European Journal of Ecology, 6.2, 2020, pp. 1-17
HERPETOFAUNAL DIVERSITY AND COMMUNITY STRUCTURE IN THE
MURCHISON FALLS-ALBERT DELTA RAMSAR SITE, UGANDA
Mathias Behangana1, Richard Magala2,10, Raymond Katumba3, David Ochanda4, Stephen
Kigoolo5, Samuel Mutebi6, Daniele Dendi7,8,9, Luca Luiselli7,8,9 and Daniel F. Hughes11
1
Department of Environmental Sciences, Makerere University, P.O. Box 7062, Kampala, Uganda;
E-mail: mbehangana@gmail.com
2
Department of Geography and Environmental Management, Liverpool University Hope
University, L16 9JD, UK; E-mail: magalarich20@gmail.com
3
Department of Wildlife and Animal Resources Management (WARM), Makerere University P.O.
Box 7062, Kampala, Uganda; E-mail: raykatumba21@gmail.com
4
Total E & P Uganda, Department of Environment and Social Affairs, P.O. Box 34867, Kampala,
Uganda; E-mail: david.ochanda@total.com
5
Biodiversity Solutions Ltd, P.O. Box, 22587, Kampala, Uganda; E-mail: skigoolo@yahoo.com
6
Q-Sourcing Servtecfor Total E & P Uganda, Department of Environment and Social Affairs, P.O.
Box 34867, Kampala, Uganda; E-mail: samuel.mutebi@external.total.com/samuelmutebi001@
gmail.com
7
Institute for Development, Ecology, Conservation, and Cooperation, via G. Tomasi di Lampedusa
33, I-00144 Rome, Italy; E-mails: d.dendi@ideccngo.org; l.luiselli@ideccngo.org
8
Department of Applied and Environmental Biology, Rivers State University of Science and Technology, P.M.B. 5080, Port Harcourt, Nigeria
9
Department of Zoology, Faculty of Sciences, University of Lomé, Lomé, Togo
10
Department of Natural Resource Ecology and Management, Iowa State University, Ames, IA,
50011, USA.
11
Department of Biology, Coe College, 1220 1st Avenue NE, Cedar Rapids, IA 52402, USA;
E-mail: dhughes@coe.edu
Abstract.
1. The area of the Murchison Falls-Albert Delta is among the most important for conservation in East Africa due
to the high species richness, and the presence of several endemic species of conservation concern.
2. Here, we report a study on the diversity patterns and community structure of the herpetofauna of this area.
3. Field studies were conducted in the Albert Nile Delta Ramsar site between 1st October 2017 and 9th September
2018. The data collection relied on Visual Encounter Surveys (VES), pitfall trapping, and dip netting. Descriptive statistics, i.e. species numbers in each transect were used as a measure of the present biodiversity, whereas
Chao1 and Chao species estimator algorithms were used to predict the potential number of species found in
each site/habitat.
4. A total of 898 individuals representing 25 reptile species belonging to four orders, 15 families, and 20 genera
were recorded during the 12 months of surveys.
5. The data shows some non-random spatial and temporal patterns whereby there is a cyclic reptilian diversity
peaking during the December-March and again towards June-August-September which are peaks of the dry
season.
6. The most frequently encountered species were Varanus niloticus, Crocodylus niloticus, Agama agama, Trachylepis maculilabris, and Lygodactylus guttularis, which accounted for almost 90% of all recorded individuals.
7. A total of 27 amphibian species, belonging to nine families and 10 genera were recorded during the period
of the survey. The diversity and abundance graphs would indicate amphibians having bimodal peaks (September-December, and March-May). The diversity seemed to dip during the dry season months – which is the
opposite case for reptiles.
Key words: Amphibia; Reptilia; diversity metrics; Albert Delta; Murchison Falls; Red List.
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Mathias Behangana et al. – Herpetofaunal Diversity
Introduction
The Murchison Falls-Albert Delta Wetland System, Uganda, is Ramsar Site No. 1640 (https://www.
ramsar.org/document/ramsar-advisory-mission-report-90-uganda-2018). It lies within the Lake Albert
Basin and falls almost entirely within the boundaries
of the Murchison Falls National Park, with an area of
about 17,293 ha, located at 01°57’N 031°42’E in Buliisa and Nwoya Districts. The site stretches from the
top of Murchison Falls, where the River Nile flows
through a rock cleft some 6m wide, to the delta at its
confluence with Lake Albert. The exception to this
is a one-kilometer wide band of land measured from
the southern bank of the Nile River running from the
western boundary of the national park to the point
at which the river joins Lake Albert. The end of the
Ramsar Site here can also be located as the western
edge of the inland delta formed by the Nile River’s
entry to Lake Albert (https://www.ramsar.org/document/ramsar-advisory-mission-report-90-uganda-2018). It is important as a spawning ground for
Lake Albert fisheries, supports globally threatened
bird species, and provides wetland habitat for biodiversity during the dry seasons. It is an area of high
species diversity and endemism making it a critical
area for wildlife at the regional scale. However, with
the impending oil exploration and production by the
government of Uganda and human pressures on the
south bank of the Nile river (Behangana et al. 2017),
the diversity and community structure of herpetofauna appears to be potentially under threat. Therefore,
understanding the diversity and community structure
of reptiles and amphibians in the Delta area of Murchison Falls is essential for supporting conservation
policies, guiding mitigation measures and, more importantly, providing data for species diversity monitoring (Brauneder et al. 2018).
