The Cyphomyrus Myers 1960 (Osteoglossiformes: Mormyridae) of the Lufira basin (Upper Lualaba: DR Congo): A generic reassignment and the description of a new species

Abstract

Within a comparative morphological framework, Hippopotamyrus aelsbroecki, only known from the holotype originating from Lubumbashi, most probably the Lubumbashi River, a left bank subaffluent of the Luapula River, is reallocated to the genus Cyphomyrus. This transfer is motivated by the fact that H. aelsbroecki possesses a rounded or vaulted predorsal profile, an insertion of the dorsal fin far anterior to the level of the insertion of the anal fin, and a compact, laterally compressed and deep body. In addition, a new species of Cyphomyrus is described from the Lufira basin, Cyphomyrus lufirae. Cyphomyrus lufirae was collected in large parts of the Middle Lufira, upstream of the Kyubo Falls and just downstream of these falls in the lower Lufira and its nearby left bank affluent, the Luvilombo River. The new species is distinguished from all its congeners, that is, firstly, from C. aelsbroecki, C. cubangoensis and C. discorhynchus, by a low number of dorsal fin rays, 27‐32 (vs. higher, 36 (37), 34 (33‐41) an 38 (38‐40), respectively) and, secondly, from C. aelsbroecki, C. cubangoensis, and C. discorhynchus by a large prepelvic distance, 41.0–43.8% LS (vs. shorter, 39.7%, 38.9–39.1% and 37.0–41.0% LS, respectively). The description of yet another new species for the Upemba National Park and the Kundelungu National Park further highlights their importance for fish protection and conservation in the area. Hence, there is an urgent need for the full integration of fish into the management plans of these parks.

Full text

SPECIAL ISSUE REGULAR PAPER
The Cyphomyrus Myers 1960 (Osteoglossiformes: Mormyridae)
of the Lufira basin (Upper Lualaba: DR Congo): A generic
reassignment and the description of a new species
Christian Mukweze Mulelenu
1,2,3,4
| Bauchet Katemo Manda
2,3,4
| Eva Decru
3,4
|
Auguste Chocha Manda
2
| Emmanuel Vreven
3,4
1
Département de Zootechnie, Faculté des
Sciences Agronomiques, Université de
Kolwezi, Kolwezi, Democratic Republic of the
Congo
2
Département de Gestion des Ressources
Naturelles Renouvelables, Unité de recherche
en Biodiversité et Exploitation durable des
Zones Humides, Université de Lubumbashi,
Lubumbashi, Democratic Republic of the
Congo
3
Vertebrate Section, Ichthyology, Royal
Museum for Central Africa, Tervuren, Belgium
4
Laboratory of Biodiversity and Evolutionary
Genomics, KU Leuven, Leuven, Belgium
Correspondence
Emmanuel Vreven, Curator of Fishes
Vertebrate Section, Royal Museum for Central
Africa (RMCA), Leuvensesteenweg 13, B-3080
Tervuren, Belgium
Email: emmanuel.vreven@africamuseum.be
Funding information
This study was made possible by a DEA
scholarship (2016–2018) to the first author
(CMM) from the Mbisa Congo project
(2013-2018), financed through a framework
agreement project between the RMCA and the
Belgian Development Cooperation.
Abstract
Within a comparative morphological framework, Hippopotamyrus aelsbroecki, only
known from the holotype originating from Lubumbashi, most probably the Lubumbashi
River, a left bank subaffluent of the Luapula River, is reallocated to the genus
Cyphomyrus. This transfer is motivated by the fact that H. aelsbroecki possesses a
rounded or vaulted predorsal profile, an insertion of the dorsal fin far anterior to the
level of the insertion of the anal fin, and a compact, laterally compressed and deep
body. In addition, a new species of Cyphomyrus is described from the Lufira basin,
Cyphomyrus lufirae.Cyphomyrus lufirae was collected in large parts of the Middle Lufira,
upstream of the Kyubo Falls and just downstream of these falls in the lower Lufira and
its nearby left bank affluent, the Luvilombo River. The new species is distinguished
from all its congeners, that is, firstly, from C. aelsbroecki,C. cubangoensis and
C. discorhynchus, by a low number of dorsal fin rays, 27-32 (vs. higher, 36 (37),
34 (33-41) an 38 (38-40), respectively) and, secondly, from C. aelsbroecki,
C. cubangoensis,andC. discorhynchus by a large prepelvic distance, 41.0–43.8% L
S
(vs.
shorter, 39.7%, 38.9–39.1% and 37.0–41.0% L
S
, respectively). The description of yet
another new species for the Upemba National Park and the Kundelungu National Park
further highlights their importance for fish protection and conservation in the area.
Hence, there is an urgent need for the full integration of fish into the management
plans of these parks.
KEYWORDS
Cyphomyrus aelsbroecki,Cyphomyrus discorhynchus, DNA barcoding, Kundelungu National
Park, morphometry, Upemba National Park
1|INTRODUCTION
The family Mormyridae is endemic to the freshwaters of Africa
(Lévêque et al., 1990). With a total of 224 valid species (Fricke et al.,
2019) and 21 genera (Sullivan et al., 2016) it is one of the largest fish
families on the continent. The family is present and well-diversified in
most continental aquatic ecosystems (Hopkins et al., 2007; Lavoué
et al., 2000; N'da et al., 2014), with the greatest species-level diversity
occurring in the Congo and Lower Guinea ichthyofaunal provinces
(Hopkins et al., 2007; Roberts, 1975; Sullivan et al., 2016). Species of
this family are African osteoglossomorph fishes of great interest for
their ability to produce electric discharges from an electric organ
located in their caudal peduncle (Hopkins, 1986; Lavoué et al., 2000).
The highest densities of electroreceptors are on the tip of the chin,
with median densities on the nasal region and low densities on the
back (Hollmann et al., 2008: Figure 8). The electric discharges pro-
duced are only a few millivolts and thus are not generated for attack
or defence as in Malapteruridae (Skelton, 1996; Sullivan et al., 2016).
Received: 9 January 2019 Accepted: 15 December 2019
DOI: 10.1111/jfb.14237
FISH
J Fish Biol. 2020;1–19. wileyonlinelibrary.com/journal/jfb © 2019 The Fisheries Society of the British Isles 1

Instead, they are used primarily in electro-location of objects in their
environment and communication, which is important for facilitating
activity at night or in murky waters (Feulner et al., 2009; Hopkins,
1986; Rich et al., 2017; Teugels et al., 2001). Each species has its own
unique electric organ discharge (EOD) and the sexes of some species
also may have different discharges (Rich et al., 2017).
The genus Cyphomyrus Myers 1960 is morphologically characterized
by: (a) a rounded or arched back; and (b) a long dorsal fin, with its origin
situated in between the pelvic- and the anal-fin origin and its posterior
end situated behind that of the anal fin (Kramer & Van der Bank, 2011;
Levin & Golubtsov, 2018; Skelton, 1996). In addition, species of
Cyphomyrus possess 27–40 rays on the dorsal fin, 22–27 rays on the anal
fin, 10–12 rays on the pectoral fin, 12, rarely 13, circumpenduncular
scales (Kramer & Van der Bank, 2011; Skelton, 1996) and 58–74 scales
along the lateral line (Boulenger, 1909). The chin has a short bulbous pro-
tuberance, which hides the essentially inferior position of the mouth
(Kramer & Van der Bank, 2011). Teeth, generally three to five in the upper
jaw and five to six in the lower jaw, are situated on the ventral midline of
both jaws. The body is compact, laterally compressed and deep.
The name Cyphomyrus is derived from the Greek prefix ‘kuphos’,
meaning hunchback, and the Greek ‘muros’, the often-used combining
form for mormyrus (Myers, 1960). The genus Cyphomyrus currently con-
tains six valid species (Fricke et al., 2019; Kramer & Van der Bank, 2011;
Levin & Golubtsov,2018). Four of these are known from the Congo basin,
more precisely C. discorhynchus (Peters 1852), C. macrops (Boulenger
1909), C. psittacus (Boulenger 1897) and C. wilverthi (Boulenger 1898)
(see Gosse, 1984). Cyphomyrus macrops and C. wilverthi are Congo basin
endemics (Gosse, 1984), while C. psittacus, described from the Wagenia
or BoyomaRapids near Kisangani (Democratic Republic of the Congo), is
also found in West Africa (Bigorne, 2003; Gosse, 1984) and C.dis-
corhynchus, described from the Lower Zambezi (Mozambique), is found
throughout the Zambezi and the Lake Malawi and Tanganyika basins
(Gosse, 1984; Scott et al., 2006; Skelton, 1996). Furthermore, C. petherici
(Boulenger 1898) is known from the White Nile, the Blue Nile, the
Murchison Nile and the Omo system (Golubtsov & Darkov, 2008;
Levin & Golubtsov, 2018). As such, its presence in the Congo basin, that
is, the Malagarazi River (De Vos et al., 2001), seems doubtful and requires
confirmation (see Levin & Golubtsov, 2018). Finally, C. cubangoensis
(Pellegrin 1936), is known from the Okavango River basin (syntypes)
and Okavango delta, the Kwando River and the Upper Zambezi River
system (Kramer & Van der Bank, 2011).
During a series of recent field expeditions (2012–2016), in the
Upper Lualaba (sensu Teugels & Thieme, 2005), covering parts of both
the Upemba National Park (UNP) and the Kundelungu National Park
(KNP), two groups of Cyphomyrus specimens were collected. Of the two
groups of specimens collected, the first is here identified as
C. discorhynchus, to date the only species reported from the Upper Lua-
laba and the lower stretch of its largest right bank tributary, the (Lower)
Lufira (see Poll, 1976). The second group consists of specimens collected
in the Middle Lufira basin and just downstream of the Kyubo Falls in the
Lower Lufira and the lower stretch of its nearby left bank affluent the
Luvilombo River. These specimens were revealed to be morphologically
clearly different from all valid Cyphomyrus species currently known from
the Congo basin, although they are similar to C. discorhynchus and
Hippopotamyrus aelsbroecki (Poll 1945), herein transferred to the genus
Cyphomyrus (see below). Furthermore, a detailed comparison of the
newly collected specimens with C. cubangoensis and C. discorhynchus,the
two species currently known from Southern Africa, was also undertaken
and revealed that the specimens from the Middle and Lower Lufira rep-
resent a new species for science.
In addition, the paper also proposes the transfer of H. aelsbroecki,
originally described as a new Gnathonemus from Lubumbashi, most
probably the Lubumbashi River, which is a left bank subaffluent of the
Luapula River, to the genus Cyphomyrus. This reallocation is motivated
based on the external morphological diagnostic characters of the
genus Cyphomyrus as identified by Kramer and Van der Bank (2011).
As such, the purpose of the present paper is to (a) motivate the trans-
fer of H. aelsbroecki to the genus Cyphomyrus and (b) formally describe
a new Cyphomyrus species for science, named C. lufirae sp. nov.
2|MATERIALS AND METHODS
2.1 |Collections
Specimens for this study were collected during recent field trips
(2012–2016) and mostly deposited in the Royal Museum for Central
Africa (RMCA) or were obtained from the Musée National d’Histoire
Naturelle (MNHN), RMCA, Zoologisches Museum Berlin (ZMB) and
Zoologisches Staatssammlung München (ZSM) collections. During
these recent field trips, fish were caught using various fishing
methods, e.g., gill nets and fykes. Freshly caught fish were
anaesthetized by an overdose of clove oil. After death, fin clips for
molecular analysis were taken and preserved in 99% ethanol. Speci-
mens for further morphological studies were preserved in 10% for-
malin before being transferred to 70% ethanol for long-term storage
at the RMCA. Permanent tags were attached to specimens bearing
a unique number by which a specimen and its fin clip are linked.
2.2 |Morphological data
A total of 90 specimens of Cyphomyrus were examined, including
64 from the Upper Lualaba basin and 26 specimens from other areas
for comparative purposes (see material examined). Among the 64 speci-
mens from the Upper Lualaba basin, there were 19 specimens from the
Kamalondo Depression and the more downstream part of the Lower
Lufira, here all identified as C. discorhynchus. These specimens were
included in the analyses because of the close similarity of
C. discorhynchus with the new species and the widespread geographical
distribution of this species (see Skelton, 1996). The other 45 specimens
from the Upper Lualaba basin were collected from the Lower and the
Middle Lufira basin, and all belong to the new species. The 26 specimens
studied for comparative purposes include the lectotype and the four
paralectotypes of C. discorhynchus.Todate,C. discorhynchus contains
two junior synonyms, C. smithersi (Määr 1962) and C. tanganicanus
2MUKWEZE MULELENU ET AL.
FISH

