.14), and further research recommended for 34% of all

thermal and hydrological changes (Allan, 2004). In this

species and 39% of threatened species. It has become

study, it is mainly odonates that are threatened directly by

apparent through this study that there is insufficient

agriculture, through loss of habitat.

information on the distributions, populations and ecology of

many freshwater species in the LMNNC. A reliable

8.4.4 Water management

information baseline on freshwater species must be

established, maintained and monitored in the future. For

Natural system modifications were recorded as a threat

fisheries, it is recommended that this is established through

to 10% of both all and threatened species (Figure 8.9).

compilation of coordinated catch and effort data from

This is driven in part by poor water management and use,

fisheries around LMNN. These data could be used both to

primarily affecting odonates that are negatively impacted

better assess the status of fisheries and to better inform Red

by abstraction of ground and surface water for agricultural

List assessments of important fishery species, such as

use. One such species is the Mulanje damsel ( Oreocnemis

chambo ( Oreochromis spp.) and kampango ( Bagrus

phoenix) (EN), which is threatened by drainage of its swamp

meridionalis).

habitats (Clausnitzer, 2018). Construction of dams is another

driver and this is a more significant threat to fishes. Dam

In many cases, the lack of information can be addressed

construction results in changed hydrology and flow regimes

through field surveys (Figure 8.15). In particular there is

132

Conservation Planning

Monitoring

Research

Research

Education & awareness

Land/water management

Land/water protection

Law & policy

Conservation Livelihood, economic & other incentives

Species management

0%

10%

20%

30%

40%

50%

60%

70%

Threatened species

All species

Figure 8.14 Percentage (%) of freshwater species (decapods, fishes,

molluscs, odonates and plants) in the Lake Malawi/Nyasa/

Niassa Catchment with each recommended conservation or research

action, coded to high level conservation or research actions of the IUCN
Red List Classification Scheme. Results are displayed separately for
threatened species (red) and all species (blue).

an urgent need to resume standardised surveys of the

capacity to do this work, there is a high risk that a number of

freshwater biodiversity of LMNN in Malawi and to extend this

these species wil deteriorate in status before they have been

into Tanzania and Mozambique. A focus for future survey

described and assessed.

and monitoring should be the sub-basins with high numbers

of DD species highlighted in Figure 8.7 and those with high

8.5.2 Conservation actions recommended

numbers of threatened species highlighted in Figure 8.6.

Land and water management was identified as the priority

There is a similarly urgent need for formal taxonomic

action, recorded for 14% of al species and 24% of threatened

description of the many still undescribed species of the

species (Figure 8.14). In most cases, management actions

LMNN, such as several hundred cichlid fishes endemic to

should be targeted at the catchment scale, employing

the lake (Konings, 2016; Snoeks, 2000). It is possible to

methods such as Integrated River Basin Management

assess undescribed species for the IUCN Red List but a

(IRBM) or Environmental Flows (E-Flows) due to the high

number of conditions need to be met, including there being

levels of hydrological connectivity within the catchment.

a conservation benefit of assessing the species and work

Given the low turnover rate for LMNN, pol utants originating

being underway to describe them (IUCN Standards and

from across the catchment will remain and accumulate

Petitions Subcommittee, 2017). Owing to the large number

within the system. These pollutants may periodically be

of undescribed species within the basin and the limited

released into the upper water column due to seasonal

upwelling, particularly in the southern arms of the LMNN

where they may seriously impact water biodiversity and have

occasional y led to large scale fish kil s.

Land and water protection was also frequently recommended,

particularly for threatened species (22%) (Figure 8.14). Key

Biodiversity Areas (KBAs) are sites contributing to the global

persistence of biodiversity and were identified through this

study for freshwater species (e.g. Figure 8.16; see Chapter

10). It is recommended that the identified KBAs are used

as a basis for the expansion of the protected area network

to better represent and conserve freshwater biodiversity.

Where freshwater species of conservation priority are

present in existing protected areas or other management

units it is important that their presence is communicated to

site managers, and that strategies for their conservation are

Figure 8.15 A field survey on the Shire River in Malawi. © Denis

Tweddle

incorporated into existing management plans.

133
Figure 8.16 Otter Point is part of the Cape Maclear Key Biodiversity
Area (KBA) in Malawi. © Hans Hil ewaert (CC BY-SA 4.0) Species
management was also frequently recommended

policies needed to ensure sustainable development and

for threatened species (20%) (Figure 8.14). The main focus is

conservation of freshwater biodiversity within the LMNNC.

for management of harvest and trade, which is of particular

relevance to the fisheries of LMNN for both human food and

ornamental use. Without enforced fisheries regulation there

Bibliography

has already been a significant decline in many important fish

stocks (see Chapter 4) and there is significant risk of further

Albrecht, C. and Clewing, C. (2018). Lanistes nasutus . The IUCN

widespread fisheries collapse, with potentially disastrous

Red List of Threatened Species 2018: e.T11286A120111577.

consequences for local livelihoods and the economies of the

Available at: http://dx.doi.org/10.2305/IUCN.UK.2018-2.

three riparian countries of the LMNNC.

RLTS.T11286A120111577.en (Accessed: 26 June 2019).

Albrecht, C., Clewing, C., Kahwa, D., Magombo, Z.L.K.,

Final y, education and awareness raising are highlighted as

Kaunda, E. and Mailosi, A. (2018). Bulinus succinoides .

being important for 3% of al species and 16% of threatened

The IUCN Red List of Threatened Species 2018:

species (Figure 8.14). In addition to raising awareness

e.T3320A122888722. Available at: http://dx.doi.

on the unique endemic assemblages, it is important to

org/10.2305/IUCN.UK.2018-2.RLTS.T3320A122888722.

communicate widely the benefits of clean and healthy

en (Accessed: 26 June 2019).

wetland systems that support high freshwater species

Allan, J.D. (2004). ‘Landscapes and Riverscapes: The

diversity and associated livelihoods. Both within and

Influence of Land Use on Stream Ecosystems’, Annual

outside the LMNNC, wetlands are often seen as wasted

Review of Ecology, Evolution, and Systematics,

land, and therefore a site for dumping waste products,

35(1), pp. 257–284. https://doi.org/10.1146/annurev.

or as the source of problematic disease vectors, such as

ecolsys.35.120202.110122

mosquitoes (Smith et al., 2014). It is vital that the importance

Bootsma, H.A. and Jorgensen, S.E. (2005). Lake Malawi/

of wetland systems is shared with a number of target groups,

Nyasa: Experiences and Lessons Learned. Kusatsu,

including local communities, private sector developers and

Japan: International Lake Environment Committee

government departments. It is of course government that

Foundation, Lake Basin Management Initiative: Experience

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Chairuca, L. (2016). Mozambique National Report. IUCN,

Konings, A. (2016). Malaŵi cichlids in their natural habitat. El

Building River Dialogue and Governance Phase 3

Paso, Texas: Cichlid Press.

(BRIDGE-3): 2016-2018.

Konings, A. (2018). Metriaclima usisyae . The IUCN Red List

Chavula, G.M.S. (2016). Malawi: Country report on the

of Threatened Species 2018: e.T120732502A120732518.

state of the environment and outlook report for Lake

Available at: http://dx.doi.org/10.2305/IUCN.UK.2018-2.

Malawi/Nyasa/Niassa. IUCN, Building River Dialogue and

RLTS.T120732502A120732518.en (Accessed: 27 June Governance Phase 3

(BRIDGE-3): 2016-2018.

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Clausnitzer, V. (2010). Al ocnemis maccleery . The IUCN Red

Sayer, C.A., Máiz-Tomé, L. and Darwall, W.R.T. (2018).

List of Threatened Species 2010: e.T180563A7648402.

Freshwater biodiversity in the Lake Victoria Basin:

International Union for Conservation of Nature. Available

Guidance for species conservation, site protection, climate

at: https://www.iucnredlist.org/species/180563/7648402

resilience and sustainable livelihoods. Cambridge, UK and

(Accessed: 4 May 2019).

Gland, Switzerland: IUCN. https://doi.org/10.2305/IUCN.

Clausnitzer, V. (2018). Oreocnemis phoenix . The IUCN Red

CH.2018.RA.2.en

List of Threatened Species 2018: e.T59947A75437856.

Smith, K.G., Barrios, V., Darwall, W.R.T. and Numa, C. (2014).

International Union for Conservation of Nature. Available

The status and distribution of freshwater biodiversity in

at: https://www.iucnredlist.org/species/59947/75437856
the Eastern Mediterranean. Cambridge, UK, Malaga,

(Accessed: 24 April 2019).

Spain and Gland, Switzerland: IUCN, The IUCN Red List

Darwall, W.R.T., Smith, K.G., Allen, D.J., Holland, R.A.,

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in African freshwaters: underwater, under threat. Gland,

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on the state of the environment and outlook report for Lake

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Malawi/Nyasa/Niassa. IUCN, Building River Dialogue and

Tweddle, D. (2018a). Labeo worthingtoni . The IUCN Red List

Governance Phase 3 (BRIDGE-3): 2016-2018.

of Threatened Species 2018: e.T57507878A58341426.

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S., McKinnon, J., Walter, R. and Pfaender, J. (2012).

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‘Alien invasion in Wal ace’s Dreamponds: records of the

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hybridogenic “flowerhorn” cichlid in Lake Matano, with

Tweddle, D. (2018b). Zaireichthys compactus . The IUCN Red

an annotated checklist of fish species introduced to the

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Malili Lakes system in Sulawesi’, Aquatic Invasions, 7(4),

Available at: http://dx.doi.org/10.2305/IUCN.UK.2018-2.

pp. 521–535. https://doi.org/10.3391/ai.2012.7.4.009

RLTS.T167987A58317895.en (Accessed: 27 June 2019).

IUCN Standards and Petitions Subcommittee (2017).

Tweddle, D. (2018c). Opsaridium microlepis : The IUCN Red

Guidelines for Using the IUCN Red List Categories and

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Criteria. Version 13.

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Kafakoma, R. (2019). State of the Lake Malawi/Nyasa/Niassa

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Ecosystem., Building River Dialogue and Governance

Phase 3 (BRIDGE-3): 2016-2018.

135

Chapter 9

Monitoring trends in the status of

freshwater biodiversity within the

Lake Malawi/Nyasa/Niassa Catchment:

the IUCN Red List Index

Sayer, C.A.1

Contents

9.1 Introduction
...........................................................................................................................
.........................................................................136

9.2 Method
...........................................................................................................................
................................................................................136

9.2.1 Calculation
...........................................................................................................................
..................................................................136

9.2.2 Red List Indices (RLIs) for the Lake Malawi/Nyasa/Niassa Catchment
................................................................................................137

9.3 Results
...........................................................................................................................
................................................................................138
9.4 Discussion
...........................................................................................................................
...........................................................................140

Species in the spotlight

...........................................................................................................................
..............................................................140

Genuine changes in Red List category

...........................................................................................................................
...............................140

Bibliography
...........................................................................................................................
...............................................................................142

9.1 Introduction

group being investigated must have been assessed for the

IUCN Red List at least twice in order to calculate the RLI. The

The IUCN Red List Index (RLI) is used to measure trends

RLI is calculated from the number of species in each Red List

in the overall extinction risk of groups of species, as an

category and the number of species changing categories

indicator of trends in the status of biodiversity (Bubb et

between assessments as a result of genuine improvement

al., 2009). Extinction is a key measure of biodiversity loss

or deterioration in status (i.e. genuine changes). Changes

and, as a result, the RLI has been adopted as a biodiversity

in category resulting from improved knowledge or revised

indicator by a number of international conservation policies

taxonomy (i.e. non-genuine changes) are excluded (Bubb et

and agreements. For example, the global RLI has been used

al., 2009).

to track progress towards the Convention on Biological

Diversity’s (CBD) 2010 Biodiversity Targets and Aichi

The RLI can be calculated using Equation 9.1 (Butchart et al.,

Biodiversity Targets, and the Millennium Development

2007):

Goals (MDGs), while subsets of the RLI have been used

ΣW

to track progress under various multilateral environmental

RLI = 1 – s c (t,s)

agreements, such as the Ramsar Convention and the

WN

Convention on Migratory Species (CMS) (Bubb et al., 2009;

EX

Butchart et al., 2005, 2010; Tittensor et al., 2014). The RLI is

Equation 9.1 Equation to calculate the IUCN Red List Index

also the official indicator for the Sustainable Development

(RLI) following Butchart et al. 2007.

Goals (SDGs) Target 15.5.

Where W

is the weight of category c for species s at time

c(t,s)

9.2 Method

t, W is the weight for the category Extinct (EX), and N is the

EX

number of assessed species excluding those considered

9.2.1 Calculation

Data Deficient (DD) in the current time period and those

considered to be EX in the year the set of species was

The RLI is based upon the categories of species extinction

first assessed. The category weights ( c) used are: Least

risk as published on the IUCN Red List. Al species within the

Concern (LC), 0; Near Threatened (NT), 1; Vulnerable (VU),

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK
136

2; Endangered (EN), 3; Critical y Endangered (CR), 4; and CR

Freshwater crabs were assessed by the IUCN Species

(Possibly Extinct) (CR(PE)), CR (Possibly Extinct in the Wild)

Survival Commission ( SSC ) Freshwater Crustacean

(CR(PEW)), Extinct in the Wild (EW) and EX, 5.

Specialist Group in both 2004 and 2008. The species native

to the LMNNC were re-assessed through this study in 2017

In simple terms, to calculate the RLI the number of species

(see Chapter 3). The RLI for freshwater crabs, therefore,

in each Red List category is first multiplied by the category

compares the status of these species in 2004, 2008 and

weight. These products are then summed and divided by

2017. One species ( Potamonautes bellarussus) had its Red

the maximum possible product (the number of species

List category back-cast to 2004 and 2008 because it was

multiplied by the maximum weight) and then subtracted from

only described in 2014 (Daniels, Phiri & Bayliss, 2014) and

1. The index produced can take any value from 0 to 1.

assessed for the first time in 2018. Six species are included
in this RLI.

A RLI value of 1 indicates that all species are LC, whereas

a RLI of 0 indicates that all species are EX. Declines in RLI

Freshwater shrimps were assessed in 2012, with the results

values over time indicate that the expected risk of extinction

published by De Grave et al. (2015), and the species native

is increasing, unchanging RLI values indicate that the

to LMNNC were re-assessed through this study in 2017

expected risk of extinction is remaining the same, and

(see Chapter 3). The RLI for freshwater shrimps, therefore,

increases in RLI values over time indicate that the expected

compares the status of these species in 2012 and 2017. All

risk of extinction is decreasing.

species had published assessments at both time points.

Three species are included in this RLI.

It is possible to disaggregate global RLIs to show trends

at finer scales, for example at national or regional scales.

Freshwater fishes and molluscs were assessed most

RLIs at sub-global scales can either be based on global

recently through this project (see Chapters 4 and 5,

or regional Red List assessments. If considering global

respectively) and previously as part of the assessment of

assessments then it is necessary to assess for each species

freshwater biodiversity across continental Africa by Darwall

within that region that underwent a genuine change in

et al. (2011). The freshwater fish and mollusc assessments

its status (as indicated by movement between Red List

conducted for this project were completed over two years

categories) whether the processes driving this change also

(2017 and 2018) and were reviewed in 2018. Therefore, 2018

occurred within the region (Bubb et al., 2009).

was chosen as the most recent time point for assessment.

The assessments presented in Darwall et al. (2011) were

9.2.2 Red List Indices (RLIs) for the Lake

completed over a number of years (2003–2009) but al were

Malawi/Nyasa/Niassa Catchment

reviewed in 2009. Therefore, 2009 was chosen as the

previous time point for assessment. The RLIs for freshwater

Individual RLIs for the Lake Malawi / Nyasa / Niassa

fishes and mol uscs, therefore, compare the status of these

Catchment (hereafter LMNNC) were calculated for each

groups in 2009 and 2018. Red List categories were back-cast

taxonomic group separately because the time frames

to 2009 for eight mollusc and 127 fish species. This was

of the Red List assessments differed across groups. It

necessary due to a combination of recent taxonomic

should be noted that this disaggregation to individual

changes (primarily for the molluscs) or because, although the

taxonomic groups reduces the sample sizes and, therefore,

species had been assessed at time points in the past, none

the robustness of the trends.

of these assessments were conducted within the previous

assessment period (2003–2009). The RLIs for freshwater

Freshwater plants were excluded from RLI calculation

fishes and molluscs consider 459 and 38 species,

because the Red List assessments of this group are not

respectively.

complete for all species native to the basin (see Chapters 2

and 7).

The majority of odonates native to the LMNNC were

assessed most recently in 2016, with the results published in

It is possible to back-cast (i.e. retrospectively adjust or

Sayer et al. (2018), and previously as part of the assessment

assign) Red List categories for newly added species, species

of freshwater biodiversity across continental Africa by

that have undergone non-genuine changes in Red List

Darwall et al. (2011). The first time point was chosen as

category, or species that were previously DD but have since

2009, as described above for Darwall et al. (2011), and the

been assigned a category that allows for their inclusion in

second time point was chosen as 2016. Therefore, the RLI

RLI calculations (Butchart et al., 2007). Red List categories

for odonates compares the status of these species in 2009

for 166 species were back-cast by the Red List assessors

and 2016. Thirty-two odonates had their Red List categories

involved in this study because they did not have published

back-cast to 2009 because, as for the fishes and mol uscs,

assessments for the start of the comparison period.

although the species had been assessed at time points in

137
the past, none of these assessments were conducted within

9.3 Results

the previous assessment period (2003–2009). This RLI

considers 155 species.

The RLIs for each taxonomic group are displayed in Figure

9.1 for ease of comparison.

A number of the freshwater fishes, molluscs and odonates

have been assessed at other time points outside of the

Figure 9.1 shows that the taxonomic group facing the highest

respective assessment periods discussed above. However,

expected rate of biodiversity loss within the LMNNC is the

these time periods are not included in the RLI calculations

fishes, with the steepest decline in RLI values (from 0.95 in

because the assessments were not comprehensive for each

2009 to 0.93 in 2018). Twenty-two fish species were uplisted

taxonomic group.

(i.e. moved to a category indicating higher relative extinction

0.98

0.96

0.94

RLI)

0.92

ndex (

ist I

Red L

0.9
0.88

0.86

0.84

2002

2004

2006

2008

2010

2012

2014

2016

2018

2020

Year

Crabs

Fishes

Mol uscs

Odonates

Shrimps

Figure 9.1 IUCN Red List Index (RLI) of species survival for freshwater
species in the Lake Malawi/Nyasa/Niassa Catchment: i) crabs (blue,
N=6); ii) fishes (red, N=417); iii) molluscs (purple, N=37); iv) odonates
(yellow, N=155); and v) shrimps (green, N=1).

N refers to the number of non-Data Deficient (DD) and extant species in

the taxonomic group in the first year of assessment. An RLI value of 1.0
equates to all species being categorised as Least Concern (LC), and
hence that none are expected to go extinct in the near future. An RLI
value of zero indicates that all species have gone Extinct (EX). Source of
Icons from the Noun Project (www.thenounproject.com): Crab by Arief
Sugiyanto; Fish by ruliani; snails by BGBOXXX; Dragonfly by Matt
Hawdon; and Shrimp by mas kamal.

138
risk; e.g. Figure 9.2) between 2009 and 2018 due to genuine

in species found in the deep lake. However, a systematic

deteriorations in their populations (see Appendix 2). This

survey is required to support this conclusion (D. Van Damme

was out of the 417 species considered (excluding one EX

and C. Albrecht pers. comm. 2019).

and 41 DD species in 2009). Fishing represents the primary

threat to freshwater fishes in the LMNNC, either through

The crabs and shrimps are the least threatened groups in

direct harvesting or indirect effects, such as degradation of

the LMNNC and their RLI values show no change in their

habitats caused by fishing methods (see Chapter 4). Four

expected risk of extinction over the time periods considered.

of the species that have undergone genuine deteriorations

The RLI value for crabs was 1.00 in 2004, 2008 and 2017, and

in their populations are highlighted in Species in the

the value for shrimps was 1.00 in both 2012 in 2017 (Figure

spotlight: Genuine changes in Red List category p. 140.

9.1). It should be noted that these RLIs are based on very

small numbers of species with only six crabs and a single

Mol uscs are the most threatened group in the LMNNC, with

shrimp (excluding two DD species) considered. However,

the lowest RLI values, and their expected rate of biodiversity

this stable trend is thought to be a realistic representation

loss is the next greatest (with a RLI value of 0.88 in 2009

of the status of freshwater decapods in the LMNNC (N.

and 0.87 in 2018; Figure 9.1). One species (out of the 37

Cumberlidge pers. comm. 2019) with the majority of species

species considered and excluding one DD species in 2009;

having widespread distributions, including occurring beyond

see Appendix 2) of mollusc ( Bellamya robertsoni; Figure

the LMNNC, and no major threats identified.

9.3) was uplisted from EN (back-cast) in 2009 to CR in 2018

due to a genuine deterioration in its population resulting

The odonates show a slightly greater level of threat than the

from habitat loss and degradation (D. Van Damme pers.

crabs and shrimps but no change in expected extinction

comm. 2019). However, this change is based on inferred

risk over the time period considered. The RLI value for

information. The last systematic col ection of mol uscs in the

odonates was 0.99 in both 2009 and 2016 based on the 155

catchment was conducted by Mandahl-Barth in the 1970s

species native to the LMNNC (of which no species were

(Mandahl-Barth, 1972). Mandahl-Barth’s data, together with

DD). However, this stable trend is not considered a realistic

the work of Brown (1994), were the primary sources for the

representation of the status of odonates in the LMNNC. The

original assessments of the LMNNC mol uscs in Darwal et

availability and quality of habitats for odonates is in decline

al. (2011). Additional col ections have been made in the 21st

due to rapid population growth and associated agriculture,

century, primarily by Albrecht, Schultheiß and Genner, but

urbanisation and industry in the LMNNC. This deterioration

these col ections have focussed on specific genera or parts

in habitat is leading to population declines in species, but at

of Lake Malawi/Nyasa/Niassa (hereafter LMNN). These

present these are not of great enough magnitude to result

additional col ections were used to inform the assessments

in Red List category changes and, therefore, not reflected

published through this study. It is thought that there is a

in the RLI. A number of species also occur in reserves and

negative trend in the mol uscs of the LMNNC that is stronger

so parts of their habitat are protected (V. Clausnitzer & K.-D.

than indicated by the RLI values and particularly severe

Dijkstra pers. comm. 2019).

