Description of Scriptaphyosemion ... - zoological Bulletin

Bonn zoological Bulletin 61 (1): 13–28 July 2012

Description of Scriptaphyosemion wieseae
(Cyprinodontiformes: Nothobranchiidae),
a new species from northern Sierra Leone

Rainer Sonnenberg1,2 & Eckhard Busch3
1 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany
2 Max-Planck-Institut für Evolutionsbiologie, Department of Evolutionary Genetics, August-Thienemann-Strasse 2,

D-24306 Plön, Germany. Corresponding author. E-mail: [email protected]
3 Diederichsstrasse 45, D-42855 Remscheid, Germany.

Abstract. Scriptaphyosemion wieseae, new species, is described from collections in northern Sierra Leone. Results of a
small DNA study indicate, that S. bertholdi from the southeastern part of the country is the closest relative. Both species
can be distinguished by male and female color pattern and male caudal fin shape.
Résumé. Scriptaphyosemion wieseae, une nouvelle espèce, est décrite du Nord de la République de Sierra Leone. Le ré-
sultat d’une petite étude moléculaire ADN montre que son espèce sœur est S. bertholdi, présente au Sud-Est du pays. Les
deux espèces se distinguent par les couleurs du mâle et de la femelle et par la forme de la nageoire caudale du mâle.
Key words. Upper Guinean ichthyofaunal province, taxonomy, DNA, phylogeny.

INTRODUCTION

The West African killifish genus Scriptaphyosemion Rad- non which may confuse species identification; however,
da & Pürzl, 1987 is relatively species rich, compared to some of the most variable species might be composed of
the closely related genera Archiaphyosemion Radda, 1977, more than one species (see e.g. figures or drawings in
Callopanchax Myers, 1933, and Nimbapanchax Sonnen- Seegers 1997; Wildekamp 1993). This combination of in-
berg & Busch, 2009. At present 18 specific names are traspecific variability, inclusion of probably different
available in Scriptaphyosemion (Eschmeyer 2011; Huber species under one taxon, and the lack of a specific diag-
2007; Lazara 2001), but only 10 (Huber 2007; Wildekamp nosis in many species descriptions contributed to some
& Van der Zee 2003) to 13 (Costa 2008; Lazara 2001) confusion in Scriptaphyosemion taxonomy (see e.g.
species are currently accepted as valid, depending on au- Scheel 1990). More than a decade ago, Murphy et al.
thor and concept (Huber 2007; Eschmeyer 2011; Lazara (1999) published the first molecular genetic phylogeny of
2001; Scheel 1990; Wildekamp 1993). Among the West Scriptaphyosemion, based on the sequences of the partial
African killifish genera, species number in Script- cytochrome b and 12S rRNA genes and including eleven
aphyosemion is only exceeded by the Upper Guinean species. This phylogeny gave a first insight into the rela-
species of the related genus Epiplatys Gill, 1862 (Costa tionships and an additional set of independent characters
2008; Huber 2007; Wildekamp & Van der Zee 2003). for further taxonomic and phylogenetic research.

In nothobranchiid killifishes, most species are diagnosed Species diversity of Scriptaphyosemion is highest in
by male color pattern (Amiet 1987; Sonnenberg 2007), Sierra Leone, Liberia, and Guinea. However, the distri-
which is, however, increasingly supplemented by molec- bution area also includes Burkina Faso, Gambia, Guinea-
ular approaches (e.g. Agnèse et al. 2009; Legros et al. Bissau, Ivory Coast, Mali, and Senegal (Huber 2007; Pau-
2005; Legros & Zentz 2007a, b; Sonnenberg 2007, 2008; gy et al. 1990; Teugels et al. 1988; Wildekamp 1993;
Sonnenberg & Blum 2005; Sonnenberg & Busch 2009, Wildekamp & Van der Zee 2003). In Sierra Leone, Script-
2010; Sonnenberg et al. 2006, 2011; Van der Zee & Son- aphyosemion species are mainly known from the coastal
nenberg 2011). In Scriptaphyosemion, female color pat- lowland, the northern and eastern killifish fauna is only
tern has also been used as character for diagnosing species little known (Etzel 1977, 1978a, b, 1984; Huber 2007;
or groups of species (Roloff 1971a). Wildekamp 1993; Wildekamp & Van der Zee 2003).

In most species of Scriptaphyosemion, interspecific vari- In 1993, the second author collected together with B.
ance in male color pattern is accompanied by intraspecif- Wiese nothobranchiids in the northern part of Sierra Leone
ic variance between and within populations, a phenome- (Busch 1995a), including a probably new species near

Received: 06.02.2012 Corresponding editor: F. Herder

Accepted: 15.06.2012

14 Rainer Sonnenberg & Eckhard Busch

Table 1. List of specimens used for the DNA analyses with locality data and GenBank accession numbers (ND2+tRNAs, mtL-
SU rDNA, nLSU rDNA). * Sequences published in Sonnenberg & Busch (2009).

Species sample Country Collection locality GenBank acc. no.

Epiplatys sexfasciatus RS1364 Gabon G 02/91 JX124276, FJ872035*, FJ872049*

Scriptaphyosemion bertholdi RS1402 Sierra Leone SL 93/51 JX124261, JX124233, JX124247

