![Scales of Chelon persicus (SMF 35897 [KAU13-125], 189 mm SL) stained with alizarin red. A: body scale from the anterior trunk above the pectoral fin; B: predorsal scale from midline anterior to the first dorsal fin; C. interorbital scale. Scale bar = 500µm.](https://www.researchgate.net/profile/Sergey-Bogorodsky/publication/363566286/figure/fig3/AS:11431281114068077@1674224043240/Scales-of-Chelon-persicus-SMF-35897-KAU13-125-189-mm-SL-stained-with-alizarin-red_Q320.jpg)
![Fresh specimens of Chelon persicus. A: SMF 35898 [KAU13-427], 66.7 mm SL, Al Wajh, Saudi Arabia, Red Sea; B: SMF uncat [QG17-83], 260 mm SL, Qatar, Arabian/Persian Gulf; C: SMF uncat [QG17-115], 275 mm SL, Qatar, Arabian/ Persian Gulf. Photos by S.V. Bogorodsky.](https://www.researchgate.net/profile/Sergey-Bogorodsky/publication/363566286/figure/fig4/AS:11431281114093273@1674224043357/Fresh-specimens-of-Chelon-persicus-A-SMF-35898-KAU13-427-667-mm-SL-Al-Wajh-Saudi_Q320.jpg)
![Detailed photographs of the head of fresh specimens of Chelon persicus. A: SMF uncat [QG17-83], 260 mm SL, Qatar, Arabian/Persian Gulf; B: detailed photographs of the head of SMF uncat [QG17-115], 275 mm SL, Qatar, Arabian/ Persian Gulf. Photos by S.V. Bogorodsky.](https://www.researchgate.net/profile/Sergey-Bogorodsky/publication/363566286/figure/fig5/AS:11431281114068078@1674224043472/Detailed-photographs-of-the-head-of-fresh-specimens-of-Chelon-persicus-A-SMF-uncat_Q320.jpg)
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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by B. Frable: 23 Aug. 2022; published: 15 Sept. 2022 1
Zootaxa 5188 (1): 001–042
https://www.mapress.com/zt/
Copyright © 2022 Magnolia Press Article
https://doi.org/10.11646/zootaxa.5188.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:24E869D4-DDB4-4113-A9E9-478904048F9D
Contributions to the taxonomy of the mugilid genus Chelon Artedi
(Teleostei: Mugilidae), with a major review of the status of C. persicus Senou,
Randall & Okiyama, 1995
PHILIPP THIEME1,2*, SERGEY V. BOGORODSKY3,4, TILMAN J. ALPERMANN3, ALAN K. WHITFIELD5,
RUI FREITAS6 & JEAN-DOMINIQUE DURAND2
1Deutsches Meeresmuseum, Katherinenberg 14-20, 18439 Stralsund, Germany.
�
phil.thieme2016@gmail.com; https://orcid.org/0000-0002-3065-1272
2IRD, MARBEC, Université de Montpellier, IRD, CNRS, cc093, Place E. Bataillon, 34095 Montpellier Cedex 05, France.
�
jean-dominique.durand@ird; https://orcid.org/0000-0002-0261-0377
3Research Institute and Natural History Museum Frankfurt, Senckenberganlage 25, 60325 Frankfurt a.M., Germany.
�
Tilman.Alpermann@senckenberg.de; https://orcid.org/0000-0002-8723-4576
4Station of Naturalists, Omsk, Russia.
�
ic187196@yandex.ru; https://orcid.org/0000-0002-8679-9735
5South African Institute for Aquatic Biodiversity (SAIAB), Private Bag 1015, Grahamstown 6140, South Africa.
�
a.whitfield@saiab.nrf.ac.za; https://orcid.org/0000-0003-1452-7367
6Institute of Engineering and Marine Sciences, Atlantic Technical University, Mindelo 163, Cabo Verde
�
rfreitas@uta.cv
*Corresponding author
Abstract
New data based on multigene phylogenetic analyses using the COI, 16S, and cytb genes and subsequent molecular
species delimitation revealed that the mugilid genus Chelon contains 11 species. Of these, two species, Chelon sp. A and
Chelon sp. B, remained unidentified in previous studies. While Chelon sp. B seemingly is a close relative to C. dumerili
(Steindachner, 1870), no more detailed information were provided for Chelon sp. A which is present along the east coast
of South Africa. Genetic analyses performed in this study, revealed that specimens from the Arabian/Persian Gulf and
the Red Sea are nested within one genetic lineage alongside Chelon sp. A. Morphological, morphometric, and meristic
analyses of specimens from all three areas demonstrated that they belong to Chelon persicus Senou, Randall & Okyiama,
1995. Chelon persicus was originally described in the genus Chelon but was later placed either in the genus Liza or
Planiliza. However, results presented herein confirm its placement in Chelon and its distribution range is extended in
the Western Indian Ocean from the type locality (Arabian/Persian Gulf) west to the Red Sea and south to South Africa.
Further, the uncertain validity of the recently described Chelon caeruleus Deef, 2018 from the Mediterranean Sea and
a previously proposed generic character distinguishing Chelon from Planiliza and Parachelon, the shape of the paired
postzygapophyses of the second vertebra, are briefly discussed.
Key words: Mugiliformes, Planiliza, phylogenetic analysis, Arabian/Persian Gulf, range extension
Introduction
In the last decade, studies on the phylogeny of the family Mugilidae provided a much-needed update on the
systematic relationships of species of different mugilid genera (Durand et al. 2012a; Durand et al. 2012b; Durand &
Borsa 2015; Xia et al. 2016). Xia et al. (2016) recognized four subfamilies (Myxinae, Mugilinae, Rhinomugilinae,
and Cheloninae) based on the most comprehensive and integrative analysis of morphological and genetic characters.
One of them is the subfamily Cheloninae which comprises six genera, Chelon Artedi, 1793, Neochelon Durand et
al., 2012, Oedalechilus Fowler, 1903, Parachelon Durand et al., 2012, Planiliza Whitely, 1945 and Pseudomyxus
Durand et al., 2012, and can be distinguished from the other subclades by a set of six morphological characters. But,
Cheloninae Xia, Durand & Fu, 2016 is objectively invalid as the name is already occupied by Cheloninae Nees,
1816, a subfamily of parasite wasps.
THIEME ET AL.
2 · Zootaxa 5188 (1) © 2022 Magnolia Press
Durand et al. (2012a) proposed to synonymize the genus Liza with Chelon and to keep the latter name for
the species as they demonstrated that Mugil capito Cuvier, 1829, the type species of the genus Liza Jordan &
Swain 1884, and Mugil chelo Cuvier, 1829, the type species of the genus Chelon, share the same common ancestor
(Durand et al. 2012a; Durand et al. 2012b). Furthermore, they suggested reinstating the genus Planiliza as the name
for all the remaining Chelon and Liza species not derived from the common ancestor of Mugil capito and Mugil
chelo (Durand et al. 2012a).
The two genera can be distinguished by an osteological character and their distribution area. The relative size
and shape of the postzygapophysis of the second vertebra is short but gently curved and hook-like in Chelon
but long and curved with a blunt tip in Planiliza (Xia et al. 2016). Species currently recognized in the genus
Chelon are distributed in the Mediterranean Sea, Eastern Atlantic Ocean and along the eastern coast of South
Africa north to Mozambique, while species currently recognized in the genus Planiliza are distributed from the
Western Indian Ocean to the West Pacific. Only few species from both genera overlap in their distribution ranges,
i.e., C. richardsonii (Smith, 1846) and C. tricuspidens (Smith, 1935), with Planiliza alata (Steindachner, 1892), P.
macrolepis (Smith, 1846) and P. melinopterus (Valenciennes, 1836), along the eastern coast of South Africa (Durand
& Whitfield 2015).
Previous genetic analyses revealed two unidentified species within the genus Chelon (Chelon sp. A and Chelon
sp. B) from South Africa (Durand et al. 2012a; Durand et al. 2012b; Durand 2015; Durand & Borsa 2015). While
Chelon sp. B is closely related to Chelon dumerili and resembles it morphologically, Chelon sp. A is seemingly more
closely related to C. richardsonii and C. tricuspidens (Durand & Borsa 2015). However, Chelon sp. A did not match
any previous species description from its distribution area, i.e., the eastern coast of South Africa.
In order to assign the unidentified species Chelon sp. A to a species name, we examined morphological
and molecular characters of additional material and compared it with available genetic datasets and previous
morphological descriptions.
One candidate species was Chelon persicus Senou, Randall & Okiyama, 1995 which had first been described
in detail from material collected in the Arabian/Persian Gulf (Senou et al. 1996). Carpenter et al. (1997) placed
this species in the genus Liza, probably assuming the absence of the genus Chelon from the Arabian/Persian Gulf.
More recently, the species was placed in the genus Planiliza (Eagderi et al. 2019; Fricke et al. 2022) because the
distribution area of C. persicus is more typical for species of this genus. In its formal description, Senou et al.
(1996) noted the presence of one pair of long spine-like postzygapophyses on the second vertebra as a characteristic
feature of Chelon persicus. However, Xia et al. (2016) identified this as a distinguishing character of Parachelon
grandisquamis (Valenciennes, 1836), the only representative of the monospecific genus Parachelon within the
subfamily Cheloninae.
In this paper, we have conducted a comparative analysis of specimens from the Arabian/Persian Gulf, Red Sea
and South Africa, combined with molecular data, to reveal which species Chelon sp. A represents. Furthermore, we
analysed the distribution ranges of all eleven species comprised in the genus Chelon.
Material and methods
Sampling
A total of 17 specimens from the Red Sea, Arabian/Persian Gulf, and Cabo Verde were sampled and sequenced in
this study (Appendix 1). A fin clip from each specimen was collected and stored in a 1.5 ml Eppendorf tube with
90–96% ethanol, and later stored at -20°C.
Genetic analyses
Genomic DNA was isolated from fin clips using the G-Spin Total DNA extraction minikit (iNtRON Biotechnology,
Jungwon-gu, Gyeonggi, Korea) following the manufacturer’s recommendations. A portion of each 16S ribosomal
RNA gene (16S), cytochrome oxidase I gene (COI), and cytochrome b gene (cytb) was amplified by polymerase
chain reaction (PCR) using the primers listed in Durand et al. (2012b). Polymerase Chain Reaction (PCR) was
conducted in a total volume of 40 µL containing 20 µL of MyTaq PCR Mastermix (Bioline, London, UK), 16 µL of
ultrapure water, 0.8 µL of BSA (Euromedex, Souffelweyersheim, France), 0.6 µL of each primer (3 M), and 2 µL of
DNA template. The conditions used during PCR reaction were as follows: initial denaturation temperature at 92°C
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 3
for 5 min followed by 35 cycles of strand denaturation at 92°C for 1 min, primer annealing at 52°C for 45 s (except
for 16S with 50°C annealing temperature), primer extension at 72°C for 1.5 min, and a final extension at 72°C for 5
min. Sanger sequencing of both strands was performed by Genoscreen (Lille, France) with the same primers used in
PCR amplification. DNA sequences were edited, aligned, and concatenated using SeaView v.4 (Gouy et al. 2010).