Figure 9.3 Bellamya robertsoni was uplisted from a back-cast

category of Endangered (EN) in 2009 to Critically Endangered

Figure 9.2 Pseudotropheus brevis was uplisted from Least

(CR) in 2018. © Naturalis Biodiversity Center Wikimedia Commons

Concern (LC) in 2006 to Endangered (EN) in 2018. © Ad Konings

(CC0)

139

9.4 Discussion

may rely on remotely sensed data sets, such as for land cover

changes.
Overall, the RLIs suggest that freshwater biodiversity

in the LMNNC is in decline and that the risk of species

Despite the unique nature and the importance of the

extinctions is increasing, in particular for the fishes. It is

freshwater species of the LMNNC, particularly of LMNN

therefore vital that conservation actions are implemented to

itself, we currently lack essential basic information on the

stop and reverse these declines where possible. Relevant

distribution and population for most of the taxonomic groups

conservation actions are detailed in the chapters for each

considered. Standardised lake or basin-wide surveys have

taxonomic group (see Chapters 3–7).

not been conducted, either at all or in recent years, and there

are no significant long-term programmes for monitoring

RLIs and the trends they depict are only as good as their data

the state of aquatic biodiversity throughout LMNN and its

inputs. Red List assessments are considered scientifically

catchment. There is much evidence for declines in water

robust as they follow a standardised method, are based

quality and loss of natural habitats through conversion

on quantitative criteria and use the best scientific data

to other land uses, but there are few data available to

available. Red List assessments also undergo a thorough

determine the impact of these environmental changes on

review process before publication. However, Red List

the freshwater species themselves. Consequently, many of

assessments for a point in time may be revised, for example

the Red List assessments are based on inferred declines in

as knowledge of species and their habitats increases,

species populations or distributions, rather than scientific

resulting in changes to the Red List categories assigned.

monitoring data. This lack of monitoring means that real-

Additional y, the Red List categories are broad in nature with

time changes in the status of freshwater biodiversity are

wide thresholds for moving between categories and, as a

not being detected. There is, therefore, an urgent need

result, RLIs should be considered only a coarse measure of

for standardised surveys of freshwater biodiversity in the

changes in the status of biodiversity over time. It should also

LMNNC, combined with the setup of long-term monitoring

be recognised that time lags often occur between changes

stations. These surveys and monitoring programmes

in the real-life situation of a species, detection of these

must identify individuals at the species level if we are to

change, and incorporation of these changes into Red List

have sufficient information to manage and conserve the

assessments (Bubb et al., 2009). Finally, in the absence of

global y unique diversity of species living in this basin. The

regular monitoring, changes in threats to species are often

results of these surveys can be used to better inform Red

hard to detect and their impacts hard to quantify over the

List assessments, which can in turn be used to help track

time periods used here to calculate RLIs. In such cases we

trends in the status of freshwater biodiversity in the LMNNC

through use of tools such as the RLI.

Species in the spotlight

Genuine changes in Red List category

Sayer, C.A.1

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ
Kampango ( Bagrus meridionalis)

Bagrus meridionalis (Figure 9.4), or kampango as it is

local y known, occurs in LMNN, the Upper Shire River and

Lake Malombe. It is an important food fish and is caught

primarily using gill nets and longlines with the majority

of harvesting occurring in the southern part of LMNN in

Malawi, although there is also a relatively new fishery in

Mozambique (Phiri et al., 2018). This species was assessed

as LC in 2006 (Kazembe, 2006a). However, since the

previous assessment, declines in utaka (open water-

dwelling cichlids) stocks have meant that commercial

Figure 9.4 A pair of kampango ( Bagrus meridionalis) at

Thumbi West Island in Malawi. This species was uplisted

fishers have increasingly started to target kampango. This

from Least Concern (LC) in 2006 to Critically Endangered

has resulted in an estimated population decline of 90%

(CR) in 2018. © Ad Konings

140
over the last decade, supporting uplisting and an assessment of CR in 2018
(Phiri et al., 2018). Kampango is, however, widespread throughout LMNN
including in many of less heavily fished areas for which data on population
status are currently unavailable. Impacts on kampango are, therefore, likely
to be localised, but provide a warning for the future of more widespread
effects from fishing. The 2018 assessment of CR is a precautionary
assessment based on the data currently available.

Corematodus shiranus

Corematodus shiranus (Figure 9.5) is a predator that specialises on eating

scales from the caudal peduncle and tails of Oreochromis (chambo) species
(e.g. Figure 9.6). It closely resembles its prey such that even chambos are
unable to distinguish between this predator and their conspecifics (Konings,
2016) and it may be caught unnoticed by fishers with its prey (Konings,
2018a). This species was assessed as LC in 2006 (Kazembe & Makocho,
2006). However, over the last ten years, populations of its prey species have
declined by 94% and al chambo species are now considered CR (Kanyerere,
Phiri & Shechonge, 2018; Konings, 2018a, 2018b; Phiri & Kanyerere,
2018). As a consequence, Corematodus shiranus was uplisted to CR in 2018
with declines suspected to match that of its prey (Konings, 2018a).

Figure 9.5 Corematodus shiranus was uplisted from Least

Concern (LC) in 2006 to Critically Endangered (CR) in 2018

based on a decline in its Oreochromis (chambo) species

prey. © George Turner

Figure 9.6 Oreochromis karongae (Critically Endangered,

CR) is predated on by Corematodus shiranus. © George

Turner

Melanochromis chipokae

Melanochromis chipokae (Figure 9.7) is known only from

Chindunga Rocks (near Chipoka) and another small reef

in the south-western arm of LMNN (Konings, 2018c). This

species was assessed as VU in 2006 (Kazembe, 2006b).

Very few individuals of this species have been observed

over the last decade. In 2017, a breeding and restocking

program was initiated and 68 individuals were released at

Chidunga Rocks. However, in October of the same year

no individuals of the species could be found at the release

site. This species is popular in the ornamental fish trade

and it is suspected that the reintroduced individuals were

col ected for this trade. Based on its restricted distribution

and past population decline due to the ornamental fish

Figure 9.7 Melanochromis chipokae was uplisted from

trade, this species was uplisted to CR in 2018 (Konings,

Vulnerable (VU) in 2006 to Critically Endangered (CR) in 2018.

2018c).

© Ad Konings

141
Sungwa ( Serranochromis robustus)

Serranochromis robustus (Figure 9.8), or sungwa as it is

local y known, is found in LMNN, Lake Malombe and the

lower reaches of inflowing rivers. Despite the species nearly

being extirpated from Lake Malombe and the Shire River in

the 1970-1980s due to destruction of its weeded habitats

by seine netting, it was assessed as LC in 2006 based on

its widespread overall range (Kazembe, 2010). However,

this threat is continuing and a new threat to this species

has emerged. Based on the impacts of the disease on its

congener S. jal ae in the Zambezi and Okovango systems,

this species is considered highly vulnerable to epizootic

ulcerative syndrome (EUS). Therefore, in 2018 this species

Figure 9.8 Sungwa ( Serranochromis robustus) at Gome in

was uplisted to CR based on expert opinion on past and

Malawi. This species was uplisted from Least Concern (LC)

future population declines (Konings & Tweddle, 2018).

in 2006 to Critically Endangered (CR) in 2018. © Ad Konings

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Sites

Shire River, Malawi. © Denis Tweddle

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144

Chapter 10

Freshwater Key Biodiversity Areas in the

Lake Malawi/Nyasa/Niassa Catchment

Sayer, C.A.1 and Darwal , W.R.T.1

Contents

10.1
Background.......................................................................................................
............................................................................................145

10.2 Methodology
...........................................................................................................................
......................................................................146

10.2.1 KBA criteria and thresholds

...........................................................................................................................
......................................146

10.2.2 Freshwater KBA delineation process

...........................................................................................................................
........................147

10.3 Results
...........................................................................................................................
...............................................................................151

10.3.1 Freshwater KBA trigger species

...........................................................................................................................
...............................151

10.3.2 Freshwater KBAs overview

...........................................................................................................................
.......................................151

10.3.3 Current levels of management and protection

...........................................................................................................................
.........151

10.3.4 Site champions

...........................................................................................................................
........................................................155
10.4
Recommendations.............................................................................................
...........................................................................................155

10.5 Next steps

...........................................................................................................................
..........................................................................157

Sites in the spotlight

...........................................................................................................................
...................................................................157

The Lake Malawi National Park World Heritage Site

...........................................................................................................................
...........158

Mpasa and the Lower Bua and Nkhotakota Wildlife Reserve

.......................................................................................................................16
0

Restoring landscapes and conserving biodiversity in Malawi

........................................................................................................................1
62

A unique fish fauna benefiting from the Nyika National Park, Vwaza Marsh
Wildlife Reserve and North Rumphi Key Biodiversity Areas ........166

Bibliography
...........................................................................................................................
...............................................................................170

10.1 Background

to this resolution (WCC 3.013), the IUCN Species Survival

Commission (SSC) and the IUCN World Commission on

Over the last four decades, a number of organisations

Protected Areas (WCPA) established a Joint Task Force

have invested in compiling information on the location

on Biodiversity and Protected Areas, which since 2012

of sites that are significant for biodiversity. Since the late

has mobilised expert input from IUCN commissions,

1970s, BirdLife International has maintained criteria for the

members, secretariat staff, conservation organisations,

identification of Important Bird Areas (IBAs) and over 13,400

academics, decision-makers, donors and the private

sites have been identified worldwide (BirdLife International,

sector to consolidate globally-agreed scientific criteria

2019). Building on this approach, other methodologies

and harmonise work for identifying KBAs (IUCN, 2016). All

have been developed (for example, Important Plant Areas

these efforts have culminated in A Global Standard for the

(IPAs), Alliance for Zero Extinction (AZE) sites and Prime

Identification of KBAs (IUCN, 2016), which can be applied

Butterfly Areas) for multiple taxonomic groups in freshwater,

robustly across taxonomic groups and all elements of

terrestrial and marine environments. These approaches

biodiversity.

general y focus on one group of species or one biome, and

use diverse assessment criteria, which has led to some

KBAs are “sites contributing significantly to the global

confusion amongst decision-makers, as well as duplication

persistence of biodiversity” (IUCN, 2016). This does

of conservation efforts (Dudley et al., 2014).

not imply that a specific site-based conservation action,

such as protected area (PA) designation, is required. Such

As a consequence, during the World Conser vation

management decisions should be based on conservation

Congress held in Bangkok Thailand in 2004, IUCN members

priorit y-set ting exercises, which combine data on

requested for IUCN “to convene a worldwide consultative

biodiversity importance with the available information on

process to agree a methodology to enable countries to

site vulnerability and the management actions needed to

identify Key Biodiversity Areas” (IUCN, 2004). In response

safeguard the biodiversity for which the site is important.

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

145

It is often desirable to incorporate other data into priority-

on the distribution, abundance, ecology and extinction

setting, such as conservation cost, opportunity for action,

risk for species in a set of taxonomic groups that are

importance for conserving evolutionar y histor y and

considered reliable indicators of the biological structure

connectivity. KBAs thus do not necessarily equate to

and functioning of freshwater ecosystems (decapods,

conservation priorities but are invaluable for informing

fishes, molluscs, odonates and freshwater plants) (see

systematic conservation planning and priority-setting,

Chapters 3–7); ii) identifying those river/lake sub-basins,

recognising that conservation priority actions may also be

as well as sites within LMNN, containing species that

outside of KBAs (IUCN, 2016).

appear to meet the KBA criteria; iii) validating (through

stakeholder consultations) KBAs within those sub-basins

Data generated through application of the KBA standard

and sites, taking into account the hydrological connectivity

are expected to have multiple uses (Dudley et al., 2014).

of the sub-basin where the KBA resides; and iv) compiling

KBAs can support the strategic expansion of PA networks

sets of additional information about each KBA to support

by governments and civil society working toward the

management of the biodiversity elements triggering the

achievement of the Aichi Biodiversity Targets (in particular

criteria. Each of these processes is covered in more detail

Targets 11 and 12), as established by the Convention on

below.

Biological Diversity (CBD; Butchart et al., 2012); serve

to inform the description or identification of sites under

international conventions (such as Wetlands of International

10.2 Methodology

Importance designated under the Ramsar Convention,

natural World Heritage Sites, and Ecologically and

10.2.1 KBA criteria and thresholds

Biologically Significant Areas (EBSAs) as described under

the CBD); contribute to development of other effective

The methodology for identification and delineation of global

area-based conservation measures (Jonas et al., 2014);

freshwater KBAs in the LMNNC followed the new global

inform private sector safeguard policies, environmental

standard for identification of KBAs (IUCN, 2016). The new

standards and certification schemes; support conservation

global KBA criteria provide quantitative thresholds for

planning and priority-setting at national and regional

identifying sites that contribute significantly to the global

levels; and provide local and indigenous communities

persistence of: A) Threatened biodiversity; B) Geograph-

with opportunities for employment, recognition, economic

ically restricted biodiversity; C) Ecological integrity; D)

investment and societal mobilisation (IUCN, 2016).

Biological processes; and E) Biodiversity through com-

prehensive quantitative analysis of irreplaceability (IUCN,

At present, freshwater KBAs have not been identified for

2016).

most parts of the world and as a result there are currently

few opportunities for conservation and development

Sites identified as potential KBAs should ideally be

managers to take account of freshwater biodiversity within

assessed against al criteria. However, not al of the criteria

the planning process (Darwal et al., 2011). Al iance for Zero

are applicable or relevant for the freshwater taxonomic

Extinction (AZE) sites (Ricketts et al., 2005), a subset of

groups considered in this study, for example because

KBAs that contain the last or only populations of globally

some criteria relate to ecosystem types and not species.

Critically Endangered (CR) or Endangered (EN) species,

Meeting any one of the criteria (or sub-criteria) is enough for

have recently (in 2018) been identified for freshwater

a site to be considered for qualification as a KBA. Species

decapods (crabs, crayfishes and shrimps; see www.

meeting the KBA thresholds and criteria are defined

zeroextinction.org) but are in urgent need of identification

as KBA trigger species. Only criteria relevant to species

for other freshwater taxa. This study aims to fill the gap on

were considered in this study and as a result, criteria A2,

freshwater KBAs within the Lake Malawi/Nyasa/Niassa

B4 and C were not used as these refer to ecosystem types.

Catchment (hereafter LMNNC), paving the way for better

Other criteria, such as B2, B3, D3 and E, were not utilised

representation of freshwater biodiversity within the PA

due to lack of adequate data. The criteria and thresholds

network and for consideration of freshwater biodiversity

employed in this study are summarised in Table 10.1.

in conservation management. It builds on initial work by

WWF to identify priority conservation areas in Lake Malawi/

Population data were not available for the majority

Nyasa/Niassa (hereafter LMNN) (Chafota et al., 2005) with

of freshwater species considered and therefore, the

the benefit of new and extensive data on the status and

percentage of the global distribution area of the species

distribution of freshwater species (see Chapters 3–8).

that occurred within each KBA was the assessment

parameter used as a proxy for the percentage of the global

The process leading to the identification and delineation of

population when considering whether a species qualified

freshwater KBAs in the LMNNC included: i) collating data

as a KBA trigger species at each site.

146

Table 10.1 Selected Key Biodiversity Area (KBA) criteria used for the
delineation of freshwater KBAs in the Lake Malawi/Nyasa/

Niassa Catchment. Adapted from IUCN, 2016.

A1: Threatened species

(a) Site regularly holds ≥0.5% of the global population AND ≥5 functional
reproductive units of a global y Critical y Endangered (CR) or Endangered
(EN) taxon

(b) Site regularly holds ≥1% of the global population AND ≥10 functional
reproductive units of a global y Vulnerable (VU) taxon (c) Site regularly
holds ≥0.1% of the global population AND ≥5 functional reproductive units
of a global y Critical y Endangered (CR) or Endangered (EN) taxon listed as
such based only on a population size reduction in the past or present

(d) Site regularly holds ≥0.2% of the global population AND ≥10 functional
reproductive units of a global y Vulnerable (VU) taxon listed as such based
only on a population size reduction in the past or present

(e) Site effectively holds the entire global population of a CR or EN taxon

B1: Individual y geographical y restricted species

Site regularly holds ≥10% of the global population size AND ≥10
reproductive units of a species

D1: Demographic aggregations

(a) Site predictably holds an aggregation representing ≥1% of the global

population size of a species, over a season, and during one or more key
stages of its life cycle

D2: Ecological refugia

Site supports ≥10% of the global population size of one or more species
during periods of environmental stress, for which historical evidence shows
that it has served as a refugium in the past and for which there is evidence to
suggest it would continue to do so in the foreseeable future 10.2.2
Freshwater KBA delineation process

b. Derive proposed site boundaries based on biological

data

10.2.2.1 Freshwater KBAs in the drainage basin of

Using the distribution maps assembled in Stage 1a above,

Lake Malawi/Nyasa/Niassa

al sub-basins (level 8 HydroBASINS; see Chapter 2) in the

The identification and delineation of freshwater KBAs in the

LMNNC that contained potential KBA trigger species were

drainage basin of LMNN fol owed a three-step process:

identified. For each sub-basin, a list of potential trigger

species present (based on the Red List distribution maps)

10.2.2.1.1 Stage 1. Desktop analysis

and the potential criteria met was produced. This analysis

The first step of the process was a desktop analysis of data

was done using Microsoft Excel and R (R Core Team, 2016).

collated through the IUCN Red List assessment process

Maps were created to show the richness of potential KBA

for the following freshwater taxonomic groups: decapods,

trigger species per sub-basin (e.g. Figure 10.1).

fishes, mol uscs, odonates and plants (see Chapters 3–7).

10.2.2.1.2 Stage 2. Stakeholder KBA validation and

The datasets collected included information on species

delineation workshop

distributions (digital shapefiles) and the IUCN Red List

A KBA validation and delineation workshop was held in Senga

Categories of species, indicating their relative extinction

Bay, Malawi in October 2018, in col aboration with regional

risk.

stakeholders including species experts, conservation NGOs

and government representatives (Figure 10.2). The aim of this

a. Assemble spatial datasets of:

workshop was to validate whether the sub-basins identified

i. Red List distribution maps for freshwater decapods,

as containing potential KBA trigger species (Figure 10.1)

fishes, mol uscs, odonates, and plants;

met the KBA criteria, and then to derive KBA site boundaries

i . Existing KBAs (including AZE sites), Ramsar sites

that were biological y relevant to these species, yet practical

and PAs.

for management (IUCN, 2016). Due to the large number of

sub-basins containing potential KBA trigger species and

KBA delineation is an iterative process that makes use

time limitations at the workshop, sub-basins were prioritised

of better and more recent data as they become available

based on the number of potential KBA trigger species with the

(IUCN, 2016). Red List assessments of the majority of

sub-basins with the highest number of potential KBA trigger

the freshwater species considered were updated in 2018

species investigated first. Workshop participants were first

through the first component of this study (see Chapters 3–7),

asked to confirm the presence of the KBA trigger species

to ensure that data are traceable to a reliable source and

within each sub-basin identified during stage 1 (desktop

sufficiently recent to give confidence that the biodiversity

analysis) and to then delineate KBA boundaries according to

elements are still present at the sites.

the fol owing procedures:

147
¯

0 25 50

100

150

200Kilometers

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

In-lake distribution of potential KBA trigger

species
Number of potential KBA trigger

species per sub-basin

1-3

6-7

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

8 - 46

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 10.1 Richness of potential Key Biodiversity Area (KBA) trigger

species in sub-basins in the Lake Malawi/Nyasa/Niassa Catchment
(richness data are classified using quantiles) with the in-lake
distribution of potential KBA trigger species.

148

a. Confirmation of KBA trigger species presence within

2007; Nel et al., 2011). PAs also often lack target actions

sub-basins

for management of freshwater biodiversity and often fail

Species presence was confirmed based on museum records

in dealing with pressures coming from outside the PA

from major col ections, coarse scale distribution records and

boundaries.

regional and international expert knowledge. The presence

of the species at the thresholds required to meet the KBA

d. Delineation of new freshwater KBAs

criteria was first validated using the level 8 HydroBASINS

When there was no spatial overlap between the proposed

boundary and then revalidated as the boundary was refined

freshwater KBA and any pre-existing KBAs or PAs, site

(see Stages 2b, 2c and 2d below). Species that did not meet

boundaries were based on the location of focal areas

the thresholds once the final boundary was delineated were

identified for the freshwater KBA trigger species (if the focal

excluded as KBA trigger species but maintained in the

area met the KBA thresholds and criteria). Focal areas are

documentation and listed under other important biodiversity

distinct sites (e.g. river headwaters, lakes, or springs)

at the site.

of particular importance for the long-term survival of

the species (e.g. spawning areas, feeding areas, or

b. Boundary delineation with respect to pre-existing

sites supporting a significant part of the population of

KBAs

a species) (see Abell et al., 2007). It was recommended

Identification and delineation of new KBAs should take into

where possible to delineate focal areas using level 12

consideration the boundaries of pre-existing KBAs because

HydroBASINS, the smal est grain size available.

many have national recognition, active conservation and

monitoring initiatives, and/or are linked to international,

The new KBA global standard acknowledges that when

national, regional legislative and policy processes (IUCN,

delineating sites that fall outside existing KBAs and PAs, it

2016). Overlapping KBA boundaries are not permitted.

is often necessary to incorporate other data on land/water

Thus, where freshwater trigger species were present in

management and catchments boundaries to derive practical

sub-basins overlapping existing KBAs, the boundary of the

site boundaries (IUCN, 2016). In the case of freshwater KBAs,

existing site was adopted if:

using sub-basins to delineate site boundaries provides clear

benefits as they represent well defined and ecologically

■ The trigger species presence within the site met the KBA

meaningful management units, they facilitate ease of data

criteria thresholds; and

storage, search and management (tabular format), account

■ The boundary was ecological y relevant for management

for hydrological connectivity, facilitate input to conservation

of the freshwater KBA trigger species.

planning software such as Marxan, and can be applied

flexibly at 12 different resolutions, the smallest being

c. Boundary delineation with respect to PAs

approximately 10 km2. In addition, there is a growing body

PAs are established and largely well recognised manage-

of environmental data being compiled specifically for the

ment units with the goal of safeguarding the biodiversity

HydroBASIN sub-basin units.

contained within them. Additional recognition of the site as a

freshwater KBA, using the existing site boundaries, can

For some species, the inherent connectivity of aquatic

bring further attention to their importance and better focus

systems presents challenges for effective management at

management towards any newly recognised freshwater

the site scale. Many aquatic species are highly mobile and

species of conservation concern. Therefore, when a

maybe widespread throughout a basin (e.g. migratory fish

freshwater trigger species fell within a sub-basin overlapping

species) and may, therefore, not occur at identifiable sites

an existing PA it was appropriate to use the PA boundary to

at globally significant population levels. Such species may

delineate the KBA if:

not benefit from site scale conservation, but from a wider

catchment management approach. However, the majority

■ The PA contained enough of the KBA trigger species to

of species within the LMNNC are not highly mobile and are

meet the threshold of significance; and

instead locally confined, for example the endemic cichlids

■ The boundary was ecological y relevant for the species.

of LMNN (e.g. Figure 10.3). Although these species would

likely benefit from being within a KBA, they are unlikely to

It is important to highlight, however, that regional-scale

be positively affected by the presence of a KBA if found just

assessments of the coverage and effectiveness of PAs

outside the boundary.

have shown them to be largely ineffective for conserving

freshwater habitats and species (Hermoso et al., 2016;

e. Complete minimum documentation requirements for

Leadley et al., 2014). For example, rivers have often been

each KBA

used to delineate the borders of PAs rather than being the

Finally, workshop participants were asked to complete

targets of conservation themselves (Abell, Allan & Lehner,

datasheets with the minimum documentation requirements

149
for each associated KBA including: a site description, list

incorporated into the KBA datasheets and boundaries.

of validated trigger species, description of threats and

KBAs that passed the review were then submitted to BirdLife

habitat types within the site, and conservation actions in

International for publication in the next update of the World

place and recommended. Al of this information is required

Database on Key Biodiversity Areas (WDKBAs).

to justify confirmation of a site as a KBA, and as guidance

for management of the KBA, site-scale monitoring, national

10.2.2.3 Freshwater KBAs in Lake Malawi/Nyasa/

conservation planning and priority-setting, and global and

Niassa

regional analyses.

