Research Article |
Corresponding author: Eva Willén ( eva.willen@slu.se ) Academic editor: Eva Decru
© 2022 Adèle Mputu, Eva Willén, Anders Wilander, Stina Drakare, Kevin Bishop, Bila-Isia Inogwabini.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Mputu A, Willén E, Wilander A, Drakare S, Bishop K, Inogwabini B-I (2022) Fish populations and biomass in headwater streams of the Lake Tumba Landscape, DR Congo, 2007–2011. Acta Ichthyologica et Piscatoria 52(3): 195-208. https://doi.org/10.3897/aiep.52.77638
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The fish biodiversity in the Congo River and its tributaries is extremely rich but the information on fish communities in the headwaters in terms of catch and biomass estimates is rare. Fishes in the running and stagnant waters in this region are of vital importance as a food resource for local residents. This study aimed to describe the fish community, catch, and biomass in the three headwater streams Bambou, Lebomo, and Bongo in the Lake Tumba Landscape (LTL) of the Democratic Republic of the Congo. Such information is of vital importance as a benchmark to understand the sustainability of the fish population for future generations of residents of the LTL. The field data were collected from 2007 through 2011, including dry and wet seasons. Here we present the results of this systematic, multi-annual study which was the first for fishes in streams of this region. In total, 50 species of 15 families were found in the nutrient-poor brown waters of these streams where high concentrations of humic acids cause a low pH. Among abundant species occurring in all three streams were the cyprinid Enteromius holotaenia (Boulenger, 1904), the mormyrid Marcusenius moorii (Günther, 1867), the alestids Clupeocharax schoutedeni Pellegrin, 1926 and Bryconaethiops boulengeri Pellegrin, 1900, and the clariid Clarias angolensis Steindachner, 1866. Bongo Stream was distinguished from the others by a rich abundance of Alestopetersius compressus (Poll et Gosse, 1963). The presence of several species at low pH (between 5.0 and 5.5) is new information that lowers the bottom of the pH interval for these species compared to earlier reports. The maximum total length (TL) of some other species was by 5–20 percentage points higher than those reported earlier. The median weight per unit effort (WPUE) in the streams varied between 30 and 115 g per hour during the dry seasons and between 18 and 86 g per hour during the wet seasons. The fish biomass in the streams varied between 0.05 and 0.7 g ⸱ m–2 with a median 0.14 g ⸱ m–2. This relatively low value compared to other tropical headwaters may be a result of the low pH and dark color of these headwaters. The results of the study serve as a reference point to which future monitoring of fish fauna can be compared for sustainable management of the LTL.
acidity, catch, Congo River, fish biomass, fish species, headwaters, humic water, multi-annual data
The Congo River and its tributaries, covering an area of about 4 000 000 km2, is the world’s second-largest drainage basin after the Amazon River. It’s ichthyofauna is extremely rich—to a large extent incompletely known (
Fishes in the running and stagnant waters are of vital importance as a food resource for the local residents of the LTL; 85% of the protein in their diet derives from fishes (Carpe n.d.). There are indications that the sustainability of the fish population in the landscape is compromised by changes in land use, and in some areas also by overfishing (
Some important fish studies have been performed on other parts of the Congo River basin. For example, the fish fauna of the largest tributary, the Lualaba River, and its associated waters has been documented by several expeditions (e.g.,
The presently reported study was undertaken on three LTL headwater streams, Bambou, Lebomo, and Bongo, to improve knowledge of community structure and diversity of the LTL fish fauna. These streams were investigated systematically over a period of 3–5 years, with a special focus on the catch and biomass data. We describe the stream fish community composition and biomass with the aim to compare variation within and between streams over years and seasons. The relations to water quality are explored, and comparisons are made with previous studies from the Congo River basin. The information presented in this paper will increase our knowledge on the fish biomass in central African waters that are relatively poor in nutrients, low in pH, and high in dissolved organic matter.
Characteristics of investigated streams. The investigated streams Bambou, Lebomo, and Bongo are situated in the Mai-Ndombe province in the central part of the Democratic Republic of the Congo (Fig.
The stream catchments were estimated from satellite images and their area ranged from 90 to 180 km2 (Table
Basic physical and chemical data for the streams investigated in the headwater streams of the Lake Tumba Landscape, DR Congo. Drainage area data based on satellite images. Median width and depth with ranges in brackets. Chemical values presented as medians with ranges in brackets.