All assembled nucleotide sequences were deposited in GenBank (see Table 1 for accession numbers).
Phylogenetic analyses
Phylogenetic relationships among species of the genus Chelon were investigated using a dataset consisting of the
concatenation of the three mitochondrial fragments (1936 base pairs) obtained for seven specimens analysed in
this study, plus all reference Chelon species (23 specimens + 4 outgroup specimens) used in previously published
phylogenetic studies that either investigated the genus or species delineation in the family Mugilidae (Durand et al.
2012a; Durand et al. 2012b; Durand & Borsa 2015) (see Table 1).
Phylogenetic analyses were conducted based on a partitioned maximum-likelihood (ML) method as implemented
in IQ-TREE (Nguyen et al. 2015). The partitions were set by gene and the best Substitution Model was identified
using ModelFinder (Kalyaanamoorthy et al. 2017). The Best-fit model according to Bayesian information criterion
scores was K2P+G4 for the 16SRNA section, HKY+F+G4 for the COI section and TPM3+F+G4 for the Cytb section.
Nodal support was assessed with Ultrafast bootstrapping (Hoang et al. 2018) based on 1,000 pseudo-replicates. The
ML analysis was conducted with the IQ-Tree web server (Trifinopoulos et al. 2016). The phylogenetic trees were
visualized and edited under FIGTREE v1.4.0. Homologous sequences of Parachelon grandisquamis were chosen
as outgroups to root the reconstructed phylogenetic trees (see Table 1).
Species delimitation analyses
The COI barcode dataset (Appendix 1) was analyzed for the purpose of delimiting species using three different
DNA-based approaches: the Automatic Barcode Gap Discovery (ABGD) algorithm (Puillandre et al. 2012), the
Bayesian Poisson Tree Processes (PTP) model (Zhang et al. 2013) and the Refined Single Linkage (RESL) as
implemented in BOLD and used to produce Barcode Index Numbers (BINs; Ratnasingham & Hebert 2013). All
approaches cluster nucleotide sequences into operational taxonomic units (OTUs) or putative species without a
priori species hypotheses. The ABGD algorithm analysis was conducted by using the online version of ABGD
(http://wwwabi.snv.jussieu.fr/public/abgd/) with the K2P (Kimura 1980) pairwise distance model; other parameters
were set as default. While species delimitation with the PTP model was estimated on the PTP webserver (http://
species.h-its.org/ptp/) with 500,000 Markov-Chain Monte Carlo (MCMC) generations and the default parameters.
Chelon species distribution
The distributional ranges of Chelon species were re-evaluated by downloading all available geographic references
of specimen barcodes assigned to Barcode Index Numbers (BINs) in the BOLD system lists for Chelon species
owing to RESL algorithm.
Morphological analyses
Eight Red Sea specimens and two Arabian/Persian Gulf specimens from the collection of the Senckenberg Research
Institute and Natural History Museum in Frankfurt (SMF) and the King Abdulaziz University Marine Museum
(KAUMM) in Jeddah as well as one specimen from the east coast of South Africa stored in the collection of the
South African Institute for Aquatic Biodiversity (SAIAB) were examined. The eight specimens from the Red Sea
previously were provisionally identified as Planiliza macrolepis.
Morphometric measurements were conducted using a digital calliper to the nearest 0.1 mm. Measurements and
meristic data were taken from all specimens following Grant & Spain (1975) and Senou et al. (1987) and are defined
as follows: Standard length (SL), distance between anterior-most tip of the upper jaw and base of the caudal fin (end
of hypural plate); Body depth, vertical distance through the origin of the first dorsal fin to the ventral margin of the
abdomen; Body width at origin of first dorsal fin, horizontal width of the body at the origin of the first dorsal fin;
Body width at origin of pectoral fins, distance between the bases of the pectoral fins; Head length, distance between
anterior-most tip of upper jaw to middle part of the posterior margin of the opercle (excluding gill membrane); Head
width, horizontal distance between posterior bony edges of the left and right orbit; Head depth, distance between the
dorsal and ventral surface of head vertical through the posterior bony edge of orbit; Snout length, distance between
THIEME ET AL.
4 · Zootaxa 5188 (1) © 2022 Magnolia Press
the anterior-most point of upper lip and anterior bony edge of orbit; Postorbital length, distance from posterior
bony edge of orbit to middle part of the posterior margin of the opercle (excluding gill membrane); Orbit diameter,
distance between anterior and posterior bony edge of orbit; Interorbital width, distance between the dorsal bony
edge of orbits; Upper jaw length, distance between anterior-most point/midpoint of upper jaw to the lateral tip of
the maxilla; Width of mouth, distance between angles of closed mouth; Thickness of upper lip, height of the upper
lip at the middle; Caudal peduncle depth, minimal distance vertical between dorsal and ventral surface of caudal
peduncle; Caudal peduncle width, maximal width horizontal at the hypural joint; Caudal peduncle length, oblique
distance between end of anal-fin base and middle of base of the caudal fin (end of hypural plate); Prefirst dorsal fin
length, distance between anterior-most point of upper jaw and origin of first dorsal-fin; Presecond dorsal fin length,
distance between anterior-most point of upper jaw and origin of second dorsal-fin; Preanal fin length, distance
between anterior-most point of upper jaw and origin of anal-fin; Prepelvic fin length, distance between anterior-
most point of upper jaw and origin of pelvic-fin; Base of second dorsal fin, distance between anterior and posterior
insertion of second dorsal fin; Base of anal fin, distance between anterior and posterior insertion of anal fin; Longest
first dorsal fin spine, length of the longest (first) spine of the erect first dorsal fin; Longest second dorsal fin ray,
length of the longest fin ray of the erect second dorsal fin; Longest anal fin ray, length of the longest fin ray of the
erect anal fin; Caudal fin length, distance between base of caudal fin and level of posterior tip of caudal fin; Pectoral
fin length, distance between origin of the pectoral fin and posterior tip of pectoral fin; Pelvic fin length, distance
between origin of the pelvic fin and posterior tip of pelvic fin; Pectoral axillary scale length, length of the elongated
scale in the pectoral axis measured from insertion of pectoral fin to posterior tip of axillary scale.
Additionally, x-ray images of one specimen, SMF 35897 [KAU13-125], were taken and the same specimen was
dissected to examine its gut. Scale samples were taken from SMF 35897 [KAU13-125] which were stained with
alizarin red. Specimens collected from Qatar (Qatar Museum uncat [QG17-83] and Qatar Museum uncat [QG17-
115]) were also photographed, examined, and barcoded.
Results
Genetic analyses
The Chelon phylogenetic tree comprised 11 evolutionary lineages that were well delineated by all methods used
to determine the number of OTUs or BINs (Fig. 1). Ten species names are available for these 11 BINs, with the
specimens from South Africa named as Chelon sp. A by Durand & Borsa (2015) belong to the same BIN/OTU as
specimens from the Red Sea and Arabian/Persian Gulf identified as Chelon persicus. The remaining unidentified
BIN/OTU, provisionally named Chelon sp. B by Durand & Borsa (2015), consists of specimens from South Africa
morphologically identified as C. dumerili but genetically highly divergent (7.5% under the K2P model) from other
West African C. dumerili, including some collected at the type locality of St Louis in Senegal (Fig. 1). These two
BINs remained phylogenetically close; the node being well supported. Other sister species relationships stressed
in the phylogenetic tree concern C. tricuspidens and C. bandialensis (Diouf, 1991), as well as C. labrosus (Risso,
1827) and C. bispinosus (Bowdich, 1825).
Chelon species’ distribution ranges inferred from DNA barcoding data mostly match three biogeographic areas:
NE Atlantic and Mediterranean Sea for C. auratus (Risso, 1810), C. ramada (Risso, 1827), C. labrosus and C.
saliens (Risso, 1810) (Fig. 2A–D); West Africa for C. bispinosus, C. bandialensis and C. dumerili (Fig. 2E– G); and
South Africa for Chelon sp. B, C. richardsonii and C. tricuspidens (Fig. 2H– J).
An exception to the above general trends is the distribution of Chelon persicus which is exclusively present in
the Western Indian Ocean (Fig. 2K) while all other Chelon species are present in the Atlantic Ocean. The known
distribution range of C. persicus recovered from DNA barcodes encompasses both the southwestern portion of the
Indian Ocean and the Arabian/Persian Gulf plus the Red Sea. Furthermore, C. ramada barcodes have also been
obtained from specimens collected in the Red Sea (Fig. 2B, Appendix 1); however, these specimens were caught
close to a drainage canal associated with an aquaculture facility and the presence of an established population of
C. ramada in the Red Sea has yet to be verified. In addition, a few specimens sampled in Senegal, Cameroon,
Angola and South Africa were identified morphologically as C. saliens (Appendix 1), extending its distribution
range southwards (Fig. 2D).
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 5
FIGURE 1. Phylogenetic relationships among species (corresponding to Table 1) belonging to the genus Chelon inferred by
the maximum-likelihood analyses of the partitioned concatenated alignment method with Kimura 2 Parameters + G nucleotide
substitution model based on the combined mitochondrial genes dataset. Branch length is proportional to inferred number of
nucleotide substitutions. Numbers at nodes represent bootstrap values in percentages. Values < 70% are not shown. The tree
was rooted with Parachelon grandisquamis. Results from species delimitation analyses based on COI gene sequences using the
RESL, ABGD and PTP algorithms, as well as species name identified for each evolutionary lineage, are mentioned on the right
of the tree. Chelon species labeled sp. “B” were morphologically identified as Chelon dumerili in South Africa and is potentially
a new species.
THIEME ET AL.
6 · Zootaxa 5188 (1) © 2022 Magnolia Press
FIGURE 2. Distribution ranges of Chelon spp. inferred from GBIF occurrence data obtained from geolocalised preserved
specimens (blue dots) as well as DNA barcodes available in BOLD (consultation February 2022) plus some unpublished data
(red dots, Appendix 1). Doubtful GBIF reference are pointed by a “?”. A: Chelon auratus BIN BOLD:AAD7360; B: Chelon
ramada BIN BOLD:AAD5443; C: Chelon labrosus BIN BOLD:AAE2670; D: Chelon saliens BIN BOLD:AAF6190; E: Chelon
bispinosus BIN BOLD:AEO1527; F: Chelon bandialensis BIN BOLD:ACC0109; G: Chelon dumerili (GBIF data correspond
to Chelon dumerili spp.) BIN BOLD:ACC0658; H: Chelon sp. B (GBIF data correspond to Chelon dumerili spp.) BIN BOLD:
AAD7493; I: Chelon richardsonii BIN BOLD:AAD7487; J: Chelon tricuspidens BIN BOLD:AAF6205; K: Chelon persicus
BIN BOLD:AAC0698.