LMNN is represented by a single HydroBASIN of 29,181

km2 and contains 300 potential KBA trigger species.

Details for potential site champions were also recorded.

Although the lake itself would meet the criteria to qualify

Site champions are individuals or organisations that are

as a KBA due to the presence of many threatened species

well placed to raise awareness of the existence of the

(to meet criterion A), restricted range or endemic species

KBAs and the issues faced with respect to threats to

(to meet criterion B), and as it is the site of many important

biodiversity, and to help implement the required actions

biological processes (to meet criterion D), we did not think

to safeguard these globally important sites.

that the lake met the definition of a ‘site’ given in the KBA

standard (“a geographical area on land/or in water with

10.2.2.2 Stage 3. Review

defined ecological, physical, administrative or management

After the workshop, the KBA datasheets and boundaries

boundaries that is actually or potentially manageable as a

were finalised by members of the IUCN Freshwater

single unit”; IUCN, 2016), given its large size and occurrence

Biodiversity Unit (FBU) to ensure the criteria and the

over multiple countries. As a result, we decided to delineate

minimum documentation requirements were met for each

KBAs within LMNN.

KBA. The KBA datasheets and boundary images were

then made available online for approximately one month

It was not possible to run a desktop analysis for potential

for final review by workshop attendees and by additional

KBAs within LMNN fol owing the process discussed above

species experts and regional stakeholders who were

because no species were mapped to defined polygons

unable to attend the workshop. Feedback from this review

within the lake. Threatened and restricted-range species

process was incorporated into the KBA datasheets and

were identified as potential KBA trigger species and their

boundaries. The KBAs were then submitted to the KBA

distributions in the lake were mapped (Figure 10.1). A list of

Secretariat for publication. Sites in Mozambique were

sites of potential KBAs within the lake was then put together

reviewed by the recently established Mozambique KBA

through consideration of the sites identified by Chafota et

National Coordination Group (NCG). NCGs have not yet

al. (2005) and in consultation with experts prior to the KBA

been established for Malawi and Tanzania and therefore,

delineation workshop. Stages 2 and 3 (see above) were

sites in these countries were reviewed by the Head of the

then fol owed from this initial list of sites. When there was no

KBA Secretariat. Feedback from this review process was

spatial overlap between the proposed freshwater KBA and

Figure 10.2 Participants of the Key Biodiversity Area (KBA) validation

and delineation workshop held in Senga Bay, Malawi in October 2018.
From left to right, front row: A. Palmer-Newton, A. van Wyk, A.
Pegado, L. Chigamane, M. Mwithokona, G. Kanyerere, P. Kaliba and A.
Shechonge; back row: C. Sayer, T. Phiri, Z. Ndhlovu, S. Sakhama, D.
Tweddle, S. Manda, M. Ngochera and W. Darwall. © Catherine Sayer

150

related to threatened biodiversity (criteria A1a, A1b, A1c,

A1d and A1e), 126 species considered as triggers based on

the criteria related to geographical y restricted biodiversity

(criterion B1) and one species considered as a trigger based

on the criteria related to biological processes (criteria D1a

and D2) (Table 10.2). Furthermore, six of these species are

also identified as AZE species (criteria A1e; Table 10.2) facing

an overwhelmingly high risk of extinction, and confirming the

urgency to develop and implement effective conservation

actions and management plans for freshwater biodiversity in

the LMNNC.

Figure 10.3 A male Petrotilapia xanthos at Gallireya Reef (part

10.3.2 Freshwater KBAs overview

of Chilumba and Youngs Bay Key Biodiversity Area, KBA). © Ad

Konings

Twenty-two important river, lake and wetland areas were

any pre-existing KBAs or management units, delineation

validated by the regional and international experts at the KBA

was based on focal areas delineated using a combination of

delineation workshop as freshwater KBAs. Six of these KBAs

expert knowledge, habitat and bathymetry data.

are also AZE sites. Of the 22 freshwater KBAs, six were in the

drainage basin of LMNN, 10 were in the lake itself (e.g. Figure

10.5), and six included both lake and inland habitats. The

10.3 Results

majority of the freshwater KBAs (18) were newly delineated

KBAs, with only four sites adopting existing KBAs (Figure

10.3.1 Freshwater KBA trigger species

10.4, Table 10.3, Appendix 3). The total area of the freshwater

KBAs is 9,440 km2, which is equivalent to 7% of the total area

The preliminary desktop analysis identified 359 potential

of the LMNNC.

KBA trigger species, out of which 143 were validated by the

regional and international experts at the KBA delineation

10.3.3 Current levels of management and

workshop (Table 10.2), meaning that their presence was

protection

confirmed within the relevant sub-basins or sites at a

threshold to trigger the KBA criteria. See Appendix 3 for

Approximately 6,600 km2 (70% of the total area) of the

the full list of validated KBA trigger species in each newly

freshwater KBAs identified fall within the boundaries of

delineated freshwater KBA.

existing management units (PAs, existing KBAs and Ramsar

sites) (Figure 10.6). Of this total, c. 6,500 km2 are within

The freshwater KBAs validated at the workshop support

existing KBAs, all of which have adopted PA boundaries:

28 species considered as triggers based on the criteria

Nyika National Park, Vwaza Marsh Wildlife Reserve,

Table 10.2 Number of trigger species per freshwater taxonomic group

and per Key Biodiversity Area (KBA) criterion. Note that some trigger
species may meet more than one of the KBA criteria and therefore, the
totals per taxonomic group are not necessarily the sum of the fol owing
rows.

Decapods

Fishes

Molluscs

Odonates

Plants

All

Trigger species

139

143

Threatened Biodiversity
0

25

28

(Criterion A)

A1a

16

18

A1b

0
9

A1c

A1d

A1e (AZE)

0
0

Geographically Restricted

124

126

Biodiversity (Criterion B)

B1

124

126

Biological Processes

1
0

(Criterion D)

D1a

D2

151
¯

0 25 50

100

150

200Kilometers

32

5
7

10

1112

1413

15

16

17 18 19

20

21

22

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

KBAs with freshwater trigger species

Adopted KBAs (non-AZE sites)

New KBAs (non-AZE sites)

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

New KBAs (AZE sites)

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap

contributors, and the GIS User Community

Figure 10.4 Key Biodiversity Areas (KBAs) with freshwater trigger

species in the Lake Malawi/Nyasa/Niassa Catchment, highlighting
Alliance for Zero Extinction (AZE) sites. Map ID numbers for KBAs
with freshwater trigger species are listed in Table 10.3. See
Supplementary Material for detailed KBA boundary images.

152

Table 10.3 Key Biodiversity Area (KBA) names, map ID numbers

Nkhotakota Wildlife Reserve and Liwonde National Park

and whether they qualify as Alliance for Zero Extinction (AZE)

(Figure 10.7) (see Sites in the spotlight: Mpasa and the

sites for all KBAs with freshwater trigger species in the Lake

Lower Bua and Nkhotakota Wildlife Reserve Key

Malawi/Nyasa/Niassa Catchment in Figures 10.4 and 10.6. See

Biodiversity Areas p. 160, and Sites in the spotlight: A

Supplementary Material for detailed KBA boundary images.

unique fish fauna benefiting from the Nyika National

Map ID KBA name

AZE site

Park, Vwaza Marsh Wildlife Reserve and North Rumphi

Lake Kyungululu

Yes

Key Biodiversity Areas p. 166). A number of additional

Kiwira Mbaka Lufiryo

No

KBAs (Makanjira, Maleri Islands, Cape Maclear, and Lake

Lower Songwe River

No
Malawi Southeast Arm KBAs) fall partially or completely

Chilumba and Youngs Bay

No

within the Lake Malawi National Park, which is also a World

Ruhuhu River Mouth

No

Heritage Site (Department of Surveys, Government of

Nyika National Park

No

Malawi, 1983) (see Sites in the spotlight : The Lake

North Rumphi

No

Malawi National Park World Heritage Site p. 158).

Vwaza Marsh Wildlife Reserve

No
However, the lake area of the Lake Malawi National Park is

Puulu-Mbamba Bay

No

not included on Protected Planet (UNEP-WCMC & IUCN,

10

Tukombo-Sanga-strip

Yes

2019 ) and so this has not been included in the area

11

Chizumulu Island and Taiwanee Reef

Yes

calculation above. The remaining c. 100 km2 represents

12

Nkwichi Bay

No

KBAs that fall within Ramsar sites. The Mozambican waters

13

Lower Bua

No
of the lake, including Chizumulu Island and Taiwanee Reef

14

Nkhotakota Wildlife Reserve

No

and Nkwichi Bay (Figure 10.8) KBAs, are al within the Lake

15

Mbenji Island

No

Niassa and its Coastal Zone Ramsar site ( Ramsar

16

Makanjira

Yes

Secretariat, 2019).

17

Maleri Islands

Yes

18

Cape Maclear

Yes

Even though close to half of the freshwater KBAs (and 70%

19

Lake Malawi Southeast Arm

No

of their total area), therefore, already have some sort of

20

Upper Shire

No

recognition, and potentially are already protected and

21

Lake Malombe

No

managed for biodiversity, it is important to highlight that in

22

Liwonde National Park

No

most cases freshwater species, with the exception of

Figure 10.5 Boadzulu Island (part of the Lake Malawi Southeast Arm
Key Biodiversity Area, KBA). © Catherine Sayer 153
¯

0 25 50

100

150

200Kilometers

32

8
9

10

1112

1413

15

16

17 18 19

20

21

22

Legend

Lake Malawi/Nyasa/Niassa

Lake Malawi/Nyasa/Niassa Catchment

KBAs with freshwater trigger species

Existing KBAs

Protected Areas

Sources: Esri, HERE, Garmin, Intermap, increment P Corp., GEBCO,

USGS, FAO, NPS, NRCAN, GeoBase, IGN, Kadaster NL, Ordnance

Ramsar Sites

Survey, Esri Japan, METI, Esri China (Hong Kong), (c) OpenStreetMap
contributors, and the GIS User Community

Figure 10.6 Key Biodiversity Areas (KBAs) with freshwater trigger

species in the Lake Malawi/Nyasa/Niassa Catchment, with existing
management units (KBAs, protected areas and Ramsar sites) overlaid.
Map ID numbers for KBAs with freshwater trigger species are listed in
Table 10.3.

154

water birds, are not often the focus of conservation and

workshop as individuals or organisations well placed to

management actions within these areas that are delineated

raise awareness of the existence of the KBAs and the issues

primarily for terrestrial species (mammals, reptiles and

faced with respect to threats to biodiversity, and to help

birds). Therefore, it is important to inform the management

implement the required actions to safeguard these global y

authorities of these PAs, existing KBAs and Ramsar sites

important sites (see Appendix 4).

about the presence of these freshwater KBA trigger species

within their site boundaries, in order that appropriate

management strategies can be adopted.

10.4 Recommendations

Approximately 2,840 km2 (30% of the total area) of the

Close to half of the number and the majority (70%) of the

freshwater KBAs identified fall outside the boundaries of

total area of the freshwater KBAs validated through this

existing management units (PAs, existing KBAs and Ramsar

study were found to lie within the boundaries of existing

sites). This suggests that significant gaps remain in the

management units (PAs, existing KBAs and Ramsar sites).

coverage of freshwater biodiversity by existing conservation

The additional recognition of these sites as global freshwater

management units. A strategic expansion of the PA network

KBAs brings them greater individual recognition and

is recommended to include some of these critical areas of

col ectively helps to highlight the urgent need to implement

conservation concern (see Sites in the spotlight: The Lake

more effective conservation actions and environmental

Malawi National Park World Heritage Site p. 158).

safeguards for freshwater biodiversity in the LMNNC. Most

of these existing management units have been delineated

10.3.4 Site champions

primarily for terrestrial species such that they will often fail

to focus on targeted management for the many restricted

Forty-nine potential KBA site champions were identified

range and threatened species living in freshwater habitats.

by stakeholders at the KBA delineation and validation

It is now a priority to inform the management authorities for

Figure 10.7 The Shire River in Liwonde National Park Key Biodiversity
Area (KBA). © Denis Tweddle 155
Figure 10.8 Nkwichi Bay Key Biodiversity Area (KBA) is within Lake
Niassa and its Coastal Zone Ramsar site. © TravelingOtter (CC

BY-SA 2.0)

these sites of the need to develop new management actions

at this point KBAs also need to be re-evaluated to ensure

that specifically focus on conservation of these globally

they still qualify. For example, there may be cases where

important freshwater species.

conservation actions have been successful and therefore,

the trigger species originally identified no longer meet

The remaining sites, representing 30% of the total area

the KBA criteria thresholds. Ultimately the process for

of freshwater KBAs, are located outside of any existing

identification of KBAs should be national y driven such that

management units and represent priority gaps in the current

all relevant parties can be directly involved, especially to

network. The location of these KBAs should inform future

facilitate any recommendations to change boundaries of

strategies for improving the representation of freshwater

existing PAs or KBAs. The work presented above represents

biodiversity within the regional PA network (see Sites in

the first steps in taking this process forwards and provides a

the spotlight: The Lake Malawi National Park World

baseline data set to inform future KBA designations.

Heritage Site p. 158) or as targets for habitat restoration

efforts where PA status might be inappropriate. It is hoped

The primary threats to freshwater species in the LMNNC, as

that the potential KBA site champions identified through this

identified through this project (see Chapters 3–8), include:

project (see Appendix 4) will help stimulate these actions

i) fishing and harvesting of aquatic resources; ii) water

by building awareness of the existence of these priority

pollution from agricultural, domestic and industrial waste

freshwater sites and the need for conservation actions.

leading to eutrophication and sedimentation; iii) habitat

degradation and soil erosion caused by deforestation,

The identification and delineation of KBAs is necessarily a

primarily for agriculture; iv) poor water management and

fluid and ongoing process responding to the provision of

use, primarily for agriculture; and v) invasive alien species.

new information and a constantly changing environment

The impacts of these types of threat tend to spread rapidly

and, thus, it is expected that this current freshwater KBA

throughout sub-basins, such that localised conservation

dataset for the LMNNC will continue to be refined and

actions restricted to limited parts of a sub-basin will often

updated. Red List categories change over time as they are

fail to provide effective solutions. It is therefore necessary

updated through the Red List reassessment process, and

to focus on management of the wider catchment within

156

which KBAs reside, taking into account both lateral and

The freshwater KBAs identified in this project wil also help

longitudinal hydrological connectivity.

support the implementation of multilateral environmental

agreements in the LMNNC, such as the Ramsar Convention,

Integrated River Basin Management (IRBM), or a similar

guiding conservation planning and priority-setting at

strategy, is an approach recommended for most freshwater

national level to: i) identify new and potential Wetlands of

KBAs to ensure effective management of both upstream and

International Importance (Ramsar sites) under Criteria 2

downstream threats, often originating outside of the KBA

to 9; ii) update existing Ramsar site management to focus

boundaries. In many cases threats can originate some

on the new freshwater trigger species found within their

distance from the KBA itself. This approach is fundamental

boundaries (e.g. Lake Niassa and its Coastal Zone Ramsar

to better coordinate conservation, management and

site); and iii) identify existing Ramsar sites meeting the

development planning of water, land and related resources

KBA criteria that are undergoing adverse changes in their

across sectors, and to maximise the economic and social

ecological character and that might be eligible for inclusion

benefits derived from water resources in an equitable

on the Montreux Record and to potentially benefit from a

manner while preserving and, where necessary, restoring

Ramsar Advisory Mission.

freshwater ecosystems.

The network of freshwater KBAs identified through this

The Environmental Flows (E-Flows) assessment method-

project will also help the countries within the LMNNC

ology is also an important tool for the conservation and

in their work towards meeting the Aichi Biodiversity

management of freshwater KBAs. E-Flows aim to maintain

Targets (in particular Targets 11 and 12) as established

the quality, quantity and timing of water flows required to

by the Convention on Biological Diversity. These two

sustain freshwater ecosystems and the human livelihoods

targets specifically address the need for species and sites

that depend on them (Dyson, Bergkamp & Scanlon, 2003).

conservation. In addition, freshwater KBAs can help identify

As a first priority E-Flows should be determined, where

freshwater ecosystem priorities for the UN Sustainable

appropriate, for all freshwater KBAs involving riverine

Development Goals (SDGs), and provide a better metric

systems and smal er lakes fed by rivers.

for measurement of Sustainable Development target

6.6 focused on protecting and restoring water-related

Although invasive alien species are currently only recorded

ecosystems, 6.5 focused on implementing integrated water

as a threat to 2% of species native to the LMNNC (see

resources management at all levels, and 15.1 focused on the

Chapter 8), they are considered to be the second greatest

conservation, restoration and sustainable use of terrestrial

threat to biodiversity global y (Bel ard, Cassey & Blackburn,

and inland freshwater ecosystems and their services (United

2016). Freshwater systems are highly vulnerable to invasive

Nations, 2016).

species due to the relative ease, plus severe and wide-

ranging consequences, of invasion ( Moorhouse &

Finally, it is expected that the list of freshwater KBAs pre-

Macdonald, 2015). Invasive species have had dramatic

sented in this report will guide conservation investment

negative effects of the freshwater biodiversity in other

priorities and inform performance standards and environ-

African Great Lakes (e.g. Nile perch ( Lates niloticus) in Lake

mental safeguard policies of financial institutions and the

Victoria; Sayer et al., 2018) and therefore, within the LMNNC

private sector to help avoid or minimise impacts of their

it is advised to trace pathways for introduction, prevent

operations in and around these critical sites for freshwater

future introductions, and to manage, or where feasible,

biodiversity in the LMNNC.

eradicate invasive species. Information on the distribution of

invasive species, their impacts, pathways of invasion and

management recommendations can be found in the Global

10.5 Next steps

Invasive Species Database (www.iucngisd.org/gisd).

Information collated through the KBA delineation and

This report and related policy brief will be circulated to

validation process should ultimately feed into the GISD,

all KBA site champions and cross-sectorial government

which is also linked to the IUCN Red List.

departments in all countries of the LMNNC. Additionally,

KBA datasheets including detailed information on the

Periodic updates of IUCN Red List assessments and

sites and their KBA trigger species are available through

monitoring of KBA sites will enable calculation of a Red List

the World Database on Key Biodiversity Areas (WDKBAs;

Index (RLI) for all freshwater species assessed (see Chapter

www.keybiodiversityareas.org) and will be made available 9) in order to

track trends in the projected overall extinction

through the Integrated Biodiversity Assessment Tool

risk of freshwater species, and so potentially helping to

( IBAT; www.ibat-alliance.org), a tool that is already inform managers on the

effectiveness of any management

well known amongst the private sector and donor

interventions.

community.

157
Sites in the spotlight

The Lake Malawi National Park World Heritage Site

Sayer, C.A.1

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

The Lake Malawi National Park World Heritage property lies at the southern
end of LMNN. It is comprised of a large terrestrial area (primarily the Cape
Maclear Peninsula), three other mainland areas, 12 islands (e.g. Figure 10.9),
and a relatively small freshwater area extending 100 m from the shoreline. It
has a land area of 87.1 km2 but covers only 7 km2 of the lake, equivalent to
0.02% of the lake’s surface area (IUCN & UNESCO, 2014).

The park was established in 1980 and inscribed on to the World Heritage list
in 1984 on the basis of three criteria: (vi ) for its exceptional natural beauty;
(ix) for its outstanding example of biological evolution, as shown by
adaptive radiation and speciation in the rocky-shore haplochromine cichlids
(mbuna); and (x) for the outstanding diversity in freshwater fishes it hosts
(IUCN & UNESCO, 2014; World Heritage Committee, 1984a).

Given the small aquatic area of the property and its inscription primarily for
its freshwater biodiversity, the Bureau of the World Heritage Committee
recommended at the time of inscription that the area of the property be
extended (World Heritage Committee, 1984b). Additional y, a
UNESCO/IUCN monitoring mission to the property in 2014 resulted in the
fol owing recommendation: “… the States Parties of Malawi, Mozambique
and Tanzania should investigate the feasibility of increasing protection for
additional areas of the shoreline and islands that have been identified as
important localities for the protection of endemic fish and evolutionary
processes throughout the lake. Where possible, these areas might be
designated as reserves or community-run ‘special use zones’ and might
ultimately be incorporated into an extended trans-national serial property”
(IUCN & UNESCO, 2014). The report also recommended that these areas
be identified based on scientific knowledge of species distributions and
ecology (IUCN & UNESCO, 2014). The World Heritage Committee, which
is the decision making body of the Convention, subsequently adopted this
mission recommendation in 2014 (Decision 38 COM 7B.92) (World
Heritage Committee, 2014).

KBAs are “sites contributing significantly to the global persistence of

biodiversity” (IUCN, 2016) and are delineated fol owing standardised
criteria based on scientific knowledge of species and/or ecosystems. The in-
lake KBAs identified through this study could, therefore, represent a
blueprint for this network of additional sites.