Character | Stream | ||
---|---|---|---|
Bambou | Lebomo | Bongo | |
Landscape type | Forest, savannah | Grassland, savannah | Mature forest |
Land use | Natural | Ranching | Logging |
Coordinates | 2.29ºS, 16.30ºE | 2.57ºS, 16.55ºE | 1.95ºS, 17.10ºE |
Bottom substrate | Sand, pebble | Sand, stone | Mud, boulder |
Elevation [m] | 440 | 380 | 340 |
Drainage area [km2] | 90 | 100 | 180 |
Number of measurements | 8 | 8 | 6 |
Width [m] | 7.6 (3.7–12.5) | 7.7 (6.3–9.9) | 8.5 (4.6–17.0) |
Depth [m] | 0.4 (0.17–1.4) | 0.7 (0.38–0.97) | 0.6 (0.18–1.15) |
Number of measurements | 10 | 10 | 4 |
pH | 5.0 (4.8–5.8) | 6.4 (6.0–6.8) | 5.7 (5.5–5.8) |
Color [mg Pt⸱ L–1] | 120 (30–120) | 50 (10–90) | 55 (40–60) |
TOC [mg ⸱ L–1] | 10.6 (4.5–12) | 3.8 (0.8–8.9) | 5.9 (4.5–6.5) |
NO2,3-N [µg ⸱ L–1] | 70 (2–91) | 52 (1–140) | 300 (280–340) |
Total P [µg ⸱ L–1] | 41 (26–54) | 35 (5–63) | 69 (61–79) |
Sampling and analysis. The study period was 2007–2011. In each stream 10 transects set approximately 200 m from each other were delineated. The total length covered was 1800 m. Five transects were placed upstream and five downstream of the existing bridges to facilitate subsequent sampling. In Bongo two transects became located upstream of the 25 m high waterfall and eight downstream. Width and depth were measured every meter by a stick at the ten fishing transects in each stream on eight occasions in Bambou and Lebomo, and six in Bongo. For each transect, a mean of three central depth values was used in order to avoid possible “pits”. Due to the heavy workload during fishing, this was done on other occasions close to fishing days.
An apparent visual observation of water color was recorded during a total of 53 days where the water was classified as white, clear, or brown as established for Amazonian waters by
Samples for water chemistry were collected at the bridge location on ten occasions in Bambou and Lebomo and on four occasions in Bongo. The samples were sent to the accredited water chemistry laboratory at the Department of Aquatic Sciences and Assessment, Swedish University of Agricultural Sciences, Uppsala, Sweden, and analyzed using standardized methods (International standards, ISO and European standards, EN).
Fishing was carried out in a way that allowed separation of catches between day (08:00–16:00 h or 18:00 h) and night (18:00–06:00 h). There was a variation in the times for setting and harvesting of the fishing gear which was set out up to five times a day at 08:00, 12:00, 14:00, 16:00, and/or 18:00 h. During the daytime, harvesting occurred two or four hours after nets were set. Night-time fishing nominally started at 18:00 h and continued until 06:00 h the following morning. The total number of fishing days was 176 of which 68 occurred during the dry seasons and 106 in wet seasons (Table
Fishing days by stream, season, and gear for the headwater streams of the Lake Tumba Landscape, DR Congo.
Stream | Year | Season | Hook | Net | Net and hook | Total fishing days |
---|---|---|---|---|---|---|
Bambou | 2007 | Wet | — | 12 | — | 12 |
Bambou | 2008 | Dry | — | 5 | — | 5 |
Bambou | 2008 | Wet | 4 | 14 | — | 18 |
Bambou | 2009 | Dry | — | 2 | — | 2 |
Bambou | 2010 | Dry | — | 7 | 3 | 10 |
Bambou | 2010 | Wet | 4 | 6 | — | 10 |
Bambou | 2011 | Wet | 3 | 7 | — | 10 |
Bambou | Total | 11 | 53 | 3 | 67 | |
Bongo | 2008 | Dry | 1 | 7 | — | 8 |
Bongo | 2008 | Wet | — | 7 | 1 | 8 |
Bongo | 2009 | Dry | 1 | 6 | — | 7 |
Bongo | 2009 | Wet | — | 6 | — | 6 |
Bongo | 2010 | Dry | — | 2 | 4 | 6 |
Bongo | 2010 | Wet | — | 3 | 5 | 8 |
Bongo | Total | 2 | 31 | 10 | 43 | |
Lebomo | 2008 | Dry | — | 3 | 4 | 7 |
Lebomo | 2008 | Wet | — | 7 | — | 7 |
Lebomo | 2009 | Dry | 7 | 5 | 1 | 13 |
Lebomo | 2009 | Wet | — | 5 | — | 5 |
Lebomo | 2010 | Dry | — | 3 | 7 | 10 |
Lebomo | 2010 | Wet | 7 | 7 | 1 | 15 |
Lebomo | 2011 | Wet | — | 4 | 5 | 9 |
Lebomo | Total | 14 | 34 | 18 | 66 | |
Total | 27 | 118 | 31 | 176 |
The number of sampling events varied between years but always included periods of the nominally dry and wet seasons. The streams were not fished during either the times of lowest or highest water levels as illustrated for Bambou Stream (Fig.