S. brueningi RS1377 Sierra Leone SL 03/04 JX124262, JX124234, JX124248

S. brueningi RS1403 Sierra Leone SL 89, Perie JX124263, JX124235, JX124249

S. cauveti RS1680 Guinea GM 97/05 JX124264, JX124236, JX124250

S. chaytori RS1379 Sierra Leone SL 93/23 Ngabu JX124265, JX124237, JX124251

S. fredrodi RS1398 Sierra Leone SL 89, Matanga JX124266, JX124238, JX124252

S. geryi RS1381 Sierra Leone SL 03/09, Robis 1 JX124267, FJ872033*, FJ872047*

S. guignardi RS1380 Guinea GM 97/13 JX124268, JX124239, JX124253

S. liberiense RS1757 Liberia L 97 JX124269, JX124240, JX124254

S. roloffi RS1376 Sierra Leone SL 03/10 JX124270, JX124241, JX124255

S. schmitti RS1755 Liberia RL 83/121 JX124271, JX124242, JX124256

S. sp. RS1378 Sierra Leone SL 03/18 JX124272, JX124243, JX124257

S. wieseae RS1382 Sierra Leone SL 03/16 JX124273, JX124244, JX124258

S. wieseae RS1686 Sierra Leone SL 03/15 JX124274, JX124245, JX124259

S. wieseae RS1687 Sierra Leone SL 03/17 JX124275, JX124246, JX124260

Lenghe Curoh. In 2003 they revisited this locality and For counts and measurements, see Sonnenberg & Busch
found two additional collection localities of this species (2009, 2010). All measurements were taken with a digi-
(Fig. 1). Preliminary crossing experiments by engaged tal caliper, partly under a dissecting microscope, to the
aquarists (Vandersmissen 2003) indicated its probable in- nearest 0.1 mm. Measurements, including those of the
compatibility with S. bertholdi (Roloff, 1965), which was head, are presented as percentages of standard length (SL).
considered to be the potentially closest relative. Here, we The head region of one specimen was scanned with a
provide a formal species description and present the phy- Scanco viva CT40 at the Max-Planck-Institut für Evolu-
logenetic position of the new species within Script- tionsbiologie in Plön, digital X-ray images were made with
aphyosemion, based on mitochondrial and nuclear DNA a Faxitron LX-60 Digital Specimen Radiography System
sequences. at the ZFMK. Fin rays of unpaired fins were counted on
digital X-ray images; the relative position of the first dor-
MATERIAL AND METHODS sal fin ray to the anal fin (D/A) is estimated following Son-
nenberg & Schunke (2010). Nomenclature of the neuro-
The species description is based on collections made in mast system on the head follows Scheel (1968), for supra-
Sierra Leone by the second author and B. Wiese during orbital (or frontal) squamation Hoedeman (1958). Male
their second visit in this area in November 2003. Speci- and female color patterns are compared with that given
mens were deposited in the Royal Museum for Central in the respective description and on pictures or drawings
Africa (MRAC, Tervuren, Belgium), the Zoologische published in the literature.
Staatssammlung München (ZSM, Munich, Germany), and
the Zoologisches Forschungsmuseum Alexander Koenig As species concept we employ the pragmatic approach
(ZFMK, Bonn, Germany). of Moritz et al. (2000) and recognize species by diagnos-
tic combinations of characters.
Bonn zoological Bulletin 61 (1): 13–28
©ZFMK

New Scriptaphyosemion from Sierra Leone 15

Fig. 1. Distribution map of Scriptaphyosemion in Sierra Leone based on data from Huber (2007) and own data of the second au-
thor. White square: S. bertholdi; black triangle: S. brueningi; white circle: S. chaytori; dark grey triangle: S. fredrodi; dark grey
square: S. geryi; light grey circle: S. guignardi; white triangle: S. liberiense; black circle: S. roloffi; black square: S. wieseae. TT
marks the type localities of S. bertholdi and S. wieseae.

A DNA dataset was prepared for 15 Scriptaphyosemion Sequences were aligned with ClustalX 1.8 (Thompson
specimens, including 11 species, and Epiplatys sexfascia- et al. 1997), checked by eye, and merged in a single align-
tus Gill, 1862 as outgroup. We sequenced the complete ment in BioEdit 7.0.5.3 (Hall 1999). As the ND2 se-
mitochondrial ND2 gene and adjacent tRNAs, and frag- quences of three species are not complete, missing sites
ments of the mitochondrial and nuclear LSU rDNA genes. were filled with N [S. liberiense (560 bp), S. schmitti (616
Parts of the sequences were published in earlier papers of bp), and S. cauveti (58 bp)].
the authors; all samples are listed with GenBank acces-
sion numbers and collection locality in Table 1. For de- The protein coding gene was translated into the corre-
tails on the DNA methods, see Sonnenberg & Busch sponding amino acid sequence to check for a functional
(2009, 2010) and Sonnenberg et al. (2007). coding sequence, and base composition of mitochondri-
al sequences was estimated with MEGA 4.1 beta (Tamu-
Abbreviations used in Table 2 and Figures 2–3: BER = ra et al. 2007). Uncorrected p-distances with pairwise ex-
S. bertholdi, BRU = S. brueningi (Roloff, 1971b), CAU clusion of missing data were calculated in MEGA 4.1 be-
= S. cauveti (Romand & Ozouf, 1995), CHY = S. chay- ta (Tamura et al. 2007). Number of constant, variable, and
tori (Roloff, 1971a), FRE = S. fredrodi (Vandersmissen informative characters were estimated with PAUP 4.0b10
et al., 1980), GER = S. geryi (Lambert, 1958), GUI = S. (Swofford 1998).
guignardi (Romand, 1981), LIB = S. liberiense
(Boulenger, 1908), ROL = S. roloffi (Roloff, 1936), SMI Data were analyzed using a maximum parsimony (MP)
= S. schmitti (Romand, 1979), Ssp = S. sp. SL 03/18, WIE and a Bayesian approach (BA). Maximum parsimony
= S. wieseae, new species, SEX = E. sexfasciatus. analyses were performed with PAUP 4.0b10 (Swofford
1998) with the exclusion of uninformative positions and

Bonn zoological Bulletin 61 (1): 13–28 ©ZFMK

16 Rainer Sonnenberg & Eckhard Busch

Fig. 2. Results of the maximum parsimony analyses. Shown is one out of three equally short trees from the analysis with the
complete dataset. Values left to nodes are the corresponding bootstrap values of all three analyses (complete dataset / third protein
coding position excluded / alignment columns with missing data excluded). Values for nodes were only given, if at least one analy-
sis has a bootstrap support above 75 %. Tree length = 1262, consistency index (CI) = 0.5166, homoplasy index (HI) = 0.4834, re-
tention index (RI) = 0.5570, rescaled consistency index (RC) = 0.2878.