Morphological analysis
The eight examined specimens collected from the Red Sea, the two specimens from the Arabian/Persian Gulf
and the specimen from South Africa were identified as Chelon persicus. The diagnosis is based on the following
characters provided by Senou et al. (1996): a comparison of morphometric measurements (Table 2) and meristic
characters (Table 3), dentition, a pair of long spine-like postzygapophyses of the second vertebra, three short and
three long unbranched pyloric caeca, ctenoid scales on the body, and predorsal scales with a single groove. We
identified additional traits, some of which were already mentioned by Senou et al. (1996), that are characteristic of
Chelon persicus.
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 7
TABLE 1. List of specimens used in phylogenetic reconstruction.
Species BIN Individual
no.
Voucher
no.
Id BOLD GenBank nos. Sampling location Date Collector(s)
(Durand & Borsa,
2015)
COI 16S rRNA cyt b
Chelon auratus BOLD:
AAD7360
168 - ANG-
BF5180-12
JQ060456 GQ252691 JQ060199 Goulette, Tunisia Apr 03 H. Blel
Chelon auratus BOLD:
AAD7360
169 - ANG-
BF5304-12
JQ060457 JQ060704 JQ060200 Agadir, Morocco - A. Pariselle
Chelon auratus BOLD:
AAD7360
171b - ANG-
BF5179-12
JQ060458 JQ060705 JQ060201 Dakhla, Western
Sahara
Mar. 2010 A. Pariselle
Chelon auratus BOLD:
AAD7360
171c - ANG-
BF5303-12
JQ060459 JQ060706 JQ060202 Khnifiss, Morocco Mar. 2010 A. Pariselle
Chelon auratus BOLD:
AAD7360
171d - ANG-
BF5178-12
JQ060460 JQ060707 JQ060203 Khnifiss, Morocco Mar. 2010 A. Pariselle
Chelon
bandialensis
BOLD:
ACC0109
54 MNHN
2009-
0790
ANG-
BF5302-12
JQ060461 JQ060708 JQ060204 Saloum Estuary,
Senegal
2008 O. Diouf
Chelon
bandialensis
BOLD:
ACC0109
56 - ANG-
BF5177-12
JQ060462 JQ060709 JQ060205 Bubaque, Bijagos
Islands, Bissau Guinea
May 2008 Y. Bettarel
Chelon
bispinosus
BOLD:
AEO1527
CbiCV-01 - MULID001-
21
OP028827 OP028829 OP028831 Porto Grande Bay,
Cape Verde
10 Dec. 2013 R. M. Freitas
Chelon
bispinosus
BOLD:
AEO1527
CbiCV-02 - MULID002-
21
OP028823 OP028828 OP028830 Porto Grande Bay,
Cape Verde
10 Dec. 2013 R. M. Freitas
Chelon dumerili BOLD:
ACC0658
73 SAIAB-
83185
ANG-
BF5301-12
JQ060463 JQ060710 JQ060206 Saloum Estuary,
Senegal
06 Mar. 2009 O. Sadio, J.
Raffray
Chelon dumerili BOLD:
ACC0658
76 - ANG-
BF5176-12
JQ060464 JQ060711 JQ060207 St Louis, Senegal 26. Jan 08 J.-D. Durand
Chelon dumerili BOLD:
ACC0658
77 - ANG-
BF5300-12
JQ060465 JQ060712 JQ060208 Fish market, Lome,
Togo
19 June 2007 J.-F. Trape
Chelon dumerili BOLD:
ACC0658
078b - ANG-
BF5175-12
JQ060466 JQ060713 JQ060209 Fish market, Bissau,
Guinea
04 May 2010 J.-D. Durand
......continued on the next page
THIEME ET AL.
8 · Zootaxa 5188 (1) © 2022 Magnolia Press
TABLE 1. (Continued)
Species BIN Individual
no.
Voucher
no.
Id BOLD GenBank nos. Sampling location Date Collector(s)
(Durand & Borsa,
2015)
COI 16S rRNA cyt b
Chelon labrosus BOLD:
AAE2670
179b - ANG-
BF5327-12
JQ060411 JQ060660 JQ060154 Khnifiss, Morocco Mar. 2010 A. Pariselle
Chelon labrosus BOLD:
AAE2670
180 - ANG-
BF5202-12
JQ060412 GQ258711 JQ060155 Goulette, Tunisia 22 Dec. 2005 H. Blel
Chelon labrosus BOLD:
AAE2670
178 - ANG-
BF5166-12
JQ060484 JQ060729 JQ060227 Agadir, Morocco - A. Pariselle
Chelon persicus BOLD:
AAC0698
161 SAIAB-
78131
ANG-
BF5162-12
JQ060492 JQ060734 JQ060235 St Lucia, South Africa 14 May 2005 A.K. Whitfield,
J.-D. Durand, J.
Panfili
Chelon persicus BOLD:
AAC0698
KAU13-
135
SMF
35897
MULID015-
22
ON943238 ON943243 ON943247 Red Sea S. Bogorodosky
Chelon persicus BOLD:
AAC0698
KAU13-
125
SMF
35897
MULID016-
22
ON943236 ON943242 ON943246 Red Sea S. Bogorodosky
Chelon persicus BOLD:
AAC0698
KAU13-
124
SMF
35897
MULID017-
22
ON943234 ON943241 ON943245 Red Sea S. Bogorodosky
Chelon persicus BOLD:
AAC0698
KAU13-
123
SMF
35897
MULID018-
22
ON943231 ON943240 ON943244 Red Sea S. Bogorodosky
Chelon persicus BOLD:
AAC0698
QG17-
083
LID-
MA3629-22
OP028368 OP028365 OP028366 Qatar, Arabian/Persian
Gulf
S. Bogorodosky
Chelon ramada BOLD:
AAD5443
173 - ANG-
BF5293-12
JQ060479 GQ258707 JQ060222 Goulette, Tunisia Apr 03 H. Blel
Chelon ramada BOLD:
AAD5443
174b - ANG-
BF5168-12
JQ060480 JQ060726 JQ060223 Zerga, Morocco Mar. 2010 A. Pariselle
Chelon
richardsonii
BOLD:
AAD7487
157 SAIAB-
64270
ANG-
BF5292-12
JQ060481 JQ060727 JQ060224 Orange River Estuary,
Namibia
12 June 2001 J. Koekemoer
Chelon
richardsonii
BOLD:
AAD7487
158 - ANG-
BF5167-12
JQ060482 JQ060728 JQ060225 Great Fish Estuary,
South Africa
12 May 2005 A.K. Whitfield,
J.-D. Durand
......continued on the next page
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 9
TABLE 1. (Continued)
Species BIN Individual
no.
Voucher
no.
Id BOLD GenBank nos. Sampling location Date Collector(s)
(Durand & Borsa,
2015)
COI 16S rRNA cyt b
Chelon saliens BOLD:
AAF6190
166 - ANG-
BF5291-12
JQ060483 GQ258709 JQ060226 Goulette, Tunisia Apr 03 H. Blel
Chelon sp. B BOLD:
AAD7493
79 - ANG-
BF5299-12
JQ060467 JQ060714 JQ060210 Great Fish Estuary,
South Africa
12 May 2005 A.K. Whitfield,
J.-D. Durand
Chelon sp. B BOLD:
AAD7493
80 - ANG-
BF5174-12
JQ060468 JQ060715 JQ060211 Great Fish Estuary,
South Africa
12 May 2005 A.K. Whitfield,
J.-D. Durand
Chelon
tricuspidens
BOLD:
AAF6205
155 - ANG-
BF5285-12
JQ060495 JQ060740 JQ060238 Sundays Estuary,
South Africa
08. Aug 07 A.K. Whitfield
Parachelon
grandisquamis
BOLD:
ACC0632
88 - ANG-
BF5296-12
JQ060473 JQ060720 JQ060216 Saloum, Sénégal 06 Mar. 2009 O. Sadio, J.
Raffray, L. Tito
Parachelon
grandisquamis
BOLD:
ACC0632
089b - ANG-
BF5171-12
JQ060474 JQ060721 JQ060217 Fish market, Bissau,
Guinea
04 May 2010 J.-D. Durand
Parachelon
grandisquamis
BOLD:
ACC0632
90 MNHN
2009-731
ANG-
BF5295-12
JQ060475 JQ060722 JQ060218 Saloum, Sénégal 06 Mar. 2009 O. Sadio
Parachelon
grandisquamis
BOLD:
ACC0632
91 SAIAB-
83182
ANG-
BF5170-12
JQ060476 JQ060723 JQ060219 Saloum, Sénégal 06 Mar. 2009 O. Sadio
THIEME ET AL.
10 · Zootaxa 5188 (1) © 2022 Magnolia Press
Chelon persicus Senou, Randall & Okiyama, 1995
Persian Mullet
Figures 3–6, Tables 2 & 3
Chelon persicus Senou et al., 1995: 235 (species account without designation of type material)—Senou et al. 1996: 72 (Doha
fish market, Qatar, Arabian/Persian Gulf; formal species description); Al-Jufaili et al. 2010: 32 (Arabian/Persian Gulf).
Liza persicus—Carpenter et al. 1997: 203 (Arabian/Persian Gulf).
Planiliza persicus—Eagderi et al. 2019: 87 (Arabian/Persian Gulf).
Chelon sp. A—Durand & Borsa 2015: 8 (South Africa).
Diagnosis (based on combined data from Senou et al. (1996) and present study). Second dorsal-fin rays 10; anal-fin
rays 9–10 (usually 10); scales finely ctenoid, 33–36 in longitudinal series; predorsal scales with a single groove;
caudal fin forked; upper lip with unicuspid, tiny, ciliform teeth in one row; ventral edge of preorbital notched at
corner of mouth, posterior end of maxilla exposed when mouth closed; tip of pelvic fin is vertical to or behind origin
of first dorsal fin; second vertebra with a pair of long, slightly curved, spine-like postzygapophyses; three short and
three long unbranched pyloric caeca; pectoral-fin base silvery, without melanophores; reaches 275 mm SL.
Material examined. SMF 35897 [sample tissue KAU13-123], 194.2 mm SL, [KAU13-124] 187.8 mm SL,
[KAU13-125] 188.9 mm SL, [KAU13-135] 200.7 mm SL, Red Sea, Saudi Arabia, Al Wajh, lagoon, st 5, 13 June
2013; KAUMM 459 [KAU13-136], 172.0 mm SL, [KAU13-137] 188.4 mm SL, [KAU13-138] 170.5 mm SL, Red
Sea, Saudi Arabia, Al Wajh, lagoon, st 5, 13 June 2013; SMF 35898 [sample tissue KAU13-427], 66.7 mm SL, Red
Sea, Saudi Arabia, Al Wajh, lagoon, st 15, 15 June 2013; Qatar Museum uncat [sample tissue QG17-83], 260 mm
SL, Arabian/Persian Gulf, Qatar, off Doha, 03 December 2017; Qatar Museum uncat [sample tissue QG17-115],
275 mm SL, Arabian/Persian Gulf, Qatar, off Doha, 10 December 2017; SAIAB 83180, 106.2 mm SL, Western
Indian Ocean, South Africa, KwaZulu-Natal Province, Cape Vidal, 15 May 2007.