Water level recordings from Bambou Stream, the Lake Tumba Landscape, DR Congo. Fishing days indicated as blue markers for the wet season and brown markers for the dry season. Black markers indicate level recording days with no fishing. Line is a cubic smoothing. The green dots and vertical lines indicate the two dates with pH-measurements.
An overview of the Congolese fish fauna has never been compiled and identification keys are lacking for the majority of regions and taxonomic groups (
Weight (catch) was calculated as a daily sum of catches for all fishing gear and species. The Weight Per Unit Effort (WPUE) and number of fish per hour (NPUE) were calculated with respect to actual fishing hours. Different subsets of fishing time and gear were used in order to obtain detailed, comparable subsets as specified for each statistical analysis.
In order to assess the total fish biomass in each stream, fishing was performed repeatedly during 4–12 days with the aim of achieving total fish depletion in the study area. The fishing gear was harvested 3–4 times per day and set up again immediately after each harvest, covering a 24-h period, until no or low catch was recorded, indicating complete or nearly complete removal of fish from the study area for the size of fish caught by the used mesh sizes.
Statistical analyses. Data handling and univariate statistical calculations were done in JMP® 14.0.0. Non-metric multidimensional scaling (NMDS) was used for visual representation and to compare the similarity in community composition for daytime (08:00–18:00 h) net-sampled fish abundances (NPUE). Only daytime catches were included in this analysis as not in all sites both day- and night-time catches were done. Differences between streams and seasons (groups) were tested with an analysis of similarities (ANOSIM,
Stream characteristics. Bambou Stream was the shallowest with a mean maximum depth of 0.5 m while the other streams were about 0.7 m deep on average (range in Table
The streams were of similar width (Table
Bambou and Bongo are tropical blackwaters with high concentrations of humic acids coloring the water deep brown (measured as total organic carbon (TOC) and water color) and causing a low pH of around 5 (Table
Fish occurrence, abundance, size, and frequency. A total of 2028 fish individuals representing 50 species were caught in this study. The majority of specimens were identified up to the species level, however, ten could be identified up to genus only (Table
Summary table for species caught in the headwater streams of the Lake Tumba Landscape, DR Congo. Total number and measured maximum total length (TL) of each species. The FishBase data (FB) on maximum (TL) or standard length (SL) are provided for comparison (
FAMILY | Number of fish sampled | TL max [cm] | FB TL max [cm] | FB SL max [cm] | ||
---|---|---|---|---|---|---|
Species | Bambou | Lebomo | Bongo | |||
PROTOPTERIDAE | ||||||
Protopterus dolloi Boulenger, 1900 | 0 | 0 | 2 | 82 | 130 | |
POLYPTERIDAE | ||||||
Polypterus ornatipinnis Boulenger, 1902 | 5 | 0 | 0 | 40 | 60 | |
Polypterus polli Gosse, 1988 | 1 | 0 | 0 | 22.2 | 32.1 | |
MORMYRIDAE | ||||||
Brienomyrus sp. | 1 | 0 | 9 | 15.5 | ||
Campylomormyrus sp. | 0 | 4 | 2 | 22 | ||
Cyphomyrus psittacus (Boulenger, 1897) | 11 | 0 | 0 | 20.5 | 30 | |
Gnathonemus petersii (Gunther, 1862) | 0 | 5 | 0 | 23.6 | 35 | |