(Boulenger 1906). Therefore, the three syntypes of C. tanganicanus,
originally described from Sumbu and a river at Msamba on the Lake
Tanganyika basin, and seven additional specimens, all originating from
some Lake Tanganyika tributaries, were included in the analysis. Note
that throughout the presentation of the results the syntypes of
C. tanganicanus and the C. discorhynchus specimens from the affluent
rivers of the Lake Tanganyika basin have been labelled separately to
illustrate the observed (geographical) differences with the
C. discorhynchus identified specimens from the Zambezi and the Upper
Lualaba. The holotype of C. smithersi (NMBZ 0833) could not be exam-
ined because it could not be obtained on loan. Therefore, five speci-
mens from Lake Kariba, type locality of C. smithersi, were included as
topotypic specimens for this species. Finally, four of the 14 syntypes of
C. cubangoensis and the holotype of C. aelsbroecki were also included.
Eleven counts and 29 measurements were taken on each specimen.
Counts follow Boden et al. (1997). For the dorsal- and anal-fin ray counts
the two simple rays preceding the segmented rays were also included in
these counts. All rays were counted at their base. The total number of
vertebrae was not used for the analyses because it was not taken on all
specimens and all species. All measurements were taken according to
Boden et al. (1997) except for the two different measurements of the
caudal peduncle depth for which only the middle caudal peduncle depth
has been taken as both measurements revealed to be highly similar. Five
measurements have been added, four according to Kramer and Van der
Bank (2011: Figure 3): (a) the post-dorsal distance (pD), (b) the distance
between pectoral and pelvic fins (PPF), (c) the length of snout 2 (LSo) and
(d) the length of snout 3 (LSc). In addition, one is taken for the first time,
that is the body depth at the level of the anterior insertion point of the
dorsal fin, which is the distance between the anterior insertion point of
the dorsal fin and the corresponding point on the ventral edge of the
body when following a vertical line perpendicular on the horizontal body
axis. All measurements were taken using callipers of 0.1 mm precision on
the left-hand side of the specimens except when this side was damaged
in which case the use of the right-hand side was mandatory.
Data were explored and analysed using multivariate principal com-
ponent analyses (PCA). Meristics and measurements were analysed
separately, with the correlation matrix used for the PCA on the raw
meristics, and the covariance matrix for the PCA on the measurements
expressed as percentages (Bookstein et al., 1985). Measurements on
the head were expressed in percentage of head length (L
H
) and mea-
surements on the body in percentage of standard length (L
S
). Sca-
tterplots were used to explore the characters that could separate the
species. Statistical analyses were executed in STATISTICA software for
Windows version 7 (Statsoft, Inc). Being invariable, the number of
pelvic-fin rays was not included in the PCA.
In the description of the new species, measurements are expressed
as proportions following Boulenger (1898) in order to allow direct com-
parison with the original descriptions and recent redescriptions (see
Kramer & Van der Bank, 2011) of its congeners. In addition, tabulated
measurements are provided in percentages alongside the description. In
the diagnosis for the new species, the provided meristic comparative
data for its congeners are based on: (a) first-hand personal observations;
and (b) the original descriptions of these species (between brackets).
This approach has been preferred to avoid reporting large amounts of
meristic intraspecific variation which might be largely due to misidentifi-
cations of subsequently collected specimens. However, for the mea-
surements only morphometric data generated within the framework of
the present study were presented, as the original descriptions lack data
on the diagnostic measurements retained.
Non-parametric Mann–Whitney U-tests with sequential
Bonferroni correction (Rice, 1989) were used for univariate compari-
sons of the raw meristic data and the percentages, in % L
H
or % L
S
for
the head and body measurements, respectively. For measurements,
specimens of similar length classes (96 < L
S
≤163) were used to pre-
vent the potential confounding effects of allometric growth.
All locality data have been translated into English. Following the
new policy of the RMCA, collection numbers MRAC A0–A9 are listed
as MRAC 2000–2009 and B0–B6 as MRAC 2010–2016.
2.3 |Molecular data
For a total of 22 samples, 651 bp of the barcode gene cytochromec
oxidase (COI) mitochondrial DNA (mtDNA) has been successfully
sequenced. DNA was extracted from fin clips using a Nucleospin
Tissue kit (Macherey-Nagel, Germany). The COI gene was amplified
using the universal M13-tailed primer cocktail for fish DNA barcoding
(Ivanova et al., 2007). The 25 μl PCR reactions included 2.5 μl of PCR
buffer (10×), 2.5 μl of dNTP (2 mM), 1.25 μl of primer cocktail (2 μM),
0.2 μl of Taq DNA polymerase (5 units per μL), 2.0 μl of extracted
DNA and 16.75 μl of mQ-H
2
O. The PCR profile was 3 min at 94C,
followed by 35 cycles of 45 s at 94C, 40 s at 51C, 1 min 30 s at
72C and ending with a final extension of 10 min at 72C (Decru et al.,
2016). Amplification was visually checked on 1.2% agarose gels, and
PCR products were purified using an ExoSAP-IT PCR Clean-up Kit
(Affymetrix, Inc.; Thermo Fisher Scientific, Waltham, MA, USA). Stan-
dard Sanger Sequencing was executed by the external company Mac-
rogen, where samples were sequenced bidirectionally using an ABI
3730XL capillary sequencer. The DNA sequences were assembled
and visually checked in CodonCode Aligner 4.2.7 (CodonCode Corpo-
ration; Centerville, MA, USA) and aligned and analysed in MEGA 7.02.
Sequences from Genbank from Genomyrus donnyi were included as
an outgroup. Unfortunately, for C. cubangoensis, no samples nor
sequences on Genbank were available. A model test was performed
and the HKY + G was chosen as the most suitable model under the
akaike information criterion. Maximum likelihood (ML) trees with
500 bootstrap replications were produced. The sequences, including
the paragenetype of the new species, were deposited in GenBank for
future reference (Supporting Information Table S1).
2.4 |Abbreviations
AIC, Akaike Information Criterion; a.s.l., above sea level; BEZHU,
Biodiversité et Exploitation durable des Zones Humides; COI, cyto-
chrome c oxidase subunit; CTB/BTC, Coopération Technique Belge/
MUKWEZE MULELENU ET AL.3
FISH