Description (based on material examined). Body relatively elongate, body depth at origin of first dorsal fin
3.4–3.7 in SL and caudal peduncle compressed. Head depth equal to or greater than head width, head length 4.1–4.3
in SL. Snout relatively blunt, subequal to orbit diameter, eye diameter 3.3–4.8 in head length. First dorsal fin with
IV spines, second fin with 9–10 (rarely 9) rays. Origin of first dorsal fin closer to base of caudal fin than to tip of
snout. Origin of second dorsal fin above first quarter of anal fin. Anal fin with III spines and 10 rays. Pectoral-fin
rays 15–17. Tip of pectoral fin not reaching vertical at origin of first dorsal fin and not behind tip of pelvic-fin spine.
If laid forward, pectoral fin not reaching anterior edge of eye. Tip of pelvic fin vertical to or behind origin of first
dorsal fin. Caudal fin forked. Three supraneurals present and one each positioned between neural spines of vertebrae
two and three, four and five, and six and seven. A pair of postero-dorsally elongated, slightly curved, and spine-like
postzygapophyses present on second vertebra. Anterior first dorsal fin pterygiophore positioned posterior to neural
spine of seventh vertebra. Anterior pterygiophores of second dorsal as well as anal fin positioned anterior to neural
and haemal spine of thirteenth vertebra, respectively. Three interdorsal pterygiophores present. Three short and
three long unbranched pyloric caeca attached to stomach.
Adipose eyelid covers only margins of eye; adipose tissue covers larger area anterior to eye reaching anterior
nostril. Preorbital fills space between eye and mouth. Preorbital serrated along antero-ventral to posterior edge
starting from above corner of mouth (Fig. 3). Ventral edge of preorbital notched at corner of mouth. Postero-ventral
corner of preorbital slightly rounded or truncate and reaching vertical behind anterior edge of eye. Corner of mouth
vertical below anterior edge of posterior nostril. Maxilla reaching below corner of mouth, its posterior tip sigmoidal
and visible when mouth closed. One row of minute, unicuspid, ciliform teeth present on upper lip. Lower lip with
thin edge, directed horizontally forward, and large symphysial knob.
Body scales finely ctenoid (Fig. 4), 33–35 scales in longitudinal series. Scales in transverse rows 11–12,
predorsal scales to origin of first dorsal fin 11–13, to origin of second dorsal fin 21–23, circumpeduncular scales
16–18 (Table 3). Predorsal scales anterior to first dorsal fin with single groove. Scales on head cycloid and arranged
irregularly. Interorbital scales large, decreasing in size posteriorly. Dorsal scales on head forward to, or nearly to,
posterior nostrils. Minute scales present on base of all fins, more prominent on the second dorsal, anal, caudal, and
pectoral fins; covering approximately proximal third of first and second dorsal fin and anal fin. Two consecutive
axillary scales lateral on each side of first dorsal fin, posterior axillary scales reaching behind base of last ray of first
dorsal fin. Axillary scales also present lateral to origin of pelvic fin reaching about halfway along pelvic-fin spine.
Very small axillary scale present above pectoral fin.
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 11
TABLE 2. Morphometric measurements of examined specimen as percentages of standard length in comparison to Chelon persicus (Senou et al. 1996).
Chelon
persicus
SAIAB
83180
SMF
35897
[KAU12-
123]
SMF
35897
[KAU13-
124]
SMF
35897
[KAU13-
125]
SMF
35897
[KAU13-
135]
KAUMM
459
[KAU13-
136]
KAUMM
459
[KAU13-
137]
KAUMM
459
[KAU13-
138]
SMF
35898
[KAU13-
427]
SMF
uncat
[QG17-
83]
SMF
uncat
[QG17-
115]
Standard length
(mm)
195.4–248.9 106.2 199.5 195.0 194.9 207.0 180.5 195.6 175.7 68.6 260.0 275.0
Total length 124–128 - 123.0 123.2 121.0 124.2 123.4 123.0 124.2 123.8 - -
Body depth
origin D1
24–28 27.8 28.3 32.0 28.3 28.5 30.5 27.8 29.2 29.6 29.9 27.0
Body width
origin D1
12.0–14.7 11.1 14.9 13.9 14.7 16.5 19.9 17.2 17.2 12.8 - -
Body width
origin PF
14.5–15.5 15.0 16.8 17.3 16.4 17.5 19.1 16.4 16.0 16.7 15.8 15.6
Head length 23–24 26.5 22.8 23.6 22.4 24.5 24.8 23.9 22.5 27.6 22.7 22.7
Head width – 16.1 13.6 13.8 13.0 12.8 14.6 13.1 13.1 14.9 14.0 -
Head depth 12.3–13.9 15.0 12.8 12.2 12.8 14.1 15.6 13.4 13.9 16.9 12.9 12.7
Snout length 6.6–7.1 6.1 5.6 5.7 6.2 6.2 7.0 5.9 6.0 6.1 5.3 5.0
Orbit diameter 5.4–6.1 7.9 5.6 5.4 5.9 5.4 5.7 5.4 5.6 8.2 5.6 5.4
Interorbital width 8–9 10.1 11.0 11.2 10.0 10.9 11.8 10.0 10.6 10.4 9.8 10.1
Upper-jaw length - 7.2 5.0 5.1 5.4 5.0 6.1 5.7 5.2 5.4 5.4 5.1
Width of mouth 7.1–7.8 9.8 8.1 8.2 8.1 8.6 9.2 8.4 8.0 10.5 - -
Thickness upper
lip
1.4–1.7 1.4 1.5 1.5 1.6 1.8 1.9 1.6 1.8 2.2 - -
Postorbital length 11.4–12.7 13.9 12.7 12.3 11.1 13.1 13.3 12.5 12.0 13.9 12.1 12.0
Caudal-peduncle
depth
11–12 12.3 12.1 12.9 12.1 12.5 12.7 12.4 12.5 11.5 11.5 12.5
Caudal-peduncle
width
- 3.1 3.3 3.0 2.9 2.9 3.7 2.9 2.6 2.6 2.7 2.8
Caudal-peduncle
length
17–19 17.1 17.9 18.5 18.7 17.1 - 19.8 17.5 15.9 17.2 17.9
......continued on the next page
THIEME ET AL.
12 · Zootaxa 5188 (1) © 2022 Magnolia Press
TABLE 2. (Continued)
Chelon
persicus
SAIAB
83180
SMF
35897
[KAU12-
123]
SMF
35897
[KAU13-
124]
SMF
35897
[KAU13-
125]
SMF
35897
[KAU13-
135]
KAUMM
459
[KAU13-
136]
KAUMM
459
[KAU13-
137]
KAUMM
459
[KAU13-
138]
SMF
35898
[KAU13-
427]
SMF
uncat
[QG17-
83]
SMF
uncat
[QG17-
115]
Predorsal length
D1
51–54 55.2 54.3 52.0 51.6 54.4 58.5 54.8 53.8 55.7 55.2 53.5
Predorsal length
D2
76–78 78.7 78.9 78.5 75.3 80.2 80.8 79.6 78.2 78.6 80.7 76.7
Preanal-fin
length
73–80 76.5 78.0 75.0 73.3 76.9 80.4 74.7 76.1 76.7 80.0 76.0
Prepelvic length 39–41 40.4 39.1 39.0 37.9 41.6 42.5 39.8 39.5 43.6 42.3 40.0
Base of D2 7.6–8.4 7.6 7.6 7.6 8.5 8.0 8.2 7.5 7.5 8.2 8.2 8.0
Base of anal fin 8.4–10.1 10.0 10.3 10.0 10.1 10.3 10.4 10.9 10.4 10.6 10.2 10.6
Longest spine
in D1
12.1–15.0 13.9 13.2 15.1 13.9 14.9 14.2 13.8 15.1 15.9 13.6 13.2
Longest D2 fin
ray
14.6–15.7 14.9 13.8 14.6 14.9 15.4 14.4 13.8 14.9 14.9 13.8 14.1
Longest anal-fin
ray
14.5–15.8 15.2 14.3 16.5 14.9 16.4 13.6 16.6 16.5 15.7 14.2 13.4
Caudal-fin length - - 23.0 23.2 24.7 24.2 23.4 23.0 24.2 23.8 23.0 22.8
Pectoral-fin
length
17.9–18.9 20.4 20.2 20.8 18.3 20.2 20.9 19.4 19.7 21.7 19.0 17.8
Pelvic-fin length 14.4–15.7 16.9 16.0 16.7 15.2 17.1 17.5 16.1 16.6 17.1 14.7 15.0
Axillary scale
length
4.9–6.0 4.4 4.5 5.6 4.4 5.3 3.8 4.9 4.9 4.1 - -
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 13
FIGURE 3. Head of Chelon persicus. A & B: SMF 35897 [KAU13-125], 189 mm SL, with important characters marked,
Asterisk = single row of minute teeth on upper jaw.
FIGURE 4. Scales of Chelon persicus (SMF 35897 [KAU13-125], 189 mm SL) stained with alizarin red. A: body scale from
the anterior trunk above the pectoral fin; B: predorsal scale from midline anterior to the first dorsal fin; C. interorbital scale.
Scale bar = 500µm.
Fish coloration shown in Figs. 5 and 6. Grey dorsally with greenish tinge, silvery below, without markings but
with faint narrow grey stripes along centre of scale rows; pectoral-fin base silvery, without melanophores; caudal
fin with narrow posterior black margin.
Distribution and habitat. Currently known from scattered localities in the Western Indian Ocean, including
the Arabian/Persian Gulf, Red Sea, Mozambique and South Africa (Fig. 2K). This species forms schools in shallow
coastal waters, including lagoons and estuaries.
Remarks. In the formal description of C. persicus it was noted that both the second dorsal and anal fin usually
comprise nine fin rays (Table 3). In the specimens from this study, we counted ten fin rays (only one specimen has
nine fin rays in the second dorsal fin). This difference may be due to the general assumption that the last fin ray
constitutes one deeply bifurcate ray or double ray. However, osteological studies on mugilids showed that two fin
rays with separate bases are present (Ghasemzadeh 2015; Thieme & Moritz 2020). We were also able to observe the
separate bases in specimen SMF 35897 [KAU13-125].
The species name was first included in the monograph on coastal fishes of Oman (Randall 1995) accompanied
by a species account (without designation of types), and was later formally described by Senou et al. (1996). We
follow the former publication and subsequent Catalog of Fishes (Fricke et al. 2022) regarding the date of first
mention of C. persicus.
THIEME ET AL.
14 · Zootaxa 5188 (1) © 2022 Magnolia Press
FIGURE 5. Fresh specimens of Chelon persicus. A: SMF 35898 [KAU13-427], 66.7 mm SL, Al Wajh, Saudi Arabia, Red Sea;
B: SMF uncat [QG17-83], 260 mm SL, Qatar, Arabian/Persian Gulf; C: SMF uncat [QG17-115], 275 mm SL, Qatar, Arabian/
Persian Gulf. Photos by S.V. Bogorodsky.
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 15
TABLE 3. Meristic characters of examined specimens in comparison to Chelon persicus (Senou et al. 1996). Asterisk =
In Senou et al. (1996) the last fin ray presumably was counted as one bifurcated ray, while in this study two separate rays
were counted.