Four KBAs identified through this study lie partially or completely within
the current boundary of the Lake Malawi National Park World Heritage
property: Makanjira, Maleri Islands, Cape Maclear, and Lake Malawi
Southeast Arm KBAs.

Figure 10.9 Mumbo Island is part of the Lake Malawi National Park
World Heritage property. © Marco Derksen (CC BY-NC 2.0) 158
Figure 10.10 Chindongo saulosi (Critically Endangered, CR)

Figure 10.11 A territorial male Aulonocara maylandi at Luwala

is a trigger species for Chizumulu Island and Taiwanee Reef

Reef in Malawi. This species is a trigger for Makanjira Key

Key Biodiversity Area (KBA). © Ad Konings

Biodiversity Area (KBA) to which it is endemic. © Ad Konings

Additionally, a number of KBAs within LMNN but outside

of the current property boundary have been delineated,

including a site in the recently established Lake Niassa

and its Coastal Zone Ramsar site in Mozambique (Ramsar

Secretariat, 2019). Opportunities to extend the World

Heritage property based on these KBAs include:

■ Inclusion of Chilumba and Youngs Bay KBA (#4 in

Figure 10.4 and Figure 10.6) on the west coast of LMNN

in Malawi. This KBA is formed of the Chilumba Peninsula

and the string of small islands and reefs offshore

(Mphanga Rocks, Luwino Reef, Katale Island, Maison

Figure 10.12 A male Metriaclima mbenjii at Mbenji Island. This

Reef, and Chirwa Island), in addition to Youngs Bay,

species is endemic to Mbenji Island Key Biodiversity Area

including Gal ireya Reef. The combined habitats of rock,

(KBA). © Ad Konings

sand and offshore reef support a high diversity of cichlid

species, including two site endemics: Labeotropheus

simoneae and Petrotilapia xanthos.

■ Inclusion of Chizumulu Island and Taiwanee Reef KBA (#11). This is a

transboundary KBA, with the islands and nearshore waters belonging to
Malawi, and the surrounding waters belonging to Mozambique. This KBA is
within the Lake Niassa and its Coastal Zone Ramsar site. It hosts 12 site
endemic species, including the Critical y Endangered (CR) Chindongo
saulosi (Figure 10.10), making this KBA an Al iance for Zero Extinction
(AZE) site.

■ Extension to include the entirety of Makanjira KBA (#16), including

Chimwalani and Luwala Reefs. These reefs are home to the CR and site
endemic cichlid Aulonocara maylandi (Figure 10.11), which also qualifies
this KBA an AZE site.

■ Inclusion of Mbenji Island KBA (#15) in the western side of LMNN in

Malawi. This KBA is home to four site endemic cichlids: Aulonocara
koningsi, Copadichromis mbenji , Metriaclima mbenji (Figure 10.12) and
Pseudotropheus galanos.

■ Inclusion of Nkwichi Bay KBA (#12), which falls within the Lake Niassa
and its Coastal Zone Ramsar site in Mozambique. This site has a rich cichlid
communities and was delineated for the CR Chambo Oreochromis
squamipinnis, an important food fish.

■ Inclusion of Puulu-Mbamba Bay KBA (#9) along the north-eastern coast

of LMNN, which extends from Puulu Island to Mbamba Bay and includes
Pomanda, Hongi Island, Mbahwa Island and Longi Island. The mix of
habitats in the KBA supports a diverse cichlid community, including four
site endemic species.

■ Inclusion of Tukombo-Sanga Strip KBA (#10) in the western side of

LMNN in Malawi. This is an AZE site for the CR

Aulonocara kandeense, and is also an important site for other cichlids and
Chambo.
It is hoped that the States Parties of Malawi, Mozambique and Tanzania wil
consider these KBAs in light of the World Heritage Committee’s request.

159

Mpasa and the Lower Bua and Nkhotakota Wildlife Reserve

Key Biodiversity Areas

Phiri, T.B.1 and Tweddle, D.2

1 Senga Bay Fisheries Research Centre, Malawi Department of Fisheries,

P.O. Box 316, Salima, Malawi 2 South African Institute for Aquatic
Biodiversity, 11 Somerset Street, Grahamstown 6140, South Africa There is
great conservation value in recognising a site as a globally important KBA,
as demonstrated here by the Nkhotakota Wildlife Reserve (NWR; Figure
10.13). Gazetted in 1938, NWR is the largest wildlife reserve in Malawi
with an area of close to 1,800 km2. It supports high faunal and floral
biodiversity. The forest cover in NWR protects the steep slopes of the Great
African Rift escarpment and thus conserves water in that part of the
catchment. One of the major affluent rivers of LMNN, the Bua River,
traverses the reserve after originating from Mchinji highlands on the
Malawi–Zambia border.

The river flows for approximately 200 km through plains that have been
extensively cleared for tobacco farming in the districts of Lilongwe,
Kasungu, Dowa and Ntchisi. The river then enters the NWR and covers a
distance of approximately 35 km over the rift val ey escarpment where
numerous rapids and deep pools exist throughout (Tweddle, 1983). Within
the reserve, the river width varies from 20 m to over 200 m, with wider
stretches divided into numerous rivulets by small islands. There are
extensive areas of reedy shoreline with other patches of weed providing
good fish cover. These important habitats are now, however, greatly reduced
as a result of river bank destabilisation caused by recent changes in the flow
regime due to massive land clearance above the reserve. There are also fast-
flowing rocky stretches interspersed with deep pools. Extensive gravel and
sandy-bottomed shallows (Figure 10.14) are present in many areas and these
are the spawning grounds of many potamodromous fish species, notably the
mpasa Opsaridium microlepis (Figure 10.15) and sanjika Opsaridium
microcephalum (Tweddle, 1983).

Conservation is an important component of management policies of the

Malawi Departments of Fisheries and of National Parks and Wildlife, but
with limited resources available for management, the ecological health of
NWR deteriorated in the past. However, in 2015 the government joined
forces with African Parks to manage the reserve. The reserve is now fenced,
law enforcement has been greatly increased, wildlife is recovering and
populations are being enhanced through reintroductions.

After leaving the NWR, an unprotected 15 km stretch of the Bua River flows
to LMNN. This part of the river is vital for conservation of potamodromous
fish species, which migrate into NWR for spawning. Notable species of
commercial and conservation significance include mpasa, which is now
listed as Vulnerable on the IUCN Red List on the basis of a predicted future
population decline, primarily due to over-fishing and habitat loss and
degradation (Tweddle, 2018).

Through this study, the stretch of the Bua River from NWR to its mouth in
LMNN was proposed as a KBA (named Lower Bua KBA) for freshwater
trigger species. The NWR is already confirmed as a KBA important for
more than 280 species of birds, but this study also identified it as important
for freshwater trigger species. Mpasa is identified as a trigger species for
both of these KBAs under criteria A1 for threatened species and D1 for
demographic aggregations (see Appendix 3).

Recently, the river has attracted a lot of attention and conservation efforts are
being implemented. A number of conservation projects have been proposed
to assist in the management and restoration of the river. Threats to the
riverine biodiversity that need attention include: over-fishing, illegal fishing
(using mosquito net, setting gill nets and fishing weir blocking the whole
breadth of the river), bad land practice (cultivating along river banks,
deforestation), damming and water abstraction for irrigation, poaching in
NWR and proposed development of a hydro-electricity power generation
station. In the short term, stimulated by recognition of the Lower Bua as a
KBA, efforts are being made to al ow migratory fish species to swim
upstream into the wildlife reserve for breeding. In collaboration with the
Malawi Department of Fisheries, research is being funded to keep track of
the mpasa catches and study ecological changes in the river. African Parks
plans to construct terraces on the Bua Irrigation Dam (Figure 10.16) when
the water level goes down in July 2019.

Additionally, Community Conservation Committees in all villages along the

15 km stretch outside of NWR have been established, mobilising resources
to support the Malawi Department of Fisheries with enforcement in critical
areas, including the river mouth, against il egal gears and fishing.

160
Figure 10.13 Nkhotakota Wildlife Reserve is a Key Biodiversity Area
(KBA) for freshwater species. © Catherine Sayer Figure 10.14 Sandy-
bottomed shallows in Nkhotakota Wildlife Reserve. © Amy Palmer-
Newton Figure 10.15 Mpasa ( Opsaridium microlepis) caught at Bua

Figure 10.16 Bua River Irrigation Dam in May 2019. © Titus B.

River mouth in May 2019. © Titus B. Phiri

Phiri

161

Restoring landscapes and conserving biodiversity in Malawi

Beatty, C.R.1

1 Forest Conservation Programme, IUCN (International Union for

Conservation of Nature), 1630 Connecticut Ave. N.W., Suite 300,
Washington, DC 20009, USA

In 2017, the Ministry of Natural Resources, Energy and Mining in Malawi

published both a national assessment of the opportunities for forest
landscape restoration (FLR) and an accompanying strategy (Ministry of
Natural Resources, Energy and Mining, Malawi, 2017). Within these
documents, Malawi outlined many objectives for landscape restoration,
including supporting food security, building landscapes that were more
resilient to changes in climate, and supporting biodiversity by choosing
restoration actions and species that would complement Malawi’s biodiversity
commitments and goals. FLR in Malawi intends to use a diversity of
restoration approaches, which include sustainability and conservation
improvements to agricultural systems, such as conservation agriculture and
agroforestry, as well as improved management of forests and woodlots,
riparian tree planting and additional restoration activities. These
interventions in degraded or deforested landscapes have the potential to
relieve species and ecosystems from the persistent and widespread pressures
and threats they face from a number of different human activities. Secondly,
this work also has the potential to enhance biodiversity in areas that are
undergoing restoration through the use of native species and ecological
restoration.
During the national FLR assessment, an audit of threatened species was
undertaken using The IUCN Red List of Threatened SpeciesTM and this
revealed a proportional y high number of threatened freshwater species. At
first these species were not included in the national FLR assessment because
many of the restoration interventions are not intended to restore freshwater
systems, and the assessment’s focus on forests and agricultural land favoured
reporting on how FLR

could support creating habitat or relieving pressures for forest-dependent

species. However, when analysts looked into the threats facing threatened
freshwater species, the primary threat for the majority of these species was
sedimentation.

This threat is commonly addressed in FLR, especial y for improving

hydropower efficiency and for conserving precious topsoil, and so this
analysis indicated that there would also be significant potential to use FLR to
respond to this major threat to freshwater species in Malawi.

Here, we demonstrate an estimate of the degree of potential degradation in

each of the terrestrial KBAs (i.e. those on land) for freshwater trigger species
discussed in this chapter and elucidate the combinations of degrading factors
experienced by each of these areas. However, since degradation analysis
through the national FLR assessment was only completed for Malawi’s
terrestrial areas, the data do not permit a degradation analysis of the aquatic
KBAs (i.e. those within LMNN).

Instead, for these aquatic KBAs we chose to analyse the watershed terrestrial
areas (i.e. aquatic KBA catchments) that supply the rivers and coastlines
where these aquatic KBAs are found (Figure 10.17). From this, we can see
not only which areas display more components of degradation, but we can
also describe in which ways the degradation criteria combine within each
KBA.

Based on the national FLR assessment, we can demonstrate the potential

degradation criteria present within each terrestrial KBA or aquatic KBA
catchment. For example, the Vwaza Marsh Wildlife Reserve (Figure 10.18)
KBA has 28
potential combinations of the nine degradation input criteria (Figure 10.19).
Here the central area of the KBA (coloured cream in Figure 10.19) has three
to four overlapping degradation criteria including fire, high poverty, low soil
fertility and low evapotranspiration. While some of these criteria may simply
be characteristic of the underlying geology (i.e. soil fertility in water-logged
soils is typical y considered to be low), others, such as the incidence of high
poverty and fire, are criteria that can help practitioners to develop strategies
to address the pressures that are threatening species here or downstream.
These estimates are based on a validated national scale analysis of
restoration opportunities but have not yet been ground-truthed. However,
they may provide insights into the types, combinations and extent of
landscape degradation criteria that potential y exist in this KBA.

Similar analyses can be completed for any of the objectives described in the
national FLR assessment, for example for food security criteria
combinations as shown in Figure 10.20 for Vwaza Marsh Wildlife Reserve
KBA. Here we see that this KBA has a significant central portion where
poverty, low evapotranspiration, lack of access to markets and non-timber
forest products (NTFP), and rainfed cropland interact as components of food
security. Taken with the additional 162
Figure 10.17 Number of coincident degradation criteria for terrestrial
Key Biodiversity Areas (KBAs) (i.e. those on land) and aquatic KBA
catchments (i.e. the watershed terrestrial areas of KBAs in Lake
Malawi/Nyasa/Niassa) in the Malawian part of the Lake
Malawi/Nyasa/Niassa Catchment. The nine degradation criteria used
can be found in Malawi’s National Forest Landscape Restoration
Assessment (2017).

163

degradation criteria above (Figure 10.19), practitioners can generate locally

driven solutions that integrate these and other finer scale concerns into KBA
management plans and landscape restoration strategies. Again, the intention
of this analysis is not necessarily to drive or suggest specific actions that
might be taken within the reserve, but to illustrate the combination of food
security factors that can be seen inside the KBA based on data that formed
part of a wel -funded and high profile landscape restoration initiative in
Malawi.

The benefit of these analyses is that they can be used in the planning and
recommendation stages with regards to the identification of KBAs and of
strategies for how to safeguard or restore biodiversity in these areas. For
example, as landscape restoration initiatives are planned that include
freshwater KBAs, or as freshwater conservation programmes are developed,
both initiatives can look for natural synergies that support both freshwater
conservation and landscape restoration or development objectives. Since
many KBAs in Malawi are included in or surrounded by areas of high
poverty, both development and conservation objectives must reconcile the
needs of people that depend on these KBAs and the needs of the KBA
trigger species. Using the analysis above, we can see that in Vwaza Marsh
Wildlife Reserve, there are a multitude of ways that the degradation or food
security data combine with each other in this landscape (Figure 10.19,
Figure 10.20). Of course, field validation of these data wil be required but, at
the planning stage, it al ows conservation, restoration and development
practitioners to explore the combinations of these criteria that cover the
greatest area and then design programmes that respond to these criteria.

Based on the degradation and food security analysis above for Vwaza Marsh
Wildlife Reserve (Figure 10.19, Figure 10.20), practitioners could explore
how fire, poverty, soil fertility, evapotranspiration, poor market access, and
irrigation for rainfed cropland might interact. In this scenario there may be
opportunities to improve livelihood conditions through conservation
agriculture, agroforestry, and ecological restoration that would support
human livelihoods and support the habitat requirements of trigger species in
this KBA. This is especial y the case for freshwater species, which are often
threatened by sedimentation. Upstream restoration of degraded landscapes
through FLR interventions can reduce water sedimentation and these
activities can be targeted to support habitat improvements for KBA trigger
species.

With a more fine-scale assessment of not only the threats facing species in
KBAs, for example through IUCN Red Listing, but also through parallel and
complementary processes, such as FLR assessments, those working on
biodiversity conservation and landscape restoration can work together to
ensure that the food security needs of people are met, while also restoring
and conserving the biodiversity of the landscapes that contain KBAs. For
more information on how biodiversity and landscape restoration are
complementary see Biodiversity Guidelines for Forest Landscape
Restoration Opportunities Assessments (Beatty, Cox & Kuzee, 2018).

Figure 10.18 Vwaza Marsh Wildlife Reserve. © Dr Thomas Wagner (CC

BY-SA 3.0)
164
Figure 10.19 Degradation criteria combinations for the Vwaza Marsh
Wildlife Reserve Key Biodiversity Area (KBA) in Malawi.

Figure 10.20 Food security criteria combinations for the Vwaza Marsh
Wildlife Reserve Key Biodiversity Area (KBA) in Malawi.

165
A unique fish fauna benefiting from the Nyika National Park, Vwaza
Marsh

Wildlife Reserve and North Rumphi Key Biodiversity Areas

Tweddle, D.1

1 South African Institute for Aquatic Biodiversity, 11 Somerset Street,

Grahamstown 6140, South Africa Origins of a unique fish fauna

The South Rukuru River system hosts a fish fauna distinct from that of al
other rivers flowing into LMNN, which share a fairly uniform riverine
fauna. Two species, Enteromius seymouri (Figure 10.21) and Labeobarbus
nthuwa (Figure 10.22), recently described from the South Rukuru system by
Tweddle & Skelton (2008) qualify both the existing Nyika National Park and
Vwaza Marsh Wildlife Reserve as KBAs for freshwater species, as well as
the newly delineated North Rumphi KBA. In addition, there are other
species in the South Rukuru, including a distinctive, undescribed fine-
spotted Enteromius species, at least one Amphilius species and a Chiloglanis
species, which are distinct from their congeners elsewhere in the LMNNC
(Tweddle & Skelton, 2008) and are currently under taxonomic investigation.
The South Rukuru is also the only river in Malawi where Clarias
liocephalus, a species typical of Upper Zambezi headwater tributaries,
occurs (Teugels, 1986; Tweddle et al., 2004). Many of these species are now
in decline and there is a need for national recognition of these global y
important KBAs in order to protect their populations, particularly those of
endemic species.

The South Rukuru River follows an unusual course in its upper reaches
(Figure 10.23). The river rises on the western slopes of the Viphya Plateau,
on the opposite side of the plateau to LMNN. It then turns north, skirting the
border with Zambia, which marks the watershed between the Luangwa River
system and the South Rukuru for approximately 150 km.

The river then turns east at the point where the drainage systems of the
Vwaza Marsh join the river. It is fed by several streams descending steeply
from the Nyika Plateau before the river cuts through the steep Njakwa Gorge
(Figure 10.24) near Rumphi and from there down to LMNN. After
descending the rift escarpment as a series of rapids cut through a heavily
faulted Karoo-age trough, and shortly before entering the lake, the South
Rukuru drops over the Wongwe and Fufu waterfal s, which form a barrier
preventing upstream movement of lake fishes (Tweddle & Skelton, 2008).

The very close proximity of the South Rukuru River to the Luangwa River
watershed on the Zambian border, and the presence of swamps and evidence
of former lakes in the area (Hopkins, 1973), suggests tectonic warping of the
land surface associated with the LMNN rifting has altered the

courses of the rivers in this area. Unconsolidated pebble

sheets up to 30 m above the present level of Lake Kazuni

(Figure 10.25, marked on Figure 10.23) probably represent

the littoral deposits of a once much more extensive lake

created by the tectonic movements that raised the rift

boundary. Prior to that, the present South Rukuru River

west of the Viphya Plateau would have been a west-flowing

Luangwa River tributary. A possible site of river capture

is marked on Figure 10.23, where the South Rukuru

Figure 10.21 Enteromius seymouri (Vulnerable, VU) is a

trigger species for Nyika National Park and Vwaza Marsh

appears likely to have been a south-flowing part of the

Wildlife Reserve and North Rumphi Key Biodiversity Areas

Luangwa tributary that arises at that site. The tectonic

(KBAs). © Denis Tweddle

warping would have resulted in the river reversing its flow

direction, cutting it off from the Luangwa system and

over-flowing from the proto-Lake Kazuni through the

Njakwa Gorge (marked on Figure 10.23) and then eroding

the river channel down to the present level (Tweddle &

Skelton, 2008). Given this geological history, it is likely

that the fish fauna is derived from the eastern escarpment

tributaries of the Luangwa River, which have not yet been

systematically sampled, and this explains the notable

differences between the South Rukuru fauna and that of all

Figure 10.22 Labeobarbus nthuwa (Near Threatened, NT) is

a trigger species for Nyika National Park and Vwaza Marsh

the other LMNN streams.

Wildlife Reserve Key Biodiversity Areas (KBAs). © Denis

Tweddle

166
Chilumba and Youngs Bay

Nyika National Park

North Rumphi

Rumphi

Vwaza Marsh Wildlife Reserve

Njakwa Gorge

Lake Kazuni

Luangwa

Val ey
Zambia

South Rukuru

Malawi

Site of possible river capture

Mzimba

Mzimba

Tukombo-Sanga-strip

0 5 10

20

30

40Kilometers

Source: Esri, DigitalGlobe, GeoEye, Earthstar Geographics, CNES/Airbus

DS, USDA, USGS, AeroGRID, IGN, and the GIS User Community

Figure 10.23 The South Rukuru River catchment. Key Biodiversity

Areas (KBAs) with freshwater trigger species in the Lake
Malawi/Nyasa/Niassa Catchment are overlaid (green hashed polygons),
including Vwaza Marsh Wildlife Reserve, Nyika National Park and
North Rumphi KBAs that are important for the conservation of
Enteromius

seymouri and Labeobarbus nthuwa.

167
Figure 10.24 The South Rukuru River passes through Njakwa Gorge. ©
Denis Tweddle

Species at risk

Early in the 20th century, the South Rukuru River at

Njakwa Gorge was described as a crystal clear sparkling

stream. However, by the 1970s the water was turbid

throughout the year as a result of silt from erosion in the

catchment caused by extensive land clearance. The

situation has continued to deteriorate. When sampled

in 1976 the Mzimba River in Mzimba town (Figure 10.26,

marked on Figure 10.23) was described as a “stream

2–4 m wide, up to 1 m deep, overhanging trees and other

Figure 10.25 Lake Kazuni. © Dr Thomas Wagner (CC BY-SA 3.0)

vegetation” (Tweddle & Willoughby, 1978). By 1992,

however, the river was wide and sandy and the only fish

Table 10.4 Samples taken by D. Tweddle when electric fishing in

caught were hiding in gabions at the road bridge. The river

the Mzimba River at Mzimba town.

continues to deteriorate, with these changes occurring as

Number of fish

a result of flash flooding due to the total deforestation of

Species

1976

1992

2010

the catchment upstream on the Viphya Plateau.

Enteromius seymouri

113

56

Enteromius kersteni

74

Jackson (1961) reported that C. liocephalus (under

Enteromius lineomaculatus

509

86

the name C. carsonii) was common in rocky habitats.

Enteromius paludinosus

90

71

Tweddle & Willoughby (1978) collected four specimens

Enteromius radiatus

from the Mzimba River in Mzimba town in 1976 but the

Enteromius sp. nov. fine spot

14

species has not been recorded since anywhere in the

Labeo cylindricus

36

South Rukuru system. Ongoing deterioration in the

Chiloglanis sp. nov.

66

river has had a major impact on the abundance and

Clarias liocephalus

distribution of most species, as il ustrated in Table 10.4.

