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

2012·9.1 MB·English

by  Rainer Sonnenberg

2
Bonn zoological Bulletin 61 (1): 13-28 July 2012
Description of Scriptaphyosemion wieseae
(Cyprinodontiformes: Nothobranchiidae),
a new species from northern Sierra Leone
Rainer Sonnenberg1 & Eckhard Busch1
' Zoologisches Forschungstnuseum AlexanderKoenig. Adenauerallee 160, D-531J3 Bonn. Germany
- Max-Planck-InstitutfirEvolutionsbiologie, DepartmentofEvolutionaiy Genetics, August-Thienemann-Strasse 2,
D-24306 Plon, 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 ofa
small DNAstudy indicate, that5. bertholdi from the southeastern part ofthe country is the closest relative. Both species
can be distinguished by male and female color pattern and male caudal fin shape.
Resume. Scriptaphyosemion wieseae, une nouvelle espece, est decrite duNord de la Republique de SierraLeone. Lere-
ADN
sultatd"unepetite etudemoleculaire montreque sonespece sceurestS. bertholdi.presente au Sud-Estdu pays. Les
deux especes se distinguent par les couleurs du male et de la femelle etpar la forme de la nageoire caudale du male.
Key words. UpperGuinean ichthyofaunal province, taxonomy, DNA, phylogeny.
INTRODUCTION
TheWestAfricankillifish genus Scriptaphyosemion Rad- non which may confuse species identification; however,
da & Piirzl, 1987 is relatively species rich, compared to some ofthe most variable species might be composed of
the closelyrelated generaArchiaphyosemion Radda, 1977, more than one species (see e.g. figures or drawings in
Callopanchax Myers, 1933, andNimbapanchax Sonnen- Seegers 1997; Wildekamp 1993). This combination ofin-
berg & Busch, 2009. At present 18 specific names are traspecific variability, inclusion of probably different
available in Scriptaphyosemion (Eschmeyer 201 Huber species under one taxon, and the lack ofa specific diag-
1;
2007; Lazara2001),butonly 10 (Huber2007; 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 201 1; Lazara (1999) published the first moleculargenetic phylogeny of
2001; Scheel 1990; Wildekamp 1993). Among the West Scriptaphyosemion, based on the sequences ofthe 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 ofthe related genus Epiplatys Gill, 1862 (Costa tionships and an additional set ofindependent characters
2008; Huber 2007; Wildekamp & Van der Zee 2003). for further taxonomic and phylogenetic research.
Innothobranchiidkillifishes, 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. Agnese et al. 2009; Legros et al. Bissau, IvoryCoast, Mali, and Senegal (Huber2007; 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 derZee 2003). In Sierra Leone, Script-
2010; Sonnenberg et al. 2006, 201 1; Van derZee & 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
ternhas also been used as characterfordiagnosing species little known (Etzel 1977, 1978a, b, 1984; Huber 2007;
&
or groups ofspecies (Roloff 1971a). Wildekamp 1993; Wildekamp Van der Zee 2003).
Inmost species ofScriptaphyosemion, interspecific vari- In 1993, the second author collected together with B.
ance in male colorpattern is accompanied by intraspecif- Wiese nothobranchiids in the northernpartofSierraLeone
ic variance between and within populations, a phenome- (Busch 1995a), including a probably new species near
Received: 06.02.2012 Corresponding editor: R Herder
Accepted: 15.06.2012
14 Rainer Sonnenberg & Eckhard Busch
Table 1. List ofspecimens 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.