Chelon
persicus
SAIAB
83180
SMF 35897
[KAU12-123]
SMF 35897
[KAU13-124]
SMF 35897
[KAU13-125]
SMF 35897
[KAU13-135]
Longitudinal scale rows 34-36 35 33 34 34 34
Dorsal-fin rays * IV + 9 IV + 10 IV + 10 IV + 10 IV + 10 IV + 10
Pectoral-fin rays 15-17 16 16 16 16 16
Anal-fin rays * III,8-9 III, 10 III,10 III,10 III,10 III,10
TV 11 - 12 12 12 -
PD1 - - 13 12 11 -
PD2 - - 23 22 21 -
CP - - 17 17 18 -
Continued.
KAUMM 459
[KAU13-136]
KAUMM 459
[KAU13-137]
KAUMM 459
[KAU13-138]
SMF 35898
[KAU13-427]
SMF uncat
[QG17-83]
SMF uncat
[QG17-115]
Longitudinal scale
rows
34 35 34 34 34 35
Dorsal-fin rays * IV + 10 IV + 9 IV + 10 IV + 10 IV + 10 IV + 10
Pectoral-fin rays 16 15 17 17 16 16
Anal-fin rays * III,10 III,10 III,10 III,10 III,10 III,10
TV 11 11 - 11 11 11
PD1 11 12 - 11 - -
PD2 21 23 - 23 - -
CP 16 18 - 18 16 -
FIGURE 6. Detailed photographs of the head of fresh specimens of Chelon persicus. A: SMF uncat [QG17-83], 260 mm SL,
Qatar, Arabian/Persian Gulf; B: detailed photographs of the head of SMF uncat [QG17-115], 275 mm SL, Qatar, Arabian/
Persian Gulf. Photos by S.V. Bogorodsky.
Comparisons. Chelon persicus can be distinguished from other Chelon species by the number of scales in the
longitudinal series (33–36), while congeners have more than 40 scales except C. dumerili which has 34–39 and
C. bandialensis which has 32–33 scales (Fischer et al. 1981; Thomson & Luther 1983; Harrison 2016). Chelon
dumerili differs from C. persicus in having multiple grooves on its predorsal scales while C. persicus only has a
single groove (Senou et al. 1996).
THIEME ET AL.
16 · Zootaxa 5188 (1) © 2022 Magnolia Press
Chelon persicus is externally similar to the sympatric Planiliza macrolepis (Smith, 1846) but differs by scales
dorsally on the head reaching forward to posterior nostrils versus scales usually extending beyond anterior nostrils
in P. macrolepis. Teeth on the upper lip of C. persicus are tiny and ciliform, usually without a second row, whereas
for P. macrolepis the upper lip has an outer row of very small, well-spaced, peg-like teeth and an irregular inner row
of small teeth, well separated from the outer row. The pectoral-fin base is whitish and unpigmented in C. persicus
but pigmented with melanophores, often with a golden sheen, in P. macrolepis.
Internally C. persicus differs from all other Chelon species in having a pair of long, spine-like postzygapophyses
on the second vertebrae, instead of a pair of short and hook-like postzygapophyses. Chelon persicus also differs
from C. auratus, C. bispinosus, C. labrosus and C. ramada by the presence of short and long pyloric caeca versus
pyloric caeca of subequal length in these species.
Discussion
Recent molecular analyses reported two cryptic species in the genus Chelon from the eastern coast of South Africa
(Durand et al. 2012a; Durand et al. 2012b; Durand & Borsa 2015). The first one, Chelon sp. B, is morphologically
and genetically very similar to Chelon dumerili but the name for that species is matter of an additional study. Three
nominal species (Mugil hoefleri Steindachner, 1882, Liza alosoides Fowler, 1903, and Mugil canaliculatus Smith,
1935) were placed in synonymy with C. dumerili (see Thomson 1997), and their types should be re-examined to
determine if they are conspecific with C. dumerili or C. sp. B. Further, Durand & Whitfield (2015) mentioned that
Liza natalensis (Castelnau, 1861) may attribute to C. sp. B and therefore the types of this species also need to be
re-examined. The genetic divergence of C. dumerili from C. sp. B could have been facilitated by the development of
the cool temperate Benguela Current on the west coast of South Africa and Namibia. This marine upwelling system
would have acted as a barrier to the coastal movements of warm water mugilid species such as C. dumerili.
In contrast, the second cryptic species, Chelon sp. A, did not match any previously described species from
this area (Durand & Borsa 2015). Molecular analyses performed in this study, using Chelon persicus specimens
collected from the Red Sea and Arabian/Persian Gulf, grouped these specimens with Chelon sp. A from South Africa
(Fig. 1, Fig. 2K). These data were confirmed by the morphological analysis, as the specimens from the Red Sea, the
Arabian/Persian Gulf as well as from South Africa revealed high conformity with the formal description of Chelon
persicus, given by Senou et al. (1996).
While the latest study placed it in the genus Planiliza (Eagderi et al. 2019) the results from our study confirm
its placement in the genus Chelon, as originally described by Senou et al. (1995, 1996). In addition, we report an
extended distribution range for Chelon persicus that is broadly distributed in the Western Indian Ocean, from its
type locality in the Arabian/Persian Gulf, west to the northern part of the Red Sea and south to the eastern coast of
South Africa. Also, it is the only species of the genus Chelon present in the Western Indian Ocean north of South
Africa (Durand & Whitfield 2015).
The genus Chelon currently includes 11 species: C. auratus, C. bandialensis, C. bispinosus, C. dumerili, C.
labrosus, C. persicus, C. ramada, C. richardsonii, C. saliens, C. sp. B, and C. tricuspidens. The four species, C.
auratus, C. bispinosus, C. labrosus and C. ramada, share a distinctive morphological character by having 5–9 pyloric
caeca of subequal length. The remaining seven species (C. bandialensis, C. dumerili, C. persicus, C. richardsonii,
C. saliens, C. sp. B, and C. tricuspidens) are characterised by having a combination of short and long pyloric caeca.
These two groups were also retrieved in the present phylogenetic analysis (Fig. 1) and the length of the pyloric
caeca (equal or unequal) seems to be a character useful to distinguish them. The 11 species of the genus Chelon are
distributed in the Baltic and the North Sea, in the Mediterranean, Black, and Caspian Sea, along the eastern coasts of
the Atlantic from Norway to South Africa (with C. bispinosus endemic to Cabo Verde Islands) as well as the Indian
Ocean coast of South Africa, the Red Sea and the Arabian/Persian Gulf (Fig. 2).
The recently described nominal species Chelon caeruleus Deef, 2018 (incorrectly named caeruleum) reportedly
also has a pair of spine-like postzygapophyses on the second vertebrae (Deef 2018). However, the description of C.
caeruleus is lacking important meristic data. Furthermore, Deef (2018) remarked that C. caeruleus is distinguished
by bearing long and short pyloric caeca but showed them in an unusual order (i.e., short and long caeca are mixed
and not arranged in separate groups), as well as the pair of spine-like neural postzygapophyses, of which one is
shorter than the other. It may be assumed that the unequal length of these postzygapophyses is an artefact and the
shorter one was broken in the dissected anterior vertebral column of the holotype (see Deef 2018: Fig. 3).
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 17
Phylogenetic analyses (based on COI) done by Deef (2018) resulted in a close relationship of C. caeruleus and
C. labrosus. In the current phylogenetic analysis, including all available genetic lineages, no extra lineages close to
C. labrosus were found (Fig. 1). The unavailability of the sequencing data on BOLD or GenBank for C. caeruleus is
a major problem. We doubt the validity of Chelon caeruleus and future samplings in the proposed distribution area
of C. caeruleus may provide better insights on this matter.
Previously, Xia et al. (2016) proposed that the morphology of the paired postzygapophyses on the second
vertebra is characteristic for the genera Chelon, Parachelon and Planiliza. While in the earlier derived Parachelon
these postzygapophyses supposedly are long and spine-like, they are long and curved with a blunt tip in Planiliza, and
short, curved and hook-like in Chelon. However, specimens examined during this study of C. persicus have a pair of
long, slightly curved, postzygapophyses with a sharp tip. Based on observations from x-rays of Chelon, Parachelon,
and Planiliza species, the categorization by Xia et al. (2016) does not capture the diversity of postzygapophyses
morphology and needs to be re-evaluated. While Parachelon can be distinguished by the lunate caudal-fin shape, an
additional study is needed to provide a distinct differential diagnosis based on morphology/osteology for the three
genera.
Acknowledgements
We are grateful to Susanne Dorow, Jennifer Steppler and Moritz Sonnewald for assistance at SMF. Thanks are also
due to Matthias Juhas and Stephanie Simon (SMF) for assisting in molecular genetic analyses and we gratefully
acknowledge the Grunelius-Möllgaard Laboratory at SMF for lab support. Recent material collected from the Red
Sea was obtained thanks to Dr Ashiq Ur Rahman and as part of scientific research cooperation between King
Abdulaziz University (KAU), Faculty of Marine Sciences (FMS), Jeddah, Saudi Arabia, and the Senckenberg
Research Institute (SRI), Frankfurt, Germany, under the framework of the Red Sea Biodiversity Project, funded by
KAU GRANT NO. “D/1/432-DSR”. We also acknowledge, with thanks, KAU and SRI for technical and financial
support as well as Ali Al-Aidaroos, Mohsen Al Sofiyani (KAU), Friedhelm Krupp (SRI and SMF) for their help in
the completion of the present study. Friedhelm Krupp (SRI and SMF) is thanked for his help in the organization of
a field trip to Qatar for the second author. In addition, we are grateful to Dirk Neumann and Ulrich Schliewen at the
Zoologische Staatssammlung München (ZMS), Munich, Germany, for x-ray images.
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MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 19
APPENDIX 1. Genetic data used in this study with reference to origin of data, species name deposited for each sample, BIN number and retrieved species name from BIN affili-
ation as well as collection place of each sample. Lat-values and lon-values in red indicate approximated origin locality of sample.
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
ANGBF3520-
12
HQ131882 44La Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
23.71 -15.953 Morocco
ANG-
BF37660-19
KT313011 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAD7360
Chelon
auratus
36.719 51.633 Iran south of Cas-
pian Sea
ANGBF4365-
12
HM208835 41La Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
30.427 -9.623 Morocco
ANGBF4402-
12
HM208836 43La Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
North East
Atlantic
Mediterranean
Sea
ANGBF4403-
12
HM208834 38La Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD7360
Chelon
auratus
Morocco
ANGBF5178-
12
JQ060460 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
28.035 -12.24 Morocco
ANGBF5179-
12
JQ060458 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
23.71 -15.953 Morocco Western Sahara
ANGBF5180-
12
JQ060456 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
36.81 10.296 Tunisia
ANGBF5303-
12
JQ060459 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
28.035 -12.24 Morocco
ANGBF5304-
12
JQ060457 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
30.427 -9.623 Morocco
ANG-
BF53069-19
LC490599 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
31.271 30.155 Egypt Mediterranean
Sea
ANG-
BF53070-19
LC490600 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
31.271 30.155 Egypt Mediterranean
Sea
ANG-
BF53071-19
LC490601 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
31.271 30.155 Egypt Mediterranean
Sea
ANG-
BF53072-19
LC490602 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
31.212 29.916 Egypt Mediterranean
Sea
......continued on the next page
THIEME ET AL.