Heteromormyrus sp. | 0 | 0 | 5 | 20.5 | ||
Hippopotamyrus sp. | 16 | 2 | 2 | 17 | ||
Marcusenius dundoensis (Poll, 1967) | 4 | 3 | 0 | 21 | 12.3 | |
Marcusenius moorii (Günther, 1867) | 16 | 6 | 94 | 20 | 21.4 | |
Mormyrops anguilloides (Linnaeus, 1758) | 0 | 1 | 0 | 37 | 150 | |
Mormyrops sp. | 16 | 3 | 25 | 17.1 | ||
Mormyrus caballus Boulenger, 1898 | 3 | 0 | 0 | 18.5 | 50 | |
Myomyrus macrodon Boulenger, 1898 | 0 | 9 | 0 | 24 | 24 | |
Petrocephalus christyi Boulenger, 1920 | 1 | 4 | 3 | 13 | 7.7 | |
Petrocephalus simus Sauvage, 1879 | 0 | 2 | 1 | 12.1 | 12 | |
Pollimyrus adspersus (Günther, 1866) | 0 | 1 | 0 | 8 | 9.8 | |
Pollimyrus sp. | 2 | 1 | 35 | 15.5 | ||
Pollimyrus tumifrons (Boulenger, 1902) | 2 | 14 | 0 | 19 | 11.2 | |
ALESTIDAE | ||||||
Alestopetersius compressus (Poll et Gosse, 1963) | 0 | 0 | 152 | 11.5 | 7.8 | |
Brycinus sp. | 0 | 0 | 1 | 6.5 | ||
Bryconaethiops boulengeri Pellegrin, 1900 | 96 | 52 | 18 | 24 | 25 | |
Clupeocharax schoutedeni Pellegrin, 1926 | 127 | 84 | 3 | 20 | 25 | |
DISTICHODONTIDAE | ||||||
Mesoborus crocodilus Pellegrin, 1900 | 3 | 3 | 0 | 29.5 | 26.5 | |
Phago boulengeri Schilthuis, 1891 | 1 | 0 | 0 | 14 | 17 | |
HEPSETIDAE | ||||||
Hepsetus microlepis (Boulenger, 1901) | 5 | 2 | 12 | 38 | 26.7 | |
CYPRINIDAE | ||||||
Enteromius holotaenia (Boulenger, 1904) | 171 | 25 | 77 | 17.5 | 12 | |
Raiamas christyi (Boulenger, 1920) | 42 | 43 | 0 | 19 | 17.7 | |
CLARIIDAE | ||||||
Channallabes apus (Günther, 1873) | 0 | 2 | 0 | 29 | 41.6 | |
Clarias angolensis Steindachner, 1866 | 22 | 7 | 146 | 40 | 35 | |
Clarias gabonensis Günther, 1867 | 1 | 3 | 0 | 21 | 36 | |
Clarias jaensis Boulenger, 1909 | 1 | 11 | 8 | 29 | 48,3 | |
Clarias platycephalus Boulenger, 1902 | 3 | 3 | 19 | 43 | 37.6 | |
Clarias sp. | 0 | 1 | 0 | 14.5 | ||
Platyclarias machadoi Poll, 1977 | 73 | 12 | 7 | 26.2 | 20.1 | |
MALAPTERURIDAE | ||||||
Malapterurus melanochir Norris, 2002 | 0 | 1 | 2 | 14 | 98 | |
Malapterurus sp. | 0 | 0 | 1 | 11 | ||
CLAROTEIDAE | ||||||
Parauchenoglanis balayi (Sauvage, 1879) | 117 | 5 | 0 | 19.5 | 39 | 31.7 |
Parauchenoglanis pantherinus (Pellegrin, 1929) | 2 | 0 | 0 | 17 | 29.2 | |
Parauchenoglanis punctatus (Boulenger, 1902) | 12 | 58 | 0 | 38 | 41.0 | |
SCHILBEIDAE | ||||||
Schilbe marmoratus Boulenger, 1911 | 0 | 69 | 35 | 21 | 21.7 | |
MOCHOKIDAE | ||||||
Euchilichthys royauxi Boulenger, 1902 | 0 | 1 | 2 | 12.5 | 22 | |
Synodontis sp. | 0 | 0 | 1 | 12.5 | ||
ANABANTIDAE | ||||||
Ctenopoma multispine Peters, 1844 | 14 | 5 | 8 | 15 | 14 | |
Microctenopoma nanum (Günther, 1896) | 0 | 0 | 1 | 9.2 | 6.7 | |
CICHLIDAE | ||||||
Hemichromis elongatus (Guichenot, 1861) | 44 | 14 | 13 | 22 | 18.7 | |
Hemichromis stellifer Loiselle, 1979 | 41 | 5 | 22 | 12.6 | 10 | |
Pelmatochromis nigrofasciatus (Pellegrin, 1900) | 0 | 2 | 0 | 7.3 | 11.6 | |
MASTACEMBELIDAE | ||||||
Mastacembelus congicus Boulenger, 1896 | 1 | 5 | 0 | 36.5 | 43.5 | |
Total number of species | 31 | 36 | 29 |
When considering the number of specimens per family caught in the three streams, the pattern varied slightly. Alestids stand out markedly with 26% of the total number of caught fish specimens belonging to just four species (Table
Major species (caught in numbers ≥20) sorted by abundance for each stream for the headwater streams of the Lake Tumba Landscape, DR Congo. Percentages are compared to the total abundance in each stream.