gaps coded as fifth character, heuristic tree searches were All trees were rooted with Epiplatys sexfasciatus as out-
performed with random addition of sequences for 1,000 group. Only nodes with bootstrap support values ≥ 75%
replicates. Bootstrap tests were performed with random or posterior probabilities ≥ 0.95 were considered as sup-
addition of sequences for 100 replicates and 10,000 boot- ported by the data. Nodes below these values were con-
strap replicates. Three different analyses were made, one sidered as unsupported by the data and not resolved.
with the complete dataset, one with the exclusion of third
protein coding positions, and a third with exclusion of re- RESULTS
gions, in which one of the above listed specimens has
missing data. Otherwise, all parameters were identical. DNA Analysis. (Figs 2–3, Table 2). The complete align-
ment of the ND2, tRNA, and both LSU sequences has a
Bayesian analyses were performed with MrBayes 3.1.2 length of 3199 bp including alignment introduced gaps for
(Huelsenbeck & Ronquist 2001), data were partitioned in a total of 16 specimens (503 bp tRNAs, 1047 bp ND2, 481
mitochondrial non protein coding regions (tRNA, mtLSU), bp mtLSU rDNA, 1168 bp nLSU rDNA). Of these, 2278
protein coding regions (ND2), and the nuclear LSU se- bp are constant, 456 bp variable but uninformative, and
quences. Number of generation was set to 10,000,000 with 465 bp parsimony informative. The ND2 sequence trans-
sampling of every 1,000 generations and the results of the lates into 349 amino acids with no unexpected stop codon,
first 10% of the generations were discarded as burn-in time the base composition of all mitochondrial genes shows the
after checking that the runs had reached the stationary typical base composition for this organelle genome with
phase at this point. Four analyses were performed: the two an anti-G bias (Zhang & Hewitt 1996). Average uncorrect-
alignments, one with the complete data set and one with ed pairwise distance for all sequences is 7.4 %, maximum
the exclusion of regions, in which one of the above men- observed distance is 16.6 % between outgroup and ingroup
tioned specimens has missing data, were both performed specimens, maximum observed distance between ingroup
with NST=2 and NST=6. specimens is 9.0 %, further details are given in Table 2.

Bonn zoological Bulletin 61 (1): 13–28 ©ZFMK

Bonn zoological Bulletin 61 (1): 13–28 Table 2. Uncorrected p-distances with pairwise exclusion of missing data for the complete dataset / the mitochondrial ND2+tRNAs / the mitochondrial LSU rDNA / the nuclear LSU rDNA.

sample BER1401 BRU1377 BRU1403 CAU1680 CAY1379 FRE1398 GER1381 GUI1380 LIB1757 ROL1376 SMI1755 Ssp1378 WIE1382 WIE1686 WIE1687

BRU1377 6.3/ 11.3/ New Scriptaphyosemion from Sierra Leone
BRU1403 2.7/ 0.9
©ZFMK CAU1680 6.2/ 11.0/ 0.4/ 0.9/
CAY1379 2.7/ 0.9 0.0/ 0.0
FRE1398 8.3/ 14.5/ 8.2/ 14.6/ 8.2/ 14.4/
GER1381 3.6/ 2.0 3.8/ 1.6 3.8/ 1.6
GUI1380 5.2/ 8.2/ 7.2/ 12.5/ 7.0/ 12.2/ 8.6/ 14.2/
LIB1757 2.5/ 2.0 3.6/ 1.4 3.6/ 1.4 4.6/ 2.8
ROL1376 5.3/ 8.9/ 6.8/ 12.1/ 6.6/ 11.7/ 8.6/ 15.1/ 6.1/ 9.9/
SMI1755 3.4/ 1.3 3.4/ 0.9 3.4/ 0.9 4.0/ 1.8 4.6/ 1.3
Ssp1378 7.5/ 12.4/ 7.9/ 13.4/ 7.7/ 13.0/ 9.0/ 15.5/ 7.7/ 12.8/ 8.2/ 13.8/
WIE1382 3.6/ 2.3 4.2/ 1.9 4.2/ 1.9 3.8/ 2.5 4.0/ 2.3 4.0/ 2.2
WIE1686 5.9/ 9.7/ 7.5/ 13.0/ 7.4/ 12.8/ 8.7/ 15.1/ 6.2/ 9.9/ 6.6/ 10.9/ 8.3/ 13.8/
WIE1687 3.4/ 1.7 4.6/ 1.3 4.6/ 1.3 4.2/ 2.1 3.8/ 2.0 5.1/ 1.4 4.4/ 2.2
SEX1364 5.6/ 11.4/ 2.1/ 4.4/ 2.1/ 4.3/ 7.1/ 14.8/ 6.4/ 12.8/ 5.6/ 11.8/ 6.5/ 12.8/ 6.5/ 12.9/
3.6/ 1.4 1.1/ 0.5 1.1/ 0.5 4.9/ 1.7 4.4/ 1.5 3.8/ 1.0 5.1/ 1.7 5.5/ 1.4
6.6/ 11.7/ 6.9/ 12.1/ 6.9/ 12.1/ 8.8/ 15.4/ 6.8/ 12.2/ 6.9/ 12.4/ 5.9/ 9.9/ 7.5/ 12.8/ 5.9/ 12.3/
2.3/ 1.5 3.8/ 0.9 3.8/ 0.9 3.8/ 2.2 2.7/ 1.1 4.0/ 0.7 2.5/ 1.7 4.0/ 1.7 4.6/ 0.8
6.5/ 11.8/ 5.9/ 11.0/ 5.9/ 11.1/ 5.4/ 9.9/ 6.4/ 10.9/ 6.0/ 10.6/ 6.3/ 11.5/ 6.7/ 11.9/ 3.9/ 7.7/ 5.7/ 10.2/
4.9/ 2.8 3.8/ 2.4 3.8/ 2.4 4.4/ 2.3 5.3/ 3.1 4.6/ 2.7 4.6/ 2.8 5.9/ 2.7 4.9/ 2.2 4.2/ 2.5
5.5/ 9.2/ 6.6/ 11.9/ 6.4/ 11.4/ 8.6/ 15.0/ 6.1/ 10.1/ 1.5/ 2.9/ 7.9/ 13.2/ 6.6/ 10.7/ 5.5/ 11.6/ 6.8/ 11.9/ 6.1/ 10.6/
3.2/ 0.9 3.2/ 0.9 4.2/ 1.8 4.4/ 1.3 0.6/ 0.0 4.2/ 2.2 5.3/ 1.4 3.6/ 1.0 4.2/ 0.7 5.3/ 2.7 3.2/ 1.3
2.8/ 4.7/ 6.4/ 11.8/ 6.2/ 11.4/ 8.2/ 14.3/ 5.3/ 8.5/ 5.2/ 8.6/ 7.3/ 12.1/ 5.8/ 9.5/ 5.2/ 10.8/ 6.8/ 11.8/ 5.8/ 10.5/ 5.2/ 8.7/
0.8/ 1.0 2.3/ 0.6 2.3/ 0.6 3.6/ 1.8 3.0/ 2.0 3.0/ 1.3 3.6/ 2.2 3.8/ 1.5 3.2/ 1.2 2.7/ 1.5 4.4/ 2.6 2.7/ 1.3
2.7/ 4.5/ 6.4/ 11.8/ 6.2/ 11.4/ 8.2/ 14.5/ 5.3/ 8.4/ 5.1/ 8.5/ 7.4/ 12.4/ 5.9/ 9.8/ 5.4/ 11.2/ 6.5/ 11.4/ 5.8/ 10.5/ 5.2/ 8.7/ 0.5/ 0.9/
1.1/ 1.0 2.5/ 0.6 2.5/ 0.6 3.4/ 1.8 3.2/ 2.0 2.7/ 1.3 3.4/ 2.2 3.6/ 1.5 3.4/ 1.2 2.5/ 1.5 4.2/ 2.6 3.0/ 1.3 0.2/ 0.0
3.0/ 5.1/ 6.5/ 11.9/ 6.2/ 11.3/ 8.1/ 14.3/ 5.3/ 8.4/ 5.2/ 8.9/ 7.6/ 12.8/ 6.1/ 10.1/ 5.4/ 11.1/ 6.7/ 11.8/ 5.8/ 10.4/ 5.3/ 9.0/ 0.7/ 1.1/ 0.7/ 1.2/
1.1/ 0.9 2.5/ 0.7 2.5/ 0.7 3.4/ 1.9 3.2/ 1.9 2.7/ 1.3 3.4/ 2.2 3.6/ 1.6 3.4/ 1.2 2.5/ 1.4 3.8/ 2.8 3.0/ 1.3 0.6/ 0.1 0.4/ 0.1
16.5/ 24.4/ 16.2/ 23.9/ 16.3/ 24.0/ 16.6/ 24.2/ 16.5/ 24.4/ 16.4/ 24.4/ 16.6/ 24.6/ 16.6/ 24.6/ 15.5/ 26.0/ 16.3/ 24.0/ 12.3/ 18.4/ 16.4/ 24.3/ 16.3/ 24.5/ 16.2/ 24.3/ 16.2/ 24.3/
13.1/ 6.7 13.5/ 6.6 13.5/ 6.6 14.8/ 7.0 13.3/ 6.7 13.3/ 6.6 13.9/ 6.6 13.9/ 6.6 13.9/ 6.7 13.5/ 6.7 12.4/ 7.0 13.7/ 6.5 12.4/ 6.6 12.4/ 6.6 12.4/ 6.6