m
n
Epiplatys sexfasciatits RS1364 Gabon (j uZ//yqi1 JA1Z4Z/0, rJo/ZUjJ , rjolZKjHy
Scriptaphyosemion bertholdi RSI402 Sierra Leone 5L yj/j JTAYlllz/4IZOo^ll, JA1Z4ZJJ, JTAY1MZA41ZA41/
1
S. brueningi RS1377 Sierra Leone JTAY11Z744Z70AZ7, JTAYlMZ^AtlZ'jl'tA, JTAY11Z944Z74408
S. brueningi RS1403 Sierra Leone oy, rene JTAYlIZl'/tlZTOAJI, JTAY11Z744Z7J3J^, JTAY.1l7ZH4z7.4tQy
lYnn
S. cauveti RSI680 Guinea ulvl yi/Vj JTAY11Z7'4+Z70A44, JAIZ'+Zj-)O*;, JiAy1iZo4aZoJ^Uo
S. chaytori RSI379 Sierra Leone JlAY1HZ/4nZOAJS, JTAYl1Z7l4Z7J37/, JTAYl1Z7'AI7Z^j11
ftCLlfL/LtI RSI ^98 Sierra Leone oy, iviatanga JTAYI1OZz4lZO0fS0^, JTAYI1Z7^4Z7j1o8, JTAY11Z744Z73^7Z
S. geiyi RS1381 oierra Leone 5L UJ/U7, KOD1S 1 JTAY1iz7f4z7^o./7, rPTj8o7/7zfulj'?jV, PrTjSo7/7zfUi',H4-7/*
S. guignardi RS1380 Iu.u11i1n1*1e/ aO (nj\lvAl yQ7I/I11j1 JTAYl1Z7<447ZOAo8, JTAYl1Z1fAZ1j1yQ, JTAY11Z744Z9J^J1
S. liberiense RSI757 Liberia L97 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 Forcounts and measurements, see Sonnenberg Busch
found two additional collection localities ofthis 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), whichwas head, are presentedaspercentages ofstandard 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 fur Evolu-
logenetic position of the new species within Script- tionsbiologie in Plon, digital X-ray imagesweremadewith
aphyosemion, based on mitochondrial and nuclear DNA a Faxitron LX-60 Digital Specimen Radiography System
sequences. at the ZFMK. Fin rays ofunpaired fins were counted on
digital X-ray images; the relative position ofthe first dor-
sal fin ray to the anal fin (D/A) is estimated following Son-
MATERIALAND METHODS &
nenberg Schunke (2010). Nomenclature ofthe neuro-
mast system on the head follows Scheel (1968), for supra-
The species description is based on collections made in orbital (or frontal) squamation Hoedeman (1958). Male
Sierra Leone by the second author and B. Wiese during and female color patterns are compared with that given
their second visit in this area in November 2003. Speci- in the respective description and on pictures or drawings
mens were deposited in the Royal Museum for Central published in the literature.
Africa (MRAC, Tervuren, Belgium), the Zoologische As species concept we employ the pragmatic approach
Staatssammlung Miinchen (ZSM, Munich, Germany), and ofMoritz et al. (2000) and recognize species by diagnos-
the Zoologisches Forschungsmuseum Alexander Koenig tic combinations ofcharacters.
(ZFMK, Bonn, Germany).
Bonn zoological Bulletin 61 (1): 13-28 ©ZFMK
0
New Scriptaphyosemion from Sierra Leone 15
fat ' 1
Sierra Leone
o
.
On
Q
Liberia
km
50 A A
Fig. 1. Distribution map ofScriptaphyosemion in Sierra Leone based on data from Huber (2007) and own data ofthe second au-
thor. White square: 5. 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 ofS. bertholdi and S. wieseae.
ADNAdatasetwasprepared for 15 Scriptaphyosemion Sequences were aligned with ClustalX 1.8 (Thompson
specimens, including 11 species, andEpiplatyssexfascia- et al. 1997), checkedbyeye, 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 ofthree species are not complete, missing sites
ments ofthe mitochondrial and nuclearLSU rDNAgenes. were filledwithN [S. liberiense (560 bp), S. schmitti (616
Parts ofthe sequences were published in earlierpapers of bp), and S. cauveti (58 bp)].
the authors; all samples are listed with GenBank acces- The protein coding gene was translated into the corre-
sion numbers and collection locality in Table 1. For de- sponding amino acid sequence to check for a functional
tails on the DNA methods, see Sonnenberg & Busch coding sequence, and base composition ofmitochondri-
MEGA
(2009, 2010) and Sonnenberg et al. (2007). al sequences was estimated with 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 ofmissing data were calculated in MEGA4.1 be-
= S. cauveti (Romand & Ozouf, 1995), CHY = S. chay- ta (Tamura et al. 2007). Numberofconstant, variable, and
tori (Roloff, 1971a), FRE = S. fredrodi (Vandersmissen informative characters were estimatedwith PAUP4.0b1
et al., 1980), GER = S. geryi (Lambert, 1958), GUI = S. (Swofford 1998).