Belgische Technische Coöperatie; CU, Cornell University; DEA,
Diplôme d'Etudes Approfondies; DRC, Democratic Republic of the
Congo; ISP, Institut Supérieur Pédagogique; KNP, Kundelungu
National Park; LH, head length; LS, standard length; LT, total length;
mtDNA, mitochondrial DNA; MNHN, Musée National d'Histoire
Naturelle, Paris, France; MRAC, Musée Royal de l'Afrique Centrale,
Tervuren, Belgium; MWU-test, Mann–Whitney U-test; NHM, Natural
History Museum, London, UK; PC, principal component; PCA, princi-
pal components analysis; RMCA, Royal Museum for Central Africa,
Tervuren, Belgium; SAIAB, South African Institute for Aquatic Biodi-
versity, Grahamstown, South Africa; UNIKOL, Université de Kolwezi;
UNILU, Université de Lubumbashi; UNP, Upemba National Park;
ZMB, Zoologisches Museum Berlin, Berlin, Germany; ZSM,
Zoologisches Staatssammlung München, München, Germany.
3|RESULTS
3.1 |Motivation for the generic transfer of
H. aelsbroecki (Pellegrin 1936) to the genus
Cyphomyrus
Hippopotamyrus aelsbroecki (Poll 1945) was originally described as a
Gnathonemus species based on a single specimen, the holotype, from
Lubumbashi, most probably the Lubumbashi River, which is a left bank
affluent of the Kafubu River, itself a left bank affluent of the Luapula
River. In his original description, Poll (1945) reports that G. aelsbroecki
belongs to the small group of Gnathonemus species with a dorsal fin
that is much longer than the anal one and clearly originates in front of
the insertion level of the latter. In his osteological and morphological
work on the genera Gnathonemus,Marcusenius,Hippopotamyrus and
Cyphomyrus, Taverne (1971) reassigned G. aelsbroecki to the genus
Hippopotamyrus Pappenheim 1906. In the same work, Taverne (1971)
also synonymised the genus Cyphomyrus with Hippopotamyrus,this
based on some osteological evidence only: (a) the shared presence of a
well-developed lateral ethmoid; (b) five circumorbital bones with the
antorbital and first infraorbital fused; (c) the same shape and arrangement
of the bones of the snout and lower jaw; and (d) the presence of five
hypural bones in the caudal skeleton. However, as G. aelsbroecki was
described on a single specimen, the holotype, and no other specimens
had been collected since, its attribution to the genus Hippopotamyrus
must have been based on external morphological evidence only. Later
on, the genus Cyphomyrus was rehabilitated by Van der Bank and Kramer
(1996) based on: (a) external morphological evidence; (b) the waveforms
of the EOD; and (c) an allozyme study on C. discorhynchus. Subsequently,
a mitochondrial DNA sequence analysis using cytochrome b was also
performed which demonstrated that the genus Cyphomyrus, including its
type species C. psittacus, is different from the sympatric Hippopotamyrus
ansorgii species complex (Kramer et al., 2004; Kramer & Swartz, 2010;
Van der Bank & Kramer, 1996). However, the generic assignment of this
species complex remains problematic and further combined mitochon-
drial and nuclear molecular evidence revealed that it requires classifica-
tion into a different genus (see Sullivan et al., 2016: Figure 5).
After morphologically examining the holotype of H. aelsbroecki
(Figure 1a,b) and considering the diagnostic characters presented by
Poll (1945), Myers (1960) and Kramer and Swartz (2010) for the genus
Cyphomyrus,H. aelsbroecki corresponds well to the diagnosis of this
genus. So, certainly the long dorsal fin, with its origin well ahead of
that of the anal fin and its posterior end just posterior to that of the
anal fin [vs. anterior and posterior end of both dorsal and anal fin, at
the same vertical level (as observed in H. castor, type species of the
genus Hippopotamyrus) or the origin of the dorsal fin even a little
behind that of the anal fin as in some other species of the genus
Hippopotamyrus] distinguishes Cyphomyrus from Hippopotamyrus
(Hopkins et al., 2007; Myers, 1960). Additional characters are the
rounded or arched predorsal profile and the deep and compact body
of Cyphomyrus (Kramer & Van der Bank, 2011; Myers, 1960) [vs.a
straight or convex predorsal profile but not vaulted with a rather elon-
gated body of moderate depth for the genus Hippopotamyrus
(Hopkins et al., 2007; Myers, 1960)]. As such, Hippopotamyrus
aelsbroecki is here transferred to the genus Cyphomyrus as it conforms
with the diagnostic characters of the genus as provided above follow-
ing Myers (1960) and Kramer and Van der Bank (2011).
3.2 |Analysis of the meristics
A first PCA was carried out on 10 meristics for all examined speci-
mens (n= 90) (Figure 2). The most important loadings on PCI were for
the number of circumpenduncular scales, the number of pectoral-fin
rays, the number of dorsal-fin rays and the number of scales between
the dorsal and anal fins. The highest loadings on PCII were for the
number of scales between the pelvic fin and the lateral line and the
number of anal-fin rays (Table 1a).
Four groups can be distinguished on this PCA (Figure 2). First, the
three groups on the left-hand side and mainly situated on the negative
side of PCI contain (a) the lectotype and the four paralectotypes of
C. discorhynchus, and the specimens of Lake Kariba (Zambezi basin)
and the Upper Lualaba (Congo basin) all also attributed to this species,
these mainly situated on the negative side of PCII, (b) the three exam-
ined syntypes of C. tanganicanus, a current junior synonym of
C. discorhynchus, as well as some additional specimens from the Lake
Tanganyika affluents, these also mostly situated on the negative side
of PCII, and (c) the four examined syntypes of C. cubangoensis, these
entirely situated on the positive side of PCII. Second, the fourth group,
that is the only one situated on the right-hand side and almost entirely
on the positive side of PCI, contains the specimens from the Middle
and Lower Lufira here recognized as a new species for science and
named C. lufirae. In addition, the holotype of C. aelsbroecki is located
near to the 0 point of PCI and slightly on the positive side of PCII.
Mann–Whitney U (MWU) tests, with sequential Bonferroni correc-
tion, were performed on all meristics included in the PCA (Table 1b).
Between C. lufirae and C. discorhynchus, eight out of the 10 meristics
were found to be highly significant (P≤0.001 after Bonferroni correc-
tion). Furthermore, between C. lufirae and C. tanganicanus,acurrent
junior synonym of C. discorhynchus, five out of the 10 meristics were
4MUKWEZE MULELENU ET AL.
FISH

found to be significantly different, three of which were highly signifi-
cantly different. In addition, between C. lufirae and C. cubangoensis,four
meristics were found to be significantly different, of which a single one
was highly significantly different.
After the MWU tests, meristic variables whose differences were
significantly or highly significantly different were also evaluated using
scatterplots to identify those that could be used as good diagnostic
characters for species identification. Four meristic characters with
slightly overlapping values separate C. lufirae from C. discorhynchus
(Figure 3a–d), and three characters with slightly overlapping values
separate C. lufirae from C. cubangoensis (Figure 3a–c). In addition,
specimens of C. lufirae have a higher number of scales between the
anterior base of the dorsal fin and the anterior base of the anal fin
compared to C. aelsbroecki (Figure 3d).
3.3 |Analysis of the measurements
A second PCA was carried out on 28 measurements in percentages
(n= 90) (Figure 4). The most important loadings on PCI were for the
prepelvic length, the prepectoral length, the length of the head, the dis-
tance between the pelvic fin and the anal fin, and the depth of the cau-
dal peduncle. The highest loadings on PCII were for the distance
between the pectoral and anal fins, the body depth at the dorsal fin
insertion and depth of the head (Table 2a).
Two major groups can be distinguished on this PCA (Figure 4).
First, a major group situated mainly on the positive side of PCI con-
tains: (a) the lectotype and the four paralectotypes of C. discorhynchus
and the specimens from Lake Kariba (Zambezi basin) and the Upper
Lualaba (Congo basin) all also attributed to this species and all mainly
situated on the negative side of PCII; (b) the three studied syntypes of
C. tanganicanus, a current junior synonym of C. discorhynchus, and some
additional specimens from the Lake Tanganyika basin affluents; (c) the
syntypes of C. cubangoensis; and (d) the holotype of C. aelsbroecki all
situated mainly or entirely on the positive side of PCII. Second, a major
(a)
1 cm
(b)
FIGURE 1 Cyphomyrus
aelsbroecki, MRAC 54990, holotype,
female, 71.4 mm L
S
, DRC:
Lubumbashi (former Elisabethville):
(a) drawing of the holotype (Poll,
1945: Figure 4) and (b) photograph of
the preserved holotype
–3
–2 –1 0
PCI
12
–2
–1
0
PCII
1
2
3
4
FIGURE 2 Scatterplot of PCI against PCII for a PCA on 10 meristics
for all examined specimens (n= 90). Cyphomyrus aelsbroecki:, holotype;
C. cubangoensis:,syntypes;C. discorhynchus:, lectotype; ,
paralectotypes; , specimens from the Kamalondo Depression; C. lufirae:
,holotype; , other specimens; C. tanganicanus (currently a junior
synonym of C. discorhynchus): ,syntypes; , other specimens from
some affluents of the Lake Tanganyika basin
MUKWEZE MULELENU ET AL.5
FISH