Bambou | Lebomo | Bongo | |||
---|---|---|---|---|---|
Species | [%] | Species | [%] | Species | [%] |
Enteromius holotaenia | 20 | Clupeocharax schoutedeni | 18 | Alestopetersius compressus | 22 |
Clupeocharax schoutedeni | 15 | Schilbe marmoratus | 15 | Clarias angolensis | 21 |
Parauchenoglanis balayi | 14 | Parauchenoglanis punctatus | 12 | Marcusenius moorii | 13 |
Bryconaethiops boulengeri | 11 | Bryconaethiops boulengeri | 11 | Enteromius holotaenia | 11 |
Platyclarias machadoi | 9 | Raiamas christyi | 9 | Schilbe marmoratus | 5 |
Hemichromis elongatus | 5 | Enteromius holotaenia | 5 | Pollimyrus sp. | 5 |
Raiamas christyi | 5 | Mormyrops sp. | 4 | ||
Hemichromis stellifer | 5 | Hemichromis stellifer | 3 | ||
Clarias angolensis | 3 |
Compared to thirteen other studies of fish fauna found in the Congo River basin (Table
Species list from the presently reported study compared with published records from other streams/rivers in the Congo River drainage area.
FAMILY | Reference study | Total No. | ||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | |
PROTOPTERIDAE | ||||||||||||||
Protopterus dolloi | – | x | x | – | x | x | x | – | – | – | – | – | x | 6 |
POLYPTERIDAE | ||||||||||||||
Polypterus ornatipinnis | – | – | x | – | x | x | – | x | – | – | – | – | – | 4 |
Polypterus polli | – | – | – | – | – | x | – | – | – | – | – | – | – | 1 |
MORMYRIDAE | ||||||||||||||
Cyphomyrus psittacus | – | x | – | x | x | x | – | x | – | – | – | – | x | 6 |
Gnathonemus petersii | L | x | x | – | x | x | – | x | x | – | x | – | x | 9 |
Marcusenius dundoensis | – | – | – | – | – | – | – | – | – | – | – | – | – | 0 |
Marcusenius moorii | B L N | x | x | – | x | x | – | x | – | x | x | x | x | 10 |
Mormyrops anguilloides | – | x | x | x | x | x | – | x | – | – | – | – | x | 7 |
Mormyrus caballus | – | x | x | x | x | x | – | x | – | – | x | – | – | 7 |
Myomyrus macrodon | – | – | – | x | – | – | – | – | – | – | – | – | – | 1 |
Petrocephalus christyi | – | x | – | – | x | x | – | x | x | – | x | – | x | 7 |
Petrocephalus simus | – | – | – | – | – | x | x | – | – | x | – | x | – | 4 |
Pollimyrus adspersus | – | – | – | – | x | x | – | – | – | – | – | – | – | 2 |
Pollimyrus tumifrons | – | – | – | – | – | x | – | – | – | – | – | – | – | 1 |
ALESTIDAE | ||||||||||||||
Alestopetersius compressus | – | – | – | – | – | x | – | – | – | – | – | – | – | 1 |
Bryconaethiops boulengeri | B | x | x | – | x | x | x | x | x | x | x | – | x | 11 |
Clupeocharax schoutedeni | – | x | x | – | – | – | – | – | – | – | – | – | x | 3 |
DISTICHODONTIDAE | ||||||||||||||
Mesoborus crocodilus | – | – | x | – | – | x | – | x | x | – | x | – | x | 6 |
Phago boulengeri | – | x | – | – | x | x | x | x | – | – | x | – | x | 7 |
HEPSETIDAE | ||||||||||||||
Hepsetus microlepis | – | – | x | – | x | x | – | – | x | x | x | – | x | 7 |
CYPRINIDAE | ||||||||||||||
Enteromius holotaenia | – | – | – | – | – | x | – | – | x | x | x(cf) | – | – | 4 |
Raiamas christyi | B(cf) | – | – | – | – | x | x | – | x | x | x | x | x | 8 |
CLARIIDAE | ||||||||||||||
Channallabes apus | B | x | x | x | x | x | – | – | – | – | x | – | x | 8 |
Clarias angolensis | – | x | x | x | x | x | – | – | – | – | x | x | x | 8 |
Clarias gabonensis | – | x | x | x | x | x | – | – | x | x | – | x | x | 9 |
Clarias jaensis | – | – | – | – | – | – | – | – | – | x | – | – | – | 1 |
Clarias platycephalus | B | x | x | – | x | x | – | – | – | – | – | x | – | 5 |
Platyclarias machadoi | – | – | – | – | – | – | – | – | – | – | – | – | – | 0 |