The need to quantitatively assess the diversity
and community structure of herpetofauna is also supported by the fact that there have been very few field
surveys on amphibians and reptiles in the Murchison
Falls-Albert Delta Wetland System, with most studies
focusing on the ecology, population biology and conservation of the Nile crocodile Crocodylus niloticus
(Hutton 1991; Baguma 1996; Thorbjarnarson & Shirley 2009; Behangana 2014; Behangana et al. 2017).
Apart from crocodiles, this site is inhabited by other
reptile species, including the Nile monitor (Varanus
niloticus) as well as the threatened Nile Soft-shelled
Turtle (Trionyx triunguis), which is considered to
be CR B1ab (i, ii, iii, iv) in Uganda’s national red
list (Behangana 2014). Other species considered as
threatened according to Uganda’s red list include Pelusios adansonii CR B1ab (ii, iii) and P. chapini CR
B1ab (ii, iii), which are also likely to be in the area of
the Delta Ramsar Area as they had been recorded in
adjacent habitats (MB, pers. obs.). According to Rhodin et al. (2017), Pelusios adansonii is not known
from Uganda, but Spawls et al. (2018) report that it
likely occurs in northwestern Uganda. As for P. chapini, Rhodin et al.’s (2017) distribution map seems
to overlap with a large portion of western Uganda
near Lake Albert and this is confirmed by Spawls et
al. (2018). On the other hand, no amphibian species
of conservation concern have been recorded in the
Murchison Falls-Albert Delta Wetland System. Thus,
the scientific knowledge on the structure and organization of the reptile and amphibian communities
of the area (species richness, dominance, evenness)
remains anecdotal.
In this paper we quantitatively analyze, for the
first time in the international scientific literature, the
species composition, diversity and community metrics of the herpetofauna in the Murchison Falls-Albert Delta Wetland System by reporting field data
collected throughout twelve months, covering the
end of the wet season in 2017 and to the beginning
of the dry season in 2018. Although the field study
was designed to provide baseline data against which
oil and gas exploration and development activities
would be assessed, nonetheless it provided noteworthy insights into the community ecology of herpetofauna in tropical East Africa.
Study Area
The Murchison Falls-Albert Delta Wetland System lies on the Victoria Nile and is between 300-500
metres in width, has a strong current flowing from
east to west, and stretches from below the falls to
an area where the river becomes shallow as it enters
Lake Albert and the Victoria Nile Delta, a distance of
over 27 kilometers (WMD/NU, 2008). The site is situated within the Murchison-Semliki Landscape with
a “tropical wet and dry or savanna climate” (Aw)
according to the Köppen climate classification (e.g.
Peel et al. 2007). The distinct dry season characteristic for this class of climate is less pronounced in the
landscape and daily maximum temperatures remain
relatively low because of its average elevation above
a thousand meter which has a cooling effect. Mean
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Mathias Behangana et al. – Herpetofaunal Diversity
annual rainfall ranges between 1350-1600 mm distributed over two distinct rainy seasons from April
to May and from October to December. The mean
annual temperature ranges between 23-29 °C and
max 31 °C (see Peel et al. 2007). Evapotranspiration
(moisture loss from evaporation and transpiration
from plants) is relatively low for an Aw climate due
to moderate maximum daily temperatures in the dry
season (<30 °C) (Peel et al. 2007) The habitat along
the river is dominated by Vossia cuspidata (Hippo
grass) and Cyperus papyrus with tree cover behind.
Some parts of the riverbanks are lined by gallery forest/woodland. Several islands are present in the river,
some of which are large, over 2 km2, or more. These
islands support a range of vegetation communities;
papyrus, swamp reeds, tall grass, a few trees while
others are of short, grazed grass which is favored as
habitat by various fauna including mammals, birds,
herpetofauna, and insects. The geographic scope of
the surveys covered the whole delta area of the Murchison Falls-Albert Nile Delta Ramsar site (Fig. 1).
Five transects were demarcated and surveyed,
each transect surveyed on a monthly basis from the
same starting to the ending point, for the entire 12
months. The various sampling sites were reached
by a small boat. Surveys covered at least five days
during each month, with two days on the land transects and three days on the river channel transects.
In addition to transect surveys, two days of pitfall
trapping along the land transects on either bank of
the river were added in each month of the survey.