group situated on the negative part of the PCI only contains the speci-
mens of the new species for science, C. lufirae.
MWU tests, with sequential Bonferroni correction, were per-
formed for all measurements included in the PCA (Table 2b). Between
C. lufirae and C. discorhynchus, 12 out of the 29 measurements were
found to be significantly different (P≤0.05 after Bonferroni correc-
tion), 10 of which were highly significantly different (P≤0.001). Fur-
thermore, between C. lufirae and C. tanganicanus, a current junior
synonym of C. discorhynchus, 11 out of the 29 measurements were
found to be significant different, seven of which were highly signifi-
cantly different. Between C. lufirae and C. cubangoensis, two out of
the 29 measurements were found to be significantly different.
After the MWU tests, variables whose differences are signifi-
cantly or highly significantly different were also evaluated using sca-
tterplots to identify those that could be used as diagnostic characters
for species identification. The differentiating characters between
C. lufirae and C. cubangoensis and C.discorhynchus, respectively, are
the caudal peduncle depth, 6.9–8.4% L
S
(vs. 6.1–6.7% and 5.7–7.0%
L
S
, respectively) and the prepelvic length, 41.0–43.8% L
S
(vs.
38.9–39.1% and 37.0–41.0% L
S
, respectively) (Figure 5).
3.4 |Molecular data: DNA barcoding
On the ML tree based on COI (mtDNA), the 12 sequences of C. lufirae
constitute a different clade, from a genetic distance of 1.2% with
C. discorhynchus (Figure 6).
3.5 |Cyphomyrus lufirae sp. nov.
Figures 7a,b and 8a, and Table 3.
3.5.1 |Holotype
MRAC 2015-005-P-145, female, 112.2 mm L
S
; DRC: UNP, Lualaba
Province, Dikulwe River (Bunkeya-Kyubo), Middle Lufira River,
Upper Lualaba (0957018.600S; 02656047.100E); Alt. 888 m above sea
level (a.s.l)., Upemba National Park Expedition 2014, 23 October
2014.
3.5.2 |Paratypes
MRAC 2015-005-P-146-148, 115.0–126.6 mm L
S
; same data as
holotype. MRAC 2015-005-P-149-150, 103.8–113.6 mm L
S
; DRC:
UNP, Lualaba Province, Dikulwe River (Kijiba Kikwepe, village Kalwa),
Middle Lufira River, Upper Lualaba (0957018.600S; 02656047.100E);
Alt. 888 m a.s.l., Upemba National Park Expedition 2014, 25 October
2014. MRAC 2015-005-P-151, 105.8 mm L
S
; same data as holotype;
22 October 2014. MRAC 2015-005-P-155, 108.6 mm L
S
; same data
as holotype; 21 October 2014. AMNH 275001 (formerly MRAC
2015-005-P-152), 92.6 mm L
S
; same data as MRAC 2015-005-P-
151. BMNH 2019.9.23.1 (formerly MRAC 2015-005-P-153),
94.5 mm L
S
; same data as MRAC 2015-005-P-151. ZSM 47511 (for-
merly MRAC 2015-005-P-154), 90.3 mm L
S
; same data as MRAC
2015-005-P-151.
3.5.3 |Additional non-type material examined
MRAC 2015-005-P-0196-0197, 93.8–103.0 mm L
S
; same locality as
holotype; Coll. C. Mukweze, B. Katemo & J. S. Kiwele, 22 October 2014.
MRAC 2015-006-P-0613, 116.1 mm L
S
; DRC: KNP, Haut-Katanga Prov-
ince, Kafila River (village Kienge), Middle Lufira River, right affluent of the
TABLE 1 (a) PC loadings for the first three PCs of the PCA performed on 10 meristics for all examined specimens (n= 90)
Variable
(a) PC loadings (b) MWU tests
PCI PCII PCIII
C. lufirae vs.
C.discorhynchus
C.lufirae vs.
C. tanganicanus
C.lufirae vs.
C.cubangoensis
Dorsal-fin rays −0.788139 −0.414011 −0.278887 ** ** NS
Anal-fin rays −0.638195 −0.574585 −0.378570 ** * NS
Pectoral-fin rays −0.863405 0.109295 −0.270942 ** NS **
Scales on lateral line −0.650228 −0.394094 0.220513 ** ** NS
Circumpeduncular scales 0.896880 0.051097 −0.012001 ** ** *
Scales between dorsal and anal fins 0.775920 −0.542607 −0.003062 ** NS *
Scales between dorsal fin and lateral line 0.628621 −0.514933 0.081631 ** NS *
Scales between pelvic fin and lateral line 0.446454 −0.628891 0.010822 NS NS NS
Teeth in upper jaw −0.634800 −0.027332 0.360443 ** NS NS
Teeth in lower jaw −0.497661 −0.215868 0.735743 NS *NS
Note (a).Cyphomyrus aelsbroecki: holotype (n= 1); C. cubangoensis: syntypes (n= 4); C. discorhynchus: lectotype (n= 1), paralectotypes (n= 4), and additional
specimens (n= 25); C. lufirae: holotype (n= 1), paratypes (n= 10) and additional specimens (n= 34); C. tanganicanus (currently a junior synonym of
C. discorhynchus): syntypes (n= 3) and additional specimens from some affluent rivers of the Lake Tanganyika basin (n= 7). Bold values indicate the most
important variables. (b) Results of Mann–Whitney U-tests with sequential Bonferroni correction for the 10 meristics. Note (b). NS, not significant;
*significant, α= 0.05; **highly significant, α= 0.001.
6MUKWEZE MULELENU ET AL.
FISH

Upper Lualaba (10330200S; 27290200E); Alt. 898 m a.s.l.; Coll. E. Abwe,
G. Kapepula & C. Kalombo, 20 August 2014. MRAC 2015-006-P-
0715-0176, 113.6–120.2 mm L
S
; same data as MRAC 2015-006-P-0613.
MRAC 2015-006-P-0717, 118.1 mm L
S
; DRC: KNP, Haut-Katanga Prov-
ince, Lufira River (village Kapiri), Middle Lufira River, right affluent of the
Upper-Lualaba (94104700S; 271302800E); 872 m a.s.l.; Coll. E. Abwe,
G. Kapepula & C. Kalombo, 19 October 2014. MRAC 2016-002-P-
0240-0244, 85.7–137.5 mm L
S
; DRC: KNP, Haut-Katanga Province, Kafila
River (Kivuko, Mwadingusha-Kienge road), Middle Lufira River, right afflu-
ent of the Upper Lualaba (103504500S; 273102600E); 875 m a.s.l.; Coll.
E. Abwe, G. Kapepula & C. Kalombo, 20 October 2015. MRAC
2016-002-P-0316, 133.3 mm L
S
; DRC: KNP, Haut-Katanga Province,
Luvua River (road bridge Sampwe-Mukana), Middle Lufira River, right
affluent of the Upper Lualaba (91902200S; 272201700E); 911 m a.s.l.;
Coll. E. Abwe, G. Kapepula & C. Kalombo, 16 August 2015. MRAC
2015-005-P-0193-0195, 100.4–124.5 mm L
S
;DRC:UNP,Haut-
Katanga Province, Lufira River (approximate downstream of Kyubo
Falls), Middle Lufira River, right affluent of the Upper Lualaba (9310300S;
27201000E); 835 m a.s.l.; Coll. C. Mukweze, B. Katemo & S. Kiwele,
18 October 2014. MRAC 2016-025-P-0237, 136.5 mm L
S
;samelocal-
ity as MRAC 2015-05-P-0193-0195; Coll. E. Vreven, B. Katemo,
M. Kasongo, J. Mulagizi & S. Kiwele, 14 August 2016. MRAC
2015-005-P-0190-0192, 72.9–85.8 mm L
S
; DRC: UNP, Haut-Katanga
Province, Luvilombo River (downstream tributary left of Lower Lufira
basin after Luvilombo's small Falls), Lower Lufira River, right affluent of
the Upper Lualaba (9310600S; 2720700E); 858 m a.s.l.; Coll. C. Mukweze,
B. Katemo & S. Kiwele, 18 October 2014. MRAC 2012-031-P-
2165-2176, 85.1–121.7 mm L
S
; DRC: KNP, Haut-Katanga Province,
Lufira River (upstream of KyuboFalls), Middle Lufira River, right affluent
of the Upper Lualaba (9310000S; 27205100E); 871 m a.s.l.; Coll.
E. Vreven, A. Chocha, M. Katemo, E. Abwe & M. Kasongo,16 September
2012. MRAC 2016–038-P-0017-0023, 113.2–125.5 mm L
S
; DRC:
11
60 10080 140120
Standard length (mm)
160 180 200
12
13
Circumpeduncular scales
14
15
16
17
(a)
60 10080 140120
Standard length (mm)
160 180 200
10
Pectoral-fin rays
11
(b)
26
60 10080 140120
Standard length (mm)
160 180 200
28
30
Dorsal-fin rays
32
34
36
38
(c)
24
60 100
80 140
120
Standard length (mm)
160 180 200
26
28
Scales between dorsal and anal fins
30
32
34
(d)
FIGURE 3 Scatterplots of the number of (a) circumpeduncular scales, (b) pectoral-fin rays, (c) dorsal-fin rays and (d) scales between dorsal and
anal fins against standard length (in mm) (n= 90). Cyphomyrus aelsbroecki:, holotype; C. cubangoensis: , syntypes; C. discorhynchus: , lectotype;
, paralectotypes; , specimens of the Kamalondo Depression; C. lufirae: , holotype; , other specimens; C. tanganicanus (currently a junior
synonym of C. discorhynchus): , syntypes; , other specimens from some affluents of the Tanganyika basin
MUKWEZE MULELENU ET AL.7
FISH

KNP, Haut-Katanga Province, Kijiba Mujingi, Kambokoto and Lubumba-
shi Luiji River (Nkumbula village), Middle Lufira River, right affluent of
the Upper Lualaba (92104900S; 272704400E); 939 m a.s.l.; C. Mukweze,
E. Abwe, P. Kiwele, L. Ngoy & J. Mulagizi, 23 October 2016. MRAC
2015-006-P-0610-0612, 80.0–91.0 mm L
S
; DRC: KNP, Haut-Katanga
Province, Lake Mpungwe (Coopelu village near Kinyonge River),
Middle Lufira River, right affluent of the Upper Lualaba (10007.00S;
271503200E); 897 m a.s.l.; Coll. E. Abwe, G. Kapepula & C. Kalombo,
25 October 2014.
3.5.4 |Diagnosis
Cyphomyrus lufirae (Figure 8a) is distinguished from all its conge-
ners, that is, C. aelsbroecki (Figure 1a,b), C. cubangoensis
(Figure 8b), C. discorhynchus (Figure 8c,d), C. macrops,C. psittacus
and C. wilverthi, by a larger caudal peduncle depth, 6.8–8.4% L
S
(vs.
smaller, 5.3%, 6.1–6.7%, 5.7–7.0%, 5.2%, 5.8% and 6.2% L
S
,
respectively), and a higher number of circumpeduncular scales,
14–16, rarely 12–13 vs. exclusively 12 in all congeners except for
C. discorhynchus, which rarely also possesses 13 circumpeduncular
scales.
In addition, C. lufirae differs from C. macrops,C. psittacus and
C. wilverthi by a lower number of dorsal fin rays, 27–32 (vs.higher,
36 (37), 34 (33–41) and 38 (38–40), respectively) and from C. aelsbroecki,
C. cubangoensis and C. discorhynchus by a longer prepelvic distance,
41.0–43.8% L
S
(vs. shorter, 39.7%, 38.9–39.1%, and 37.0–41.0% L
S
,
respectively).
3.5.5 |Description
Based on the holotype, 10 paratypes and 34 additional specimens.
Proportional measurements and meristics are given in Table 3 and
general appearance is shown in Figure 8. The maximum recorded
size of this species was 137.5 mm L
S
.Deepbody,2.8–3.5 times in
L
S
, laterally compressed. Predorsal profile convex. Head broadly
rounded, 3.4–3.9 times in L
S
. Snout rounded and projecting beyond
mouth, 4.1–5.2 times in L
H
. Mouth small and inferior. Chin with a
short bulbous protuberance, nostrils closer to eye than mouth.
Interorbital space 3.0–4.2 times in L
H
.Dorsalandanalfinsobliquely
orientated. Dorsal fin length equal or slightly longer than head, its
origin situated anterior to anal-fin origin, and its base longer than
that of the anal fin. Distance between pelvic and anal fins relatively
equidistant to pectoral-fin length. Distal end of pectoral fin pointed,
shorter than head length. Caudal peduncle 2.4–3.2 times as long
as deep.
3.5.6 |Colouration in life
Overall body coloration of C. lufirae dull silver, tinged with black and
brown and yellowish overtone especially on head. Yellowish colora-
tion more intense in specimens from downstream Kyubo Falls and
from Luvilombo River. Fins greyish to black yellowish at their base
(Figure 8a). The silver colouration in particular is sometimes less
intense or translucent for pelvic and caudal fins. With an oblique
black band from anterior base of dorsal-fin up to third or fourth
scale row below the lateral line. The oblique black band is less con-
spicuous in darker coloured specimens, but is more discernible in
the yellowish coloured specimens of Lower Lufira, and Luvilombo
River.
3.5.7 |Colouration in alcohol
Specimens from the Middle Lufira chocolate to blackish whereas
those from Lower Lufira yellowish or brown. Oblique black band often
becoming more distinguishable than in life. Fins greyish to black or
translucent.
3.5.8 |Distribution, habitat diversity and ecology
Cyphomyrus lufirae is known from the Middle Lufira (Figures 9a,b and
10) and from the Lower Lufira, from the foot of the Kyubo Falls
(Figure 9c) and from its nearby left bank affluent the Lower Luvilombo
River (Figure 9d). The species is currently thus considered to be
endemic to the Middle and Lower Lufira basin.
The species was recorded in river sections with moderate current
at all localities on the Lower and Middle Lufira. The depth of the water
was variable, about 50 cm in the Luvilombo River, about 5 m in the
–3
–3 –2 –1 0
PCI
13
2
–2
–1
0
PCII
1
2
3
FIGURE 4 Scatterplot of PCI against PCII for a PCA on
measurements (in %) for all examined specimens (n= 90).
Cyphomyrus aelsbroecki:, holotype; C. cubangoensis: , syntypes;
C. discorhynchus:, lectotype; , paralectotypes of Lower Zambezi
basin; , specimens of Kamalondo Depression; C. lufirae: ,
holotype; , other specimens; C. tanganicanus (currently a junior
synonym of C. discorhynchus): , syntypes; , other specimens from
some affluents of the Tanganyika basin
8MUKWEZE MULELENU ET AL.
FISH