MALAPTERURIDAE | ||||||||||||||
Malapterurus melanochir | – | – | x | – | – | x | – | – | – | – | – | – | x | 3 |
CLAROTEIDAE | ||||||||||||||
Parauchenoglanis balayi | – | – | – | x | x | x | – | – | – | x | – | – | – | 4 |
Parauchenoglanis pantherinus | – | – | – | – | – | – | – | – | – | x | – | – | – | 1 |
Parauchenoglanis punctatus | N | x | x | – | x | x | – | x | – | – | x | x | x | 9 |
SCHILBEIDAE | ||||||||||||||
Schilbe marmoratus | – | x | x | – | x | x | – | x | x | – | x | – | x | 8 |
MOCHOKIDAE | ||||||||||||||
Euchilichthys royauxi | – | – | – | – | – | x | – | – | – | – | – | – | – | 1 |
ANABANTIDAE | ||||||||||||||
Ctenopoma multispine | – | – | – | – | – | – | – | – | – | – | – | – | – | 0 |
Microctenopoma nanum | B L | x | x | – | x | x | x | – | – | x | x | – | x | 9 |
CICHLIDAE | ||||||||||||||
Hemichromis elongatus | B N | – | x | x | x | x | – | x | x | x | x | x | – | 10 |
Hemichromis stellifer | B | – | – | x | x | – | – | – | x | – | x | x | – | 6 |
Pelmatochromis nigrofasciatus | – | – | x | – | x | x | – | – | – | – | x | – | x | 5 |
MASTACEMBELIDAE | ||||||||||||||
Mastacembulus congicus | – | – | – | – | x | x | – | x | – | – | x | – | – | 4 |
Total number | 10 | 17 | 20 | 10 | 24 | 32 | 6 | 14 | 11 | 12 | 19 | 9 | 20 |
As expected, larger species tended to be caught with a hook rather than with a net. There was a difference in total length (TL) between catches with the two gears (Wilcoxon P < 0.0001). The median TL was 15.3 cm for hook fishing and 12.5 cm for nets. The largest species caught only by the hook was Protopterus dolloi in the Bongo stream with a maximum TL of 82 cm. Large species caught by net and reaching lengths of about 40 cm were Polypterus ornatipinnis Boulenger, 1902, Hepsetus microlepis (Boulenger, 1901), and Parauchenoglanis punctatus (Boulenger, 1902). There was a lack of small individuals due to the available fishing gear. The shortest fish caught was 6.3 cm TL.
The maximum TL of three species in this study exceeded the maximum TL recorded in FishBase (
Presently reported species from headwater streams of the Lake Tumba Landscape, DR Congo with lengths considerably larger than maximum total length (TL) recorded in FishBase (FB,
Species | No. of fish caught | Median TL [cm] | Max TL [cm] | FB TL (SL) [cm] | Max TL/FB TL |
---|---|---|---|---|---|
Clarias angolensis | 175 | 19.5 | 40 | 35 | 1.1 |
Enteromius holotaenia | 273 | 12.0 | 17.5 | na (12) | na |
Raiamas christyi | 85 | 15.6 | 23.8 | 17.7 | 1.3 |
Pollimyrus tumifrons | 16 | 16.0 | 19 | 11.2 | 1.7 |
The high waterfall in Bongo most probably impeded some species from moving upstream. Only six species were found upstream from the 25 m high waterfall: Alestopetersius compressus, Clarias angolensis, Clarias platycephalus Boulenger, 1902, Enteromius holotaenia, Marcusenius moorii (Günther, 1867), Mormyrops sp. They were all found downstream of the waterfall as well.
Visual water color and fish occurrence. Enteromius holotaenia, Hemichromis elongatus (Guichenot, 1861), Cyphomyrus psittacus (Boulenger, 1897), Bryconaethiops boulengeri, and Platyclarias machadoi were species caught in the largest numbers during periods with brown water. Species almost totally lacking during brown water phases were Alestopetersius compressus in Bongo, Schilbe marmoratus Boulenger, 1911 and Parauchenoglanis punctatus in Bongo and Lebomo, as well as Marcusenius moorii in all streams. Species not caught during periods of brown water were roughly equally abundant during white and clear water phases.