Details of the study transects are given in supplementary materials Table S1. All Delta Channel transects
start from around Pipeline Crossing North (PCN) to
the lakeside (Table S1). The Delta Channel transects
covered all navigable areas where the small boat
could reach including the floating islands overlooking Panyimur and Wanseko. Not all the waters of the
Figure 1. Map of the Murchison Falls-Lake Albert Delta Wetland System showing the study transects. Symbols: LS =
Land Transect South (LTS), DS = Delta Channel South (DCS), DM = Delta Channel Mid (DCM), DN = Delta Channel
North (DCN), LN = Land Transect North (LTN)
3
Mathias Behangana et al. – Herpetofaunal Diversity
Ramsar site were surveyed because some areas were
very shallow (some channels of the main ones and
the open waters at the river mouth lakewards) and not
accessible even by small boat.
Methods
2.1. Species identification
Identification of herpetofauna followed Schiøtz
(1999), Spawls et al. (2002; 2006; 2018), and Channing & Howell (2006). AmphibiaWeb (2017) and
The Reptile Database (Uetz & Hošek 2018) were
also used. The taxonomy of several African species
is still under debate. Concerning Agama agama, it
should be remarked that the taxonomy of this species
complex is still unresolved, and that, according to
some authorities, other species may occur in Uganda
(Leaché et al. 2016; Spawls et al. 2018). The conservation status of the herpetofauna is reported using the
IUCN Red Listing (IUCN, 2018) and the Ugandan
Red List (WCS, 2016).
2.2. Field protocol
Three methods were applied in the field study:
Visual Encounter Surveys (VES), Pitfall Trapping,
and Dip Netting. All these methods were applied
only during daytime, owing to health and safety issues raised by the funder with respect to night surveys. Ground-truthing marked out one kilometer
stretches for future reference along each transect.
All coordinates for sampling locations and routes
during fieldwork were marked using a portable GPS
(Garmin) and expressed in UTM-WGS 84 system.
Every amphibian or reptile individual recorded was
thus referenced to its current location and to these
sections of transects.
The surveys were habitat based. Key amphibian and reptilian habitats were stratified for ease of
sampling along each of the five transects sampled.
The key habitats for amphibians focused on for the
purposes of the surveys included lentic habitats and
vegetated wetlands. Suitable habitats for reptiles
included vegetated riverbanks, rocky outcrops, big
trees and woodlands. Edges of riverbanks were carefully monitored for any sun-basking reptiles.
Visual Encounter Surveys (VES, sensu Heyer
et al. 1994) were carried out on foot and by boats.
Potential retreat sites (stones, woods, cover objects)
were inspected in order to find any concealed specimens. Anuran species whose audio calls were known
by the main author were used to locate and record the
associated species. Each transect would be split into
sections depending on the dominant vegetation types
and for land transects, each section walked for one
hour while documenting any herpetofauna observed
while boats were used for water transects. Two boats
were used during the survey – a large boat with a 315
Horse Power engine in the open waters and as support
to a smaller one with a 15 Horse Power engine that
was used to access the shallower waters and sometimes coming to the edges of the river banks. Daytime counts of reptiles were conducted along the three
channels of the Ramsar Delta area by use of a small
engine boat because of its maneuverability and ability to access the shallower waters. All surveys started
from a marked point upstream. A group of three researchers would sit at locations on the starboard side
(right side when looking forward toward the bow) of
the boat and look out for any herpetofauna. On sighting an individual or hearing a call, a GPS recording
of the location of that individual was made at a perpendicular distance from the boat, the distance estimated, and photos of the habitats and species taken
where possible. The survey started in early mornings
between 800–900 hrs of the sampling day, moving
slowly downstream surveying one side of the channel, up to the mouth of the channel lakewards. At the
turn around point, the survey continued on the opposite side of the channel until the start point, with daily surveys usually ending from 1300–1400. Reptiles
basking on the banks of the river or floating on vegetation or in the river were counted and the activities
they were involved in documented.
A standardized dip-net was used to scoop through
aquatic habitats to sample for aquatic species and
tadpoles. Specimens of aquatic species or tadpoles
caught by this method, if not identifiable in the field
were preserved for later identification in the lab.
At selected sites, pitfall traps were set up with
a drift fence in the study area along land transects
to sample any surface-dwelling herpetofauna (Dodd
1991; Mitchell et al. 1993; Heyer et al. 1994, Handley & Varn 1994; Msuya 2001). Each drift fence
consisted of eleven 20-liter plastic buckets placed at
an interval of 10 m, covering a total length of 100 m.
The buckets were placed in holes dug in the substrate
using a hoe or pick-ax, such that their rim was level
with the ground.
A 100-meter-long and 0.5-meter-high drift fence
of black polythene supported vertically by wooden laths was set in an alternating manner with the
4
Mathias Behangana et al. – Herpetofaunal Diversity
(a)
16
14
14
12
10
10
9
10
10
10
9
8
7
8
7
6
No. of Species
buckets in the line, to permit detection of directional movements of species. The pitfall traps were inspected twice a day. This method was tried out once
and abandoned because of logistical constraints; the
array needed monitoring overnight to protect equipment from being stolen.
Pseudoreplication was avoided by surveying
a single site only once during the surveys. So, we
would exclude that the same individuals were observed more than once during our study. Opportunistic records are herein defined as those made outside
the sampling time but that occurred in the surrounding area to be impacted by the project. These opportunistic records helped complete the checklist of the
amphibians and reptiles as much as possible.