20 · Zootaxa 5188 (1) © 2022 Magnolia Press
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
ANG-
BF53073-19
LC490603 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
31.212 29.916 Egypt Mediterranean
Sea
ANG-
BF53074-19
LC490604 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
31.212 29.916 Egypt Mediterranean
Sea
ANG-
BF53075-19
LC490605 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
31.422 31.805 Egypt Mediterranean
Sea
ANG-
BF53076-19
LC490606 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
31.422 31.805 Egypt Mediterranean
Sea
ANG-
BF53077-19
LC490607 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
31.422 31.805 Egypt Mediterranean
Sea
ANG-
BF53149-19
MG574945 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
36.719 51.633 Iran Caspian sea
DNATR043-
12
TR043EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
Turkey
DNATR840-
13
TR840EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR841-
13
TR841EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR842-
13
TR842EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR843-
13
TR843EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
......continued on the next page
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 21
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
DNATR844-
13
TR844EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR845-
13
TR845EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR846-
13
TR846EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR847-
13
TR847EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR848-
13
TR848EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR849-
13
TR849EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR850-
13
TR850EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR851-
13
TR851EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR852-
13
TR852EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR853-
13
TR853EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
......continued on the next page
THIEME ET AL.
22 · Zootaxa 5188 (1) © 2022 Magnolia Press
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
DNATR854-
13
TR854EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR855-
13
TR855EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR856-
13
TR856EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR857-
13
TR857EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR858-
13
TR858EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
DNATR859-
13
TR859EK Ankara Universi-
ty Biotechnology
Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.625 26.653 Turkey
ERDK010-21 BRCD-748 Bandirma Sheep
Breeding Re-
search Institute
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
40.428 27.633 Turkey
FFMBH2305-
14
Ex51A8 2012-1059 Istanbul Univer-
sity, Department
of Biology
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
41.03 28.959 Turkey
FFMBH2306-
14
Ex51A9 2012-1059 Istanbul Univer-
sity, Department
of Biology
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
41.03 28.959 Turkey
FFMBH2324-
14
Ex51D7 2012-1044 Istanbul Univer-
sity, Department
of Biology
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
40.668 28.112 Turkey
......continued on the next page
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 23
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
FFMBH288-
14
Ex06A3 50558 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.395 21.435 Greece Mediterranean
FFMBH289-
14
Ex06A5 50560 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.395 21.435 Greece Mediterranean
FFMBH290-
14
Ex06A6 50561 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.395 21.435 Greece Mediterranean
FFMBH291-
14
Ex06A7 50562 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.395 21.435 Greece Mediterranean
FFMBH292-
14
Ex06A8 50563 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.395 21.435 Greece Mediterranean
FFMBH293-
14
Ex06A9 50564 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
38.395 21.435 Greece Mediterranean
GBGC9237-
09
EU715468 26Lau Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
41.847 3.124 Spain
GBGC9238-
09
EU715467 25Lau Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
41.847 3.124 Spain
GBGC9239-
09
EU715466 24Lau Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
41.847 3.124 Spain
......continued on the next page
THIEME ET AL.
24 · Zootaxa 5188 (1) © 2022 Magnolia Press
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
GBGC9290-
09
EU392235 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
42.438 11.194 Italy Tyrrhenian Sea
GBGC9291-
09
EU392234 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
46.21 36.582 Russia Azov Sea
GBGCA7461-
15
KM046915 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
39.904 8.454 Italy
GB-
MIN121563-
17
KY176517 Mined from
GenBank, NCBI
BOLD:
AAD7360
Chelon
auratus
36.667 29.1 Turkey
GB-
MIN122185-
17
LC114152 Mined from
GenBank, NCBI
BOLD:
AAD7360
Chelon
auratus
36.719 51.633 Iran Caspian Sea
GB-
MIN124277-
17
KT313010 cas2 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAD7360
Chelon
auratus
36.719 51.633 Iran Caspian Sea
GB-
MIN129358-
17
KT313012 cas4 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAD7360
Chelon
auratus
36.719 51.633 Iran Caspian Sea
GB-
MIN94739-17
KT313013 cas5 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAD7360
Chelon
auratus
36.719 51.633 Iran Caspian Sea
GBM-
NB10933-20
MN610438 Mined from
GenBank, NCBI
Chelon
auratus
BOLD:
AAD7360
Chelon
auratus
42.013 35.39 Turkey Black Sea
ANGBF4370-
12
HM208825 MNHN 2009-0790-
94
Mined from
GenBank, NCBI
Chelon ban-
dialensis
BOLD:
ACC0109
Chelon
bandialensis
14.094 -16.51 Senegal Saloum estuary
ANGBF4371-
12
HM208823 2Cba Mined from
GenBank, NCBI
Chelon ban-
dialensis
BOLD:
ACC0109
Chelon
bandialensis
Senegal and
Bijagos
ANGBF4408-
12
HM208824 MNHN 2009-0790-
94
Mined from
GenBank, NCBI
Chelon ban-
dialensis
BOLD:
ACC0109
Chelon
bandialensis
Senegal Saloum estuary
......continued on the next page
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 25
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
ANGBF5177-
12
JQ060462 Mined from
GenBank, NCBI
Chelon ban-
dialensis
BOLD:
ACC0109
Chelon
bandialensis
11.284 -15.833 Guinea-Bis-
sau
Bijagos Islds
ANGBF5302-
12
JQ060461 MNHN 2009-0790 Mined from
GenBank, NCBI
Chelon ban-
dialensis
BOLD:
ACC0109
Chelon
bandialensis
14.094 -16.51 Senegal
MULID001-
21
CbiCV-01 OP028827 Present study Chelon
bispinosus
BOLD:
AEO1527
Chelon
bispinosus
16.897 -25.004 Cape Verde
MULID002-
21
CbiCV-02 OP028823 Present study Chelon
bispinosus
BOLD:
AEO1527
Chelon
bispinosus
16.897 -25.004 Cape Verde
MULID003-
21
CbiCV-03 OP028824 Present study Chelon
bispinosus
BOLD:
AEO1527
Chelon
bispinosus
16.897 -25.004 Cape Verde
MULID004-
21
CbiCV-04 OP028826 Present study Chelon
bispinosus
BOLD:
AEO1527
Chelon
bispinosus
16.897 -25.004 Cape Verde
MULID005-
21
CbiCV-05 OP028825 Present study Chelon
bispinosus
BOLD:
AEO1527
Chelon
bispinosus
16.897 -25.004 Cape Verde
ANGBF3509-
12
HQ131883 10Ldu Mined from
GenBank, NCBI
Chelon
dumerili
BOLD:
ACC0658
Chelon
dumerili
11.869 -15.571 Bissau
Guinea
Bissau
ANGBF4369-
12
HM208827 9Ldu Mined from
GenBank, NCBI
Chelon
dumerili
BOLD:
ACC0658
Chelon
dumerili
West Africa
ANGBF4407-
12
HM208826 8Ldu Mined from
GenBank, NCBI
Chelon
dumerili
BOLD:
ACC0658
Chelon
dumerili
West Africa
ANGBF5175-
12
JQ060466 078b Mined from
GenBank, NCBI
Chelon
dumerili
BOLD:
ACC0658
Chelon
dumerili
11.869 -15.571 Guinea-Bis-
sau
Bissau
ANGBF5176-
12
JQ060464 3016811 Mined from
GenBank, NCBI
Chelon
dumerili
BOLD:
ACC0658
Chelon
dumerili
16.02 -16.508 Senegal St Louis
ANGBF5300-
12
JQ060465 3016935 Mined from
GenBank, NCBI
Chelon
dumerili
BOLD:
ACC0658
Chelon
dumerili
6.146 1.278 Togo Lome
ANGBF5301-
12
JQ060463 3016936 Mined from
GenBank, NCBI
Chelon
dumerili
BOLD:
ACC0658
Chelon
dumerili
13.783 -16.486 Senegal Saloum estuary
......continued on the next page
THIEME ET AL.
26 · Zootaxa 5188 (1) © 2022 Magnolia Press
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
ANG-
BF37639-19
KY176703 Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
36.64 29.1 Turkey Mugla
ANGBF4364-
12
HM208837 45Cla Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
North East
Atlantic,
Mediterra-
nean Sea
ANGBF4401-
12
HM208838 46Cla Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
North East
Atlantic,
Mediterra-
nean Sea
ANGBF5166-
12
JQ060484 Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
30.427 -9.623 Morocco
ANGBF5202-
12
JQ060412 Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
36.81 10.296 Tunisia
ANGBF5327-
12
JQ060411 Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
28.035 -12.24 Morocco
BIM407-15 BIM F-017 SMNHTAU P.
15491
Tel Aviv Univer-
sity
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
32.803 35.029 Israel Levant Basin
BIM879-21 BIM F-117 SMNHTAU P. Tel Aviv Univer-
sity, Steinhardt
Museum of
Natural History
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
33.015 35.089 Israel Levant Basin
FFMBH1449-
14
Ex22A7 1892 Fischsammlung
Jorg Freyhof
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
33.889 -6.999 Morocco Iqem
FFMBH1450-
14
Ex22A8 1892 Fischsammlung
Jorg Freyhof
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
33.889 -6.999 Morocco Iqem
FFMBH1451-
14
Ex22A9 1892 Fischsammlung
Jorg Freyhof
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
33.889 -6.999 Morocco Iqem
......continued on the next page
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 27
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
FFMBH1868-
14
Ex37E4 50575 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
38.395 21.435 Greece Mediterranean
FFMBH285-
14
Ex06A11 50566 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
38.395 21.435 Greece Mediterranean
FFMBH286-
14
Ex06A12 50567 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
38.395 21.435 Greece Mediterranean
FFMBH294-
14
Ex06B1 50568 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
38.395 21.435 Greece Mediterranean
FFMBH295-
14
Ex06B10 50577 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
38.395 21.435 Greece Mediterranean
FFMBH296-
14
Ex06B2 50569 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
38.395 21.435 Greece Mediterranean
FFMBH297-
14
Ex06B3 50570 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
38.395 21.435 Greece Mediterranean
GBGC9231-
09
EU715474 32Chl Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
42.023 3.174 Spain
......continued on the next page
THIEME ET AL.