Influence of pH on fish occurrence. Twenty-seven species in the streams occurred at pH 5, seven species were added when the pH was 5.5, and six species at pH 6 (Table
Fish species list with lowest recorded pH from the presently reported study from headwater streams of the Lake Tumba Landscape, DR Congo confronted with the data on pH range from FishBase (
FAMILY | Lowest pH | pH from FishBase |
---|---|---|
Species | ||
PROTOPTERIDAE | ||
Protopterus dolloi | 5.5 | na |
POLYPTERIDAE | ||
Polypterus ornatipinnis | 5.0 | na |
Polypterus polli | 5.0 | na |
MORMYRIDAE | ||
Cyphomyrus psittacus | 5.0 | 6.8–7.2 |
Gnathonemus petersii | 6.0 | 6.0–8.0 |
Marcusenius dundoensis | 5.0 | na |
Marcusenius moorii | 5.0 | na |
Mormyrops anguilloides | 6.0 | na |
Mormyrus caballus | 5.0 | na |
Myomyrus macrodon | 6.0 | na |
Petrocephalus christyi | 5.0 | na |
Petrocephalus simus | 5.5 | na |
Pollimyrus adspersus | 6.0 | na |
Pollimyrus tumifrons | 5.0 | na |
ALESTIDAE | ||
Alestopetersius compressus | 5.5 | na |
Bryconaethiops boulengeri | 5.0 | na |
Clupeocharax schoutedeni | 5.0 | 6.0–7.5 |
DISTICHODONTIDAE | ||
Mesoborus crocodilus | 5.0 | na |
Phago boulengeri | 5.0 | na |
HEPSETIDAE | ||
Hepsetus microlepis | 5.0 | na |
CYPRINIDAE | ||
Enteromius holotaenia | 5.0 | 6.0–6.5 |
Raiamas christyi | 5.0 | 6.5–7.0 |
CLARIIDAE | ||
Channallabes apus | 6.0 | 6.0–8.0 |
Clarias angolensis | 5.0 | 7.0–9.0 |
Clarias gabonensis | 5.0 | 7.0–9.0 |
Clarias jaensis | 5.0 | na |
Clarias platycephalus | 5.0 | na |
Platyclarias machadoi | 5.0 | na |
MALAPTERURIDAE | ||
Malapterurus melanochir | 5.5 | na |
CLAROTEIDAE | ||
Parauchenoglanis balayi | 5.0 | na |
Parauchenoglanis pantherinus | 5.0 | na |
Parauchenoglanis punctatus | 5.0 | na |
SCHILBEIDAE | ||
Schilbe marmoratus | 5.5 | 6.5–7.5 |
MOCHOKIDAE | ||
Euchilichthys royauxi | 5.5 | na |
ANABANTIDAE | ||
Ctenopoma multispine | 5.0 | 6.0–7.5 |
Microctenopoma nanum | 5.5 | na |
CICHLIDAE | ||
Hemichromis elongatus | 5.0 | na–7.0 |
Hemichromis stellifer | 5.0 | na |
Pelmatochromis nigrofasciatus | 6.0 | na |
MASTACEMBELIDAE | ||
Mastacembelus congicus | 5.0 | na |
Diurnal differences. Comparisons of NPUE and WPUE between day and night catches were inconclusive (Wilcoxon non-parametric test). The majority of species were caught during the day-time and Hemichromis elongatus, Alestopetersius compressus, Mormyrops sp., Marcusenius moorii, and Platyclarias machadoi were always day-time catches. Only one species was solely caught during the night, and this was Myomyrus macrodon Boulenger, 1898 in Lebomo. No other species were regularly caught in significant quantities during night-time in any stream, but all those caught at night belonged to the family Mormyridae.
Fish catches. A compilation of net catches in each stream between years and seasons with day-time fishing effort is given in Table
Compilation of comparable catch statistics of the presently reported study from headwater streams of the Lake Tumba Landscape, DR Congo during daytime net-fishing period (08.00–16.00/18.00) with all 10 nets (transects).
Stream | No. of fishing days | NPUE [no. ⸱ h–1] | WPUE [g ⸱ h–1] | |||
---|---|---|---|---|---|---|
Dry | Wet | Dry | Wet | Dry | Wet | |
Bambou | 12 | 18 | 1.4 | 2.6 | 30 | 86 |
Lebomo | 26 | 25 | 0.9 | 0.5 | 38 | 18 |
Bongo | 19 | 17 | 2.8 | 1.0 | 115 | 69 |
Total | 57 | 60 |
ANOSIM analysis of community composition based on NPUE data from daytime net fishing clustered the streams separately from each other (ANOSIM P = 0.001, Global R = 0.583, Fig.
Estimation of fish biomass. The median fish biomass for all streams and occasions combined was 0.14 g ⋅ m–2. The biomass varied from 0.05 to 0.21 g ⋅ m–2 in Bambou, from 0.14 to 0.70 g ⋅ m–2 in Bongo, and from 0.05 to 0.43 g ⋅ m–2 in Lebomo (Table
Estimation of fish biomass of the presently reported study from three headwater streams of the Lake Tumba Landscape, DR Congo.