5
6
4
2
0
Month of Survey
(b)
16
14
13
14
13
14
13
13
12
Results
3.1. Distribution and diversity
of reptile species
A total of 898 individuals representing 25 reptile
species belonging to four orders (i.e. Chelonia, Sauria, Serpentes, and Crocodylia), 15 families, and 20
genera were recorded during the 12 months of surveys (Table S2). In terms of monthly variation in the
diversity of observed species, October 2017 showed
the highest reptilian diversity with 14 species, followed by January, March, July, and September 2018
(10 species each) (Fig. 2a) (raw data in Table S3).
The most frequently observed species were: Varanus niloticus (n = 361), Crocodylus niloticus (n
= 191), Agama agama (n = 172 individuals), and
Trachylepis maculilabris (n = 78) and each of them
was recorded every month of the survey. These four
species accounted for about 89.3% of all individuals
recorded in the study area.
10
No. of Species
2.3. Statistical analyses
The data were sorted according to the transect
and herpetofauna group using an excel spreadsheet.
The cleaned data were exported to Microsoft access
to perform more robust data filtering techniques to
determine the species number in each group and transect. Biodiversity pro software was used to predict
species occurrence and generating a species accumulation curve for each herpetofaunal group. To assess
the diversity and species richness between sites, the
functions “diversity” and “specnumber” in the Vegan R package (Oksanen, 2019) were used, whereas
the “renyi” diversity function was used for graphical
representation of diversities between the study sites.
11
11
12
9
8
8
7
6
4
2
0
Month of Survey
Figure 2: Species richness for reptiles (a) and amphibians
(b) over the 12-month survey period in the Murchison
Falls-Lake Albert Delta Wetland System.
3.1.1. Temporal and spatial species diversity and
richness for reptiles
The month of June 2018 had the highest combined number of individual reptiles sighted of all
months (105 individuals), followed by the months
of January and February (90 and 94 respectively),
while October (46), November and March (53 each)
showed the least combined number of individuals
sighted. Across transects, Land Transect South (LTS)
showed the highest species richness and diversity (21
species and 5.23 Simpson diversity index), followed
by Land Transect North (LTS) (17spp, 3.72 Simpson diversity index), while delta mid-channel trailed
(4 spp, Simpson diversity index 1.78 (Fig. 3a, (Tab.
S5). Although Land Transect South had the highest
species richness and diversity, it trailed below Delta
Channel North in terms of relative species abundance
by 12.7%. This was attributed to the numerous observations of Varanus niloticus which altered the over-
5
Mathias Behangana et al. – Herpetofaunal Diversity
Figure 3a: Renyi diversities for reptiles in 5 sites (transects).
(Where 1 = Delta Channel Mid (DCM), 2 = Delta Channel North (DCN), 3 = Delta Channel South (DCS), 4 = Land Transect North (LTN), 5 = Land Transect South (LTS). Renyi diversities for reptiles in 5 sites (transects). Land Transect North
and Land Transect South are more diverse than the remaining three sites. Diversity index Values at each site are represented
by dots while the median and the extremes are represented by the lines. When the sensitivity parameter on the x – axis is 0
the corresponding value is the species number at a given site, at 1, the conforming value is exponential Shannon (expH’)
and the corresponding value at sensitivity parameter 2 is the 2 is the inverse Simpson (1/D) while the conforming value of
Inf is the inverse relative dominance (1/P1).
Figure 3b: Renyi diversities for amphibians in 5 sites (transects).
Land Transect South and land Transect North are more diverse than all other sites. Diversity index Values at each site are
represented by dots while the median and the extremes are represented by the lines. When the sensitivity parameter on
the x – axis is 0 the corresponding value is the species number at a given site, at 1, the conforming value is exponential
Shannon (expH’) and the corresponding value at sensitivity parameter 2 is the 2 is the inverse Simpson (1/D) while the
conforming value of Inf is the inverse relative dominance (1/P1).
6
Mathias Behangana et al. – Herpetofaunal Diversity
all evenness. Land Transect North (LTN) and Delta
Channel Mid (DCM) had the lowest relative abundances (13.9% and 14.7% respectively)
A species accumulation curve for reptiles plotted showed that a plateau phase was not yet reached
(Fig. 4 a). This suggests that the reptilian diversity of
the surveyed sites had not been exhausted, and with
more time and/or effort and more habitats surveyed,
more species would have been added. Species diversity estimators Chao1 predicted the occurrence of 25
species, Chao2 predicted 36 species, Jacknife1 up to
32 species while Jacknife2 predicted 36 species at its
highest peak. The most conservative estimators for
this analysis were Chao1 and Jacknife1, predicting
up to 32 species. Chao1 and Jacknife1 graphs were
also still rising, meaning the species estimated could
still increase with more surveys.
3.2. Distribution and diversity
of amphibian species
A total of 315 individuals from 27 amphibian
species, all of order Anura, belonging to nine fam-
(a)
(b)
Figure 4: Species accumulation curve for reptiles (a) and amphibians (graphic (b) over the 12 months of survey in the
Murchison Falls-Lake Albert Delta Wetland System.