28 · Zootaxa 5188 (1) © 2022 Magnolia Press
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
GBGC9232-
09
EU715473 31Chl Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
42.023 3.174 Spain
GBGC9233-
09
EU715472 30Chl Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
42.023 3.174 Spain
GBGC9292-
09
EU392233 EU392233 Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
42.438 11.194 Italy Tyrrhenian Sea
GBGCA6004-
13
JX185139 LBP6848 36408 Mined from
GenBank, NCBI
Mugil
cephalus
BOLD:
AAE2670
Chelon
labrosus
GBGCA6005-
13
JX185138 LBP6848 36410 Mined from
GenBank, NCBI
Mugil
cephalus
BOLD:
AAE2670
Chelon
labrosus
GBGCA6006-
13
JX185137 LBP6848 36407 Mined from
GenBank, NCBI
Mugil
cephalus
BOLD:
AAE2670
Chelon
labrosus
GBGCA6007-
13
JX185136 LBP6848 36409 Mined from
GenBank, NCBI
Mugil
cephalus
BOLD:
AAE2670
Chelon
labrosus
GBGCA6008-
13
JX185135 LBP6848 36406 Mined from
GenBank, NCBI
Mugil
cephalus
BOLD:
AAE2670
Chelon
labrosus
GBGCA8500-
15
KJ128451 NRM:2358::
NRM:52307
Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
58.057 16.818 Sweden
GBGCA8507-
15
KJ128452 NRM:5488::
NRM:50609
Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
58.006 11.761 Sweden Hakefjorden
GBM-
NA15464-19
JF911706 Mined from
GenBank, NCBI
BOLD:
AAE2670
Chelon
labrosus
Morocco
GB-
MTG2594-16
NC_017883 Mined from
GenBank, NCBI
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
Morocco
MLFPI111-09 MLFPM7 University of
Minho
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
41.69 -8.85 Portugal Viana do Cas-
telo
MLFPI112-09 MLFPM8 University of
Minho
Chelon
labrosus
BOLD:
AAE2670
Chelon
labrosus
41.69 -8.85 Portugal Viana do Cas-
telo
......continued on the next page
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 29
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
MULID015-
22
ON943238 SMF 35879
[KAU13-135]
present study Chelon
persicus
BOLD:
AAC0698
Chelon per-
sicus
25.611 36.775 Saudi Ara-
bia
Red Sea
MULID016-
22
ON943236 SMF 35879
[KAU13-125]
present study Chelon
persicus
BOLD:
AAC0698
Chelon per-
sicus
25.611 36.775 Saudi Ara-
bia
Red Sea
MULID017-
22
ON943234 SMF 35879
[KAU13-124]
present study Chelon
persicus
BOLD:
AAC0698
Chelon per-
sicus
25.611 36.775 Saudi Ara-
bia
Red Sea
MULID018-
22
ON943231 SMF 35879
[KAU13-123]
present study Chelon
persicus
BOLD:
AAC0698
Chelon per-
sicus
25.611 36.775 Saudi Ara-
bia
Red Sea
MULID019-
22
ON943237 SMF 35898
[KAU13-427]
Present study Chelon
persicus
BOLD:
AAC0698
Chelon per-
sicus
25.611 36.775 Saudi Ara-
bia
Red Sea
LIDMA3629-
22
OP028368 QG17-083 present study Chelon
persicus
BOLD:
AAC0698
Chelon per-
sicus
25.281 51.529 Qatar Arabian/Per-
sian Gulf
LIDMA3630-
22
OP028367 QG17-115 Present study Chelon
persicus
BOLD:
AAC0698
Chelon per-
sicus
25.281 51.529 Qatar Arabian/Per-
sian Gulf
ANG-
BF28478-19
KX017695 Mined from
GenBank, NCBI
BOLD:
AAC0698
Chelon per-
sicus
ANGBF5162-
12
JQ060492 SAIAB-78131 Mined from
GenBank, NCBI
Chelon sp.
A_JDD
BOLD:
AAC0698
Chelon per-
sicus
-28.368 32.431 South Africa
DSFSE709-08 ADC08 Smith
222.5 #3
South African
Institute for
Aquatic Biodi-
versity
Liza mac-
rolepis
BOLD:
AAC0698
Chelon per-
sicus
-22.835 35.551 Mozam-
bique
DSFSG1035-
13
ADC2013
222.5 #4
South African
Institute for
Aquatic Biodi-
versity
Liza mac-
rolepis
BOLD:
AAC0698
Chelon per-
sicus
-28.835 32.028 South Africa
HVDBF159-
10
HvdB-10-
2010-64
HvdB-10-2010-64 University of
Johannesburg
BOLD:
AAC0698
Chelon per-
sicus
-33.72 25.845 South Africa Eastern Cape
......continued on the next page
THIEME ET AL.
30 · Zootaxa 5188 (1) © 2022 Magnolia Press
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
HVDBM879-
12
KZN_318 KZN_318 University of
Johannesburg
BOLD:
AAC0698
Chelon
persicus
-28.403 32.203 South Africa
SAIAB223-06 SAIAB
78130-T252
78130 South African
Institute for
Aquatic Biodi-
versity
Liza sp. BOLD:
AAC0698
Chelon
persicus
-31.597 29.607 South Africa Eastern Cape
TZMSC566-
06
Smith 222.6
#2
South African
Institute for
Aquatic Biodi-
versity
Liza sp. BOLD:
AAC0698
Chelon
persicus
-28.95 31.78 South Africa KwaZulu-Natal
TZSAL186-
04
L-III-B7-M-1 Research Col-
lection of Allan
Connell
Chelon
richardsonii
BOLD:
AAC0698
Chelon
persicus
-30.431 30.851 South Africa KwaZulu-Natal
ANG-
BF37643-19
KF676639 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
39.933 8.512 Italy
ANG-
BF37644-19
LC424371 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
31.422 31.805 Egypt Mediterranean
Sea
ANG-
BF37645-19
LC424372 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
31.422 31.805 Egypt Mediterranean
Sea
ANG-
BF37647-19
LC424374 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
31.271 30.155 Egypt Mediterranean
Sea
ANG-
BF37648-19
LC424375 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
31.271 30.155 Egypt Mediterranean
Sea
ANG-
BF37649-19
LC424376 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
31.271 30.155 Egypt Mediterranean
Sea
ANG-
BF37652-19
LC424379 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
31.212 29.916 Egypt Mediterranean
Sea
ANG-
BF37653-19
LC424380 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
31.212 29.916 Egypt Mediterranean
Sea
......continued on the next page
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 31
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
ANG-
BF37655-19
LC424382 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
31.422 31.805 Egypt Mediterranean
Sea
ANG-
BF37656-19
LC424383 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
31.422 31.805 Egypt Mediterranean
Sea
ANG-
BF37657-19
LC424384 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
31.422 31.805 Egypt Mediterranean
Sea
ANG-
BF37661-19
KX929964 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAD5443
Chelon
ramada
31.422 31.805 Egypt Mediterranean
Sea
ANG-
BF37712-19
KX929971 Mined from
GenBank, NCBI
Mugil
cephalus
BOLD:
AAD5443
Chelon
ramada
31.422 31.805 Egypt Mediterranean
Sea
ANG-
BF37713-19
KX929972 Mined from
GenBank, NCBI
Mugil
cephalus
BOLD:
AAD5443
Chelon
ramada
31.422 31.805 Egypt Mediterranean
Sea
ANGBF4363-
12
HM208839 50Lra Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
North East
Atlantic,
Mediterra-
nean Sea
ANGBF4400-
12
HM208840 52Lra Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
Morocco
ANGBF5168-
12
JQ060480 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
34.852 -6.275 Morocco
ANGBF5293-
12
JQ060479 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
36.81 10.296 Tunisia
ANG-
BF53030-19
LC487189 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
30.46 31.17 Egypt Nile Bassin
ANG-
BF53031-19
LC487190 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
30.46 31.17 Egypt Nile Bassin
ANG-
BF53033-19
LC487192 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
30.46 31.17 Egypt Nile Bassin
......continued on the next page
THIEME ET AL.
32 · Zootaxa 5188 (1) © 2022 Magnolia Press
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
ANG-
BF53034-19
LC487193 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
30.46 31.17 Egypt Nile Bassin
BIM408-15 BIM F-018 SMNHTAU P.
15490
Tel Aviv Univer-
sity
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
32.803 35.029 Israel Levant Basin
FCFPW046-
06
FCFOPB060-
07
Universidade de
Lisboa, Museu
Bocage
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
39.92 -9 Portugal
FCFPW047-
06
FCFOPB060-
08
Universidade de
Lisboa, Museu
Bocage
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
39.92 -9 Portugal
FCFPW157-
06
FCFOPB077-
03
Universidade de
Lisboa, Museu
Bocage
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
41.03 -9.98 Portugal
FCFPW177-
06
FCFOPB083-
05
Universidade de
Lisboa, Museu
Bocage
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
41.25 -8.98 Portugal
FCFPW178-
06
FCFOPB083-
06
Universidade de
Lisboa, Museu
Bocage
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
41.25 -8.98 Portugal
FFMBH1301-
14
Ex19E6 ex1540 Fischsammlung
Jorg Freyhof
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
37.528 31.269 Turkey Koeprue
FFMBH1476-
14
Ex22D10 1935 Fischsammlung
Jorg Freyhof
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
33.889 -6.999 Morocco Iqem
FFMBH1477-
14
Ex22D11 1935 Fischsammlung
Jorg Freyhof
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
33.889 -6.999 Morocco Iqem
FFMBH1483-
14
Ex22D6 1931 Fischsammlung
Jorg Freyhof
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
34.465 -5.511 Morocco Iqem
......continued on the next page
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 33
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
FFMBH2307-
14
Ex51B10 2012-1058 Istanbul Univer-
sity, Department
of Biology
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
41.03 28.959 Turkey Istanbul
GBGC9236-
09
EU715469 EU715469 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
GBGC9285-
09
EU392240 EU392240 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
42.441 11.194 Italy
GBGCA2887-
13
KC349864 KC349864 Mined from
GenBank, NCBI
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
Italy
GB-
MIN121443-
17
KX929967 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAD5443
Chelon
ramada
31.212 29.916 Egypt Mediterranean
Sea
GB-
MIN126118-
17
KX929965 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAD5443
Chelon
ramada
31.212 29.916 Egypt Mediterranean
Sea
GB-
MIN126119-
17
KX929966 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAD5443
Chelon
ramada
31.212 29.916 Egypt Mediterranean
Sea
GB-
MIN126121-
17
KX929974 Mined from
GenBank, NCBI
Mugil
cephalus
BOLD:
AAD5443
Chelon
ramada
31.212 29.916 Egypt Mediterranean
Sea
GB-
MIN96726-17
KX929973 Mined from
GenBank, NCBI
Mugil
cephalus
BOLD:
AAD5443
Chelon
ramada
31.212 29.916 Egypt Mediterranean
Sea
LKCOX143-
19
Lr001 Hebrew Univer-
sity of Jerusa-
lem, Rehovot
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
32.821 35.592 Israel
LKCOX144-
19
Lr011 Hebrew Univer-
sity of Jerusa-
lem, Rehovot
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
32.821 35.592 Israel
......continued on the next page
THIEME ET AL.