guignardi (Romand, 1981), LIB = S. liberiense Data were analyzedusing a maximum parsimony (MP)
(Boulenger, 1908), ROL = S. roloffi (Roloff, 1936), SMI and a Bayesian approach (BA). Maximum parsimony
=S. schmitti(Romand, 1979), Ssp =S. sp. SL03/18, WIE analyses were performed with PAUP 4.0b10 (Swofford
= S. wieseae, new species, SEX = E. sexfasciatus. 1998) with the exclusion ofuninformative positions and
Bonn zoological Bulletin 61 (1): 13-28 ©ZFMK
16 Rainer Sonnenberg & Eckhard Busch
BER
1402
99 / 98 / 82 j- W1E 1382
100 / 100/ 95 l wie 1687
j
i-WIE 1686
CHY
1379
911121
I
"1
GUI 1380
100/ 100/ 100 Ssp 1378
FRE 1398
100 100 HOOT BRU 1377
/
96/91/53 100/lOOj L BRl 1403
1 LIB 1757
ROL
100 85 69 1376
/ / —
GER
1381
CAU
1680
SMI 1755
—
SEX 1364
50
Fig. 2. Results ofthe maximum parsimony analyses. Shown is one out ofthree equally short trees from the analysis with the
complete dataset. Values left to nodes are the corresponding bootstrap values ofall three analyses (complete dataset/ thirdprotein
coding position excluded/alignment columns with missing data excluded). Values fornodes were only given, ifat leastone 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 rootedwithEpiplatyssexfasciatus as out-
performed with random addition ofsequences 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 ofsequences 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 withthe exclusion ofthird
protein coding positions, and a third with exclusion ofre-
gions, in which one of the above listed specimens has RESULTS
missing data. Otherwise, all parameters were identical.
Bayesian analyses were performedwith MrBayes 3.1.2 DNAAnalysis. (Figs 2-3, Table 2). The complete align-
(Huelsenbeck & Ronquist 2001), datawere partitioned in ment ofthe ND2, tRNA, and both LSU sequences has a
mitochondrial non protein codingregions (tRNA, mtLSU), length of3199 bp including alignment introducedgaps for
protein coding regions (ND2), and the nuclear LSU se- atotal of16 specimens (503 bp tRNAs, 1047 bpND2,481
quences. Numberofgenerationwas setto 10,000,000 with bp mtLSU rDNA, 1168 bp nLSU rDNA). Ofthese, 2278
sampling ofevery 1,000 generations andthe results ofthe bp are constant, 456 bp variable but uninformative, and
first 10% ofthe generations werediscarded asbum-intime 465 bp parsimony informative. The ND2 sequence trans-
after checking that the runs had reached the stationary lates into 349 amino acids withno unexpected stop codon,
phase atthispoint. Fouranalyses wereperformed: the two thebase composition ofall mitochondrial genes shows the
alignments, one with the complete data set and one with typical base composition for this organelle genome with
the exclusion ofregions, in which one ofthe above men- ananti-G bias (Zhang & Hewitt 1996).Averageuncorrect-
tioned specimens has missing data, were both performed edpairwise distance forall sequences is 7.4 %, maximum
with NST=2 and NST=6. observed distance is 16.6 %between outgroup and ingroup
specimens, maximum observed distancebetween ingroup
specimens is 9.0 %, further details are given in Table 2.