7
Mathias Behangana et al. – Herpetofaunal Diversity
ilies and 10 genera were recorded during the 12
months of surveys (Table S4). Phrynobatrachus bullans and Phrynabtrachus natalensis were the most
dominant species with a relative abundance of 34.3%
and 15.6% respectively while Hyperolius microps,
Hyperolius viridiflavus bayoni, Phrynobatrachus sp.
1, Ptychadena cf. aequiplicata, Sclerophrys pusilla and Xenopus victorianus had a very low relative
abundance (0.3%) since each was recorded once in
the study sites hence (Table S8). In general, only nine
species accounted for 91% of all individuals recorded in the study area. An indication of rarity in the
study area, where many species are not common, but
a few are abundant.
3.2.2. Temporal and spatial species diversity and
richness for amphibians
Species abundance varied markedly across the
year. April 2018 emerged with the highest number
of individuals of amphibians (791), followed by December, September and November (587, 540, and
404 respectively) (Table S5). A few species were recorded every month during the study period namely
- Phrynobatrachus bullans, Phrynobatrachus sp. 2,
and Phrynobatrachus natalensis, together with Ptychadena nilotica. Both Hoplobatrachus occipitalis
and Sclerophrys vittata were recorded for 11 months.
The months of March and April 2018 had the highest
species richness with 14 species each, followed by
November and December 2017, February and September 2018 (13 spp each), then June (12 spp), May
and August (11 spp each) (Fig. 2b, Table S5). LTS
registered the highest species diversity, richness and
relative abundance followed by LTN while DCM had
the lowest species diversity and richness but registered a higher relative abundance than the DCN (Tab.
S6). This points to the fact that in DCN, one species
(Phrynobatrachus bullans) was more common and
very abundant than the remaining three species.
A species accumulation curve plotted for amphibians showed that the plateau phase was not yet
reached (Fig. 3b). Species estimators Chao1, Chao2,
Jacknife1, and Jacknife2 put the maximum number
of species in the survey area up to 88, with Chao1
agreeing with the current number of 27, while Chao2
gives an erratic number of 88 species; Jacknife1 and
Jacknife2 predict up to 38 and 48 species respectively. Chao1 and Jacknife1 predicted a maximum of 38
amphibian species in the study area.
Discussion
Our study revealed that, as a general pattern, species diversity was highest at either end on the land
transects, and lowest in the Delta Channel. Therefore,
there was variation in species richness and diversity
in both amphibians and reptiles in the various sectors
of the study area. The occurrence of the least number of species in mid-channel was expected as most
species we encountered do not utilize open water
habitats. Species diversity was highest towards Land
Transect South (LTS) and second-highest towards
Land Transect North (LTN). The reason could be
due to an increase in diversity as one moves towards
land with anthropogenic disturbance thus favoring
common and abundant species for Land Transect
South (LTS) while the reverse is true as one moves
toward the park which is a natural habitat that favors
the more specialized species. There is literature evidence that species richness is correlated with habitat
heterogeneity at the local scale (Lundholm & Larson
2003; Báldi 2008). Stein et al. (2014) demonstrated
that habitat heterogeneity is a major driver of species
richness across taxa, biomes, and spatial scales. For
reptiles, the species estimators predict up to 32 reptile species in the Delta Ramsar Area, thus despite the
long research period (12 months) and the use of several independent survey methodologies, there may
still be a considerable amount of species that remain
undetected. The same was true, and even more evident, with regard to our surveys for amphibians (38
species predicted versus 27 species detected). Thus,
our data showed that, at least in the high biodiverse
areas of East Africa where surveys are logistically
difficult, the recorded herpetofaunal community metrics can be substantially biased because of suboptimal species detection.
The seasonal reptile metrics across the months
had a straightforward pattern: diversity peaked in the
dry season, but an abundance of observed individuals peaked in the wet season. In amphibians, there
were two peaks around the wet seasons (i.e. between
October-December, and March-May). The diversity
reaches low numbers during the dry season months.
Whereas the diversity for reptiles seemed to be influenced by major habitat and time-of-day factors, the
diversity for amphibians seemed to be influenced by
the proximity to riverbanks. The diversity of amphibians was highest along transects from either bank because of the proximity to water and increased diversity of habitats. On the other hand, unlike for reptiles
8
Mathias Behangana et al. – Herpetofaunal Diversity
where January is the peak of reptilian diversity, the
month which is the peak of the dry season recorded
the lowest amphibian diversity. The diversity for amphibians starts to go down in May with the passing of
the peak of the rainy season.
4.1. Species of Conservation Concern and
Critical Habitat Species
Most of the species recorded according to IUCN
(2020) are Not Evaluated (NE) or Least Concern
(LC) (Table S2 and S4). The species Trionyx triunguis (regionally vulnerable -VuA4bcd) (van Dijk et
al, 2017) and (Critically threatened nationally - CR
B1ab (i, ii, iii, iv)), Kinixys belliana, Trachylepis perrotetti, Crocodylus niloticus, Letheobia cf. sudanensis and Philothamnus bequaerti are reptilian species
whose populations should be monitored to detect any
eventual decline. All amphibian species reported are
of Least Concern (LC) at national and global levels.