17

18 Rainer Sonnenberg & Eckhard Busch

Fig. 3. Results of the Bayesian analyses. Shown is the consensus tree of the analysis with the complete dataset and NST=6 (A)
and the consensus tree of the dataset with excluded alignment columns with missing data and NST=6 (B). Values left to nodes are
the corresponding posterior probability values, except for values marked with * which are on the right side of the corresponding
node (complete dataset, NST=2 / complete dataset, NST=6 in A; alignment columns with missing data excluded, NST=2 / align-
ment columns with missing data excluded, NST=6 in B). Values for nodes were only given, if at least one analysis has a posteri-
or probability above 0.95.

Bonn zoological Bulletin 61 (1): 13–28 ©ZFMK

New Scriptaphyosemion from Sierra Leone 19

All three maximum parsimony analyses result in near- species. The genetic distance values (Table 2) show a sim-
ly the same tree topology and differ only in branch length ilar divergence as between S. liberiense and S. bruenin-
and bootstrap support values; here, only one of three short- gi.
est trees of the complete dataset is presented (Fig. 2). The
Bayesian analyses for the complete and the reduced Scriptaphyosemion wieseae, new species
datasets show slightly different tree topologies, independ- (Fig. 4, Table 3)
ent if the analyses were performed with NST=6 or 2. Scriptaphyosemion sp. Lenghe Curoh: Busch (1995a),
Shown are the results for the complete and the reduced Vandersmissen (2003)
dataset of the NST=6 analyses, on the nodes posterior
probabilities for both analyses for each dataset are present- Holotype. ZFMK 47680, male, 36.1 mm SL, Sierra
ed if the values are equal to or above 0.95 (Fig. 3 A+B). Leone, Lenghe Curoh, Sanghi or Sanigi River, field num-
ber SL 03/16 (9°28,04’ N, 11°40,58’ W), E. Busch and B.
Scriptaphyosemion schmitti and S. cauveti are support- Wiese, 18. Nov. 2003.
ed as basal to all other species of the genus (Figs 2–3).
As S. schmitti was placed in the study of Murphy et al. Paratypes. ZFMK 47681–47692, 8 males, 4 females,
(1999) within a clade containing the other Script- 19.5–34.7 mm SL, paratopotypes, collected with the holo-
aphyosemion species studied, we checked for possible type.
causes of this discrepancy. As the sequences studied here MRAC B2-06-P-1–2, 1 male, 22.9 mm SL, 1 female, 21.5
have no overlap with those of Murphy et al. (1999), we mm SL, paratopotypes, collected with the holotype.
compared a cytochrome b fragment of our sample with ZFMK 47693–47707, 7 males, 8 females, 18.8–41.3 mm
the data deposited in GenBank. However, both samples SL, Sierra Leone, Fadugu, at the bridge over the Mabakah
differ in only one nucleotide (results not shown), and a River, field number SL 03/15 (9°23,52’ N, 11°45,76’ W),
Blast search with our analysed sequences (see Table 1) on- E. Busch and B. Wiese, 18. Nov. 2003.
ly returned the highest similarity with other Script- ZSM 41596 (2), 1 male, 28.4 mm SL, 1 female, 22.3 mm
aphyosemion sequences. To exclude that the basal posi- SL, collected with ZFMK 47693–47707.
tion of S. schmitti is an artifact due to the missing sequence
data, we also performed the analyses with the reduced Non-type DNA samples. RS1686, 1 male, DNA vouch-
dataset, which did not contain missing data, but gave the er, preserved in 96 % ethanol, collected with ZFMK
same results (Figs 2–3). However, further samples should 47693–47707, field number SL 03/15; RS1382, 2 males,
be studied for a confirmation of the basal position, espe- 1 female, DNA voucher, preserved in 96 % ethanol, col-
cially as this node is supported neither by bootstrap nor lected with the holotype; RS1687, 1 male, DNA vouch-
posterior probability values. er, preserved in 96 % ethanol, Sierra Leone, Kabala, col-
lected in rice swamp, field number SL 03/17 (9°35’ N,
Our results indicate a closer relationship between S. 11°33’ W), E. Busch and B. Wiese, 18. Nov. 2003.
liberiense and S. brueningi, S. roloffi and S. geryi, S. fred-
rodi and a probably undescribed species (see under Dis- Diagnosis. Scriptaphyosemion wieseae is distinguished
cussion), and S. bertholdi and S. wieseae, new species. A from its congeners by a unique combination of male and
closer relationship between S. chaytori and S. guignardi female coloration characters. Males have a blue body col-
occurs in all resulting trees (Figs 2–3), but is well sup- or, red dots on side forming irregular, interrupted red
ported only in the Bayesian analyses of the complete stripes, especially on caudal peduncle a more reticulated
dataset. Both species were in addition grouped with S. pattern, unpaired and pelvic fins centre blue with a nar-
roloffi and S. geryi in the analyses of the reduced dataset, row blue margin and a broad red submargin, on anal and
but not in the maximum parsimony analyses and the dorsal fin a small number of basal red blotches, centre of
Bayesian analyses of the complete dataset (Figs 2–3). caudal fin with some red blotches, pectoral transparent to
orange with a blue margin; females with a dark brown
All analyses support a monophyletic group formed by midlateral stripe, interrupted on caudal peduncle, body on
all Scriptaphyosemion except of S. schmitti and S. cau- lower half of side and ventrally orange vs. a mainly red
veti (Figs 2–3). Another well supported subgroup is lateral body color in male S. cauveti; head and anterior
formed by S. bertholdi, S. wieseae, new species, S. chay- body yellowish to brownish, unpaired fin margins white
tori, S. guignardi, S. fredrodi, and S. sp. (see under Dis- or yellow, and pectoral fn margin yellow to orange in male
cussion); only in the analyses of the reduced dataset S. S. schmitti; males with more or less regular oblique red
roloffi and S. geryi were also included in this group (Fig. bars on side, at least on the caudal peduncle, fin margins,
2–3). To this subgroup, S. brueningi and S. liberiense, especially of caudal and pectoral, often yellow, and fe-
probably is the sister group, a hypothesis only significant- males with dark brown streaks or blotches, similar to the
ly supported in the Bayesian analyses (Fig. 3).