Bonn zoological Bulletin 61 (1): 13-28 ©ZFMK
i
New Scriptaphyosemion from Sierra Leone 17
t-
90 NO
NO CN NO
W
CN
NO ri
NO
oo NO
NO CN d NO
1—1
CN
dC^ d NO ri
o NO
Odn d d NO
idn CdN d d ri
o
cn r- ro CN NO
oo oo On
CN (~ CN p ro O NO
>n CN >n ro in ro
NdO 10.5/ NO 10.5/ NO 10.4/ oo p
CN CN CN
ro
CN oq oq CN oq oq ri CN
NO co in in in ro
o
dCN in On oq in in oq n
cn CN
m ro >n
CN oq 00 r~ in r-~ in NO ro
in NO NO CN NO CN NO CN
o
CN ^ r- CN Cd- 10.8/ CN 11.2/ CN 11.1/ CN NCNO
CN
On
On NO ON CN CN CN ro
>n ^J- ro' in ro in ro
OriN oCNq dC- 9.5/ 9.8/ m 10.1/ NO C-Nr NNOO
m O NO ON
<n in NO oq ON NO '—J NO NO CO
no <n r~ NO NO ro mi ro NO ro
oq oo in CN OO NO
ro CN ri On r-- oq ro CN CN ri CN ri CN CN NO
ri On ri ri ri ri
NO ON
ro in ON >n ro NO ON ro ro NO NO 5; NO ro
00 NO <n in ri NO 'T in r-' ro ro ro
OdN CN d d ON CN NO CO in ro ON ro CN NNOO
CN CN CN OO oo OO
CN o NO ON © ©" NO in CN CN p CN t-- NO
OO no in NO a-' NO <*' «n ro >n ri >n CN
9.9/ ro 12.8, ro 9.9/ o oroi in CCNN OdN 10.1, Od 8.5/ o 8.4/ o 8.4/ On CN NO
CN CN ro CN CN
NO r- O CN oq ?F 00 t-- CO d ro p ro CN ro CN
NO p~ NO ro NO NO CN NO <n NO in ro >n ro in ro
CN in 00 p ro m ro p
-r oq in oq >n in in r~ in CN ON ro in ro oq oq On rj
ri ri ri ri ON ri
NO NO o p oo CN ON oq 00 NO CN no CN NnOo oo
00 OO On ro oo 00 ro in 00 00 ro 00 ro 00 ro
14.4, NO 12.2, 11.7, OdN 13.0, ON 12.8, ro 4.3/ >dn 12.1, OdN * 2— oq NdO dNO ro d ^C1N- NNOO
ri
CN OO p NO NO r-- CN NO On OO On oq <r CN CN ro CN >n CN in rNoO
oo ro ro NO ro r-' ri NO ro >n ro NO NO ri NO ri NO CN
0.9/ od 14.6, NO 12.5, 12.1, OdN 13.4, ON 13.0, ro 4.4/ md 12.1, OdN 11.0, 11.9, OdN 00 NdO oq NdO 11.9, rd-- rCNo NNOO
ri
© CN
o d CN oq CN NO 00 On CN in NO ON oo ON 00 NO CN ro in in in NO
OO ro' r-' ro NO ro r--' ri NO ro in ro NO ro NO ri NO ri NO ri
ro OdN p OdN in o CN o ON ro ri ro pi r- >n oq oq CN OdN O in O— r-t CN
ri 00 ri oo ri ON CN On in
W ro CN ro CN in ro in NO ON O NO NO ro in ON oq —<i p
l£> ri NO ri 00 ro >n ri in ro ro in ro in ro NO ri NO ri ri i—i ro
o
rrr--o reTo 0N0O OrrN-o 0Or0No 0r0O o0r0o rir-> NrrO-o IITf) 0r-0 r0rio0 N0NO0O 0N(0O- NrrrOo
B B B 2 i—i i-H ro U u 1-H X
Qt as < ow o -- o s a. Id
aa oa at »2 GO
Bonn zoological Bulletin 61(1): 13-28 ©ZFMK
18 Rainer Sonnenberg & Eckhard Busch
BER
1402
1.0/1.0 WIE 1382
WIE 1686
-1.0/1i"o£ WIE 1687
CHY
1379
1.0/1.0
1.0/1.0*
GUI 1380
1— Ssp 1378
1.0/1.0
-FRE
1398
ROL
1376
0.96/0.99 1.0/1.0* GER
1381
^j-BRU
1377
1.0/
1.0/1.0 —I- BRU 1403
LIB 1757
CAU
1680
SMI 1755
SEX 1364
0.02
B
BER
1402
r
i.o/i.o
WIE 1687
i.o/To|r W> IE 1382
WIE 1686
0.98/0.98
GUI 1380
0.95/0.89* CHY
1379
ROL
1376
GER
0/1.0 1381
0.97/0.99
Ssp 1378
-Fre 1398
1.0/1.O0 r B»iRU 1377
1.0/1.0p-bBiRU 1403
LIB 1757
CAU
1680
SMI 1755
SEX 1364
0.02
Fig. 3. Results ofthe Bayesian analyses. Shown is the consensus tree ofthe analysis with the complete dataset and NST=6 (A)
and the consensus tree ofthe dataset with excluded alignment columns with missing data andNST=6 (B). Values left to nodes are
the corresponding posterior probability values, except for values marked with * which are on the right side ofthe 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, ifat least one analysis has a posteri-
or probability above 0.95.