Clarias gariepinus

Amphilius cf. uranoscopus

6
Zaireichthys sp.

168

Figure 10.26 Mzimba River at Mzimba town. © Denis Tweddle

Hope for the future – protected areas for fishes

The causes of this degradation in habitats appear

insurmountable in Malawi because of the high, and still

increasing, human population and consequent forest

clearance for agriculture. However, conservation actions

targeted to those areas where catchments, and hence

stream health, can stil be protected may bring success.

In this case, the Nyika National Park (Figure 10.27) protects

Figure 10. 27 The Nyika Nat ional Park protect s t he

the headwaters of several streams that originate on the

headwaters of several streams that originate on the plateau.

plateau, notably the South Rumphi and Runyina streams.

© firesika (CC BY-NC-ND 2.0)

Both of these are known to still support populations of

E. seymouri and L. nthuwa, and gravel spawning habitat is also secure within
the park boundaries. Enteromius seymouri still had a fairly healthy
population in the upper reaches of the South Rukuru when last sampled in
2010 (D. Tweddle unpublished data), but experience from the Mzimba River
suggests that this population is not secure. The headwaters on the Viphya
continue to be stripped of their woodland cover and so the Nyika streams are
vital strongholds for the survival of this species. A separate population of E.
seymouri occurs in the Kaziwiziwi stream that also flows from the Nyika
National Park KBA and is a tributary of the North Rumphi River (Tweddle
& Skelton, 2008). The North Rumphi is one of the few remaining healthy
river systems in the LMNNC because of the protection of its upper reaches
in the Nyika National Park, and it is also now a KBA for freshwater species.
The Vwaza Marsh Wildlife Reserve conserves extensive marsh habitat and
smal streams feeding Lake Kazuni. These wetlands stil support the smal
species in the river system, including the undescribed fine-spotted
Enteromius species (D. Tweddle unpublished data, sampled in 2010).

These two protected areas, now recognised as global y important sites

(KBAs) for freshwater fishes, as wel as the newly delineated North Rumphi
KBA, need to focus their management on conserving this unique fish fauna
of the South Rukuru.

Protected areas, traditional y set up to protect terrestrial species, can provide

significant benefits to freshwater species as demonstrated here.

169

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171

Chapter 11

How to find and access data on

freshwater species and sites

Sayer, C.A.1 and Palmer-Newton, A.F.1

Contents

11.1 The IUCN Red List of Threatened SpeciesTM

...........................................................................................................................
.....................172

11.1.1 What does a species Red List assessment include?

..........................................................................................................................
172

11.1.2 How do I search the IUCN Red List website to find a species?
..........................................................................................................173
11.1.3 How do I download a species Red List assessment and map?
.........................................................................................................175

11.2 Global Biodiversity Information Facility (GBIF)

...........................................................................................................................
...................175

11.3 World Database of Key Biodiversity Areas (WDKBAs)

...........................................................................................................................
.......176

11.3.1 How do I search the WDKBAs to find a KBA?

...........................................................................................................................
.........176

11.3.2 How do I download search results from the WDKBAs?

.....................................................................................................................178

11.4 Integrated Biodiversity Assessment Tool (IBAT)

...........................................................................................................................
.................178

The data col ated here on species (Red List data) and applied

presence, origin and seasonality in countries, as wel as the

to identify important sites for biodiversity (Key Biodiversity

extent of occurrence (EOO) and area of occupancy (AOO), if

Areas, KBAs) can be accessed through various online

available.

sources. Here we explain where and how to access the data.

Distribution map: A map of the species distribution,

primarily based on HydroBASINS (river and lake sub-basins)

11.1 The IUCN Red List of Threatened

for freshwater species.

SpeciesTM

Population: Population information, including on global

The IUCN Red List of Threatened SpeciesTM is the most

population size and trends, and population structure (e.g.

comprehensive information source on the extinction risk of

number, sizes and trends of subpopulations).

species of animals, plants and fungi. On the IUCN Red List

website (www.iucnredlist.org) you can read and download Habitat and

ecology: Information on suitable habitats

Red List assessments of species, including associated

and ecological requirements, including details relevant to

spatial data on species distributions.

extinction risk (e.g. particular life cycles, growth patterns

or behaviours that make species susceptible to specific

11.1.1 What does a species Red List

threats).

assessment include?

Threats: Information on the main threats currently affecting

Taxonomy: The higher taxonomy, scientific name, and

or likely to affect the taxon, including information on the

taxonomic authority, including information on any major

cause, scale, and the stress each threat places on the

synonyms or recent taxonomic changes.

species.

Assessment information: The IUCN Red List category

Use and trade: Information on any use and/or trade of the

assigned to a species, indicating its relative extinction risk,

species, including legal and illegal hunting and collection,

and the criteria used to assign this category together with a

and for local, national and international trade.

justification for why the species qualifies as such.

Conservation actions: Conservation and research actions

Geographic range: Distribution information, including on

in place and needed for the species.

1 Freshwater Biodiversity Unit, Global Species Programme, IUCN

(International Union for Conservation of Nature), David Attenborough
Building, Pembroke Street, Cambridge, CB2 3QZ, UK

172
11.1.2 How do I search the IUCN Red List

11.1.2.2 Using the advanced search

website to find a species?

There are many ways to refine a search on the Red List

Website. To do this, click on ‘Advanced’ next to the search

There are a number of ways to search for species Red List

bar (Figure 11.2) to display a list of search filters on the left

assessments on the IUCN Red List website.

of the screen. These filters include taxonomy, habitat types,

location, threats and more. You can also access this page

11.1.2.1 Searching by species name

directly at: www.iucnredlist.org/search.

To search by name, type the scientific or common name

of your species of interest into the search bar at the top of

the homescreen. If the species has been assessed on the

IUCN Red List it will appear below the search bar in the

results window. Click on the species box to open its Red List

Figure 11.2 Where to find the advanced search function on the

IUCN Red List website (www.iucnredlist.org).

assessment. If searching by common name please note that

the species wil only appear if the common name has been

recorded in the Red List database.

Clicking on the arrow next to the filter name wil open options

for more specific filter layers for that category.

Example: a user wants to find information on the chambo

species Oreochromis squamipinnis and so enter s

To apply a filter, select the corresponding tick box. Multiple

‘Oreochromis squamipinnis’ into the search box. They find

filters can be applied at any one time. The current filters

the species from the results and click on the species box to

wil be displayed at the top of the results window. Matching

open the assessment (Figure 11.1).

results wil appear in the results window on the right. Click on

a species box to open its Red List assessment. Make sure to

Not al species have photos on the Red List. In these cases,

clear al previous filters before starting a new search.

the species box wil appear as in Figure 11.1 but without an

image above the species name.

Example: a user is interested in species found in wetland

habitats. They click on ‘Habitats’ from the filter options and

then tick the box to the left of ‘Wetlands (inland)’. They can

then view all the assessed wetland species in the results

window (Figure 11.3).

Figure 11.3 Using the advanced search function on the IUCN

Red List website (www.iucnredlist.org) to select for species found in

wetlands.

Example: the same user then decides they wish to search

for all species in wetland habitats in the Afrotropics. They

click on the ‘Biogeographical Realms’ drop down options

and then tick the box to the left of ‘Afrotropical’. They can

then view all assessed wetland species recorded in the

Afrotropical realm in the results window (Figure 11.4).

Figure 11.1 Using the search function on the IUCN Red List

website (www.iucnredlist.org) to find the Red List assessment of chambo

( Oreochromis squamipinnis). Photo of Oreochromis

squamipinnis. © Ad Konings

173
Figure 11.4 Using the advanced search function on the IUCN Red List
website (www.iucnredlist.org) to select for species found in wetlands in
the Afrotropics.

Figure 11.5 Where to find the map search function on the IUCN Red
List website (www.iucnredlist.org).

11.1.2.3 Using the map search

and draw around the park boundary, clicking the first point

Species Red List assessments can also be found by

to finish the polygon and start the search. They can view all
searching within a region of interest on a map, although

species with published distribution maps that intersect the

the results will only display species that have a published

polygon in the results list on the left. In this case, there are

distribution map. To perform this search, click on ‘Advanced’

next to the search bar, then click on ‘Map’ from the tool bar

(Figure 11.5).

The region of interest can be drawn on the map as a polygon

using the polygon tool. This will search for any species

mapped to occur within the polygon drawn. The search

area can also be defined by placing a point on the map

using the point tool. This wil search for any mapped species

distributions that intersect the point within a 25 km radius

circle. The search results will be displayed in the results

window to the left of the map. Click on a species box from the

list to select and open its Red List assessment.

Example: a user wants to retrieve information on species

assessed and occurring within the Nyika National Park in

Figure 11.6 Using the map search function with the polygon

Malawi. They select the polygon tool while in the map search
tool on the IUCN Red List website (www.iucnredlist.org).

174

1,145 species with distribution range overlapping the Nyika

11.1.3 How do I download a species Red List

National Park (Figure 11.6). If a species distribution range

assessment and map?

overlaps the area selected it means the area is within the

known distribution range of the species but, as many species

Once you have found a species Red List assessment, you

have patchy distributions, it may not necessarily be present

can use the ‘Download’ button on the right to download the

within the area selected.

assessment in PDF form or the spatial data as a shapefile

(Figure 11.8).

Example: the same user then wants to perform a more

specific search within the park. They select the point tool and

Spatial data for selected taxonomic groups can also be

place a point in the north of the park. They can then view all

downloaded in bulk here: www.iucnredlist.org/resources/

species with published distribution maps that intersect the

spatial-data-download.

point in a 25 km radius circle (1,127 species) in the results list

on the left (Figure 11.7).

You wil need to sign up for an account on the IUCN Red List

website to download any spatial data and state how you

plan to use them before access is provided. The data are

made freely available for non-commercial use, to help inform

conservation planning and other non-commercial decision-

making processes. For commercial use of the data, visit

IBAT: www.ibat-al iance.org (see below for more details). For ful terms and
conditions of use see here: www.iucnredlist.

org/terms/terms-of-use.

11.2 Global Biodiversity Information

Facility (GBIF)

The Global Biodiversity Information Facility (GBIF) (www.

gbif.org) aims to provide open access to data about species

Figure 11.7 Using the map search function with the point tool on

on Earth. The point data records col ected through this study

the IUCN Red List website (www.iucnredlist.org).

are available to view and download on GBIF here: www.gbif.

org/dataset/1ccf6240-9a89-4c0e-bf1a-3e4ddd6197ba.

Click on the ‘Occurrences’ box to get details of the individual

It is also possible to filter the species list produced by the

records (including species name, country, coordinates, date

map search using the filters of the advanced search, as

and basis) and click on ‘Download’ to download the dataset

discussed above.

(Figure 11.9). You wil need to sign up for an account on GBIF

to download the dataset.

Figure 11.8 Downloading information from the IUCN Red List website
(www.iucnredlist.org). Photo of Oreochromis squamipinnis.

© Ad Konings

175

Figure 11.9 Viewing the dataset of point records on the Global

Biodiversity Information Facility (GBIF) website (www.gbif.org) with
where to view details of individual records and download the dataset.

11.3 World Database of Key

11.3.1.1 Using the map search

Biodiversity Areas (WDKBAs)

Selecting the ‘map search’ function will open map search

page, which can also be accessed directly here: www.

KBAs are sites contributing to the global persistence of

keybiodiversityareas.org/site/mapsearch. On the map biodiversity. On the

World Database of Key Biodiversity

search page you can either: i) use the ‘zoom to country’

Areas (WDKBAs) website (www.keybiodiversityareas.org) box to select and

zoom to a country of interest, or i ) use the

you can look at the boundaries of KBAs and follow links

navigation arrows to manual y zoom to your area of interest.

to find out more information about the trigger biodiversity

KBA boundaries appear on the map as brown polygons

elements they contain.

(Figure 11.11).

11.3.1 How do I search the WDKBAs to find a KBA?

Example: a user is interested in KBAs in Malawi. They select

There are two ways to search for a KBA on the WDKBAs:

‘Malawi’ in the dropdown box next to ‘zoom to country’ on

i) map search, and ii) text search. To access both of these

the map search page, and the map zooms to Malawi. KBAs

options, first click on ‘Search’ on the top toolbar and then

in Malawi and the surrounding area visible on the map are

select the relevant search from the dropdown (Figure 11.10).

shown as brown polygons (Figure 11.12).

Figure 11.10 Where to find the search functions on the World Database
of Key Biodiversity Areas (www.keybiodiversityareas.org).

176
Figure 11.11 Using the ‘zoom to county’ tool or the navigation arrows on
the map search function on the World Database of Key Biodiversity
Areas (www.keybiodiversityareas.org).

Figure 11.12 Using the map search function on the World Database of
Key Biodiversity Areas (www.keybiodiversityareas.org) to look at KBAs
in and around Malawi.

To find out more information about a KBA, click on its

on the map and then on its name in the pop up box that

polygon and a pop up with the KBA name will appear. Click

appears (Figure 11.13). This opens the summary KBA for the

on its KBA name to be taken to a summary page on the KBA

Nkhotakota Wildlife Reserve KBA. Here they fol ow the links

with basic information, such as the country of occurrence

to find out more information on the KBA (Figure 11.14).

and coordinates of the site, and links to further information

on external websites. To find out more information on al KBA

11.3.1.2 Using the text search

trigger species present click on the link ‘IBAT for Research

Selecting the ‘text search’ function will open map search

and Conservation Planning’. To find out more information

page, which can also be accessed directly here: www.

on any avian KBA trigger species present click on the link

keybiodiversityareas.org/site/search. On the text search

‘BirdLife DataZone IBA factsheet’.

page you can either: i) search for KBAs by continent using

the ‘region’ search, i ) search for KBAs by country using the

Example: the same user is particularly interested in

‘country/territory’ search, or iii) search for KBAs by name

Nkhotakota Wildlife Reserve KBA. They click on its polygon

using the ‘site name’ search (Figure 11.15). Please note that

177
Figure 11.13 Opening the summary page on the Nkhotakota Wildlife
Reserve KBA from the map search function on the World Database of
Key Biodiversity Areas (www.keybiodiversityareas.org).

Figure 11.14 The summary page of the Nkhotakota Wildlife Reserve

KBA on the World Database of Key Biodiversity Areas (www.

keybiodiversityareas.org).

the ‘region’ search can be used to filter the list of countries

need to provide your email address and agree to the terms

in the ‘country/territory’ search. Once you have entered your

and condition of use to download the data. The data are

search terms, click ‘Search’ to run the search and the results

made freely available for non-commercial use, to help

will be listed on a new page. Clicking on any of the KBA site

inform conservation planning and other non-commercial

names listed opens the summary page for the relevant KBA

decision-making processes. For commercial use of the

(e.g. Figure 11.14).

data, visit IBAT: www.ibat-al iance.org (see below for more details). For ful
terms and conditions of use see here: www.

Example: a user is interested in KBAs found in Tanzania.

keybiodiversityareas.org/info/dataterms.

They first select ‘Africa’ in the regions list and then select

‘Tanzania’ from the countries list (Figure 11.16). They then

click ‘Search’ and a list of KBAs in Tanzania appears on a new

11.4 Integrated Biodiversity

page (Figure 11.17).

Assessment Tool (IBAT)

11.3.2 How do I download search results from

The Integrated Biodiversity Assessment Tool ( IBAT )

the WDKBAs?

(www.ibat-alliance.org) hosts data from three key global biodiversity

datasets: The IUCN Red List of Threatened

Search results can be downloaded using the ‘Download

SpeciesTM, the World Database on Protected Areas (WDPA)

CSV file’ link at the top of the page (Figure 11.17). You will
and the WDKBAs.

178
Figure 11.15 Text search options on the World Database of Key
Biodiversity Areas (www.keybiodiversityareas.org).

Figure 11.16 Searching for KBAs in Tanzania using the text search
function on the World Database of Key Biodiversity Areas (www.

keybiodiversityareas.org).

Figure 11.17 Results of the text search for KBAs in Tanzania on the
World Database of Key Biodiversity Areas (www.

keybiodiversityareas.org) highlighting the link to download the search

results.

179
On IBAT you can:

■ Search and interact with the data map, applying data

■ Select a site of interest (Figure 11.18) and produce reports

layers on sites of KBAs, protected areas, and range rarity

on the fol owing:

of species.

• Species occurring within a set distance of the site,

■ Read and download country profiles, which can be used

based on data from the IUCN Red List

to support revisions of National Biodiversity Strategy and

• Freshwater species occurring within a set distance

Action Plans (NBSAPs).

upstream and downstream of the site, based on data

from the IUCN Red List

IBAT has a subscription service (free plans are available for

• Protected areas occurring within a set distance of the

non-commercial use) and al ows access to the Red List data

site, based on data from the WDPA

for commercial use.

• KBAs occurring within a set distance of the site,

based on data from the WDKBAs

Figure 11.18 Example of a site profile for Lake Malombe in Malawi on

the Integrated Biodiversity Assessment Tool (IBAT) website (www.ibat-
alliance.org).

180

Appendix 1. IUCN Red List assessment results

Decapods

Red List

Endemic to

Family

Binomial

Category
Red List Criteria

LMNNC

Shrimps

ATYIDAE

Caridina kaombeflutilis

DD

N/A

Yes

ATYIDAE

Caridina malawensis

DD

N/A

Yes

ATYIDAE

Caridina togoensis

LC

N/A

Crabs

POTAMONAUTIDAE

Potamonautes bel arussus

LC

N/A

POTAMONAUTIDAE

Potamonautes choloensis

LC

N/A

POTAMONAUTIDAE

Potamonautes lirrangensis

LC

N/A

POTAMONAUTIDAE

Potamonautes montivagus

LC

N/A

POTAMONAUTIDAE

Potamonautes obesus

LC

N/A

POTAMONAUTIDAE

Potamonautes suprasulcatus
LC

N/A

Fishes

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

ALESTIDAE

Brycinus imberi

LC

N/A

ALESTIDAE

Hemigrammopetersius barnardi

LC

N/A

AMPHILI DAE

Amphilius uranoscopus
LC

N/A

AMPHILI DAE

Zaireichthys compactus

DD

N/A

Yes

AMPHILI DAE

Zaireichthys lacustris

LC

N/A

Yes

AMPHILI DAE

Zaireichthys maravensis

LC

N/A

AMPHILI DAE

Zaireichthys monomotapa

LC

N/A
ANGUILLIDAE

Anguil a bengalensis

NT

N/A

BAGRIDAE

Bagrus meridionalis

CR

A2d

Yes

CICHLIDAE

Abactochromis labrosus

LC

N/A

Yes

CICHLIDAE

Alticorpus geoffreyi

LC

N/A

Yes

CICHLIDAE
Alticorpus macrocleithrum

LC

N/A

Yes

CICHLIDAE

Alticorpus mentale

LC

N/A

Yes

CICHLIDAE

Alticorpus peterdaviesi

LC

N/A

Yes

CICHLIDAE

Alticorpus profundicula

DD

N/A

Yes

CICHLIDAE
Aristochromis christyi

LC

N/A

Yes

CICHLIDAE

Astatotilapia cal iptera

LC

N/A

CICHLIDAE

Aulonocara aquilonium

LC

N/A

Yes

CICHLIDAE

Aulonocara auditor

DD

N/A

Yes

CICHLIDAE

Aulonocara baenschi
CR

B1ab(v)+2ab(v)

Yes

CICHLIDAE

Aulonocara brevinidus

LC

N/A

Yes

CICHLIDAE

Aulonocara ethelwynnae

NT

B1a+2a

Yes

CICHLIDAE

Aulonocara gertrudae

LC

N/A

Yes

CICHLIDAE

Aulonocara guentheri
EN

B1ab(v)

Yes

CICHLIDAE

Aulonocara hueseri

LC

N/A

Yes

CICHLIDAE

Aulonocara jacobfreibergi

LC

N/A

Yes

CICHLIDAE

Aulonocara kandeense

CR

B1ac(iv)+2ac(iv)

Yes

CICHLIDAE

Aulonocara koningsi
LC

N/A

Yes

CICHLIDAE

Aulonocara korneliae

LC

N/A

Yes

181

Appendix 1. IUCN Red List assessment results. Fishes, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

CICHLIDAE

Aulonocara maylandi

CR
B1ac(iv)+2ac(iv)

Yes

CICHLIDAE

Aulonocara nyassae

NT

B1a

Yes

CICHLIDAE

Aulonocara rostratum

LC

N/A

Yes

CICHLIDAE

Aulonocara saulosi

LC

N/A

Yes

CICHLIDAE

Aulonocara stonemani

LC
N/A

Yes

CICHLIDAE

Aulonocara stuartgranti

LC

N/A

Yes

CICHLIDAE

Aulonocara trematocephalum

DD

N/A

Yes

CICHLIDAE

Buccochromis heterotaenia

LC

N/A

Yes

CICHLIDAE

Buccochromis lepturus

LC
N/A

Yes

CICHLIDAE

Buccochromis nototaenia

LC

N/A

Yes

CICHLIDAE

Buccochromis rhoadesii

LC

N/A

Yes

CICHLIDAE

Buccochromis spectabilis

LC

N/A

Yes

CICHLIDAE

Caprichromis liemi

LC
N/A

Yes

CICHLIDAE

Caprichromis orthognathus

LC

N/A

Yes

CICHLIDAE

Champsochromis caeruleus

LC

N/A

Yes

CICHLIDAE

Champsochromis spilorhynchus

EN

A2ad

Yes

CICHLIDAE

Chilotilapia euchilus

LC
N/A

Yes

CICHLIDAE

Chilotilapia rhoadesii

LC

N/A

Yes

CICHLIDAE

Chindongo ater

NT

B1a+2a

Yes

CICHLIDAE

Chindongo bel icosus

LC

N/A

Yes

CICHLIDAE

Chindongo cyaneus

NT
B1a+2a

Yes

CICHLIDAE

Chindongo demasoni

VU

D1+2

Yes

CICHLIDAE

Chindongo elongatus

NT

B1a

Yes

CICHLIDAE

Chindongo flavus

NT

B1a+2a

Yes

CICHLIDAE

Chindongo heteropictus

LC
N/A

Yes

CICHLIDAE

Chindongo longior

LC

N/A

Yes

CICHLIDAE

Chindongo minutus

LC

N/A

Yes

CICHLIDAE

Chindongo saulosi

CR

B1ab(v)+2ab(v)