The phylogenetic results show that S. bertholdi is the
well supported sister species to S. wieseae, new species,
and indicates no closer relationship to other mentioned

Bonn zoological Bulletin 61 (1): 13–28 ©ZFMK

20 Rainer Sonnenberg & Eckhard Busch

Fig. 4. A) Male of Scriptaphyosemion wieseae, collection locality SL 03/16, not preserved. B) Female of Scriptaphyosemion wieseae,
collection locality, SL 03/16, not preserved.

red oblique bars of males on side, often with a dark blotch ularly distributed dark brown or red dots on side in S.
at the root of the caudal in S. geryi and S. roloffi; males chaytori; males with dark red blotches on side, caudal fin
with broad red blotches, often forming broad irregular ver- margins whitish to light blue or yellow, pectoral fin mar-
tical stripes, caudal fin margins yellow, often with a small gin white, females with dark brown blotches or oblique
white submargin followed by a broader red band, females short streaks on dorsal half of caudal peduncle and a dark
with irregularly distributed dark brown dots or short blotch on posterior end of caudal peduncle at dorsal base
oblique streaks or blotches dorsally on the caudal pedun- of caudal fin in S. fredrodi; males with light blue to blue-
cle, a weak anterior dark lateral band, and a dark blotch green body color, many irregularly distributed small red
on the posterior dorsal end of the caudal peduncle at the streaks or dots, submarginal stripes of unpaired fins dark
caudal fin base in S. brueningi; often yellow, sometimes red to nearly black, margins light blue or whitish, some-
whitish caudal fin margins in males, females with irreg- times on dorsal and often on anal absent, females with red

Bonn zoological Bulletin 61 (1): 13–28 ©ZFMK

New Scriptaphyosemion from Sierra Leone 21

Table 3. Morphometrics of Scriptaphyosemion wieseae, new species. Values in the table are based on 15 specimens, for which
all measurements and counts were taken (7 of each male and female paratypes). All measurements in percentages of standard length
(SL) except standard length in mm. TL = total length, HL = length of head, PD = predorsal fin distance, pPED = prepectoral fin
distance, pPD = prepelvic fin distance, pAD = preanal fin distance, DB = greatest body depth, DC = caudal peduncle depth, CL =
length of caudal peduncle, BD = base of dorsal fin, BA = base of anal fin, E = eye diameter, I = interorbital width, SD = standard
deviation.

Holotype Paratypes ♂ mean Paratypes ♀ mean all Types mean all Types range all Types SD

SL 36.0 25.1 24.7 25.6 21.5–36.0 4.0
TL 119.7 124.4 122.5 123.2 119.7–127.0 2.2
HL 26.9 1.3
PD 26.3 26.6 64.9 26.7 25.1–28.9 2.0
pPED 66.0 65.1 27.9 65.0 61.2–67.8 1.1
pPD 27.7 27.4 46.6 27.6 25.9–29.8 1.3
pAD 45.3 47.2 57.5 46.8 44.7–49.4 1.5
DB 57.3 57.8 19.3 57.7 54.9–59.8 0.8
DC 19.5 19.8 11.4 19.6 18.7–21.7 0.5
CL 12.3 11.9 22.5 11.7 10.9–12.3 0.7
BD 23.3 22.4 15.7 22.5 21.1–23.6 1.0
BA 17.6 16.3 19.1 16.1 14.4–17.6 1.0
E 21.1 20.4 19.8 17.9–21.5 0.5
I 5.4 6.5 0.6
11.2 6.5 9.6 6.4 5.4–7.3
9.5 9.7 8.6–1.2