Stream | Year | Season | Gear | Fishing days | Fishes caught | Stream width [m] | Biomass [g ⸱m–2] |
---|---|---|---|---|---|---|---|
Bambou | 2010 | Dry | n | 4 | 40 | 7.23 | 0.115 |
Bambou | 2010 | Wet | n | 5 | 44 | 8.03 | 0.212 |
Bambou | 2011 | Wet | n | 4 | 22 | 8.79 | 0.045 |
Bongo | 2010 | Dry | nh | 5 | 166 | 7.22 | 0.707 |
Bongo | 2010 | Wet | nh | 8 | 33 | 9.37 | 0.135 |
Lebomo | 2008 | Dry | n | 7 | 105 | 8.02 | 0.253 |
Lebomo | 2009 | Dry | nh | 5 | 55 | 7.75 | 0.137 |
Lebomo | 2009 | Wet | n | 10 | 22 | 7.56 | 0.066 |
Lebomo | 2010 | Dry | nh | 10 | 31 | 8.01 | 0.083 |
Lebomo | 2010 | Wet | n | 5 | 21 | 8.18 | 0.054 |
Lebomo | 2011 | Wet | nh | 9 | 90 | 7.79 | 0.425 |
Median | 40 | 0.135 |
Species distributions in the streams. During the presently reported study, we identified 31, 36, and 29 species in Bambou, Lebomo, and Bongo, respectively. Thanks to intense and prolonged fishing during both day and night these values are likely to be close to the total number of species present that could be caught with the net meshes and hook sizes used (Fig.
Many of the fish species of our study have been found in similar streams in the region (Table
Platyclarias machadoi, present in all of our study streams, but in particularly large numbers in Bambou, needs a special comment as it is not recorded in the studies compiled in Table
A species caught in all three streams although in a smaller number as well as in several other rivers of lower and middle Congo is Hepsetus microlepis. In the investigations of the Salonga area and N’sele region it is named Hepsetus odoe as it precedes the revision of the genus and the revalidation of Hepsetus microlepis (see
Fish size. Headwater streams are usually environmentally impoverished in relation to downstream watercourses as suggested by the river continuum concept (
Some of the common species, however, in the presently reported study, reached larger sizes than recorded in FishBase (
Tolerance of acidity. The majority of fish thrive at a narrow range of a near neutral pH (
Fish biomass aspects. Three earlier publications from Africa with areal fish biomass data were found (
One cause of lower biomass in our study is the lack of small specimens in the catch. Da Silva Gonçalves and de Souza Braga (2012) published a table relating the weight of caught species and SL. Based on this study we calculate that we may have missed 70% of fish biomass. That would raise the median biomass from 0.14 to 0.20 g ⸱ m–2. But this only explains a minor part of the difference. An important feature of all the South American streams, in addition to a different fish fauna, was a circumneutral pH and generally less colored water. Most likely the low pH values of our study streams affect the fish especially when it comes to species presence and physiological responses to a low pH. A secondary effect may be the brown-colored, nutrient-poor water (cf. Table
Only one previous study of the fish biomass for African headwaters has been found. Malaisse (1976) reported values of 1.3, 26.1, and 31.7 kg ⸱ km–1 (i.e., not per area) in the reaches of a tributary to the Kwa River (
Overfishing risk. Overfishing could be a major threat to the sustainability of continued fishing of the fish populations (
This study presents a comprehensive investigation of the fish community composition for several years in three headwater streams in the Congo basin drainage area. Alestids, cyprinids, clariids, and mormyrids were the most common fish families and, in total, 50 species representing 15 families were found. The reason for the low richness compared to other tributaries is most probably because they are small headwater streams. Ten species were found in lower pH waters than those reported in FishBase (
The Central Africa Regional Program for the Environment (CARPE) of the US Agency of International Development financially supported the fieldwork via WWF DRC. However, views expressed in this paper reflect neither those of the US Government nor those of WWF. The work was possible thanks to field support by Batsabaru, Basime, Leasau, Bokuango, and Nzabi Nono. Special thanks are forwarded to taxonomists at the Royal Museum for Central Africa in Tervuren, Belgium and at the Laboratory of Biodiversity and Evolutionary Genomics in KU, Leuven, Belgium for support in species determinations based on photo material. We also thank Asad Jamil, who prepared the satellite images for area determination by Stefan Hellgren and Andreas Rudh. Mikael Östlund assisted in the preparations of the graphs. Christian Demandt was responsible for the physical-chemical determinations, all done at the Department of Aquatic Sciences and Assessment, SLU, Sweden. We also appreciate the valuable comments from Eva Decru and three reviewers.