Yes

CICHLIDAE

Chindongo socolofi

LC
N/A

Yes

CICHLIDAE

Copadichromis atripinnis

LC

N/A

Yes

CICHLIDAE

Copadichromis azureus

NT

B1a+2a

Yes

CICHLIDAE

Copadichromis borleyi

LC

N/A

Yes

CICHLIDAE

Copadichromis chizumuluensis

LC
N/A

Yes

CICHLIDAE

Copadichromis chrysonotus

LC

N/A

Yes

CICHLIDAE

Copadichromis cyaneus

LC

N/A

Yes

CICHLIDAE

Copadichromis cyanocephalus

NT

B1a

Yes

CICHLIDAE

Copadichromis diplostigma

NT
B1a

Yes

CICHLIDAE

Copadichromis geertsi

LC

N/A

Yes

CICHLIDAE

Copadichromis ilesi

LC

N/A

Yes

CICHLIDAE

Copadichromis insularis

LC

N/A

Yes

CICHLIDAE

Copadichromis jacksoni

LC
N/A

Yes

CICHLIDAE

Copadichromis likomae

LC

N/A

Yes

CICHLIDAE

Copadichromis mbenji

LC

N/A

Yes

CICHLIDAE

Copadichromis melas

LC

N/A

Yes

CICHLIDAE

Copadichromis mloto

DD
N/A

Yes

182

Appendix 1. IUCN Red List assessment results. Fishes, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

CICHLIDAE

Copadichromis nkatae

CR

B2ab(v)

Yes

CICHLIDAE

Copadichromis parvus

LC

N/A
Yes

CICHLIDAE

Copadichromis pleurostigma

LC

N/A

Yes

CICHLIDAE

Copadichromis pleurostigmoides

LC

N/A

Yes

CICHLIDAE

Copadichromis quadrimaculatus

LC

N/A

Yes

CICHLIDAE

Copadichromis trewavasae

LC

N/A
Yes

CICHLIDAE

Copadichromis trimaculatus

LC

N/A

Yes

CICHLIDAE

Copadichromis verduyni

LC

N/A

Yes

CICHLIDAE

Copadichromis virginalis

NT

A2bd

Yes

CICHLIDAE

Coptodon rendal i

LC

N/A
CICHLIDAE

Corematodus shiranus

CR

A2d

Yes

CICHLIDAE

Corematodus taeniatus

LC

N/A

Yes

CICHLIDAE

Ctenopharynx intermedius

LC

N/A

Yes

CICHLIDAE

Ctenopharynx nitidus

LC

N/A

Yes
CICHLIDAE

Ctenopharynx pictus

LC

N/A

Yes

CICHLIDAE

Cyathochromis obliquidens

LC

N/A

Yes

CICHLIDAE

Cynotilapia afra

LC

N/A

Yes

CICHLIDAE

Cynotilapia aurifrons

LC

N/A

Yes
CICHLIDAE

Cynotilapia axelrodi

LC

N/A

Yes

CICHLIDAE

Cynotilapia chilundu

VU

D1

Yes

CICHLIDAE

Cynotilapia zebroides

LC

N/A

Yes

CICHLIDAE

Cyrtocara moori

VU

A2a

Yes
CICHLIDAE

Dimidiochromis compressiceps

LC

N/A

Yes

CICHLIDAE

Dimidiochromis dimidiatus

LC

N/A

Yes

CICHLIDAE

Dimidiochromis kiwinge

LC

N/A

Yes

CICHLIDAE

Dimidiochromis strigatus

LC

N/A

Yes
CICHLIDAE

Diplotaxodon aeneus

LC

N/A

Yes

CICHLIDAE

Diplotaxodon altus

LC

N/A

Yes

CICHLIDAE

Diplotaxodon argenteus

LC

N/A

Yes

CICHLIDAE

Diplotaxodon ecclesi

LC

N/A

Yes
CICHLIDAE

Diplotaxodon greenwoodi

LC

N/A

Yes

CICHLIDAE

Diplotaxodon limnothrissa

LC

N/A

Yes

CICHLIDAE

Diplotaxodon longimaxil a

LC

N/A

Yes

CICHLIDAE

Diplotaxodon macrops

LC

N/A

Yes
CICHLIDAE

Docimodus evelynae

LC

N/A

Yes

CICHLIDAE

Docimodus johnstoni

LC

N/A

Yes

CICHLIDAE

Exochochromis anagenys

LC

N/A

Yes

CICHLIDAE

Fossorochromis rostratus

LC

N/A

Yes
CICHLIDAE

Genyochromis mento

LC

N/A

Yes

CICHLIDAE

Gephyrochromis lawsi

LC

N/A

Yes

CICHLIDAE

Gephyrochromis moori

LC

N/A

Yes

CICHLIDAE

Haplochromis tweddlei

DD

N/A

CICHLIDAE
Hemitaeniochromis brachyrhynchus

LC

N/A

Yes

CICHLIDAE

Hemitaeniochromis spilopterus

LC

N/A

Yes

CICHLIDAE

Hemitaeniochromis urotaenia

LC

N/A

Yes

183

Appendix 1. IUCN Red List assessment results. Fishes, cont’d

Red List

Endemic to

Family

Binomial
Category

Red List Criteria

LMNNC

CICHLIDAE

Hemitilapia oxyrhynchus

LC

N/A

Yes

CICHLIDAE

Iodotropheus sprengerae

NT

B1a+2a

Yes

CICHLIDAE

Iodotropheus stuartgranti

LC

N/A

Yes

CICHLIDAE

Labeotropheus artatorostris
LC

N/A

Yes

CICHLIDAE

Labeotropheus chlorosiglos

LC

N/A

Yes

CICHLIDAE

Labeotropheus fuel eborni

LC

N/A

Yes

CICHLIDAE

Labeotropheus simoneae

LC

N/A

Yes

CICHLIDAE

Labeotropheus trewavasae
LC

N/A

Yes

CICHLIDAE

Labidochromis caeruleus

LC

N/A

Yes

CICHLIDAE

Labidochromis chisumulae

LC

N/A

Yes

CICHLIDAE

Labidochromis flavigulis

LC

N/A

Yes

CICHLIDAE

Labidochromis freibergi
LC

N/A

Yes

CICHLIDAE

Labidochromis gigas

LC

N/A

Yes

CICHLIDAE

Labidochromis heterodon

LC

N/A

Yes

CICHLIDAE

Labidochromis ianthinus

LC

N/A

Yes

CICHLIDAE

Labidochromis joanjohnsonae
NT

B1a

Yes

CICHLIDAE

Labidochromis lividus

LC

N/A

Yes

CICHLIDAE

Labidochromis maculicauda

LC

N/A

Yes

CICHLIDAE

Labidochromis mathotho

DD

N/A

Yes

CICHLIDAE

Labidochromis mbenji
LC

N/A

Yes

CICHLIDAE

Labidochromis mylodon

LC

N/A

Yes

CICHLIDAE

Labidochromis pal idus

LC

N/A

Yes

CICHLIDAE

Labidochromis shiranus

LC

N/A

Yes

CICHLIDAE

Labidochromis strigatus
LC

N/A

Yes

CICHLIDAE

Labidochromis textilis

LC

N/A

Yes

CICHLIDAE

Labidochromis vel icans

LC

N/A

Yes

CICHLIDAE

Labidochromis zebroides

EN

Yes

CICHLIDAE

Lethrinops albus
LC

N/A

Yes

CICHLIDAE

Lethrinops altus

LC

N/A

Yes

CICHLIDAE

Lethrinops argenteus

LC

N/A

Yes

CICHLIDAE

Lethrinops auritus

LC

N/A

Yes

CICHLIDAE

Lethrinops christyi
LC

N/A

Yes

CICHLIDAE

Lethrinops furcifer

LC

N/A

Yes

CICHLIDAE

Lethrinops gossei

LC

N/A

Yes

CICHLIDAE

Lethrinops leptodon

LC

N/A

Yes

CICHLIDAE

Lethrinops lethrinus
LC

N/A

Yes

CICHLIDAE

Lethrinops longimanus

LC

N/A

Yes

CICHLIDAE

Lethrinops longipinnis

LC

N/A

Yes

CICHLIDAE

Lethrinops lunaris

LC

N/A

Yes

CICHLIDAE

Lethrinops macracanthus
DD

N/A

Yes

CICHLIDAE

Lethrinops macrochir

LC

N/A

Yes

CICHLIDAE

Lethrinops macrophthalmus

LC

N/A

Yes

CICHLIDAE

Lethrinops marginatus

LC

N/A

Yes

CICHLIDAE

Lethrinops micrentodon
DD

N/A

Yes

CICHLIDAE

Lethrinops microdon

DD

N/A

Yes

184

Appendix 1. IUCN Red List assessment results. Fishes, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

CICHLIDAE

Lethrinops microstoma

LC
N/A

Yes

CICHLIDAE

Lethrinops mylodon

LC

N/A

Yes

CICHLIDAE

Lethrinops parvidens

LC

N/A

Yes

CICHLIDAE

Lethrinops stridei

DD

N/A

Yes

CICHLIDAE

Lethrinops turneri

LC
N/A

Yes

CICHLIDAE

Lichnochromis acuticeps

LC

N/A

Yes

CICHLIDAE

Mchenga conophoros

CR

B1ab(v)

Yes

CICHLIDAE

Mchenga cyclicos

NT

B1a+2a

Yes

CICHLIDAE

Mchenga eucinostomus

LC
N/A

Yes

CICHLIDAE

Mchenga flavimanus

LC

N/A

Yes

CICHLIDAE

Mchenga inornata

DD

N/A

Yes

CICHLIDAE

Mchenga thinos

LC

N/A

Yes

CICHLIDAE

Melanochromis auratus

LC
N/A

Yes

CICHLIDAE

Melanochromis baliodigma

LC

N/A

Yes

CICHLIDAE

Melanochromis chipokae

CR

A2a; B2ab(v)

Yes

CICHLIDAE

Melanochromis dialeptos

LC

N/A

Yes

CICHLIDAE

Melanochromis heterochromis

LC
N/A

Yes

CICHLIDAE

Melanochromis kaskazini

LC

N/A

Yes

CICHLIDAE

Melanochromis lepidiadaptes

CR

B1ab(v)

Yes

CICHLIDAE

Melanochromis loriae

LC

N/A

Yes

CICHLIDAE

Melanochromis melanopterus

LC
N/A

Yes

CICHLIDAE

Melanochromis mossambiquensis

NT

B1a+2a

Yes

CICHLIDAE

Melanochromis mpoto

LC

N/A

Yes

CICHLIDAE

Melanochromis robustus

NT

B1a+2a

Yes

CICHLIDAE

Melanochromis simulans

LC
N/A

Yes

CICHLIDAE

Melanochromis vermivorus

NT

B1a+2a

Yes

CICHLIDAE

Melanochromis wochepa

NT

B2a

Yes

CICHLIDAE

Metriaclima aurora

LC

N/A

Yes

CICHLIDAE

Metriaclima barlowi

LC
N/A

Yes

CICHLIDAE

Metriaclima benetos

NT

B1a

Yes

CICHLIDAE

Metriaclima cal ainos

LC

N/A

Yes

CICHLIDAE

Metriaclima chrysomal os

NT

B1a

Yes

CICHLIDAE

Metriaclima cyneusmarginatum

NT
B1a

Yes

CICHLIDAE

Metriaclima emmiltos

LC

N/A

Yes

CICHLIDAE

Metriaclima estherae

LC

N/A

Yes

CICHLIDAE

Metriaclima fainzilberi

LC

N/A

Yes

CICHLIDAE

Metriaclima flavicauda

VU
D1

Yes

CICHLIDAE

Metriaclima flavifemina

LC

N/A

Yes

CICHLIDAE

Metriaclima glaucos

NT

B1a

Yes

CICHLIDAE

Metriaclima greshakei

NT

B1a+2a

Yes

CICHLIDAE

Metriaclima hajomaylandi

LC
N/A

Yes

CICHLIDAE

Metriaclima koningsi

CR

B1ab(v)+2ab(v)

Yes

CICHLIDAE

Metriaclima lanisticola

LC

N/A

Yes

CICHLIDAE

Metriaclima lombardoi

LC

N/A

Yes

CICHLIDAE

Metriaclima lundoense

NT
B1a+2a

Yes

185

Appendix 1. IUCN Red List assessment results. Fishes, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

CICHLIDAE

Metriaclima mbenji

LC

N/A

Yes

CICHLIDAE

Metriaclima midomo

NT

B1a+2a
Yes

CICHLIDAE

Metriaclima mossambicum

LC

N/A

Yes

CICHLIDAE

Metriaclima nigrodorsalis

LC

N/A

Yes

CICHLIDAE

Metriaclima nkhunguense

LC

N/A

Yes

CICHLIDAE

Metriaclima pambazuko

LC

N/A
Yes

CICHLIDAE

Metriaclima phaeos

LC

N/A

Yes

CICHLIDAE

Metriaclima pulpican

LC

N/A

Yes

CICHLIDAE

Metriaclima pyrsonotos

LC

N/A

Yes

CICHLIDAE

Metriaclima sciasma

LC

N/A
Yes

CICHLIDAE

Metriaclima tarakiki

LC

N/A

Yes

CICHLIDAE

Metriaclima usisyae

CR

B1ab(v)+2ab(v)

Yes

CICHLIDAE

Metriaclima xanstomachus

NT

B1a+2a

Yes

CICHLIDAE

Metriaclima xanthos

LC

N/A
Yes

CICHLIDAE

Metriaclima zebra

LC

N/A

Yes

CICHLIDAE

Mylochromis anaphyrmus

LC

N/A

Yes

CICHLIDAE

Mylochromis balteatus

LC

N/A

Yes

CICHLIDAE

Mylochromis chekopae

LC

N/A
Yes

CICHLIDAE

Mylochromis ensatus

LC

N/A

Yes

CICHLIDAE

Mylochromis epichorialis

LC

N/A

Yes

CICHLIDAE

Mylochromis ericotaenia

LC

N/A

Yes

CICHLIDAE

Mylochromis formosus

LC

N/A
Yes

CICHLIDAE

Mylochromis gracilis

LC

N/A

Yes

CICHLIDAE

Mylochromis guentheri

LC

N/A

Yes

CICHLIDAE

Mylochromis incola

LC

N/A

Yes

CICHLIDAE

Mylochromis labidodon

LC

N/A
Yes

CICHLIDAE

Mylochromis lateristriga

LC

N/A

Yes

CICHLIDAE

Mylochromis melanonotus

LC

N/A

Yes

CICHLIDAE

Mylochromis melanotaenia

LC

N/A

Yes

CICHLIDAE

Mylochromis mola

LC

N/A
Yes

CICHLIDAE

Mylochromis mol is

LC

N/A

Yes

CICHLIDAE

Mylochromis obtusus

LC

N/A

Yes

CICHLIDAE

Mylochromis plagiotaenia

LC

N/A

Yes

CICHLIDAE

Mylochromis sphaerodon

LC

N/A
Yes

CICHLIDAE

Mylochromis spilostichus

LC

N/A

Yes

CICHLIDAE

Mylochromis subocularis

LC

N/A

Yes

CICHLIDAE

Naevochromis chrysogaster

LC

N/A

Yes

CICHLIDAE

Nimbochromis fuscotaeniatus

VU

A2a
Yes

CICHLIDAE

Nimbochromis linni

LC

N/A

CICHLIDAE

Nimbochromis livingstoni

LC

N/A

Yes

CICHLIDAE

Nimbochromis polystigma

LC

N/A

Yes

CICHLIDAE

Nimbochromis venustus

LC

N/A

Yes
CICHLIDAE

Nyassachromis boadzulu

EN

A2d; B1ab(v)+2ab(v)

Yes

CICHLIDAE

Nyassachromis breviceps

CR

B2ab(v)

Yes

CICHLIDAE

Nyassachromis leuciscus

DD

N/A

Yes

186

Appendix 1. IUCN Red List assessment results. Fishes, cont’d

Red List

Endemic to

Family
Binomial

Category

Red List Criteria

LMNNC

CICHLIDAE

Nyassachromis microcephalus

LC

N/A

Yes

CICHLIDAE

Nyassachromis nigritaeniatus

LC

N/A

Yes

CICHLIDAE

Nyassachromis prostoma

LC

N/A

Yes

CICHLIDAE
Nyassachromis purpurans

LC

N/A

Yes

CICHLIDAE

Nyassachromis serenus

LC

N/A

Yes

CICHLIDAE

Oreochromis chungruruensis

CR

B1ab(v)+2ab(v)

Yes

CICHLIDAE

Oreochromis karongae

CR

A2d

Yes

CICHLIDAE
Oreochromis lidole

CR

A2d

Yes

CICHLIDAE

Oreochromis shiranus

LC

N/A

CICHLIDAE

Oreochromis squamipinnis

CR

A2d

Yes

CICHLIDAE

Otopharynx antron

LC

N/A

Yes

CICHLIDAE

Otopharynx argyrosoma
LC

N/A

Yes

CICHLIDAE

Otopharynx auromarginatus

LC

N/A

Yes

CICHLIDAE

Otopharynx brooksi

LC

N/A

Yes

CICHLIDAE

Otopharynx decorus

LC

N/A

Yes

CICHLIDAE

Otopharynx heterodon
LC

N/A

Yes

CICHLIDAE

Otopharynx lithobates

LC

N/A

Yes

CICHLIDAE

Otopharynx ovatus

LC

N/A

Yes

CICHLIDAE

Otopharynx pachycheilus

VU

D2

Yes

CICHLIDAE

Otopharynx selenurus
DD

N/A

Yes

CICHLIDAE

Otopharynx speciosus

LC

N/A

Yes

CICHLIDAE

Otopharynx spelaeotes

LC

N/A

Yes

CICHLIDAE

Otopharynx tetraspilus

LC

N/A

Yes

CICHLIDAE

Otopharynx tetrastigma
LC

N/A

Yes

CICHLIDAE

Pal idochromis tokolosh

LC

N/A

Yes

CICHLIDAE

Petrotilapia chrysos

LC

N/A

Yes

CICHLIDAE

Petrotilapia flaviventris

LC

N/A

Yes

CICHLIDAE

Petrotilapia genalutea
LC

N/A

Yes

CICHLIDAE

Petrotilapia microgalana

LC

N/A

Yes

CICHLIDAE

Petrotilapia mumboensis

LC

N/A

Yes

CICHLIDAE

Petrotilapia nigra

LC

N/A

Yes

CICHLIDAE

Petrotilapia palingnathos
LC

N/A

Yes

CICHLIDAE

Petrotilapia pyroscelos

LC

N/A

Yes

CICHLIDAE

Petrotilapia tridentiger

LC

N/A

Yes

CICHLIDAE

Petrotilapia xanthos

NT

B1a+2a

Yes

CICHLIDAE

Placidochromis acuticeps
DD

N/A

Yes

CICHLIDAE

Placidochromis acutirostris

DD

N/A

Yes

CICHLIDAE

Placidochromis argyrogaster

LC

N/A

Yes

CICHLIDAE

Placidochromis boops

LC

N/A

Yes

CICHLIDAE

Placidochromis borealis
DD

N/A

Yes

CICHLIDAE

Placidochromis chilolae

DD

N/A

Yes

CICHLIDAE

Placidochromis communis

LC

N/A

Yes

CICHLIDAE

Placidochromis domirae

DD

N/A

Yes

CICHLIDAE

Placidochromis ecclesi
LC

N/A

Yes

CICHLIDAE

Placidochromis electra

LC

N/A

Yes

187

Appendix 1. IUCN Red List assessment results. Fishes, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

CICHLIDAE

Placidochromis elongatus

LC
N/A

Yes

CICHLIDAE

Placidochromis fuscus

DD

N/A

Yes

CICHLIDAE

Placidochromis hennydaviesae

LC

N/A

Yes

CICHLIDAE

Placidochromis intermedius

LC

N/A

Yes

CICHLIDAE

Placidochromis johnstoni

LC
N/A

Yes

CICHLIDAE

Placidochromis koningsi

DD

N/A

Yes

CICHLIDAE

Placidochromis lineatus

DD

N/A

Yes

CICHLIDAE

Placidochromis longimanus

LC

N/A

Yes

CICHLIDAE

Placidochromis longirostris

LC
N/A

Yes

CICHLIDAE

Placidochromis longus

DD

N/A

Yes

CICHLIDAE

Placidochromis lukomae

LC

N/A

Yes

CICHLIDAE

Placidochromis macroceps

DD

N/A

Yes

CICHLIDAE

Placidochromis macrognathus

LC
N/A

Yes

CICHLIDAE

Placidochromis mbunoides

LC

N/A

Yes

CICHLIDAE

Placidochromis milomo

LC

N/A

Yes

CICHLIDAE

Placidochromis minor

DD

N/A

Yes

CICHLIDAE

Placidochromis minutus

DD
N/A

Yes

CICHLIDAE

Placidochromis msakae

DD

N/A

Yes

CICHLIDAE

Placidochromis nigribarbis

DD

N/A

Yes

CICHLIDAE

Placidochromis nkhatae

DD

N/A

Yes

CICHLIDAE

Placidochromis nkhotakotae

DD
N/A

Yes

CICHLIDAE

Placidochromis obscurus

LC

N/A

Yes

CICHLIDAE

Placidochromis ordinarius

LC

N/A

Yes

CICHLIDAE

Placidochromis orthognathus

DD

N/A

Yes

CICHLIDAE

Placidochromis pal idus

DD
N/A

Yes

CICHLIDAE

Placidochromis phenochilus

EN

A2a; B2ab(v)

Yes

CICHLIDAE

Placidochromis platyrhynchos

LC

N/A

Yes

CICHLIDAE

Placidochromis pol i

LC

N/A

Yes

CICHLIDAE

Placidochromis rotundifrons

LC
N/A

Yes

CICHLIDAE

Placidochromis trewavasae

LC

N/A

Yes

CICHLIDAE

Placidochromis turneri

LC

N/A

Yes

CICHLIDAE

Placidochromis vulgaris

DD

N/A

Yes

CICHLIDAE

Protomelas annectens

LC
N/A

Yes

CICHLIDAE

Protomelas fenestratus

LC

N/A

Yes

CICHLIDAE

Protomelas insignis

LC

N/A

Yes

CICHLIDAE

Protomelas kirki

LC

N/A

Yes

CICHLIDAE

Protomelas labridens

LC
N/A

Yes

CICHLIDAE

Protomelas macrodon

DD

N/A

Yes

CICHLIDAE

Protomelas marginatus

LC

N/A

Yes

CICHLIDAE

Protomelas ornatus

LC

N/A

Yes

CICHLIDAE

Protomelas pleurotaenia

LC
N/A

Yes

CICHLIDAE

Protomelas similis

LC

N/A

Yes

CICHLIDAE

Protomelas spilonotus

LC

N/A

Yes

CICHLIDAE

Protomelas taeniolatus

LC

N/A

Yes

CICHLIDAE

Protomelas triaenodon

LC
N/A

Yes

188

Appendix 1. IUCN Red List assessment results. Fishes, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