or dark brown dots on side, on some specimens nearly lin- dal peduncle, greatest body depth approximately anteri-
ear arranged, on others more irregular, in S. guignardi; or to origin of pelvic fin.
mostly yellow or whitish caudal fin margins in males, ir-
regular dark blotches on dorsal half of side in females, ac- Snout slightly pointed, mouth directed upwards, lower
cording to the first description (Boulenger 1908), in S. jaw longer than upper jaw, posterior end of rictus at the
liberiense. level of the dorsal third to centre of eye. Teeth curved and
unicuspid, on dentary an outer row of larger teeth, inner
Scriptaphyosemion wieseae males (Fig. 4A) differ from rows more or less regular with smaller teeth. Premaxilla
its closest relative, S. bertholdi (Fig. 5A), by coarser and with an outer row of larger teeth and a small number of
often denser red pigmentation on body and fins and broad- short teeth irregularly on the inner part. Closed frontal neu-
er dots or short streaks in the caudal fin, caudal fin trun- romast system within one groove, preopercular neuromast
cate vs. small dots on body and fins, especially in the cau- system with 5 pores.
dal fin, caudal rounded in S. bertholdi males. Females of
S. wieseae (Fig. 4B) have a midlateral more or less com- Scales cycloid, body and head completely scaled, ex-
plete dark stripe, often interrupted posterior to origin of cept for ventral surface of head. Frontal squamation of G-
anal fin on caudal peduncle, ventral half of sides below type; scales on lateral line 27–30 + 2–3 scales on caudal
midlateral dark stripe often intense orange in wild caught fin base. Transverse rows of scales above pelvic fin 9; cir-
specimens or orange-brown in aquarium kept specimens cumpeduncular scale row 12–14.
vs. irregularly distributed dark brown dots on scales on
dorsal half of side, pale brownish-orange on anterior side Tips of dorsal, anal, and, in some males, also caudal fin
in females of S. bertholdi (Fig. 5B). slightly pointed, but no filamentous extensions, caudal fin
truncate. Fins in females are generally smaller and more
Description. See Figure 4 for overall appearance and rounded. Dorsal and anal fin origin posterior to mid-body.
Table 3 for morphometric and meristic values of the type First dorsal fin ray posterior to origin of anal fin, situat-
series. Scriptaphyosemion wieseae shows a strong sexu- ed above anal fin ray 4–6. Dorsal fin with 12–13 rays, anal
al dimorphism with males more colorful, usually larger, with 16–17 rays, caudal fin with 25–28 fin rays. Pectoral
and with slightly extended fin rays in unpaired fins. Body fin rounded, fin origin on ventral half of side, just poste-
cylindrical, elongate, and laterally slightly compressed. rior to operculum, directed posteriorly, not reaching pelvic
Dorsal profile slightly convex, ventral profile anteriorly fin origin. Pelvic fin small, origin ventrally, directed pos-
slightly convex, slightly concave to nearly straight on cau- teriorly, not or just reaching anal fin origin. Total number
of vertebrates 29–30, of which 13–14 have pleural ribs and
15–16 have haemal spines.

Bonn zoological Bulletin 61 (1): 13–28 ©ZFMK

22 Rainer Sonnenberg & Eckhard Busch

Fig. 5. A) Male of Scriptaphyosemion bertholdi, SL 93/51, Sierra Leone, Victoria, not preserved. B) Female of Scriptaphyosemion
bertholdi, SL 93/51, Sierra Leone, Victoria, not preserved.

Live coloration. Males. (Fig. 4A) Lateral head blue with hue, a light, reflecting ring around pupil. Dorsal from head
a slightly irregular red pattern typical for the endemic West to dorsal fin brownish, scales with some blue in centre and
African genera Archiaphyosemion, Callopanchax, Nimba- irregular red margins, forming an interrupted reticulated
panchax, and Scriptaphyosemion and the related Epiplatys pattern. Flanks, unpaired fins and pelvic fins blue, dorsal
(Van der Zee 2002). Lips dark grey, red line around mouth and caudal fin centre often with a yellow hue, giving a
along upper and lower lip, small dark grey dot at chin, iris more greenish impression of fin color. Red pigmentation
of eye grey, dorso-anterior with blueish or blue-greenish on side in three to four very irregular stripes, consisting

Bonn zoological Bulletin 61 (1): 13–28 ©ZFMK

New Scriptaphyosemion from Sierra Leone 23

of red blotches, especially on caudal peduncle looking the Atlantic ocean through the southern costal area (Hu-
more like an interrupted reticulation. Unpaired and ber 2007; Wildekamp 1993; Wildekamp & Van der Zee
pelvic fins with a narrow blue margin, sometimes slight- 2003), whereas S. wieseae inhabits the upper tributaries
ly lighter than color of fin centre and body; the blue mar- of the Seli or Rokel River, which enter the Atlantic ocean
gin is followed by a broad red submargin. On base of dor- more to the north (Fig. 1). The geographically closest
sal, anal, pelvic, and centre of caudal fin a small number Scriptaphyosemion species with regard to the known col-
of red blotches, pectoral fin transparent with a blue mar- lection localities of S. wieseae is S. chaytori (Fig. 1), to
gin and a, more or less complete, red submargin. the north in Guinea it is S. guignardi (Busch 1995a;
Wildekamp & Van der Zee 2003; own unpublished col-
Females. (Fig. 4B) Body brownish, dorsally darker, on lection data)
ventral half of side and belly orange. Pigmentation on head
similar to males, but mostly darker brown instead of red. Etymology. The new species is named after Barbara
Midlateral dark brown stripe, interrupted and slightly ir- Wiese, who accompanied the second author on most of
regular on caudal peduncle, scales in this dark stripe some- his research trips in West Africa and was the first who rec-
times with small red scale margins; centre of unpaired and ognized it as a new Scriptaphyosemion species.
pelvic fins brown-orange, with a very narrow blueish mar-
gin, and a faint, narrow red submarginal, except on cau- DISCUSSION
dal fin. Above submarginal on anal and pelvic fins a small
area with blueish or greenish hue, centre of dorsal fin with Scriptaphyosemion wieseae is distinguished from the
small red dots, basally larger than marginally. Pectoral fin closely related S. bertholdi (Figs 2–3) by several diagnos-
transparent. tic characters, and is probably separated by strong genet-
ic incompatibilities (Vandersmissen 2003). As proposed
Color in ethanol. Males. Body brownish, darker on back by Roloff (1971a), female color characters can be useful
and lighter on belly, red pigmentation pattern of live spec- for distinguishing between some Scriptaphyosemion
imens is now carmine or dark brown, on ventral side of species, here by the diagnostic lateral color pattern differ-
head a small grey blotch on chin. Fin color light brown, ences between S. wieseae (Fig. 4B) and S. bertholdi (Fig.
unpaired fins with a whitish margin and a carmine or dark 5B). However, female pigmentation pattern can be ob-
brown sub-margin, some carmine or brown dots, pelvic scured by several factors, as e.g. stress, and vary in inten-
fins with a dark brown margin. sity (Roloff 1971a; own observations).

Females. Similar to males body brown with a darker The combination of high variability in male coloration
dorsal and lighter ventral part, on side the same dark between isolated populations of one species and the of-
brown pigmentation as in live specimens, on ventral side ten only small diagnostic color pattern differences between
of head a grey blotch as in males. Fins light brown, on species can make species identification in Script-
dorsal several small brown dots. aphyosemion difficult (Roloff 1971a; Scheel 1990).