Bonn zoological Bulletin 61 (1): 13-28 ©ZFMK
New Scriptaphyosemion from Sierra Leone L9
All three maximum parsimony analyses result in near- species. The genetic distance values (Table 2) show asim-
ly the same tree topology and differonly in branch length ilar divergence as between S. liberiense and 5. bruenin-
and bootstrap support values; here, only one ofthree short- gi-
esttrees ofthe complete dataset is presented (Fig. 2). The
Bayesian analyses for the complete and the reduced
datasets show slightly different tree topologies, independ- Scriptaphyosemion wieseae, new species
ent if the analyses were performed with NST=6 or 2. (Fig. 4, Table 3)
Shown are the results for the complete and the reduced Scriptaphyosemion sp. Lenghe Curoh: Busch (1995a),
dataset of the NST=6 analyses, on the nodes posterior Vandersmissen (2003)
probabilities forbothanalyses foreach datasetarepresent-
ZFMK mm
ed ifthe values are equal to or above 0.95 (Fig. 3 A+B). Holotype. 47680, male, 36.1 SL, Sierra
Scriptaphyosemion schmitti andS. cauveti are support- Leone, Lenghe Curoh, Sanghi or Sanigi River, fieldnum-
ed as basal to all other species ofthe genus (Figs 2-3). ber SL03/16 (9°28,04'N, 1 1°40,58'W), E. Busch and B.
As S. schmitti was placed in the study of Murphy et al. Wiese, 18. Nov. 2003.
(1999) within a clade containing the other Script-
aphyosemion species studied, we checked for possible Paratypes. ZFMK 47681-47692, 8 males, 4 females,
mm
causes ofthis discrepancy. As the sequences studied here 19.5-34.7 SL, paratopotypes, collectedwiththe holo-
have no overlap with those ofMurphy et al. (1999), we type.
MRAC mm
compared a cytochrome b fragment of our sample with B2-06-P-1-2, 1 male, 22.9 SL, 1 female, 21.5
mm
the data deposited in GenBank. However, both samples SL, paratopotypes, collected with the holotype.
ZFMK mm
differ in only one nucleotide (results not shown), and a 47693^17707, 7 males, 8 females, 18.8-41.3
Blast searchwithouranalysed sequences (seeTable 1) on- SL, Sierra Leone, Fadugu, atthe bridge overthe Mabakah
ly returned the highest similarity with other Script- River, field number SL 03/15 (9°23,52'N, 11°45,76' W),
aphyosemion sequences. To exclude that the basal posi- E. Busch and B. Wiese, 18. Nov. 2003.
tionofS. schmittiis an artifactdue tothe missing sequence ZSM 41596 (2), 1 male, 28.4 mm SL, 1 female, 22.3 mm
data, we also performed the analyses with the reduced SL, collected with ZFMK 47693^17707.
dataset, which did not contain missing data, but gave the
sameresults (Figs 2-3). However, furthersamples should Non-type DNA samples. RSI686, 1 male, DNA vouch-
% ZFMK
be studied for a confirmation ofthe basal position, espe- er, preserved in 96 ethanol, collected with
cially as this node is supported neither by bootstrap nor 47693^17707, field number SL 03/15; RS1382, 2 males,
DNA %
posterior probability values. 1 female, voucher, preserved in 96 ethanol, col-
Our results indicate a closer relationship between S. lected with the holotype; RSI687, 1 male, DNA vouch-
%
liberiense andS. brueningi, S. roloffiandS. getyi, S.fred- er, preserved in 96 ethanol, Sierra Leone, Kabala, col-
rodi and a probably undescribed species (see under Dis- lected in rice swamp, field number SL 03/17 (9°35' N,
cussion), and S. bertholdi and S. wieseae, new species.A n°33' W), E. Busch and B. Wiese, 18. Nov. 2003.