CICHLIDAE

Protomelas virgatus

LC

N/A

Yes

CICHLIDAE

Pseudocrenilabrus philander

LC

N/A
CICHLIDAE

Pseudotropheus benetos

LC

N/A

Yes

CICHLIDAE

Pseudotropheus brevis

EN

B1ab(i )+2ab(i )

Yes

CICHLIDAE

Pseudotropheus crabro

LC

N/A

Yes

CICHLIDAE

Pseudotropheus cyaneorhabdos

CR

A2a; B2ab(v)

Yes
CICHLIDAE

Pseudotropheus elegans

LC

N/A

Yes

CICHLIDAE

Pseudotropheus fuscus

LC

N/A

Yes

CICHLIDAE

Pseudotropheus galanos

NT

B1a

Yes

CICHLIDAE

Pseudotropheus interruptus

NT

B1a

Yes
CICHLIDAE

Pseudotropheus johanni

LC

N/A

Yes

CICHLIDAE

Pseudotropheus livingstoni

LC

N/A

Yes

CICHLIDAE

Pseudotropheus lucerna

LC

N/A

Yes

CICHLIDAE

Pseudotropheus perileucos

LC

N/A

Yes
CICHLIDAE

Pseudotropheus perspicax

LC

N/A

Yes

CICHLIDAE

Pseudotropheus purpuratus

LC

N/A

Yes

CICHLIDAE

Pseudotropheus tursiops

NT

B1a

Yes

CICHLIDAE

Pseudotropheus wil iamsi

NT

B1a

Yes
CICHLIDAE

Rhamphochromis brevis

LC

N/A

Yes

CICHLIDAE

Rhamphochromis esox

VU

A2d

Yes

CICHLIDAE

Rhamphochromis ferox

DD

N/A

Yes

CICHLIDAE

Rhamphochromis longiceps

VU

A2d

Yes
CICHLIDAE

Rhamphochromis woodi

LC

N/A

Yes

CICHLIDAE

Sciaenochromis ahli

LC

N/A

Yes

CICHLIDAE

Sciaenochromis benthicola

LC

N/A

Yes

CICHLIDAE

Sciaenochromis fryeri

LC

N/A

Yes
CICHLIDAE

Sciaenochromis psammophilus

LC

N/A

Yes

CICHLIDAE

Serranochromis robustus

CR

A2c+3cde

CICHLIDAE

Stigmatochromis macrorhynchos

LC

N/A

Yes

CICHLIDAE

Stigmatochromis melanchros

LC

N/A

Yes

CICHLIDAE
Stigmatochromis modestus

LC

N/A

Yes

CICHLIDAE

Stigmatochromis pholidophorus

LC

N/A

Yes

CICHLIDAE

Stigmatochromis pleurospilus

DD

N/A

Yes

CICHLIDAE

Stigmatochromis woodi

LC

N/A

Yes

CICHLIDAE
Taeniochromis holotaenia

LC

N/A

Yes

CICHLIDAE

Taeniolethrinops cyrtonotus

LC

N/A

Yes

CICHLIDAE

Taeniolethrinops furcicauda

LC

N/A

Yes

CICHLIDAE

Taeniolethrinops laticeps

LC

N/A

Yes

CICHLIDAE
Taeniolethrinops macrorhynchus

LC

N/A

Yes

CICHLIDAE

Taeniolethrinops praeorbitalis

LC

N/A

Yes

CICHLIDAE

Tilapia sparrmani

LC

N/A

CICHLIDAE

Tramitichromis brevis

LC

N/A

Yes

CICHLIDAE

Tramitichromis intermedius
LC

N/A

Yes

CICHLIDAE

Tramitichromis lituris

LC

N/A

Yes

CICHLIDAE

Tramitichromis trilineatus

LC

N/A

Yes

189

Appendix 1. IUCN Red List assessment results. Fishes, cont’d

Red List

Endemic to

Family

Binomial

Category
Red List Criteria

LMNNC

CICHLIDAE

Tramitichromis variabilis

LC

N/A

Yes

CICHLIDAE

Trematocranus brevirostris

LC

N/A

Yes

CICHLIDAE

Trematocranus labifer

DD

N/A

Yes

CICHLIDAE

Trematocranus microstoma

EN
A2ab

Yes

CICHLIDAE

Trematocranus pachychilus

DD

N/A

Yes

CICHLIDAE

Trematocranus placodon

LC

N/A

Yes

CICHLIDAE

Tropheops biriwira

NT

B1a+2a

Yes

CICHLIDAE

Tropheops gracilior

LC
N/A

Yes

CICHLIDAE

Tropheops kamtambo

LC

N/A

Yes

CICHLIDAE

Tropheops kumwera

NT

B1a+2a

Yes

CICHLIDAE

Tropheops macrophthalmus

LC

N/A

Yes

CICHLIDAE

Tropheops microstoma

NT
B1a+2a

Yes

CICHLIDAE

Tropheops modestus

NT

B1a

Yes

CICHLIDAE

Tropheops novemfasciatus

LC

N/A

Yes

CICHLIDAE

Tropheops romandi

LC

N/A

Yes

CICHLIDAE

Tropheops tropheops

LC
N/A

Yes

CICHLIDAE

Tyrannochromis macrostoma

LC

N/A

Yes

CICHLIDAE

Tyrannochromis nigriventer

LC

N/A

Yes

CLARI DAE

Bathyclarias atribranchus

LC

N/A

Yes

CLARI DAE

Bathyclarias eurydon

LC
N/A

Yes

CLARI DAE

Bathyclarias filicibarbis

LC

N/A

Yes

CLARI DAE

Bathyclarias foveolatus

LC

N/A

Yes

CLARI DAE

Bathyclarias longibarbis

LC

N/A

Yes

CLARI DAE

Bathyclarias nyasensis

LC
N/A

Yes

CLARI DAE

Bathyclarias rotundifrons

LC

N/A

Yes

CLARI DAE

Bathyclarias worthingtoni

LC

N/A

Yes

CLARI DAE

Clarias gariepinus

LC

N/A

CLARI DAE

Clarias liocephalus

LC

N/A
CLARI DAE

Clarias ngamensis

LC

N/A

CLARI DAE

Clarias stappersii

LC

N/A

CLARI DAE

Clarias theodorae

LC

N/A

CYPRINIDAE

Engraulicypris ngalala

LC

N/A

CYPRINIDAE

Engraulicypris sardel a

LC

N/A
Yes

CYPRINIDAE

Enteromius arcislongae

LC

N/A

CYPRINIDAE

Enteromius atkinsoni

LC

N/A

CYPRINIDAE

Enteromius bifrenatus

LC

N/A

CYPRINIDAE

Enteromius eutaenia

LC

N/A

CYPRINIDAE

Enteromius innocens

LC
N/A

CYPRINIDAE

Enteromius kersteni

LC

N/A

CYPRINIDAE

Enteromius lineomaculatus

LC

N/A

CYPRINIDAE

Enteromius litamba

DD

N/A

Yes

CYPRINIDAE

Enteromius macrotaenia

LC

N/A

CYPRINIDAE

Enteromius paludinosus
LC

N/A

CYPRINIDAE

Enteromius radiatus

LC

N/A

CYPRINIDAE

Enteromius seymouri

VU

A3c

Yes

190

Appendix 1. IUCN Red List assessment results. Fishes, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC
CYPRINIDAE

Enteromius toppini

LC

N/A

CYPRINIDAE

Enteromius trimaculatus

LC

N/A

CYPRINIDAE

Enteromius zanzibaricus

LC

N/A

CYPRINIDAE

Labeo cylindricus

LC

N/A

CYPRINIDAE

Labeo mesops

CR

A2ac+3cd
CYPRINIDAE

Labeo worthingtoni

EX

N/A

Yes

CYPRINIDAE

Labeobarbus johnstoni

LC

N/A

CYPRINIDAE

Labeobarbus latirostris

DD

N/A

Yes

CYPRINIDAE

Labeobarbus nthuwa

NT

B1a

Yes

CYPRINIDAE
Opsaridium microcephalum

LC

N/A

CYPRINIDAE

Opsaridium microlepis

VU

A3cd

CYPRINIDAE

Opsaridium tweddleorum

DD

N/A

MASTACEMBELIDAE

Mastacembelus shiranus

LC

N/A

MOCHOKIDAE

Synodontis njassae

LC

N/A

MORMYRIDAE
Cyphomyrus discorhynchus

LC

N/A

MORMYRIDAE

Hippopotamyrus ansorgi

LC

N/A

MORMYRIDAE

Marcusenius livingstoni

LC

N/A

MORMYRIDAE

Marcusenius macrolepidotus

LC

N/A

MORMYRIDAE

Mormyrops anguil oides

LC

N/A

MORMYRIDAE
Mormyrus longirostris

LC

N/A

MORMYRIDAE

Petrocephalus catostoma

LC

N/A

NOTHOBRANCHI DAE

Nothobranchius kirki

VU

B1ab(i i)+2ab(i i)

NOTHOBRANCHI DAE

Nothobranchius wattersi

NT

B1b(i i)+2b(i i)

POECILI DAE

Micropanchax johnstoni

LC

N/A

SCHILBEIDAE
Pareutropius longifilis

LC

N/A

Molluscs

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

AMPULLARI DAE

Lanistes el ipticus

LC

N/A

AMPULLARI DAE

Lanistes nasutus

CR

B1ab(i ,i i,iv,v)+2ab(i ,

i i,iv,v)
Yes

AMPULLARI DAE

Lanistes nyassanus

VU

B1ab(i i)

Yes

AMPULLARI DAE

Lanistes ovum

LC

N/A

AMPULLARI DAE

Lanistes solidus

NT

B1ab(i i)

Yes

BITHYNI DAE

Gabbiel a stanleyi

VU

B1ab(i i)

Yes
CYRENIDAE

Corbicula africana

LC

N/A

IRIDINIDAE

Aspatharia subreniformis

LC

N/A

IRIDINIDAE

Chambardia nyassaensis

LC

N/A

IRIDINIDAE

Chambardia petersi

LC

N/A

IRIDINIDAE

Chambardia wahlbergi

LC

N/A
IRIDINIDAE

Mutela alata

LC

N/A

LYMNAEIDAE

Radix natalensis

LC

N/A

PLANORBIDAE

Africanogyrus coretus

LC

N/A

PLANORBIDAE

Biomphalaria angulosa

LC

N/A

PLANORBIDAE

Biomphalaria pfeifferi

LC

N/A
191

Appendix 1. IUCN Red List assessment results. Molluscs, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

PLANORBIDAE

Bulinus forskali

LC

N/A

PLANORBIDAE

Bulinus globosus

LC

N/A

PLANORBIDAE

Bulinus nyassanus

LC
N/A

Yes

PLANORBIDAE

Bulinus succinoides

EN

B1ab(i i)

Yes

PLANORBIDAE

Bulinus truncatus

LC

N/A

PLANORBIDAE

Gyraulus costulatus

LC

N/A

PLANORBIDAE

Lentorbis junodi

LC

N/A

PLANORBIDAE
Segmentorbis angustus

LC

N/A

PLANORBIDAE

Segmentorbis kanisaensis

LC

N/A

SPHAERI DAE

Eupera ferruginea

LC

N/A

SPHAERI DAE

Pisidium pirothi

LC

N/A

SPHAERI DAE

Pisidium reticulatum

LC

N/A

SPHAERI DAE
Sphaerium bequaerti

DD

N/A

THIARIDAE

Melanoides polymorpha

LC

N/A

Yes

THIARIDAE

Melanoides tuberculata

LC

N/A

UNIONIDAE

Coelatura hypsiprymna

LC

N/A

UNIONIDAE

Coelatura mossambicensis

LC

N/A
UNIONIDAE

Nyassunio nyassaensis

LC

N/A

Yes

VIVIPARIDAE

Bel amya capil ata

LC

N/A

VIVIPARIDAE

Bel amya ecclesi

CR

B2ab(i i)

Yes

VIVIPARIDAE

Bel amya jeffreysi

CR

B2ab(i ,i i)

Yes

VIVIPARIDAE
Bel amya robertsoni

CR

B1ab(i,i i)

Yes

Odonates

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

AESHNIDAE

Anaciaeschna triangulifera

LC

N/A

AESHNIDAE

Anax chloromelas

LC

N/A
AESHNIDAE

Anax ephippiger

LC

N/A

AESHNIDAE

Anax imperator

LC

N/A

AESHNIDAE

Anax speratus

LC

N/A

AESHNIDAE

Anax tristis

LC

N/A

AESHNIDAE

Gynacantha bul ata

LC

N/A
AESHNIDAE

Gynacantha immaculifrons

LC

N/A

AESHNIDAE

Gynacantha manderica

LC

N/A

AESHNIDAE

Gynacantha vesiculata

LC

N/A

AESHNIDAE

Gynacantha vil osa

LC

N/A

AESHNIDAE

Heliaeschna trinervulata

LC

N/A
AESHNIDAE

Pinheyschna rileyi

LC

N/A

AESHNIDAE

Zosteraeschna el ioti

LC

N/A

CALOPTERYGIDAE

Phaon iridipennis

LC

N/A

CHLOROCYPHIDAE

Platycypha caligata

LC

N/A

COENAGRIONIDAE

Aciagrion africanum

LC

N/A
COENAGRIONIDAE

Aciagrion steeleae

LC

N/A

COENAGRIONIDAE

Africal agma elongatum

LC

N/A

COENAGRIONIDAE

Africal agma glaucum

LC

N/A

192

Appendix 1. IUCN Red List assessment results. Odonates, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

COENAGRIONIDAE

Africal agma sinuatum

LC

N/A

COENAGRIONIDAE

Africal agma subtile

LC

N/A

COENAGRIONIDAE

Agriocnemis exilis

LC

N/A

COENAGRIONIDAE

Agriocnemis gratiosa

LC

N/A

COENAGRIONIDAE

Agriocnemis victoria

LC
N/A

COENAGRIONIDAE

Azuragrion nigridorsum

LC

N/A

COENAGRIONIDAE

Ceriagrion coral inum

LC

N/A

COENAGRIONIDAE

Ceriagrion glabrum

LC

N/A

COENAGRIONIDAE

Ceriagrion kordofanicum

LC

N/A

COENAGRIONIDAE

Ceriagrion sakeji

LC
N/A

COENAGRIONIDAE

Ceriagrion suave

LC

N/A

COENAGRIONIDAE

Ischnura senegalensis

LC

N/A

COENAGRIONIDAE

Proischnura subfurcata

LC

N/A

COENAGRIONIDAE

Pseudagrion acaciae

LC

N/A

COENAGRIONIDAE

Pseudagrion assegai

LC
N/A

COENAGRIONIDAE

Pseudagrion coelestis

LC

N/A

COENAGRIONIDAE

Pseudagrion commoniae

LC

N/A

COENAGRIONIDAE

Pseudagrion fisheri

LC

N/A

COENAGRIONIDAE

Pseudagrion gamblesi

LC

N/A

COENAGRIONIDAE

Pseudagrion glaucescens

LC
N/A

COENAGRIONIDAE

Pseudagrion hageni

LC

N/A

COENAGRIONIDAE

Pseudagrion hamoni

LC

N/A

COENAGRIONIDAE

Pseudagrion helenae

LC

N/A

COENAGRIONIDAE

Pseudagrion inconspicuum

LC

N/A

COENAGRIONIDAE

Pseudagrion kersteni

LC
N/A

COENAGRIONIDAE

Pseudagrion makabusiense

LC

N/A

COENAGRIONIDAE

Pseudagrion massaicum

LC

N/A

COENAGRIONIDAE

Pseudagrion nubicum

LC

N/A

COENAGRIONIDAE

Pseudagrion salisburyense

LC

N/A

COENAGRIONIDAE

Pseudagrion sjoestedti

LC
N/A

COENAGRIONIDAE

Pseudagrion spernatum

LC

N/A

COENAGRIONIDAE

Pseudagrion sublacteum

LC

N/A

COENAGRIONIDAE

Pseudagrion sudanicum

LC

N/A

CORDULI DAE

Hemicordulia africana

LC

N/A

GOMPHIDAE

Crenigomphus hartmanni

LC
N/A

GOMPHIDAE

Gomphidia quarrei

LC

N/A

GOMPHIDAE

Ictinogomphus ferox

LC

N/A

GOMPHIDAE

Lestinogomphus angustus

LC

N/A

GOMPHIDAE

Microgomphus nyassicus

LC

N/A

GOMPHIDAE

Nepogomphoides stuhlmanni

VU
B1ab(i )+2ab(i )

GOMPHIDAE

Notogomphus dendrohyrax

LC

N/A

GOMPHIDAE

Notogomphus praetorius

LC

N/A

GOMPHIDAE

Paragomphus cognatus

LC

N/A

GOMPHIDAE

Paragomphus elpidius

LC

N/A

GOMPHIDAE

Paragomphus genei

LC
N/A

193

Appendix 1. IUCN Red List assessment results. Odonates, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

GOMPHIDAE

Paragomphus magnus

LC

N/A

GOMPHIDAE

Paragomphus nyasicus

LC

N/A

GOMPHIDAE

Paragomphus sabicus
LC

N/A

GOMPHIDAE

Phyl ogomphus selysi

LC

N/A

LESTIDAE

Lestes amicus

LC

N/A

LESTIDAE

Lestes dissimulans

LC

N/A

LESTIDAE

Lestes ictericus

LC

N/A

LESTIDAE

Lestes pal idus

LC

N/A

LESTIDAE

Lestes plagiatus

LC

N/A

LESTIDAE

Lestes tridens

LC

N/A

LESTIDAE

Lestes uncifer

LC

N/A

LESTIDAE

Lestes virgatus

LC

N/A

LIBELLULIDAE

Acisoma inflatum
LC

N/A

LIBELLULIDAE

Acisoma variegatum

LC

N/A

LIBELLULIDAE

Aethiothemis solitaria

LC

N/A

LIBELLULIDAE

Aethriamanta rezia

LC

N/A

LIBELLULIDAE

Atoconeura biordinata

LC

N/A

LIBELLULIDAE

Brachythemis lacustris
LC

N/A

LIBELLULIDAE

Brachythemis leucosticta

LC

N/A

LIBELLULIDAE

Bradinopyga cornuta

LC

N/A

LIBELLULIDAE

Chalcostephia flavifrons

LC

N/A

LIBELLULIDAE

Crocothemis brevistigma

LC

N/A

LIBELLULIDAE

Crocothemis divisa
LC

N/A

LIBELLULIDAE

Crocothemis erythraea

LC

N/A

LIBELLULIDAE

Crocothemis sanguinolenta

LC

N/A

LIBELLULIDAE

Crocothemis saxicolor

LC

N/A

LIBELLULIDAE

Diplacodes lefebvri

LC

N/A

LIBELLULIDAE

Diplacodes luminans
LC

N/A

LIBELLULIDAE

Diplacodes pumila

LC

N/A

LIBELLULIDAE

Hadrothemis scabrifrons

LC

N/A

LIBELLULIDAE

Hemistigma albipunctum

LC

N/A

LIBELLULIDAE

Nesciothemis farinosa

LC

N/A

LIBELLULIDAE

Notiothemis jonesi
LC

N/A

LIBELLULIDAE

Olpogastra lugubris

LC

N/A

LIBELLULIDAE

Orthetrum abbotti

LC

N/A

LIBELLULIDAE

Orthetrum brachiale

LC

N/A

LIBELLULIDAE

Orthetrum caffrum

LC

N/A

LIBELLULIDAE

Orthetrum chrysostigma
LC

N/A

LIBELLULIDAE

Orthetrum guineense

LC

N/A

LIBELLULIDAE

Orthetrum hintzi

LC

N/A

LIBELLULIDAE

Orthetrum icteromelas

LC

N/A

LIBELLULIDAE

Orthetrum julia

LC

N/A

LIBELLULIDAE

Orthetrum machadoi
LC

N/A

LIBELLULIDAE

Orthetrum macrostigma

LC

N/A

LIBELLULIDAE

Orthetrum stemmale

LC

N/A

194

Appendix 1. IUCN Red List assessment results. Odonates, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

LIBELLULIDAE
Orthetrum trinacria

LC

N/A

LIBELLULIDAE

Palpopleura deceptor

LC

N/A

LIBELLULIDAE

Palpopleura jucunda

LC

N/A

LIBELLULIDAE

Palpopleura lucia

LC

N/A

LIBELLULIDAE

Palpopleura portia

LC

N/A

LIBELLULIDAE
Pantala flavescens

LC

N/A

LIBELLULIDAE

Porpax risi

LC

N/A

LIBELLULIDAE

Rhyothemis fenestrina

LC

N/A

LIBELLULIDAE

Rhyothemis semihyalina

LC

N/A

LIBELLULIDAE

Tetrathemis pol eni

LC

N/A

LIBELLULIDAE
Thermochoria jeanneli

LC

N/A

LIBELLULIDAE

Tholymis til arga

LC

N/A

LIBELLULIDAE

Tramea basilaris

LC

N/A

LIBELLULIDAE

Tramea limbata

LC

N/A

LIBELLULIDAE

Trithemis aconita

LC

N/A

LIBELLULIDAE
Trithemis annulata

LC

N/A

LIBELLULIDAE

Trithemis arteriosa

LC

N/A

LIBELLULIDAE

Trithemis bifida

LC

N/A

LIBELLULIDAE

Trithemis donaldsoni

LC

N/A

LIBELLULIDAE

Trithemis dorsalis

LC

N/A

LIBELLULIDAE
Trithemis furva

LC

N/A

LIBELLULIDAE

Trithemis hecate

LC

N/A

LIBELLULIDAE

Trithemis kirbyi

LC

N/A

LIBELLULIDAE

Trithemis pluvialis

LC

N/A

LIBELLULIDAE

Trithemis stictica

LC

N/A

LIBELLULIDAE
Trithemis werneri

LC

N/A

LIBELLULIDAE

Trithetrum navasi

LC

N/A

LIBELLULIDAE

Urothemis assignata

LC

N/A

LIBELLULIDAE

Urothemis edwardsii

LC

N/A

LIBELLULIDAE

Zygonoides fuel eborni

LC

N/A

LIBELLULIDAE
Zygonyx natalensis

LC

N/A

LIBELLULIDAE

Zygonyx torridus

LC

N/A

MACROMI DAE

Phyl omacromia africana

LC

N/A

MACROMI DAE

Phyl omacromia congolica

LC

N/A

MACROMI DAE

Phyl omacromia contumax

LC

N/A

MACROMI DAE
Phyl omacromia kimminsi

LC

N/A

MACROMI DAE

Phyl omacromia monoceros

LC

N/A

MACROMI DAE

Phyl omacromia picta

LC

N/A

PLATYCNEMIDIDAE

Al ocnemis abbotti

NT

N/A

PLATYCNEMIDIDAE

Al ocnemis maccleeryi

CR

B1ab(i i)+2ab(i i)

Yes
PLATYCNEMIDIDAE

Al ocnemis marshal i

LC

N/A

PLATYCNEMIDIDAE

Al ocnemis montana

EN

B1ab(i i)+2ab(i i)

PLATYCNEMIDIDAE

Elattoneura cel ularis

LC

N/A

PLATYCNEMIDIDAE

Elattoneura glauca

LC

N/A

PLATYCNEMIDIDAE

Mesocnemis singularis

LC

N/A
195

Appendix 1. IUCN Red List assessment results.