Distribution and habitat. Scriptaphyosemion wieseae is The recent introduction of molecular methods in taxo-
currently known from three localities in the upland of nomic and phylogenetic studies of nothobranchiid killi-
northern Sierra Leone between the Sula and Wara Wara fishes added an additional and independent set of charac-
mountains (Fig. 1); all of these are located within the Seli ters and helped to resolve complicated cases of species lev-
or Rokel River basin. At the collection localities Fadugu el taxonomy (e.g. Legros & Zentz 2007b; Legros et al.
(SL 03/15) and Lenghe Curoh (SL 03/16), S. wieseae is 2005; Sonnenberg 2007, 2008). This approach can also be
present in small rivers, at the former place together with applied to resolve the taxonomy of Scriptaphyosemion by
Epiplatys cf. njalaensis, Archiaphyosemion guineense, un- clarifying the relationships between species and pinpoint-
determined characids, gobies, and cyprinids, probably ing diagnostic characters in terms of morphology and col-
Barbus leonensis, B. stigmatopygus and a small undeter- or pattern.
mined Barbus specimen; at the latter place it was collect-
ed together with A. guineense. At Kabala (SL 03/17), on- Our present results are largely congruent with those of
ly a small number of mostly juvenile specimens were col- Murphy et al. (1999), if only the well supported nodes are
lected in large rice swamps. The second author visited this considered. Differences e.g. in the placement of S. rolof-
area two times, in 1993 and 2003, and noticed an inten- fi are based on different opinions on the species level tax-
sified agricultural land usage. onomy. Incongruences in the placement of some species
and low support of several nodes might be based on in-
Collections of the second author during four collection sufficient information content in the dataset.
trips between 1989 and 2003 suggest that there is no over-
lap of the distribution areas of S. bertholdi and S. wieseae The perceived variability of male coloration in the ab-
(Fig. 1). Scriptaphyosemion bertholdi inhabits southeast- sence of other diagnostic characters and different species
ern Sierra Leone and is found in river basins which enter concepts led to varied opinions concerning species valid-

Bonn zoological Bulletin 61 (1): 13–28 ©ZFMK

24 Rainer Sonnenberg & Eckhard Busch

ity and number between authors (e.g. Costa 2008; Huber biospecies concept of Mayr (1963) is discussed in litera-
2007; Lazara 2001; Scheel 1968, 1990; Wildekamp 1993). ture (e.g. Otte & Endler 1989), and recent examples of hy-
Even decades after Roloff (1971a) and Scheel’s (1990) brid species and mitochondrial introgressions between
comments on the taxonomic difficulties, the species lev- species prove the possibility of fertile hybrids between dis-
el taxonomy of Scriptaphyosemion is still far from being tinct species (e.g. Nolte et al. 2005; Schliewen & Klee
resolved. Here we provide an overview of species names 2004; Schwarzer et al. 2011; Sonnenberg 2007). At the
in Scriptaphyosemion and their synonyms, with short com- moment we follow Costa (2008) in regarding them as three
ments on species whose taxonomy needs a revision. We separate species.
mainly follow the species level taxonomy as proposed by
Costa (2008); the species concept employed is that of The description of Scriptaphyosemion liberiense is
Moritz et al. (2000). based on preserved females (Boulenger 1908). It has a
large distribution in coastal Liberia (Wildekamp & Van der
The following species, S. bertholdi, S. brueningi, S. Zee 2003). Aphyosemion calabaricus Ahl, 1936, described
chaytori, S. fredrodi, and S. schmitti have comparatively on specimens from an ornamental fish import without se-
smaller distribution areas, and usually show less variabil- cure collection locality, and Epiplatys melantereon
ity in the diagnostic characters than the species listed be- Fowler, 1950 were considered as junior synonyms to S.
low. They are, with the exception of S. fredrodi, concor- liberiense (Costa 2008; Eschmeyer 2011; Huber 2007).
dantly considered as valid species by most authors (Cos- However, S. liberiense in its current usage is rather vari-
ta 2008; Huber 2007; Lazara 2001; Wildekamp & Van der able in male phenotype between different populations (Et-
Zee 2003). In addition, the studied samples of these zel 1992, 1993; Milkuhn 2007, 2008; Wildekamp 1993).
species show considerable genetic distance with regard to Roloff (1971b) proposed that S. calabaricum females dif-
their closest relatives within the molecular phylogenies fer in coloration pattern from S. liberiense as described
(Figs 2–3, Murphy et al. 1999), supporting their status as by Boulenger (1908) and might represent a different
separate species. species. A careful study of S. liberiense over its distribu-
tion range will be necessary to search for potentially sep-
Scriptaphyosemion geryi is distributed from northern arating stable diagnostic characters between groups of
Sierra Leone to the Senegal (Wildekamp 1993; populations and to decide about the status of S. calabar-
Wildekamp & Van der Zee 2003). Within this large area, icum. At the moment we tentatively consider this species
differences in coloration pattern between populations are as a synonym of S. liberiense.
known (e.g. Cauvet 2008; Etzel & Berkenkamp 1979;
Wildekamp 1993), indicating that probably more than one Scriptaphyosemion roloffi was accidentally described by
species is currently included under this name. Roloff (1936), the intended description by Ahl (1937) fol-
lowed a year later. Both authors gave a detailed color pat-
Scriptaphyosemion guignardi was the first of three for- tern description of the new species, drawings or black and
mally described species of a species complex (Romand white pictures, the latter ones from E. Roloff or W. Har-
1981, 1982; Seegers 1982). According to the DNA study vey, were published by different authors (e.g. Henzelmann
of Murphy et al. (1999), these three species form a mono- 1937; Mayer 1937; Ott 1997). By later authors (e.g. Et-
phyletic group. Costa (2008) listed them as valid species, zel 1978c; Scheel 1968, 1990; Wildekamp 1993) the iden-
but Etzel & Berkenkamp (1989a, b) and Etzel & Vander- tity of S. roloffi was confused due to the inclusion of dif-
smissen (1984), based on crossing experiments, regard S. ferent populations, which do not show the diagnostic char-
nigrifluvi and S. banforense as junior synonyms of S. guig- acters of the specimens described by Roloff (1936) and
nardi. A short comparison of the published cytochrome b Ahl (1937) and probably represent different species. Et-
sequences of Murphy et al. (1999) indicate considerable zel & Berkenkamp (1980a, b, c) presented the results of
genetic divergence between the three studied specimens crossing experiments between different populations,
(4.9–10.8 %) which is in the range of intraspecific diver- which were assumed to be S. roloffi, but most probably
gence values in killifishes (Murphy et al. 1999; Sonnen- included at least one currently undescribed species, judged
berg 2007, 2008; Sonnenberg & Blum 2005; Van der Zee from the published drawings. In 1979, S. etzeli was de-
& Sonnenberg 2011; own unpublished data). In addition, scribed by Berkenkamp, based on specimens which show
these species share some coloration characters, which in- the same coloration characters as given in the description
dicates their close phylogenetic relationship. Still they are of S. roloffi (Ahl 1937; Henzelmann 1937; Mayer 1937;
diagnosable by male coloration characters. Studies em- Roloff 1936). The synonymy of S. etzeli with S. roloffi in
ploying crossing experiments (e.g. Etzel & Berkenkamp the sense of Roloff (1936) and Ahl (1937) was discussed
1980a, b, c, 1989a, b; Etzel & Vandersmissen 1984; Ro- by Busch (1995b, 1996) and Ott (1997). We are of the
mand 1986; Scheel 1968, 1990) show, that between many opinion that S. etzeli is a junior synonym of S. roloffi,
populations different levels of genetic incompatibility ex- based on the diagnostic coloration characters of males and
ist. However, the conclusion that successful crossings up females, and that S. roloffi in the current usage
to the third generation indicate that they are conspecific (Berkenkamp 1979; Etzel & Berkenkamp 1980a, b, c; Et-
is viewed with strong hesitation. The difficulties of the