pool on the Dikulwe River, and about 10 m just downstream of the
Kyubo Falls on the Lower Lufira River.
The following physico-chemical parameters of the water were
recorded in the Lower and Middle Lufira on 19–23 October 2014
and 11–18 August 2016 during the morning, between 8 and
10 a.m.: temperature 23.6–26.6C; pH 7.9–8.3 except for the type
locality where it is acidic, 5.3–6.2; dissolved oxygen 5.82–
7.19 mg l
−1
(73.1–94.1%) and conductivity 8.0–468.4 μScm
−1
but
reaching more than double the type locality or 778.9 and
913.2 μScm
−1
.
3.5.9 |Etymology
The specific name, lufirae, is a Latin adjective referring to the Lufira
River to which the new species seems endemic.
3.5.10 |Local knowledge
Cyphomyrus lufirae is called ‘sengwa’in Sanga (a Bantu language),
which refers to the general morphology of the mormyrids of this
TABLE 2 (a) PC loadings for the first two PCs of the PCA performed on 29 measurements for all examined specimens (n= 90), (b) Results of
Mann–Whitney U-tests with sequential Bonferroni correction for the 29 measurements
Variables transformed in percentages
(a) PC loadings (b) MWU tests
PCI PCII
C.lufirae vs.
C.discorhynchus
C.lufirae vs.
C.tanganicanus
C.lufirae vs.
C.cubangoensis
Standard length ––NS NS NS
Body depth −0.63247 −0.56270 NS ** NS
Body depth at the insertion of the dorsal fin −0.44469 −0.69368 NS *NS
Caudal peduncle depth −0.70181 0.16343 ** ** NS
Caudal peduncle length −0.20532 0.56598 *NS NS
Predorsal distance −0.149171 −0.47962 NS NS NS
Post-dorsal distance −0.08316 −0.03777 NS NS NS
Preanal distance −0.54930 −0.53307 NS ** NS
Prepelvic length −0.91935 0.03779 ** ** *
Prepectoral length −0.84823 0.28055 ** ** NS
Dorsal-fin length 0.27961 −0.59114 ** NS NS
Anal-fin length 0.51849 −0.17853 *NS NS
Pelvic-fin length −0.14617 −0.14964 NS NS NS
Pectoral-fin length −0.05223 −0.06372 NS NS NS
Distance between pectoral and pelvic fin −0.73046 −0.39380 ** ** *
Distance between pelvic and anal fin 0.46313 −0.68396 ** NS NS
Distance between pectoral and anal fin 0.10584 −0.72570 NS NS NS
Head length −0.74601 0.23323 ** * NS
Head depth −0.23301 −0.64009 NS NS NS
Head width 0.00807 −0.39557 NS NS NS
Interorbital width −0.40923 0.12271 ** NS NS
Eye diameter 0.07407 −0.15632 NS NS NS
Post-orbital length 0.32712 0.00471 NS NS NS
Snout length 1 (SNL) −0.21003 −0.21622 NS NS NS
Length of snout 2 (LSo) −0.14582 −0.16867 NS NS NS
Length of snout 3 (LSc) −0.27010 −0.11845 NS NS NS
Distance between nostrils 0.08866 0.33204 NS *NS
Distance between nostril and eye −0.61675 −0.01456 ** ** NS
Length of the gill opening 0.59918 −0.40354 ** * NS
Note (a).C. aelsbroecki: (holotype) (n= 1); C. cubangoensis: syntypes (n= 4); C. discorhynchus: lectotype (n= 1), paralectotypes (n= 4), and additional
specimens (n= 25); C. lufirae: holotype (n= 1), paratypes (n= 10) and additional specimens (n= 34); C. tanganicanus (currently a junior synonym of
C. discorhynchus): syntypes (n= 3), and additional specimens from some affluent rivers of the Lake Tanganyika basin (n= 7). Bold values indicate the most
important variables. Note (b). NS, not significant; *, significant, α= 0.05; **, highly significant, α= 0.001.
MUKWEZE MULELENU ET AL.9
FISH

5
60 10080 140120
Standard length (mm)
160 200180 220
6
Caudal peduncle depth (% LS)
7
8
9
(a)
36
60 10080 140120
Standard length (mm)
160 200180 220
38
39
37
Prepelvic length (% LS)
41
40
43
44
42
45
(b)
FIGURE 5 Scatterplots of (a) caudal peduncle depth (% L
S
) and (b) prepelvic length (% L
S
) against standard length (in mm) (n= 90).
Cyphomyrus aelsbroecki:, holotype; C. cubangoensis: , syntypes; C. discorhynchus: , lectotype; , paralectotypes of the Lower Zambezi basin; ,
specimens from the Kamalondo Depression; C. lufirae:, holotype; , other specimens; C. tanganicanus (currently a junior synonym of
C. discorhynchus): , syntypes; , other specimens from some affluents of the Tanganyika basin
Imbiri River KT193209 Genyomyrus donnyi
Kalule Nord River MN207892 Cyphomyrus discorhynchus
Mwanza River MN207888 Cyphomyrus discorhynchus
Kamalondo Depression MN207906 Cyphomyrus discorhynchus
Mwanza River MN207896 Cyphomyrus discorhynchus
Kamalondo Depression MN207893 Cyphomyrus discorhynchus
Kamalondo Depression MN207894 Cyphomyrus discorhynchus
Kamalondo Depression MN207905 Cyphomyrus discorhynchus
Kalule Nord River MN207895 Cyphomyrus discorhynchus
Kamalondo Depression MN207907 Cyphomyrus discorhynchus
Luapula River MN207904 Cyphomyrus discorhynchus
Lufira basin MN207902 Cyphomyrus lufirae
Lufira basin MN207901 Cyphomyrus lufirae
Lufira basin MN207899 Cyphomyrus lufirae
Lufira basin MN207887 Cyphomyrus lufirae
Lufira basin MN207886 Cyphomyrus lufirae
Lufira basin MN207889 Cyphomyrus lufirae
Lufira basin MN207890 Cyphomyrus lufirae (paragenetype)
Lufira basin MN207891 Cyphomyrus lufirae
Lufira basin MN207903 Cyphomyrus lufirae
Lufira basin MN207897 Cyphomyrus lufirae
Lufira basin MN207900 Cyphomyrus lufirae
Lufira basin MN207898 Cyphomyrus lufirae
100
0.0100
43
52
91
34
94
Imbiri River KT193208 Genyomyrus donnyi
FIGURE 6 Maximum likelihood
tree based on the mitochondrial
cytochrome coxidase subunit I (COI)
sequences. See text for details on the
tree reconstruction method and
Supporting Information Table S1 for
specimen voucher data and Genbank
accession numbers. Statistical node
support (500 bootstrap replications) is
visualized at each node. The clade of
the new species C. lufirae is
highlighted in grey. The scale bar
refers to the branch lengths,
measured in the number of
substitutions per site
10 MUKWEZE MULELENU ET AL.
FISH