closer relationship between S. chaytori and S. guignardi
occurs in all resulting trees (Figs 2-3), but is well sup- Diagnosis. Scriptaphyosemion wieseae is distinguished
ported only in the Bayesian analyses of the complete from its congeners by a unique combination ofmale and
dataset. Both species were in addition grouped with S. female coloration characters. Males have a blue body col-
roloffi and S. geryi in the analyses ofthe reduced dataset, or, red dots on side forming irregular, interrupted red
but not in the maximum parsimony analyses and the stripes, especially on caudal peduncle a more reticulated
Bayesian analyses ofthe complete dataset (Figs 2-3). pattern, unpaired and pelvic fins centre blue with a nar-
All analyses support a monophyletic group formed by row blue margin and a broad red submargin, on anal and
all Scriptaphyosemion except ofS. schmitti and S. cau- dorsal fin a small number ofbasal red blotches, centre of
veti (Figs 2-3). Another well supported subgroup is caudal fin with some red blotches, pectoral transparentto
formed by S. bertholdi, S. wieseae, new species, S. chay- orange with a blue margin; females with a dark brown
tori, S. guignardi, S.fredrodi, and S. sp. (see under Dis- midlateral stripe, interrupted on caudal peduncle, bodyon
cussion); only in the analyses of the reduced dataset S. lower halfof side and ventrally orange vs. a mainly red
roloffi and S. geryi were also included in this group (Fig. lateral body color in male S. cauveti; head and anterior
2-3). To this subgroup, S. brueningi and S. liberiense, body yellowish to brownish, unpaired fin margins white
probably isthe sistergroup, ahypothesis only significant- oryellow, andpectoral fn margin yellowto orange inmale
ly supported in the Bayesian analyses (Fig. 3). S. schmitti; males with more or less regular oblique red
The phylogenetic results show that S. bertholdi is the bars on side, at least on the caudal peduncle, fin margins,
well supported sister species to S. wieseae, new species, especially of caudal and pectoral, often yellow, and fe-
and indicates no closer relationship to other mentioned males with dark brown streaks or blotches, similar to the
Bonn zoological Bulletin 61 (1): 13-28 ©ZFMK
20 Rainer Sonnenberg & Eckhard Busch
Fig.4. A)Male ofScriptaphyosemion wieseae,collection locality SL03/16, notpreserved. B)FemaleofScriptaphyosemion wieseae,
collection locality, SL 03/16, not preserved.
red oblique bars ofmales on side, often with a darkblotch ularly distributed dark brown or red dots on side in S.
at the root ofthe caudal in S. geryi and S. roloffi; males chaytori; males with dark red blotches on side, caudal fin
withbroadred blotches, often formingbroad irregularver- 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 followedby abroaderred band, females short streaks on dorsal halfofcaudal peduncle and a dark
with irregularly distributed dark brown dots or short blotch on posterior end ofcaudal peduncle at dorsal base
oblique streaks orblotches dorsally on the caudal pedun- ofcaudal 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 ofthe caudal peduncle at the streaks or dots, submarginal stripes ofunpaired 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
1
New Scriptaphyosemion from Sierra Leone 21
Table 3. Morphometries ofScriptaphyosemion wieseae, new species. Values in the table are based on 15 specimens, for which
all measurementsandcountsweretaken (7 ofeach maleand female paratypes).All measurements in percentagesofstandard length
(SL) except standard length in mm. TL = total length, HL = length ofhead, 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 ofcaudal peduncle, BD = base ofdorsal fin, BA= base ofanal fin, E = eye diameter, I = interorbital width, SD = standard
deviation.
11U1UIyI-* 1 Ml«ljUC3 ( ) lllvdll PUaillrUaillt1vVnJ'p1sv Q_[_ mlleltattnll atillll T1vJnUpVsJ m111pK.a*In1 atillll TUvVnft1o1\si r1aUlnl*f*lpt a(liln Tvynnile\s.v SDJ
36 n 25.1 24.7 25.6 21 5-36 0 4.0
TI 119.7 124.4 122.5 123.2 119 7-127 0 2.2
HL 26.3 26.6 26.9 26.7 25.1-28.9 1.3
PD 66.0 65.1 64.9 65.0 61.2-67.8 2.0
pPED 27.7 27.4 27.9 27.6 25.9-29.8 1.1
pPD 45.3 47.2 46.6 46.8 44.7-49.4 1.3
pAD 57.3 57.8 57.5 57.7 54.9-59.8 1.5
DB 19.5 19.8 19.3 19.6 18.7-21.7 0.8
DC 12.3 11.9 11.4 11.7 10.9-12.3 0.5
CL 23.3 22.4 22.5 22.5 21.1-23.6 0.7
BD 17.6 16.3 15.7 16.1 14.4-17.6 1.0
BA 21.1 20.4 19.1 19.8 17.9-21.5 1.0
E 5.4 6.5 6.5 6.4 5.4-7.3 0.5
I 11.2 9.5 9.6 9.7 8.6-1.2 0.6
ordarkbrown 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 ofpelvic fin.