However, Sclerophrys vittatus, Phrynomantis microps and Ptychadena schillukorum should be monitored
because they are potentially vulnerable to declines.
In terms of Critical Habitats (CH; sensu Stefan
et al. 2013), banks with woody vegetation and papyrus dominated habitats, particularly along the Delta
Channel North (DCN) are in greatest need of protection before and during any activities in the oil and gas
industry. The ecotones between water and the land
are critical habitats for herpetofaunal conservation
and should be protected when carrying out any oil
and gas activity.
Acknowledgments
The research was funded by Total E&P Uganda
B.V. under a service contract to Biodiversity Solutions Limited (BSL). We thank the Uganda National
Council of Science and Technology (UNCST) for
granting research permits and the Uganda Wildlife
Authority (UWA) for field support and permits to
conduct this research in Murchison Falls National
Park. We also acknowledge the field assistance by
Environmental and Biodiversity Field Officers (EBFOs), Community Liaison Officers (CLOs), UWA
Rangers, and all other staff who contributed to the
success of this research in one way or another. Great
thanks to Michele Menegon from Trento Science
Museum (Italy) who was very helpful in species id
confirmation using DNA barcoding We are indebted
to Aaron Bauer, Eli Greenbaum, and two anonymous
referees for useful comments on the submitted draft.
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11
Mathias Behangana et al. – Herpetofaunal Diversity
Online Supplementary Materials
Table S1. Transects surveyed for herpetofauna with their start and end points, at the study area.
Code
Location
Land Transect North
LN
Circumferenced by road from Hip- From N2.26994 E31.37618
po pool, along Shoebill track to
Buligi track on Pipeline Crossing to N2.25520 E31.49560
North (PCN)
Land Transect South
LS
From Wanseko town outskirts (Ka- From N2.18679 E31.38752
tanga) to Kasinyi village - delineat- N2.22285 E31.45043
ed by road parallel to the river
to 10
North DN
Channel transect where river hugs From N2.25303 E31.48686
the Northern bank
N2.27092 E31.37122
to 20
Delta Channel Mid Tran- DM
sect
Longest distance starts where the From N2.22975 E31.44260
river splits into two main channels
N2.23835 E31.36091
to 10
Delta Channel
Transect
Channel transect where river hugs From N2.24807 E31.49746
the Southern bank
N2.18350 E31.37459
to 20
Delta Channel
Transect
South DS
Coordinates
Approximate
Length (km)
Transect Name
20
NB: The distances only refer to the navigable parts of the river by the small boat.
12
Table S2: Reptilian species recorded in the Albert Delta Ramsar Area between October 2017 and September 2018 with IUCN Status
Family
Species
Common name
Global Status
Country Status
Chelonia
Testudinidae
Kinixys belliana
Bell’s Hinge-back Tortoise
Not Evaluated (NE)
Near Threatened (NT)
Chelonia
Trionychidae
Trionyx triunguis
Nile Soft-shelled Turtle
Vulnerable (VU)
Critically Threatened
(CR C2a(i, ii)+D)
Chelonia
Pelomedusidae
Pelomedusa neumanni
Marsh terrapin
Least Concern (LC)
Least Concern (LC)
Sauria
Agamidae
Agama agama
Common Agama
Least Concern (LC)
Least Concern (LC)
Sauria
Geckonidae
Lygodactylus guttularis
Chevron-throated Dwarf
Gecko
Not Evaluated (NE)
Least Concern (LC)
Sauria
Chameleonidae
Chamaeleo gracilis
Gracile Chameleon
Least Concern (LC)
Least Concern (LC)
Sauria
Scincidae
Mochlus sundevallii
Sundevall’s Writhing
Skink
Least Concern (LC)
Least Concern (LC)
Sauria
Scincidae
Trachylepis maculilabris
Speckle-lipped Skink
Not Evaluated (NE)
Least Concern (LC)
Sauria
Scincidae
Trachylepis perrotetti
Taita Mabuya
Not Evaluated (NE)
Data Deficient (DD)
Sauria
Gerrhosauridae
Broadleysaurus major
Tawny Plated-lizard
Not Evaluated (NE)
Vulnerable (Vu)
Sauria
Varanidae
Varanus exanthematicus
Western Savanna Monitor
Least Concern (LC)
Data Deficient (DD)
Sauria
Varanidae
Varanus niloticus
Nile Monitor
Not Evaluated (NE)
Least Concern (LC)
Crocodylia
Crocodylidae
Crocodylus niloticus
Nile Crocodile
Least Concern (LC)
Near Threatened (NT)
Serpentes
Typhlopidae
Letheobia sp cf. sudanensis
Sudan Beaked Snake
Not Evaluated (NE)
Data Deficient (DD)
Serpentes
Colubridae