genus as opposed to those from the genus Hippopotamyrus and occur-
ring in the same river, which are called ‘mbubu’in the same language.
This species is highly appreciated by the local inhabitants due to the
high fat content of its meat. They prefer to eat it smoked.
4|DISCUSSION
Within the Lufira basin, a new Cyphomyrus species, here named
C. lufirae, has been discovered. Among all currently known conge-
nerics, C. lufirae is morphologically most similar to C. cubangoensis (see
Figure 8b) and C. discorhynchus (see Figure 8c,d). However, it differs
from both by a deeper caudal peduncle (Figure 5a) and a larger
prepelvic distance (Figure 5b). Further, C. lufirae is the only species of
Cyphomyrus known with generally 14–16 circumpeduncular scales,
very rarely only 12–13, one and two individual(s) respectively [vs.
12 for all other known Cyphomyrus species, although rarely 13 in
some specimens of C. discorhynchus (Kramer & Van der Bank, 2011)].
Finally, C. lufirae usually also has fewer scales on the lateral line,
60–66 [vs.63–73 for C. discorhynchus, with 63–73 scales for speci-
mens of the Upper Lualaba, 67–72 for those of Lake Kariba, 66–70
for those of the Lower Zambezi and 66–70 for those of the Tangan-
yika basin (C. tanganicanus: at present a junior synonym of
C. discorhynchus)]. While C. cubangoensis is not known from the Congo
basin and only reported from the Okavango and Upper Zambezian
basins (Skelton, 2001; Skelton, 2019), C. discorhynchus is considered
to have a wide geographical distribution covering the Zambezi and
part of the Congo basin sensu lato (Skelton, 2001; Scott et al., 2006).
Also within the Congo basin, C. discorhynchus has a wide distribution
range as it has been reported from the Lake Tanganyika basin
(Worthington & Ricardo, 1936) up to the Upper Lualaba, that is, the
Kamalondo Depression and the Lower Lufira, as well as the Upper
Kasai, at Dilolo (±10410S; 22200E) (Poll, 1976). This widespread dis-
tribution is partially the result of the synonymisation of
C. tanganicanus, originally described from Lake Tanganyika
(Boulenger, 1906), with C. discorhynchus by Boulenger (1909) in his
‘Catalog of the Fresh-water fishes of Africa’. Unfortunately he did not
provide any justification for this decision. However, examination of
the three syntypes of C. tanganicanus and seven additional specimens
from some Lake Tanganyika tributaries revealed meristic (Figure 2)
and morphometric (Figure 4) differences with C. discorhynchus. The
separation of C. discorhynchus and C. tanganicanus on the PCA
scatterplot for the meristic data is mainly attributed to differences in
the number of pectoral fin rays, as specimens of the former species
collected from the Zambezi and Upper Lualaba always had 11 rays,
whereas specimens of the latter nominal species collected from the
Tanganyika basin always had 10 rays (with the exception of one speci-
men that had 11). These specimens also showed some marginal differ-
ences in body depth and preanal distance (see Figure 11a,b). These
preliminary observations could either be interpreted as representing
intraspecific variation within C. discorhynchus, or an indication that
C. tanganicanus could be a valid species. We have, however, refrained
(a)
1 cm
2 cm
(b)
FIGURE 7 Cyphomyrus lufirae,
holotype, MRAC 2015-05-P-145,
female, 112.2 mm L
S
, DRC: Dikulwe
River (bridge Bunkeya-Kyubo), Middle
Lufira River: (a) drawing of lateral
view and (b) photograph of the
preserved specimen
MUKWEZE MULELENU ET AL.11
FISH

from making any taxonomic decisions for C. discorhynchus and its
junior synonym, C. tanganicanus, pending further examination of a
more representative sample size as well as generation of additional
data including genetics and/or EODs.
The low genetic divergence in COI sequences of C. lufirae and the
other congeners considered in the present study is consistent with pat-
terns recorded for other morphologically divergent mormyrid species.
For example, Kramer et al. (2004) recorded mtDNA cyt b sequence
divergence values of 0.6–1.7% between H. szaboi and H. ansorgii,
Kramer and Wink (2013) reported divergences of 0.4–1.6% between
Marcusenius altisambesi and M. multiquamatus,whileKrameret al.
(2013) reported a divergence value of 1.2% between Pollimyrus mar-
ianne and P. cuandoensis. More recently, Sullivan et al. (2016) presented
some species pairs within the genera Campylomormyrus Bleeker, 1974,
Marcusenius Gill, 1862, Petrocephalus Marcusen, 1854, and Cyphomyrus,
which had COI sequence divergences less than 1%.
(a)
(b)
1 cm
2 cm
(c)
(d)
FIGURE 8 Photographs of: (a) life
specimen of C. lufirae, MRAC
2015-006-P-0613, female, 116.1 mm
L
S
, DRC: Kafila River (village Kienge),
Middle Lufira River, right affluent of
the Upper-Lualaba. Photo: Expedition
KNP 2015; (b) preserved specimen of
C. cubangoensis (syntype), MRAC
138760 (ex. MNHN 1936–65),
83.2 mm L
S
; Angola: Cubango
(Okavango) basin; (c) life specimen of
C. discorhynchus, MRAC 2016-03-P-
0148, male, mm L
S
, DRC: Haut
Lomami Province: UNP: Lake
Lukanga, Kamalondo Depression.
Photo: Expedition UNP 2015; and
(d) preserved specimen
of C. discorhynchus (lectotype), ZMB
3674, female, 163.7 mm L
S
, Lower
Zambezi
12 MUKWEZE MULELENU ET AL.
FISH

TABLE 3 Measurements and counts for C. aelsbroecki,C.cubangoensis,C. discorhynchus (lectotype + paralectotype and other specimens), C. tanganicanus (synonym of C. discorhynchus) and
C. lufirae
C. aelsbroecki C. cubangoensis C. discorhynchus C. tanganicanus (synonym of C. discorhynchus)C. lufirae
s(n=4)
l
pl (n= 4) All specimens (n= 25) s (n= 3) All specimens (n= 10) All specimens (n= 45)
h Mean ± SD Range 163.4 Mean ± SD Range Mean ± SD Range Mean ± SD Range Mean ± SD Range h Mean ± SD Range
Standard length (mm) 71.4 82.9 ± 0.3.3 79.1–87.2 163.7 170.4 ± 30.9 132.8–205.8 122.3 96.5–183.6 133.3 ± 12.0 125.2–147.1 116.1 ± 16.8 85.7–147.1 112.2 107.7 ± 16.2 72.9–137.5
Percentage of standard length
Body depth 26.3 27.7 ± 1.8 25.9–30.2 33.2 29.9 ± 2.2 28.1–33.0 31.3 ± 1.5 28.7–33.7 27.0 ± 0.9 26.8–28.1 26.9 ± 1.5 24.6–29.6 32.5 31.7 ± 1.9 28.4–36.9
Body depth at the
insertion of the
dorsal fin
26.5 27.6 ± 1.2 26.9–29.5 32.5 31.2 ± 2.1 29.3–34.2 31.2 ± 1.6 28.8–35.0 29.6 ± 1.0 28.5–30.4 28.5 ± 1.5 25.3–30.4 31.5 31.0 ± 1.5 28.3–34.8
Caudal peduncle depth 5.3 6.3 ± 0.2 6.1–6.7 6.6 6.8 ± 0.1 6.6–6.9 6.3 ± 0.3 5.7–7.0 6.9 ± 0.0 6.8–7.0 6.9 ± 0.2 6.3–7.1 7.6 7.4 ± 0.4 6.9–8.4
Caudal peduncle
length
21.4 23.1 ± 0.7 22.3–24.2 20.2 20.8 ± 0.4 20.5–21.4 20.8 ± 0.9 18.7–22.3 22.1 ± 0.9 21.1–23.0 21.8 ± 0.5 21.1–23.0 21.9 21.8 ± 1.0 19.8–24.0
Predorsal distance 57.1 57.9 ± 0.8 56.7–58.6 60.6 58.8 ± 1.2 58.0–60.7 58.5 ± 1.1 56.9–61.6 59.0 ± 1.3 57.9–60.5 57.7 ± 1.5 55.3–60.5 57.9 58.1 ± 1.1 55.6–60.5
Post-dorsal distance 46.1 46.8 ± 1.1 45.6–48.0 45.2 47.1 ± 0.6 46.4–47.9 47.6 ± 1.3 44.4–50.3 47.0 ± 1.4 45.3–48.2 47.8 ± 1.5 45.3–50.5 47.8 47.9 ± 1.1 44.6–50.5
Preanal distance 62.1 61.3 ± 0.4 60.8–61.7 62.8 63.0 ± 1.5 61.7–65.1 63.0 ± 1.0 61.2–64.7 60.9 ± 0.2 60.6–61.0 60.8 ± 1.1 58.8–62.7 63.9 63.3 ± 1.2 60.7–66.1
Prepelvic length 39.7 39.0 ± 0.0 38.9–39.1 40.5 39.0 ± 2.0 37.0–40.9 40.0 ± 1.0 37.4–41.4 39.4 ± 1.1 38.2–40.4 39.2 ± 1.4 37.7–42.5 41.7 42.1 ± 0.8 40.2–43.9
Prepectoral length 23.9 23.7 ± 0.4 23.2–24.3 22.9 22.1 ± 0.6 21.5–22.9 23.2 ± 0.6 22.3–24.6 22.8 ± 1.3 21.5–24.1 22.9 ± 0.8 21.5–24.1 24.6 24.9 ± 0.8 22.9–27.2
Dorsal-fin length 27.2 26.7 ± 0.7 26.1–27.7 29.4 30.6 ± 1.0 29.3–31.6 29.9 ± 1.5 27.1–33.3 27.7 ± 0.7 27.2–28.5 28.4 ± 1.2 27.1–30.2 29.3 28.3 ± 1.0 24.8–31.1
Anal-fin length 19.9 18.7 ± 0.9 17.9–20.0 20.7 20.5 ± 0.4 20.2–21.1 19.6 ± 1.0 17.8–22.6 19.7 ± 0.6 19.1–20.3 20.0 ± 1.3 18.5–22.3 19.7 18.9 ± 1.0 17.1–21.3
Pelvic-fin length 12.1 13.1 ± 0.3 12.8–13.5 12.5 12.7 ± 0.3 12.3–12.9 13.0 ± 0.6 12.2–14.4 12.6 ± 0.6 12.0–13.3 13.0 ± 0.6 12.0–14.0 12.4 12.9 ± 0.6 11.1–14.2
Pectoral-fin length 21.3 22.4 ± 0.6 21.5–22.9 20.9 21.0 ± 1.4 19.4–22.4 21.5 ± 1.1 19.5–24.1 21.4 ± 0.4 21.2–21.9 21.5 ± 0.8 20.3–22.9 21.2 21.2 ± 1.1 19.1–23.2
Distance between
pectoral and pelvic
fin
16.0 16.1 ± 0.5 14.5–18.2 18.6 18.6 ± 0.8 17.4–19.3 17.9 ± 0.7 16.6–19.5 17.1 ± 0.4 16.7–17.6 17.2 ± 0.7 15.8–18.4 18.4 19.1 ± 0.7 17.2–20.5
Distance between
Pelvic and anal fin
22.5 21.8 ± 0.5 21.1–22.2 20.9 23.9 ± 0.6 23.2–24.7 23.2 ± 0.9 21.9–25.1 22.7 ± 0.7 22.1–23.6 21.8 ± 1.1 20.4–23.6 19.9 21.5 ± 1.1 19.0–23.5
Distance between
pectoral and anal fin
37.6 37.4 ± 1.0 36.1–38.5 41.0 41.5 ± 1.2 40.8–43.5 39.7 ± 1.0 37.6–41.5 39.6 ± 0.7 38.8–40.4 38.6 ± 1.4 35.5–40.9 36.8 38.9 ± 1.6 35.3–42.4
Head length 25.8 26.2 ± 0.6 25.4–26.9 24.5 23.6 ± 0.9 22.6–24.7 25.9 ± 0.8 24.8–27.9 25.1 ± 1.0 23.9–25.7 25.4 ± 1.0 23.9–27.6 26.5 27.0 ± 0.9 25.3–28.9
Percentage of head length
Head depth 83.0 87.3 ± 2.7 84.9–91.1 102.7 96.9 ± 4.4 91.3–100.7 96.2 ± 5.2 84.8–103.8 90.0 ± 1.8 87.9–91.3 89.1 ± 4.7 81.5–97.0 99.2 94.1 ± 6.1 84.6–114.8
Head width 44.3 46.0 ± 0.9 44.7–46.8 48.2 44.2 ± 1.7 42.3–46.2 47.2 ± 2.0 47.2–50.9 47.0 ± 0.1 46.8–47.2 46.7 ± 1.6 43.1–48.7 44.2 46.0 ± 3.0 38.8–53.2
Interorbital width 28.5 25.3 ± 0.8 24.0–25.9 26.9 26.2 ± 1.3 25.0–27.7 25.6 ± 1.2 22.1–28.0 26.8 ± 0.9 26.0–27.8 26.8 ± 1.1 24.7–29.0 26.9 28.3 ± 2.3 23.2–32.6
Eye diameter 24.1 22.5 ± 0.3 22.3–22.9 22.5 22.2 ± 1.5 20.8–24.3 23.6 ± 1.2 21.6–25.3 24.2 ± 1.0 22.9–24.8 24.0 ± 1.0 22.4–25.5 23.2 23.1 ± 1.4 20.1–26.5
(Continues)
MUKWEZE MULELENU ET AL.13
FISH