mostly yellow or whitish caudal fin margins in males, ir- Snout slightly pointed, mouth directed upwards, lower
regulardarkblotches on dorsal halfofside in females, ac- jaw longer than upperjaw, posterior end ofrictus at the
cording to the first description (Boulenger 1908), in S. level ofthe dorsal third to centre ofeye. Teeth curved and
liberiense. unicuspid, on dentary an outer row oflarger teeth, inner
Scriptaphyosemion wieseaemales (Fig. 4A) differfrom rows more or less regular with smaller teeth. Premaxilla
its closest relative, S. bertholdi (Fig. 5A), by coarser and with an outer row oflarger teeth and a small number of
often denserredpigmentationonbody and fins andbroad- shortteeth irregularly onthe innerpart. Closed frontal neu-
er dots or short streaks in the caudal fin, caudal fin trun- romast system within one groove, preopercularneuromast
catevs. small dots on body and fins, especially in the cau- system with 5 pores.
dal fin, caudal rounded in S. bertholdi males. Females of Scales cycloid, body and head completely scaled, ex-
S. wieseae (Fig. 4B) have a midlateral more or less com- cept forventral surface ofhead. Frontal squamation ofG-
plete dark stripe, often interrupted posterior to origin of type; scales on lateral line 27-30 + 2-3 scales on caudal
anal fin on caudal peduncle, ventral halfof sides below fin base. Transverserows ofscales above pelvic fin 9; cir-
midlateral dark stripe often intense orange in wild caught cumpeduncular scale row 12-14.
specimens or orange-brown in aquarium kept specimens Tips ofdorsal, anal, and, in some males, also caudal fin
vs. irregularly distributed dark brown dots on scales on slightlypointed, but no filamentous extensions, caudal fin
dorsal halfofside, pale brownish-orange on anterior side truncate. Fins in females are generally smaller and more
in females ofS. bertholdi (Fig. 5B). rounded. Dorsal and anal fin origin posteriorto mid-body.
First dorsal fin ray posterior to origin ofanal fin, situat-
Description. See Figure 4 for overall appearance and ed above anal fin ray4-6. Dorsal finwith 12-13 rays, anal
Table 3 for morphometric and meristic values ofthe type with 16-17 rays, caudal fin with 25-28 fin rays. Pectoral
series. Scriptaphyosemion wieseae shows a strong sexu- fin rounded, fin origin on ventral halfofside,just poste-
al dimorphism with males more colorful, usually larger, riorto operculum, directedposteriorly, notreachingpelvic
andwith slightly extended finrays in unpaired fins. Body fin origin. Pelvic fin small, origin ventrally, directed pos-
cylindrical, elongate, and laterally slightly compressed. teriorly, not orjust reaching anal fin origin. Total number
Dorsal profile slightly convex, ventral profile anteriorly ofvertebrates 29-30, ofwhich 13-14havepleuralribs and
slightlyconvex, slightly concave to nearly straight on cau- 15-16 have haemal spines.
Bonn zoological Bulletin 61 (1): 13-28 ©ZFMK
22 Rainer Sonnenberg & Eckhard Busch
Fig. 5. A) Male ofScriptaphyosemion bertholdi, SL93/51, SierraLeone,Victoria, notpreserved. B) Female ofScriptaphyosemion
bertholdi, SL 93/51, Sierra Leone, Victoria, not preserved.
Live coloration. Males. (Fig. 4A) Lateral headblue with hue, a light, reflecting ring aroundpupil. Dorsal from head
aslightly irregularredpatterntypical forthe endemic West to dorsal fin brownish, scales with some blue in centre and
AfricangeneraArchiaphyosemion, Callopanchax,Nimba- irregular red margins, forming an interrupted reticulated
panchax, andScriptaphyosemion andthe relatedEpiplatvs pattern. Flanks, unpaired fins and pelvic fins blue, dorsal
(Van derZee 2002). Lips darkgrey, red line around mouth and caudal fin centre often with a yellow hue, giving a
alongupper and lower lip, small dark grey dot at chin, iris more greenish impression offin color. Red pigmentation
ofeye 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