Plants

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

ACANTHACEAE

Hygrophila abyssinica

LC

N/A

ACANTHACEAE

Hygrophila auriculata

LC

N/A

ACANTHACEAE

Hygrophila pobeguini
LC

N/A

ALISMATACEAE

Burnatia enneandra

LC

N/A

ALISMATACEAE

Caldesia parnassifolia

LC

N/A

AMARANTHACEAE

Alternanthera sessilis

LC

N/A

AMARANTHACEAE

Centrostachys aquatica

LC

N/A

AMARANTHACEAE

Pandiaka carsoni
LC

N/A

APIACEAE

Afroligusticum linderi

LC

N/A

APIACEAE

Afrosciadium nyassicum

DD

N/A

APIACEAE

Centel a asiatica

LC

N/A

APIACEAE

Hydrocotyle manni

LC

N/A

APONOGETONACEAE

Aponogeton abyssinicus
LC

N/A

ARACEAE

Culcasia falcifolia

LC

N/A

ARACEAE

Lemna aequinoctialis

LC

N/A

ARACEAE

Lemna minor

LC

N/A

ARACEAE

Pistia stratiotes

LC

N/A

ARACEAE

Spirodela polyrhiza
LC

N/A

ARACEAE

Wolffia arrhiza

LC

N/A

ARACEAE

Wolffiel a welwitschi

LC

N/A

ASPLENIACEAE

Asplenium boltoni

LC

N/A

ASPLENIACEAE

Asplenium gemmiferum

LC

N/A

ASTERACEAE

Adenostemma caffrum
LC

N/A

ASTERACEAE

Crassocephalum uvens

DD

N/A

ASTERACEAE

Ethulia conyzoides

LC

N/A

ASTERACEAE

Helichrysum tithonioides

DD

N/A

Yes

ASTERACEAE

Senecio peltophorus

LC

N/A

ASTERACEAE
Sphaeranthus africanus

LC

N/A

ASTERACEAE

Vernonia tolypophora

DD

N/A

ATHYRIACEAE

Athyrium newtoni

LC

N/A

BLECHNACEAE

Blechnum attenuatum

LC

N/A

BLECHNACEAE

Blechnum australe

LC

N/A

BLECHNACEAE
Blechnum punctulatum

LC

N/A

CERATOPHYLLACEAE

Ceratophyl um demersum

LC

N/A

COMMELINACEAE

Aneilema aequinoctiale

LC

N/A

COMMELINACEAE

Commelina benghalensis

LC

N/A

COMMELINACEAE

Commelina diffusa

LC

N/A

COMMELINACEAE
Murdannia simplex

LC

N/A

CONVOLVULACEAE

Ipomoea aquatica

LC

N/A

CRASSULACEAE

Crassula hedbergi

LC

N/A

CYATHEACEAE

Cyathea dregei

LC

N/A

CYATHEACEAE

Cyathea manniana

LC

N/A

CYATHEACEAE
Cyathea thomsoni

LC

N/A

CYPERACEAE

Bolboschoenus glaucus

LC

N/A

196

Appendix 1. IUCN Red List assessment results. Plants, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

CYPERACEAE

Carex brassii

EN

B1ab(i i)+2ab(i i)
CYPERACEAE

Carex cognata

LC

N/A

CYPERACEAE

Carex echinochloe

LC

N/A

CYPERACEAE

Carex ludwigi

LC

N/A

CYPERACEAE

Carex lycurus

LC

N/A

CYPERACEAE

Carex petitiana

LC

N/A
CYPERACEAE

Cyperus albiceps

LC

N/A

CYPERACEAE

Cyperus alopecuroides

LC

N/A

CYPERACEAE

Cyperus alternifolius

LC

N/A

CYPERACEAE

Cyperus amabilis

LC

N/A

CYPERACEAE

Cyperus articulatus

LC

N/A
CYPERACEAE

Cyperus ascocapensis

LC

N/A

CYPERACEAE

Cyperus assimilis

LC

N/A

CYPERACEAE

Cyperus atribulbus

LC

N/A

CYPERACEAE

Cyperus aureobrunneus

LC

N/A

CYPERACEAE

Cyperus clavinux

LC

N/A
CYPERACEAE

Cyperus compressus

LC

N/A

CYPERACEAE

Cyperus denudatus

LC

N/A

CYPERACEAE

Cyperus derreilema

LC

N/A

CYPERACEAE

Cyperus dichrostachyus

LC

N/A

CYPERACEAE

Cyperus difformis

LC

N/A
CYPERACEAE

Cyperus diloloensis

LC

N/A

CYPERACEAE

Cyperus distans

LC

N/A

CYPERACEAE

Cyperus dives

LC

N/A

CYPERACEAE

Cyperus elegantulus

LC

N/A

CYPERACEAE

Cyperus erectus

LC

N/A
CYPERACEAE

Cyperus exaltatus

LC

N/A

CYPERACEAE

Cyperus flavescens

LC

N/A

CYPERACEAE

Cyperus haspan

LC

N/A

CYPERACEAE

Cyperus hystricoides

LC

N/A

CYPERACEAE

Cyperus isolepis

LC

N/A
CYPERACEAE

Cyperus kerni

LC

N/A

CYPERACEAE

Cyperus laevigatus

LC

N/A

CYPERACEAE

Cyperus lanceolatus

LC

N/A

CYPERACEAE

Cyperus latifolius

LC

N/A

CYPERACEAE

Cyperus laxespicatus

LC

N/A
CYPERACEAE

Cyperus lipomonostachyus

DD

N/A

CYPERACEAE

Cyperus macranthus

LC

N/A

CYPERACEAE

Cyperus macrostachyos

LC

N/A

CYPERACEAE

Cyperus melas

LC

N/A

CYPERACEAE

Cyperus mundi

LC

N/A
CYPERACEAE

Cyperus muricatus

LC

N/A

CYPERACEAE

Cyperus nigricans

LC

N/A

CYPERACEAE

Cyperus nitidus

LC

N/A

CYPERACEAE

Cyperus papyrus

LC

N/A

197

Appendix 1. IUCN Red List assessment results. Plants, cont’d

Red List

Endemic to
Family

Binomial

Category

Red List Criteria

LMNNC

CYPERACEAE

Cyperus pectinatus

LC

N/A

CYPERACEAE

Cyperus pelophilus

LC

N/A

CYPERACEAE

Cyperus persquarrosus

LC

N/A

CYPERACEAE

Cyperus polystachyos

LC
N/A

CYPERACEAE

Cyperus prieurianus

LC

N/A

CYPERACEAE

Cyperus proteus

LC

N/A

CYPERACEAE

Cyperus pseudokyl ingioides

LC

N/A

CYPERACEAE

Cyperus pulchel us

LC

N/A

CYPERACEAE

Cyperus pumilus

LC
N/A

CYPERACEAE

Cyperus ridleyi

LC

N/A

CYPERACEAE

Cyperus rotundus

LC

N/A

CYPERACEAE

Cyperus sanguinolentus

LC

N/A

CYPERACEAE

Cyperus sesquiflorus

LC

N/A

CYPERACEAE

Cyperus spissiflorus

DD
N/A

CYPERACEAE

Cyperus squarrosus

LC

N/A

CYPERACEAE

Cyperus tenuiculmis

LC

N/A

CYPERACEAE

Cyperus tenuispica

LC

N/A

CYPERACEAE

Eleocharis caduca

LC

N/A

CYPERACEAE

Eleocharis nigrescens

LC
N/A

CYPERACEAE

Fuirena ciliaris

LC

N/A

CYPERACEAE

Fuirena leptostachya

LC

N/A

CYPERACEAE

Fuirena pubescens

LC

N/A

CYPERACEAE

Fuirena stricta

LC

N/A

CYPERACEAE

Fuirena umbel ata

LC
N/A

CYPERACEAE

Fuirena welwitschi

LC

N/A

CYPERACEAE

Isolepis costata

LC

N/A

CYPERACEAE

Isolepis fluitans

LC

N/A

CYPERACEAE

Rhynchospora candida

LC

N/A

CYPERACEAE

Rhynchospora rugosa

LC
N/A

CYPERACEAE

Schoenoplectiel a articulata

LC

N/A

CYPERACEAE

Schoenoplectiel a roylei

LC

N/A

CYPERACEAE

Schoenoplectus confusus

LC

N/A

CYPERACEAE

Schoenoplectus corymbosus

LC

N/A

CYPERACEAE

Scleria catophyl a

LC
N/A

CYPERACEAE

Scleria distans

LC

N/A

CYPERACEAE

Scleria dregeana

LC

N/A

CYPERACEAE

Scleria flexuosa

LC

N/A

CYPERACEAE

Scleria foliosa

LC

N/A

CYPERACEAE

Scleria glabra

LC
N/A

CYPERACEAE

Scleria gracil ima

LC

N/A

CYPERACEAE

Scleria greigi folia

LC

N/A

CYPERACEAE

Scleria lagoensis

LC

N/A

CYPERACEAE

Scleria nyasensis

LC

N/A

CYPERACEAE

Scleria pooides

LC
N/A

CYPERACEAE

Scleria racemosa

LC

N/A

198

Appendix 1. IUCN Red List assessment results. Plants, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

CYPERACEAE

Scleria rehmanni

LC

N/A

CYPERACEAE

Scleria richardsiae
EN

B2ab(i i)

CYPERACEAE

Scleria suaveolens

LC

N/A

DIDYMOCHLAENACEAE

Didymochlaena truncatula

LC

N/A

DROSERACEAE

Drosera affinis

LC

N/A

DROSERACEAE

Drosera burkeana

LC

N/A

DROSERACEAE

Drosera dielsiana
LC

N/A

DROSERACEAE

Drosera indica

LC

N/A

DROSERACEAE

Drosera madagascariensis

LC

N/A

DRYOPTERIDACEAE

Ctenitis cirrhosa

LC

N/A

DRYOPTERIDACEAE

Ctenitis lanuginosa

LC

N/A

DRYOPTERIDACEAE

Elaphoglossum acrostichoides
LC

N/A

DRYOPTERIDACEAE

Elaphoglossum chevalieri

LC

N/A

DRYOPTERIDACEAE

Elaphoglossum hybridum

LC

N/A

DRYOPTERIDACEAE

Elaphoglossum spathulatum

LC

N/A

DRYOPTERIDACEAE

Polystichum transvaalense

LC

N/A

DRYOPTERIDACEAE

Polystichum zambesiacum
LC

N/A

EQUISETACEAE

Equisetum ramosissimum

LC

N/A

ERIOCAULACEAE

Eriocaulon teusczi

LC

N/A

ERIOCAULACEAE

Eriocaulon zambesiense

DD

N/A

EUPHORBIACEAE

Caperonia stuhlmanni

LC

N/A

EUPHORBIACEAE

Cephalocroton mol is
LC

N/A

FABACEAE

Aeschynomene afraspera

LC

N/A

FABACEAE

Aeschynomene elaphroxylon

LC

N/A

FABACEAE

Aeschynomene indica

LC

N/A

FABACEAE

Aeschynomene pfundi

LC

N/A

FABACEAE

Kotschya africana
LC

N/A

FABACEAE

Neptunia oleracea

LC

N/A

GERANIACEAE

Geranium vagans

LC

N/A

GLEICHENIACEAE

Dicranopteris linearis

LC

N/A

GUNNERACEAE

Gunnera perpensa

LC

N/A

HALORAGACEAE

Myriophyl um spicatum
LC

N/A

HYDROCHARITACEAE

Lagarosiphon cordofanus

LC

N/A

HYDROCHARITACEAE

Lagarosiphon muscoides

LC

N/A

HYDROCHARITACEAE

Najas horrida

LC

N/A

HYDROCHARITACEAE

Najas marina

LC

N/A

HYDROCHARITACEAE

Ottelia exserta
LC

N/A

HYDROCHARITACEAE

Ottelia fischeri

LC

N/A

HYDROCHARITACEAE

Ottelia ulvifolia

LC

N/A

HYDROCHARITACEAE

Val isneria spiralis

LC

N/A

HYDROSTACHYACEAE

Hydrostachys insignis

DD

N/A

HYDROSTACHYACEAE

Hydrostachys polymorpha
LC

N/A

IRIDACEAE

Gladiolus bel us

DD

N/A

ISOETACEAE

Isoetes schweinfurthi

LC

N/A

JUNCACEAE

Juncus oxycarpus

LC

N/A

199

Appendix 1. IUCN Red List assessment results. Plants, cont’d

Red List

Endemic to

Family

Binomial
Category

Red List Criteria

LMNNC

LENTIBULARIACEAE

Genlisea hispidula

LC

N/A

LENTIBULARIACEAE

Utricularia gibba

LC

N/A

LENTIBULARIACEAE

Utricularia reflexa

LC

N/A

LINDERNIACEAE

Lindernia parviflora

LC

N/A

LINDERNIACEAE
Lindernia rotundifolia

LC

N/A

LYTHRACEAE

Ammannia baccifera

LC

N/A

LYTHRACEAE

Nesaea ondongana

LC

N/A

LYTHRACEAE

Nesaea radicans

LC

N/A

LYTHRACEAE

Trapa natans

LC

N/A

MARSILEACEAE
Marsilea minuta

LC

N/A

MENYANTHACEAE

Nymphoides brevipedicel ata

LC

N/A

MENYANTHACEAE

Nymphoides indica

LC

N/A

NEPHROLEPIDACEAE

Nephrolepis undulata

LC

N/A

NYMPHAEACEAE

Nymphaea lotus

LC

N/A

NYMPHAEACEAE
Nymphaea nouchali

LC

N/A

ONAGRACEAE

Ludwigia abyssinica

LC

N/A

ONAGRACEAE

Ludwigia leptocarpa

LC

N/A

ONAGRACEAE

Ludwigia octovalvis

LC

N/A

ONAGRACEAE

Ludwigia stolonifera

LC

N/A

ORCHIDACEAE
Cynorkis brevicalcar

DD

N/A

ORCHIDACEAE

Holothrix johnstoni

DD

N/A

ORCHIDACEAE

Satyrium shirense

DD

N/A

PIPERACEAE

Piper capense

LC

N/A

POACEAE

Cenchrus macrourus

LC

N/A

POACEAE
Echinochloa frumentacea

LC

N/A

POACEAE

Leersia hexandra

LC

N/A

POACEAE

Leptochloa fusca

LC

N/A

POACEAE

Panicum nymphoides

DD

N/A

POACEAE

Phragmites mauritianus

LC

N/A

POACEAE
Sacciolepis africana

LC

N/A

POACEAE

Vossia cuspidata

LC

N/A

POLYGONACEAE

Persicaria limbata

LC

N/A

POLYGONACEAE

Persicaria senegalensis

LC

N/A

POLYPODIACEAE

Stenogrammitis oosora

LC

N/A

PONTEDERIACEAE
Heteranthera cal ifolia

LC

N/A

PONTEDERIACEAE

Monochoria africana

LC

N/A

POTAMOGETONACEAE

Potamogeton crispus

LC

N/A

POTAMOGETONACEAE

Potamogeton octandrus

LC

N/A

POTAMOGETONACEAE

Potamogeton pusil us

LC

N/A

POTAMOGETONACEAE
Potamogeton richardi

LC

N/A

POTAMOGETONACEAE

Potamogeton schweinfurthi

LC

N/A

POTAMOGETONACEAE

Stuckenia pectinata

LC

N/A

PTERIDACEAE

Adiantum capil us-veneris

LC

N/A

PTERIDACEAE

Aspidotis schimperi

LC

N/A

PTERIDACEAE
Ceratopteris cornuta

LC

N/A

200

Appendix 1. IUCN Red List assessment results. Plants, cont’d

Red List

Endemic to

Family

Binomial

Category

Red List Criteria

LMNNC

PTERIDACEAE

Ceratopteris thalictroides

LC

N/A

PTERIDACEAE

Cheilanthes leachi

LC

N/A
PTERIDACEAE

Pteris dentata

LC

N/A

ROSACEAE

Alchemil a el enbecki

LC

N/A

RUBIACEAE

Breonadia salicina

LC

N/A

SALVINIACEAE

Azol a nilotica

LC

N/A

SALVINIACEAE

Azol a pinnata

LC

N/A
SALVINIACEAE

Salvinia hastata

LC

N/A

SAPOTACEAE

Synsepalum brevipes

LC

N/A

SAPOTACEAE

Synsepalum passargei

LC

N/A

SELAGINELLACEAE

Selaginel a goudotiana

LC

N/A

SELAGINELLACEAE

Selaginel a mitteni

LC

N/A
THELYPTERIDACEAE

Cyclosorus interruptus

LC

N/A

THELYPTERIDACEAE

Pneumatopteris unita

LC

N/A

THELYPTERIDACEAE

Thelypteris bergiana

LC

N/A

THELYPTERIDACEAE

Thelypteris chaseana

LC

N/A

THELYPTERIDACEAE

Thelypteris confluens

LC

N/A
THELYPTERIDACEAE

Thelypteris friesii

LC

N/A

THELYPTERIDACEAE

Thelypteris oppositiformis

LC

N/A

TYPHACEAE

Typha domingensis

LC

N/A

VERBENACEAE

Phyla nodiflora

LC

N/A

XYRIDACEAE

Xyris atrata

DD

N/A
XYRIDACEAE

Xyris makuensis

LC

N/A

201

Appendix 2. Species considered in the Red List Index (RLI) for which
genuine changes in Red List Category were recorded

Taxonomic

Start

Category at Published or

End

Category at Direction of

group

Binomial

year

start

back-cast

year

end

change

Fishes
Aulonocara guentheri

2009

LC

Published

2018

EN

Deterioration

Fishes

Aulonocara kandeense

2009

VU

Published

2018

CR

Deterioration

Fishes

Aulonocara maylandi

2009

VU

Published
2018

CR

Deterioration

Fishes

Bagrus meridionalis

2009

LC

Published

2018

CR

Deterioration

Fishes

Champsochromis spilorhynchus

2009

LC

Published

2018

EN

Deterioration

Fishes
Chindongo saulosi

2009

VU

Published

2018

CR

Deterioration

Fishes

Copadichromis azureus

2009

LC

Published

2018

NT

Deterioration

Fishes

Corematodus shiranus

2009

LC

Published
2018

CR(PE)

Deterioration

Fishes

Mchenga conophoros

2009

VU

Published

2018

CR

Deterioration

Fishes

Melanochromis chipokae

2009

VU

Published

2018

CR

Deterioration

Fishes
Melanochromis lepidiadaptes

2009

VU

Published

2018

CR

Deterioration

Fishes

Nimbochromis fuscotaeniatus

2009

LC

Published

2018

VU

Deterioration

Fishes

Oreochromis karongae

2009

EN

Published
2018

CR

Deterioration

Fishes

Oreochromis squamipinnis

2009

EN

Published

2018

CR

Deterioration

Fishes

Pseudotropheus brevis

2009

LC

Published

2018

EN

Deterioration

Fishes
Pseudotropheus cyaneorhabdos

2009

VU

Published

2018

CR

Deterioration

Fishes

Rhamphochromis esox

2009

LC

Back-cast

2018

VU

Deterioration

Fishes

Rhamphochromis longiceps

2009

NT

Back-cast
2018

VU

Deterioration

Fishes

Serranochromis robustus

2009

LC

Published

2018

CR

Deterioration

Fishes

Trematocranus microstoma

2009

LC

Published

2018

EN

Deterioration

Fishes
Tropheops biriwira

2009

LC

Back-cast

2018

NT

Deterioration

Fishes

Tropheops kumwera

2009

LC

Back-cast

2018

NT

Deterioration

Mol uscs

Bel amya robertsoni

2009

EN

Back-cast
2018

CR

Deterioration

202

Appendix 3. Key Biodiversity Area freshwater trigger species

The table below outlines each of the KBAs and their freshwater KBA
trigger species. For each KBA, the table indicates whether this is new or
adopted (i.e. follows the boundary of an existing KBA). Additionally, the
table indicates the validated trigger species for each KBA with details
on: scientific name, taxonomic group, Red List Category, Red List
Criteria, and KBA Criteria met:

A1: Threatened species

(a) Site regularly holds ≥0.5% of the global population AND ≥5 functional
reproductive units of a globally Critically Endangered (CR) or Endangered
(EN) taxon

(b) Site regularly holds ≥1% of the global population AND ≥10 functional
reproductive units of a globally Vulnerable (VU) taxon (c) Site regularly
holds ≥0.1% of the global population AND ≥5 functional reproductive units
of a global y Critical y Endangered (CR) or Endangered (EN) taxon listed as
such based only on a population size reduction in the past or present (d) Site
regularly holds ≥0.2% of the global population AND ≥10 functional
reproductive units of a globally Vulnerable (VU) taxon listed as such based
only on a population size reduction in the past or present

(e) Site effectively holds the entire global population of a CR or EN taxon

B1: Individually geographically restricted species

Site regularly holds ≥10% of the global population size AND ≥10
reproductive units of a species
D1: Demographic aggregations

(a) Site predictably holds an aggregation representing ≥1% of the global

population size of a species, over a season, and during one or more key
stages of its life cycle

D2: Ecological refugia

Site supports ≥10% of the global population size of one or more species
during periods of environmental stress, for which historical evidence shows
that it has served as a refugium in the past and for which there is evidence to
suggest it would continue to do so in the foreseeable future Freshwater

KBA Criteria

Taxonomic KBA trigger

Red List

Red List

KBA Name

Type

group

species

Category

Criteria

A1a A1b A1c A1d A1e

B1

D1a D2

Fishes