TABLE 3 (Continued)
C. aelsbroecki C. cubangoensis C. discorhynchus C. tanganicanus (synonym of C. discorhynchus)C. lufirae
s(n=4)
l
pl (n= 4) All specimens (n= 25) s (n= 3) All specimens (n= 10) All specimens (n= 45)
h Mean ± SD Range 163.4 Mean ± SD Range Mean ± SD Range Mean ± SD Range Mean ± SD Range h Mean ± SD Range
Post-orbital length 57.7 59.4 ± 1.5 58.1–61.3 59.7 58.5 ± 0.8 57.4–59.3 60.0 ± 1.4 56.7–63.3 60.8 ± 3.0 57.4–63.5 60.2 ± 2.8 54.9–63.5 62.1 58.7 ± 1.9 55.1–63.9
Snout length 1 (SNL) 21.3 21.5 ± 0.9 20.5–22.5 23.1 21.3 ± 1.0 20.0–22.3 21.0 ± 1.0 18.9–23.3 20.6 ± 0.5 20.2–21.3 20.6 ± 1.2 18.8–22.7 19.2 21.6 ± 1.3 18.9–24.6
Length of snout
2 (LSo)
39.5 43.5 ± 1.3 41.7–45.0 42.5 41.1 ± 1.6 39.7–42.7 42.6 ± 1.4 40.2–45.5 42.7 ± 1.7 41.7–44.7 42.9 ± 1.3 41.5–44.7 41.6 42.6 ± 1.7 37.4–45.9
Length of snout 3 (LSc) 29.2 31.8 ± 0.4 31.3–32.3 31.8 29.8 ± 2.6 26.0–31.8 30.7 ± 1.3 28.0–32.9 30.1 ± 0.7 29.5–30.9 30.5 ± 0.9 29.5–31.8 29.5 31.2 ± 1.8 25.3–34.3
Distance between
nostrils
3.3 3.8 ± 0.2 3.5–4.1 4.0 3.4 ± 0.2 3.2–3.6 3.4 ± 0.3 2.3–4.0 4.1 ± 0.2 3.9–4.3 4.1 ± 0.2 3.5–4.5 3.9 3.7 ± 0.3 2.6–4.3
Distance between
nostril and eye
3.5 5.8 ± 0.4 5.3–6.2 6.3 5.8 ± 0.7 4.9–6.7 5.3 ± 0.7 3.6–6.4 4.4 ± 0.3 4.0–4.7 5.0 ± 0.5 4.0–5.7 5.5 6.3 ± 0.5 5.0–7.3
Length of the gill
opening
33.0 34.7 ± 1.0 33.7–35.9 36.4 38.2 ± 1.5 36.3–39.5 36.8 ± 1.3 34.1–39.4 36.9 ± 1.1 36.1–38.2 36.3 ± 2.4 32.3–40.4 29.3 33.3 ± 2.1 29.2–37.3
Counts Median Range Median Range Median Range Median Range Median Range
Dorsal-fin rays 32 31 31 32 34 31–37 34 31–36 33 31–34 34 31–34 31 30 27–32
Anal-fin rays 26 23 22–23 25 25 24–26 24 23–26 24 23–25 25 23–25 24 23 21–25
Pelvic-fin rays 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6
Pectoral-fin rays 10 11 11 11 11 11 11 11 10 10 11 10–11 10 10 10
Scales on lateral line 65 65 64–68 67 68 66–70 67 63–73 68 66–70 66 66–70 64 64 60–66
Circumpeduncular
scales
12 12 12 12 12 12 12 12 12 12 12 12 16 15 12–16
Scales between dorsal
and anal fins
29 28 25–28 28 29 28–30 29 28–31 30 30–31 29 30–31 31 31 20–34
Scales between dorsal
fin and lateral line
15 15 13–15 15 15 15–16 16 15–17 16 16 15 16–17 16 17 16–19
Scales between pelvic
fin and lateral line
15 15 13–15 16 16 15–16 15 14–17 17 15–17 16 15–17 18 16 15–18
Teeth in upper jaw 3 5 4–55 5 5 5 4–75 5 54–534 2–5
Teeth in lower jaw 6 6 6 6 6 6 6 5–77 7–866–865 3–7
Total number of
vertebrae
41 41 40 39–40 39 39 38–39
Note. h, holotype; l, lectotype; pl., paralectotype; s, syntype; SD, standard deviation. Ranges for all specimens includes the type specimens.
14 MUKWEZE MULELENU ET AL.
FISH

(a) (b)
(c) (d)
FIGURE 9 Habitat diversity for C. lufirae: (a) Dikulwe River upstream of the road Bunkeya-Kyubo, type locality; average water depth with
sections shaped small pools (095703200S; 265603300E), deep water (5 m), 23 October 2014; (b) Dikulwe River, just downstream of the bridge
Bunkeya-Kyubo (095701800S; 265604700E), during fyke installation in shallow water (<0.5 m), 23 October 2014; (c) Lufira River, downstream of
the Kyubo Falls on the Lower Lufira (093100200S; 270200900E), deep water (about 5–10 m), 19 October 2014; and (d) Luvilombo River below its
falls and upstream of its confluence with the Lower Lufira (093005800S; 270200200E), shallow water (<0.5 m), 19 October 2014
25°30’
–11°0’
–10°30’
–10°0’
–9°30’
–9°0’
–8°30’
26°0’ 26°30’ 27°0’ 27°30’ 28°0’ 28°30’
KNP
UNP
Lufira
Kyubo Falls
Fungwe
Kalule Nord
Dikulwe
Kafila
Luiji
Lubudi-Sampwe
hunng area
Lualaba
29°0’
75 km50250
N
FIGURE 10 Distribution of
C. lufirae in the Lufira basin, the
major right bank affluent of the
Upper Lualaba. UNP and KNP: Core
zone, dark grey area with full
contour lines; annex zone, dotted,
dark grey area with contour in
broken lines; Lubudi-Sampwe
(LS) hunting area, dashed, light grey
area hatched with full contour lines.
C. lufirae:, type locality; , other
localities; C. discorhynchus:,
distribution of the species in the
study area
MUKWEZE MULELENU ET AL.15
FISH