Bonn zoological Bulletin 61 (1): 13–28 ©ZFMK

New Scriptaphyosemion from Sierra Leone 25

zel 1978a, c; Etzel et al. 2004; Huber 2007; Wildekamp Scriptaphyosemion melantereon (Fowler, 1950): ANSP 72125,
1993) probably contains at least one undescribed species. Holotype, Liberia, Robertsport; ANSP 72126–27, Paratypes, col-
lected with holotype.
The incongruence of the placement of S. roloffi between Scriptaphyosemion roloffi (Roloff, 1936): ZMB 21071, Lecto-
our study (Figs 2–3) and Murphy et al. (1999) is caused type, Sierra Leone, tribuary of Port Loko Creek, ca. 40 km east-
by different taxonomic concepts. Their S. etzeli is in our northeast of Freetown, ZMB 31472, Paralectotypes, collected
view S. roloffi and the S. roloffi sample in their study most with Lectotype; ZFMK 47786–47798, Sierra Leone, Malai, col-
probably is an undescribed species. The voucher of Script- lection locality SL 03/11; ZFMK 47799–47813, Sierra Leone,
aphyosemion sp. in our DNA analyses (Figs 2–3) belongs Lungi Lol, collection locality SL 99/10; ZFMK 12678-80 (as S.
to the group of populations currently considered by oth- etzeli), Sierra Leone, Port Loko district, Kirma.
er authors as conspecific with S. roloffi. However, it does Scriptaphyosemion schmitti (Romand, 1979): ZFMK
not show the diagnostic coloration characters of this 47814–47815, Liberia, Juarzon, aquarium bred specimens.
species and belongs to a different genetic lineage, closer Scriptaphyosemion sp. aff. roloffi: ZFMK 47816–47819, Sier-
to S. fredrodi, and might probably be an undescribed ra Leone, Devil Hole, collection locality SL 03/18; ZFMK
species related to that labeled as S. roloffi in Murphy et 47820–47825, Sierra Leone, Kundubu, collection locality SL
al. (1999). Scriptaphyosemion hastingsi (Wright, 1972), 93/6; ZFMK 12656-58, Sierra Leone, Magbenta; ZFMK 14808-
also was an accidental description, but should be restud- 11, Sierra Leone, 15 km north Bo.
ied by new collections with regard to the species complex
currently included in S. roloffi, to which it is currently con- Acknowledgements. EB thanks Mohamed Basiru and family
sidered as a synonym (e.g. Eschmeier 2011; Huber 2007; (Brama Village, Sierra Leone) and the people of Brama Village
Wildekamp 1993). We tentatively consider it as a syn- for hospitality, organization and help during field work at four
onym, but it might turn out to be a valid name for a species research trips in Sierra Leone, Wilhelm Kugelmann
currently included in S. roloffi. (Dinkelscherben, Germany) for his support in keeping and breed-
ing many species collected on several journeys, and Barbara
For the two species names, Roloffia caldal Cahalan, Wiese (Remscheid, Germany) for her support on many research
1973 and Aphyosemion muelleri Scheel, 1968, we follow trips in West Africa since years. RS thanks Peter Bartsch and
other authors (Eschmeyer 2011; Huber 2007; Wildekamp Christa Lamour (MfN, Berlin, Germany), and Oliver Crimmen
1993), who consider them as invalid or not available, as and James Maclaine (NHM, London, Great Britain) for kind ac-
they were not mentioned by Costa (2008). cess to the collection under their care, Klaus Busse and Fabian
Herder (ZFMK, Bonn, Germany) for access to the collection and
COMPARATIVE MATERIAL the X-ray device, Kyle Luckenbill, John Lundberg, and Mark
Sabaj Perez (ANSP, Philadelphia, USA) for data, X-rays, and
Scriptaphyosemion bertholdi (Roloff, 1965): ZFMK photos of type specimens under their care, Diethard Tautz (MPI,
47708–47714, Sierra Leone, Victoria, collection locality SL Plön, Germany) for support with lab space and access to the mi-
93/51. cro Computer Tomograph, Heinz Otto Berkenkamp (Wil-
Scriptaphyosemion brueningi (Roloff, 1971b): ZFMK helmshaven, Germany), Gerhard Schreiber (Lüdenscheid, Ger-
47715–47725, Sierra Leone, Perie, collection locality SL 89 many), Thomas Litz (Attenweiler, Germany) for support with
Perie; ZFMK 47726–47735, Sierra Leone, Tawai, collection lo- Literature. We thank Günther Fleck (Saint-Gaudens, France) for
cality SL 03/2. the translation of the Abstract, Anja C. Schunke (MPI, Plön, Ger-
Scriptaphyosemion calabaricum (Ahl, 1936): ZMB 21039, Lec- many) for comments and corrections on an earlier version of the
totype, Nigeria (probably from Liberia); ZMB 31494, Paralec- manuscript, and Fabian Herder and two anonymous reviewers
totype, collected with the Lectotype. for valuable comments.
Scriptaphyosemion cauveti (Romand & Ozouf-Costaz, 1995):
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