Crotaphopeltis degeni
Yellow-flanked Snake
Not Evaluated (NE)
Least Concern (LC)
Serpentes
Colubridae
Crotaphopeltis hotamboeia
White-lipped Herald
Snake
Not Evaluated (NE)
Least Concern (LC)
Serpentes
Colubridae
Philothamnus bequaerti
Uganda Green Snake
Not Evaluated (NE)
Data Deficient (DD)
Serpentes
Colubridae
Philopthamnus semivariegatus
Variegated Bush Snake
Not Evaluated (NE)
Least Concern (LC)
Serpentes
Colubridae
Psammophis mossambicus
Olive Sand Snake
Not Evaluated (NE)
Least Concern (LC)
Serpentes
Colubridae
Psammophis sibilans
Hissing Sand Snake
Not Evaluated (NE)
Least Concern (LC)
Serpentes
Colubridae
Hapsidophrys smaragdina
Emerald Snake
Not Evaluated (NE)
Least Concern (LC)
Serpentes
Pythonidae
Python sabae
African Python
Not Evaluated (NE)
Least Concern (LC)
Serpentes
Elapide
Dendroaspis jamesonii
Jameson’s Green Mamba
Not Evaluated (NE)
Least Concern (LC)
Serpentes
Elapide
Naja subfulva
Forest Cobra
Not Evaluated (NE)
Least Concern (LC)
Serpentes
Viperidae
Bitis arietans
Puff Adder
Not Evaluated (NE)
Least Concern (LC)
Mathias Behangana et al. – Herpetofaunal Diversity
13
Order
Table S3: Reptilian species recorded in the Albert Delta Ramsar Area between October 2017 and September 2018
Species/Month of Survey
Oct17
DecNov-17 17
Agama agama
1
Bitis arietans
1
1
Jan18
1
Feb18
AprMar-18 18
1
1
1
1
1
1
May-18
Jun18
1
1
Chamaeleo gracilis
Crocodylus niloticus
1
1
1
1
1
1
1
1
1
Jul18 Aug-18
1
1
1
1
Broadleysaurus major
1
4
1
1
1
1
2
2
1
1
1
1
1
5
1
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
7
4
1
1
Psammophis mossambicus
1
Psammophis sudanensis
1
Python sabae
1
Trachylepis maculilabris
1
1
1
1
Trachylepis perrotetti
1
1
1
1
1
1
2
1
Varanus exanthematicus
1
2
1
1
1
1
1
1
1
3
1
1
12
4
Trionyx triunguis
Total
11
1
1
Philothamnus bequaerti
Varanus niloticus
1
1
1
Pelomedusa neumanni
Philopthamnus semivariegatus
1
1
1
Mochlus sundevallii
Naja subfulva
12
1
1
Letheobia sp cf. sudanensis
Lygodactylus guttularis
1
1
Hapsidophrys smaragdina
Kinixys belliana
12
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
12
14
8
9
10
9
10
5
7
1
10
6
10
99
Mathias Behangana et al. – Herpetofaunal Diversity
Dendroaspis jamesonii
1
1
Crotaphopeltis degeni
Crotaphopeltis hotamboeia
Sep18 Total
14
Mathias Behangana et al. – Herpetofaunal Diversity
Table S4: Amphibian species recorded in the Albert Delta Ramsar Area between October 2017 and September 2018 with
IUCN Status.
Family
Species
Common name
Global Status
Country Status
Pyxicephalidae
Amietia nutti
Least Concern (LC)
Least Concern (LC)
Bufonidae
Sclerophrys gutturalis
Guttural Toad
Least Concern (LC)
Least Concern (LC)
Bufonidae
Sclerophrys regularis
Common Toad
Least Concern (LC)
Least Concern (LC)
Bufonidae
Sclerophrys vittata
Lake Victoria Toad
Data Deficient (DD) Least Concern (LC)
Flat-backed Toad
Bufonidae
Sclerophrys pusilla
Least Concern (LC)
Data Deficient
Dicroglossidae
Hoplobatrachus occipitalis Crowned Bullfrog
Least Concern (LC)
Least Concern (LC)
Hemisotidae
Hemisus marmoratus
Shovel-nosed Frog
Least Concern (LC)
Least Concern (LC)
Hyperoliidae
Afrixalus quadrivittatus
Four-lined Spiny Reed Frog
Least Concern (LC)
Least Concern (LC)
Hyperoliidae
Hyperolius microps
Sharp-headed Long Reed
Frog
Least Concern (LC)
Least Concern (LC)
Hyperoliidae
Hyperolius cinnamomeoventris species complex
Cinnamon-bellied Reed Frog
Least Concern (LC)
Least Concern (LC)
Hyperoliidae
Hyperolius kivuensis
Kivu Reed Frog
Least Concern (LC)
Least Concern (LC)
Hyperoliidae
Hyperolius v. bayoni
Bayoni’s Reed Frog
Least Concern (LC)
Least Concern (LC)
Hyperoliidae
Hyperolius v. viridiflavus
Common Reed Frog
Least Concern (LC)
Least Concern (LC)
Phrynobatrachidae
Phrynobatrachus bullans
Bubbling puddle frog
Least Concern (LC)
Data Deficient (DD)
Phrynobatrachidae
Phrynobatrachus sp. 1
Phrynobatrachidae
Phrynobatrachus
natalensis
Natal Dwarf Puddle Frog
Least Concern (LC)
Least Concern (LC)
Phrynobatrachidae
Phrynobatrachus sp. 2
Phrynobatrachidae
Phrynobatrachus sp.3
Microhylidae
Phrynomantis microps
West African Rubbber Frog
Least Concern (LC)
Data Deficient (DD)
Ptychadenidae
Ptychadena anchietae
Anchieta’s Ridged Frog
Least Concern (LC)
Least Concern (LC)
Ptychadenidae
Ptychadena sp.