Based on currently available data, C. discorhynchus and C. lufirae
appear to be allopatrically distributed within the Lufira River.
Cyphomyrus discorhynchus has only been recorded from river sections
below the Kamalondo Depression whereas C. lufirae mainly occurs
above the Kyubo Falls. This suggests that the series of rapids and
pools that separate the upper and lower reaches of the Lower Lufira
River could have formed a barrier that prevented the dispersal of
C. discorhynchus into the Upper Lufira. This is consistent with Poll's
(1976) observations of the faunal discontinuities between the
Lualaba-Upemba and the Lufira rivers.
Preliminary findings from ongoing studies suggest that the Mid-
dle Lufira contains some species within the genera Hippopotamyrus,
Marcusenius and Pollimyrus that could also be endemic to this region
(CMM personal observation, 2016), which highlights the conservation
importance of this region. Currently, C. lufirae is considered endemic
to the Middle Lufira and the upper part of the Lower Lufira basin.
Unfortunately, application of the existing fish protection regulations
is largely deficient in both the UNP and KNP. As a result, anthropo-
genic threats to their fish diversity are increasing (Abell et al., 2008;
Thieme et al., 2005), and this especially since their large mammal
fauna has already been largely decimated (Hasson, 2015; Malaisse,
1997). In addition, the distribution of the new species extends into
the Lubudi-Sampwe hunting area and the annex area of the KNP,
where threats due to increasing human encroachment and settlement
are also rising. These are generally associated with increases in
unsustainable fishing practices such as the use of mosquito nets,
overfishing and habitat destruction as encountered mainly in the
Kamalondo Depression of the UNP (Brown & Abell, 2005) but which
also prevail in the Lufira Depression, the vast floodplain of the Middle
Lufira where the new species is present (Melli & Micha, 2015). How-
ever, recently (CMM personal observation, 2017), the rigorous imple-
mentation of the sustainable management of fisheries resources
by the Institut Congolais pour la Conservation de la Nature has
considerably reduced the use of prohibited fishing nets in the Lufira
Depression. For the rest of the distribution of the new species in
the KNP there are also strong threats to the fish fauna of the
rivers surrounded by villages, especially those located at the foot
of the western slope of the Kundelungu Plateau where fishing
with the use of ichthyotoxic plants, such as the so-called Buba
(Tephrosia vogelii Hooker et al. 1849), is widespread. During these
practices fishing nets with small mesh sizes (about 10 mm) are
used, which block the up- and downstream migration of fishes in
the river which tend to escape the application of the ichthytoxines
applied. Finally, as far as there is no good surveillance of the fish-
ing activities, the effective protection and conservation of the ich-
thyofauna of both the UNP and the KNP remain uncertain. It is
therefore hoped that the discovery of yet another new fish spe-
cies for this area further stresses the importance of both parks for
fish protection and conservation.
5|COMPARATIVE MATERIAL EXAMINED
5.1 |Cyphomyrus aelsbroecki (Poll 1945)
MRAC 54990, 71.4 mm L
S
; DRC: Haut-Katanga Province, Lubumbashi
(former Elisabethville), 1937, R.P Van Aelsbroecki.
5.2 |Cyphomyrus cubangoensis (Pellegrin 1936):
MRAC P-138760 (ex. MNHN 1936–65), Syntype, 83.2 mm L
S
; Angola:
Cubango (Okavango) basin. Dr. Monard (Swiss missions in Angola),
1928–29, 1932–33. MNHN 1936–0062-0064, 3 syntypes,
82.1–87.2 mm L
S
; Angola: Cubango (Okavango) basin. Dr. Monard
(Swiss missions in Angola), 1936.
24
80 100 140120
Standard length (mm)
160 200
180 220
26
Body depth (% LS)
30
28
34
32
36
(a)
58
80 100 140120
Standard length (mm)
160 200180 220
60
59
Preanal distance (% LS)
62
61
65
64
63
66
(b)
FIGURE 11 Scatterplots of (a) body depth (% L
S
) and (b) preanal distance (% L
S
) against standard length (in mm) (n= 40).
Cyphomyrus discorhynchus:, lectotype; , paralectotypes of the Lower Zambezi basin; , specimens from the Kamalondo Depression;
C. tanganicanus (currently a junior synonym of C. discorhynchus): , syntypes; , other specimens from some affluents of the Tanganyika basin
16 MUKWEZE MULELENU ET AL.
FISH

5.3 |Cyphomyrus discorhynchus (Peters 1852)
ZMB 3674, Lectotype, 163.7 mm L
S
; Mozambique: Lower Zambezi
River; W.C. Peters; Lectotype selected by Seegers (1996: 69). ZMB
3673, 3675–3676 & 23,374, four Paralectotypes, 132.8–205.8 mm
L
S
; Mozambique: Lower Zambezi River; W.C. Peters; Paralectotypes.
ZSM Zam10, 101.3 mm L
S
, Mozambique: Middle Zambezi just below
the Victoria Falls near the beginning of Batoka Gorge: rapid, about
(17560S; 25510E), 7 December 1996; Van der Bank and B. Kramer.
MRAC 2016-003-P-0149-0155, 106.2–137.1 mm L
S
; DRC: UNP,
Haut-Katanga Province, lake Lukanga (Katelwa fishing camp),
Kamalondo Depression, Upper Lualaba (82004400S; 262802500E); Alt.
563 m a.s.l.; C. Mukweze, B. Katemo, E. Muyambo & D. Mfwana,
11 September 2015. MRAC 2016-003-P-0135-0140, 96.5–131.7 mm
L
S
; DRC: UNP, Haut-Katanga Province, Lake Mulenda (Kyabu village),
Kamalondo Depression, Upper Lualaba (84602300S; 26204300E); Alt.
571 m a.s.l.; C. Mukweze, B. Katemo, E. Muyambo & D. Mfwana,
2 September 2015. MRAC 2016-003-P-0141-0145, 107.0–128.0 mm
L
S
; DRC: UNP, Haut-Katanga Province, Lake Upemba (specimens pur-
chased at Misebo's market), Kamalondo Depression, Upper Lualaba.
About (8350100S; 263004300E); Alt. 561 m a.s.l.; C. Mukweze,
B. Katemo, E. Muyambo & D. Mfwana, 11 September 2015.
5.4 |Cyphomyrus macrops (Boulenger 1909)
BMNH 1899.6.27.12, holotype, 174.0 mm L
S
; DRC: Upper Congo.
5.5 |Cyphomyrus psittacus (Boulenger 1897)
BMNH 1897.9.30.26, holotype, 112.7 mm L
S
; DRC: Congo River,
Stanley Falls (0300N; 25120E), W. Bentley.
5.6 |Cyphomyrus smithersi (Määr 1962) synonym
of Cyphomyrus discorhynchus (Peters, 1852)
MRAC 183658–60, 120.7–132.9 mm L
S
, southeastern Africa, Rhode-
sia (Zimbabwe): Chipepo River above Kariba Lake (16490S; 27500E),
01 May 1965; Matthes. MRAC 183662–63, 110.2–183.6 mm L
S
,
southeastern Africa, South Rhodesia (Zimbabwe): Sibilobilo (clear
darea), Kariba Lake, about (16470S; 28110E), 24 April 1965; Matthes.
5.7 |Cyphomyrus tanganicanus (Boulenger 1906)
synonym of Cyphomyrus discorhynchus (Peters, 1852)
BMNH 1906.9.8.3-4, Syntypes, 125.2–127.7 mm L
S
, Sumbu and river
at Msamba, Lake Tanganyika, 1906, W. Cunnington. BMNH
1906.9.8.5, Syntype, 147.1 mm L
S
, Sumbu and river at Msamba, Lake
Tanganyika, 1906, W. Cunnington. MRAC 1992.081.P.0071, 106.4 mm
L
S
, Tanzania: Ulwile Island, northern shore, Lake Tanganyika (7270S;
30340E), 1992; Expedition 1992. MRAC 1992-081-P-0163, 101.0 mm,
Tanzania: Ulwile Island, northern shore, Lake Tanganyika (7270S;
30340E), 1992; Expedition 1992. MRAC P-126349, 85.7 mm L
S
,DRC:
Uvira, Lake Tanganyika (3240S; 2980E), 1958; Expedition 1992. MRAC
91-034-P-0180-0183, 115.2–124.9 mm L
S
, Burundi: Bujumbura, Lake
Tanganyika (specimens purchased at the market) (3230S; 29220E),
October 12, 1994; L. De Vos. MRAC P-190342, 108.6 mm, Zambia:
Liemba jetty (Mpulungu), Lake Tanganyika (8460S; 3170E), 11 July
1967; H. Matthes.
5.8 |Cyphomyrus wilverthi (Boulenger 1898)
MRAC 133, syntype, 239.9 mm L
S
; DRC: Upoto, Upoto 1896,
Wilverthi.
ACKNOWLEDGEMENTS
This study was made possible by a DEA scholarship (2016–2018) to the
first author (CMM) from the Mbisa Congo project (2013-2018), financed
through a framework agreement project between the RMCA and the
Belgian Development Cooperation. We would like to thank Johanna
Kapp (ZMB) for the loan of the lectotype and the four paralectotypes of
C. discorhynchus under her care, Gabsi Zora and Patrice Pruvost (MNHN)
for their warm welcome during our study visit at the MNHN which
enabled the study of some of the syntypes of C. cubangoensis, Dirk Neu-
mann and Ulrich Schliewen (ZSM) for the loan of specimens under their
care, Katrien Dierickx (RMCA) for the excellent photographs of the types
and providing us with the GenBank accession numbers, Carl D. Hopkins
(CU), an anonymous reviewer and the assistant editor Albert Chakona
(SAIAB) for constructive comments on the manuscript, and the Project
‘Développement de la Pêche Artisanale et de l'Aquaculture au Katanga’/
PRODEPAAK (NN/3000769) CTB/BTC project (2008-2013), especially
Jean-Pierre Marquet, for financial and logistical support to the Katanga
Expedition 2012, which enabled the collecting of specimens belonging to
the new species. The University of Kolwezi (UNIKOL) and the University
of Lubumbashi (UNILU) are acknowledged, respectively, for accepting a
temporary lay-off to the UNILU of the first author and for accepting him
as a DEA student. Thanks also to Muriel van Nuffel (RMCA-Scientific
training and hospitality coordinator), Gert Boden (RMCA-FishBase),
Tobias Musschoot (RMCA-FishBase) and Kisekelwa Tchalondawa (ISP-
Bukavu) for their dedication in helping out the first author during his study
visits to the RMCA and Emmanuel Abwe (BEZHU-UNILU) for having col-
lected some of the specimens belonging to the new species in the KNP.
AUTHORS' CONTRIBUTIONS
CMM, BMK and EV were responsible for the fieldwork, fish identifica-
tion and study design, and wrote the first and subsequent revised ver-
sions of the manuscript. In addition, EV supervised the research and
critically revised the final version of the manuscript. ED was mainly
responsible for the molecular work, sequence data analyses and writ-
ing and supervision of the paragraphs related to this part of the paper.
All authors discussed, read and rewrote parts of the manuscript, and
approved its final version.
MUKWEZE MULELENU ET AL.17
FISH

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SUPPORTING INFORMATION
Additional supporting information may be found online in the
Supporting Information section at the end of this article.
How to cite this article: Mukweze Mulelenu C, Katemo
Manda B, Decru E, Chocha Manda A, Vreven E. The
Cyphomyrus Myers 1960 (Osteoglossiformes: Mormyridae) of
the Lufira basin (Upper Lualaba: DR Congo): A generic
reassignment and the description of a new species. J Fish Biol.
2020;1–19. https://doi.org/10.1111/jfb.14237
MUKWEZE MULELENU ET AL.19
FISH