cf.aequiplicata
Victoria Grassland Frog
Least Concern (LC)
Data Deficient (DD)
Ptychadenidae
Ptychadena nilotica
Nile Grass Frog
Least Concern (LC)
Least Concern (LC)
Ptychadenidae
Ptychadena oxyrhynchus
Sharp-nosed Ridged Frog
Least Concern (LC)
Least Concern (LC)
Ptychadenidae
Ptychadena porosissima
Grassland Ridged Frog
Least Concern (LC)
Least Concern (LC)
Ptychadenidae
Ptychadena Sp.
Ptychadenidae
Ptychadena schillukorum
Sudan Grassland Frog
Least Concern (LC)
Data Deficient (DD)
Pipidae
Xenopus victorianus
Lake Victoria Clawed Frog
Least Concern (LC)
Least Concern (LC)
15
Mathias Behangana et al. – Herpetofaunal Diversity
Table S5: Species richness and diversity of reptiles in the Delta Area of the Murchison Falls-Albert Delta Ramsar Site,
Uganda
Delta area or
transect
Simpson reciprocal
index
Species richness
Relative abundance
(%)
Pielou’s J evenness
Shannon index
DCM
1.78
4
13.9
0.37
0.74
DCN
3.37
8
31.5
0.89
1.33
DCS
1.90
15
20.9
0.70
0.82
LTN
3.72
17
14.7
0.90
1.74
LTS
5.23
21
18.8
0.85
2.01
Table S6: Species richness and diversity of amphibians in the Delta Area of the Murchison Falls-Albert Delta Ramsar
Site, Uganda
Delta area or
transect
Simpson reciprocal
index
Species richness
Relative abundance
(%)
Pielou’s J evenness
Shannon index
DCM
1.29
04
14.44
0.28
0.51
DCN
3.21
12
7.68
0.66
1.74
DCS
2.76
12
23.29
0.54
1.45
LTN
8.06
19
21.30
0.77
2.44
LTS
8.09
23
33.28
0.77
2.46
16
Table S8: Amphibian species recorded in the Albert Delta Ramsar Area between October 2017 and September 2018
0.03
Hyperolius viridiflavus bayoni
1
0.03
1
0.03
1
0.03
1
0.03
1
0.03
3
0.08
3
0.08
7
0.19
10
0.27
13
0.34
13
0.34
15
0.40
17
0.45
20
0.53
38
1.01
34
96
2.54
19
98
2.60
34
109
2.89
1
32
148
3.92
2
31
178
4.72
8
182
4.82
1
1
Phrynobatrachus sp.
1
Ptychadena. cf aequiplicata
1
Sclerophrys pusilla
1
Xenopus victorianus
3
Ptychadena sp
3
Sclerophrys gutturalis
7
Phrynomantis microps
10
Phrynobatrachus sp1
1
Amietia angolensis
Ptychadena oxyrhynchus
1
12
2
10
15
Hemisus marmoratus
2
Ptychadena anchietae
3
9
3
20
Ptychadena schillukorum
2
29
7
2
2
24
3
26
Hyperolius cinnamomeoventris species
complex
Hyperolius viridiflavus viridiflavus
Sclerophrys regularis
3
1
22
37
6
4
1
Hyperolius kivuensis
6
30
1
15
2
Afrixalus quadrivittatus
12
45
1
7
3
4
6
3
1
14
7
16
24
7
3
55
17
17
9
1
104
4
18
1
7
2
50
5
1
20
5
36
182
4.82
17
Hoplobatrachus occipitalis
3
13
24
Ptychadena porosissima
2
19
25
Sclerophrys vittata
1
5
50
Phrynobatrachus sp. 2
4
35
135
10
10
10
8
37
15
4
11
42
321
8.51
Ptychadena nilotica
5
29
33
9
18
20
126
60
29
20
30
50
429
11.37
Phrynobatrachus natalensis
24
17
84
8
22
11
155
79
57
7
20
107
591
15.66
Phrynobatrachus bullans
21
259
88
114
30
66
180
80
117
95
127
117
1294
34.30
Total
62
404
587
149
124
147
791
330
278
159
202
540
3773
100.00
Mathias Behangana et al. – Herpetofaunal Diversity
1
1
Hyperolius microps