34 · Zootaxa 5188 (1) © 2022 Magnolia Press
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
LKCOX145-
19
Lr012 Hebrew Univer-
sity of Jerusa-
lem, Rehovot
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
32.821 35.592 Israel
LKCOX146-
19
Lr013 Hebrew Univer-
sity of Jerusa-
lem, Rehovot
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
32.821 35.592 Israel
LKCOX147-
19
Lr002 Hebrew Univer-
sity of Jerusa-
lem, Rehovot
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
32.821 35.592 Israel
LKCOX148-
19
Lr003 Hebrew Univer-
sity of Jerusa-
lem, Rehovot
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
32.821 35.592 Israel
LKCOX149-
19
Lr004 Hebrew Univer-
sity of Jerusa-
lem, Rehovot
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
32.821 35.592 Israel
LKCOX150-
19
Lr005 Hebrew Univer-
sity of Jerusa-
lem, Rehovot
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
32.821 35.592 Israel
LKCOX151-
19
Lr008 Hebrew Univer-
sity of Jerusa-
lem, Rehovot
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
32.821 35.592 Israel
TLM002-17 TLM002 Agricultural Re-
search Organi-
zation, Volcani
Center
Chelon
ramada
BOLD:
AAD5443
Chelon
ramada
Israel
MULID010-
22
Mug-RedS-01 ON943232 Present study Chelon
ramada
Chelon
ramada
20.158 40.252 Saudi Ara-
bia
Red Sea
MULID011-
22
Mug-RedS-02 ON943233 Present study Chelon
ramada
Chelon
ramada
20.158 40.252 Saudi Ara-
bia
Red Sea
......continued on the next page
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 35
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
MULID012-
22
Mug-RedS-03 ON943235 Present study Chelon
ramada
Chelon
ramada
20.158 40.252 Saudi Ara-
bia
Red Sea
MULID013-
22
Mug-RedS-04 ON943230 Present study Chelon
ramada
Chelon
ramada
20.158 40.252 Saudi Ara-
bia
Red Sea
MULID014-
22
Mug-RedS-05 ON943239 Present study Chelon
ramada
Chelon
ramada
20.158 40.252 Saudi Ara-
bia
Red Sea
ANGBF5167-
12
JQ060482 Mined from
GenBank, NCBI
Chelon
richardsonii
BOLD:
AAD7487
Chelon
richardsonii
-33.49 27.126 South Africa
ANGBF5292-
12
JQ060481 SAIAB-64270 Mined from
GenBank, NCBI
Chelon
richardsonii
BOLD:
AAD7487
Chelon
richardsonii
-28.584 16.47 Namibia
DSFSE019-07 Smith 222.7
#3.1
South African
Institute for
Aquatic Biodi-
versity
Chelon
richardsonii
BOLD:
AAD7487
Chelon
richardsonii
-33.911 18.466 South Africa
DSFSE021-07 Smith 222.7
#5.1
South African
Institute for
Aquatic Biodi-
versity
Chelon
richardsonii
BOLD:
AAD7487
Chelon
richardsonii
-33.911 18.466 South Africa
HVDB080-09 HvdB11 87253 University of
Johannesburg,
Department of
Zoology
BOLD:
AAD7487
Chelon
richardsonii
-22.9 14.533 Namibia
HVDB335-10 HvdB-04-
2010-52
South African
Institute for
Aquatic Biodi-
versity
BOLD:
AAD7487
Chelon
richardsonii
-33.985 23.548 South Africa
HVDB336-10 HvdB-04-
2010-53
South African
Institute for
Aquatic Biodi-
versity
BOLD:
AAD7487
Chelon
richardsonii
-33.985 23.548 South Africa
......continued on the next page
THIEME ET AL.
36 · Zootaxa 5188 (1) © 2022 Magnolia Press
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
HVDB338-10 HvdB-04-
2010-55
South African
Institute for
Aquatic Biodi-
versity
BOLD:
AAD7487
Chelon
richardsonii
-33.985 23.548 South Africa
HVDB339-10 HvdB-04-
2010-56
South African
Institute for
Aquatic Biodi-
versity
BOLD:
AAD7487
Chelon
richardsonii
-33.985 23.548 South Africa
HVDB340-10 HvdB-04-
2010-57
South African
Institute for
Aquatic Biodi-
versity
BOLD:
AAD7487
Chelon
richardsonii
-33.985 23.548 South Africa
HVDB348-10 HvdB-04-
2010-65
University of
Johannesburg
BOLD:
AAD7487
Chelon
richardsonii
-33.985 23.548 South Africa
HVDB349-10 HvdB-04-
2010-66
South African
Institute for
Aquatic Biodi-
versity
BOLD:
AAD7487
Chelon
richardsonii
-33.985 23.548 South Africa
HVDB350-10 HvdB-04-
2010-67
South African
Institute for
Aquatic Biodi-
versity
BOLD:
AAD7487
Chelon
richardsonii
-33.985 23.548 South Africa
HVDBF158-
10
HvdB-10-
2010-63
HvdB-10-2010-63 University of
Johannesburg
BOLD:
AAD7487
Chelon
richardsonii
-33.72 25.845 South Africa Eastern Cape
HVDBF191-
11
HvdB-03-11-
2010-01
HvdB-03-11-2010-
01
University of
Johannesburg
BOLD:
AAD7487
Chelon
richardsonii
-22.831 14.541 Namibia
HVDBF192-
11
HvdB-03-11-
2010-02
HvdB-03-11-2010-
02
University of
Johannesburg
BOLD:
AAD7487
Chelon
richardsonii
-22.831 14.541 Namibia
HVDBF193-
11
HvdB-03-11-
2010-03
HvdB-03-11-2010-
03
University of
Johannesburg
BOLD:
AAD7487
Chelon
richardsonii
-22.831 14.541 Namibia
......continued on the next page
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 37
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
HVDBF194-
11
HvdB-03-11-
2010-04
HvdB-03-11-2010-
04
University of
Johannesburg
BOLD:
AAD7487
Chelon rich-
ardsonii
-22.831 14.541 Namibia
SAIAB684-12 MM12-245 South African
Institute for
Aquatic Biodi-
versity
BOLD:
AAD7487
Chelon
richardsonii
South Africa Eastern Cape
SAIAB703-13 MM12-053_
rerun
SAIAB 187089 South African
Institute for
Aquatic Biodi-
versity
BOLD:
AAD7487
Chelon
richardsonii
-34.411 19.278 South Africa Western Cape
SAIAB725-13 MM12-134_
rerun
SAIAB 186710 South African
Institute for
Aquatic Biodi-
versity
BOLD:
AAD7487
Chelon
richardsonii
-34.238 18.852 South Africa Western Cape
TZMSC123-
05
ADC222.7-1 Smith222.7-1 South African
Institute for
Aquatic Biodi-
versity
Chelon
richardsonii
BOLD:
AAD7487
Chelon
richardsonii
-34.64 19.29 South Africa Western Cape
TZMSC124-
05
ADC222.7-2 Smith222.7-2 South African
Institute for
Aquatic Biodi-
versity
Chelon
richardsonii
BOLD:
AAD7487
Chelon
richardsonii
-34.64 19.29 South Africa Western Cape
UKFBI834-08 KUT 6496 University of
Kansas, Biodi-
versity Research
Center
Chelon
richardsonii
BOLD:
AAD7487
Chelon
richardsonii
-33.6 26.9 South Africa Eastern Cape
ANG-
BF37659-19
KF962679 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
39.933 8.512 Italy
ANGBF4366-
12
HM208833 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
36.81 10.296 Tunisia
......continued on the next page
THIEME ET AL.
38 · Zootaxa 5188 (1) © 2022 Magnolia Press
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
ANGBF5291-
12
JQ060483 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
36.81 10.296 Tunisia
ANG-
BF53086-19
LC490616 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
31.271 30.155 Egypt Mediterranean
Sea
ANG-
BF53087-19
LC490617 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
31.212 29.916 Egypt Mediterranean
Sea
ANG-
BF53088-19
LC490618 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
31.212 29.916 Egypt Mediterranean
Sea
ANG-
BF53150-19
MG574946 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
36.675 51.601 Iran Caspian Sea
ERDK007-21 BRCD-388 Bandirma Sheep
Breeding Re-
search Institute
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
40.428 27.633 Turkey
FFMBH2315-
14
Ex51C7 2012-1057 Istanbul Univer-
sity, Department
of Biology
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
41.03 28.959 Turkey
FFMBH283-
14
Ex06A1 50556 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
38.395 21.435 Greece Mediterranean
FFMBH284-
14
Ex06A10 50565 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
38.395 21.435 Greece Mediterranean
FFMBH287-
14
Ex06A2 50557 Zoologisches
Forschungsmu-
seum Alexander
Koenig
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
38.395 21.435 Greece Mediterranean
......continued on the next page
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 39
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
GBGC9234-
09
EU715471 29Lsal Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
34.858 -6.28 Morocco
GBGC9235-
09
EU715470 28Lsal Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
34.858 -6.28 Morocco
GBGC9284-
09
EU392241 EU392241 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
42.438 11.194 Italy
GB-
MIN126273-
17
KY176520 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
36.603 36.17 Turkey Gulf of Iskend-
erun
GB-
MIN131962-
17
KY176519 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
36.651 29.113 Turkey
GB-
MIN96844-17
KY176518 Mined from
GenBank, NCBI
Chelon
saliens
BOLD:
AAF6190
Chelon
saliens
36.651 29.113 Turkey
ANGBF5174-
12
JQ060468 Mined from
GenBank, NCBI
Chelon sp.
B_JDD
BOLD:
AAD7493
Chelon sp. B -33.49 27.126 South Africa
ANGBF5299-
12
JQ060467 Mined from
GenBank, NCBI
Chelon sp.
B_JDD
BOLD:
AAD7493
Chelon sp. B -33.49 27.126 South Africa
HVDBF142-
10
HvdB-10-
2010-47
HvdB-10-2010-47 University of
Johannesburg
BOLD:
AAD7493
Chelon sp. B -33.72 25.845 South Africa Eastern Cape
HVDBF143-
10
HvdB-10-
2010-48
HvdB-10-2010-48 University of
Johannesburg
BOLD:
AAD7493
Chelon sp. B -33.72 25.845 South Africa Eastern Cape
HVDBF156-
10
HvdB-10-
2010-61
HvdB-10-2010-61 University of
Johannesburg
BOLD:
AAD7493
Chelon sp. B -33.72 25.845 South Africa Eastern Cape
HVDBM872-
12
KZN_302 KZN_302 University of
Johannesburg
BOLD:
AAD7493
Chelon sp. B -28.403 32.203 South Africa
......continued on the next page
THIEME ET AL.
40 · Zootaxa 5188 (1) © 2022 Magnolia Press
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
SAIAB217-06 SAIAB
78097-T312
78097 South African
Institute for
Aquatic Biodi-
versity
Chelon
dumerili
BOLD:
AAD7493
Chelon sp. B -31.594 29.605 South Africa Eastern Cape
SAIAB270-06 SAIAB
78293-T443
78293 South African
Institute for
Aquatic Biodi-
versity
Chelon
dumerili
BOLD:
AAD7493
Chelon sp. B -31.69 29.411 South Africa Eastern Cape
SAIAB780-13 TM12-215 South African
Institute for
Aquatic Biodi-
versity
BOLD:
AAD7493
Chelon sp. B -33.497 27.135 South Africa Eastern Cape
TZMSA337-
04
ADC222.3-1 South African
Institute for
Aquatic Biodi-
versity
Chelon
dumerili
BOLD:
AAD7493
Chelon sp. B -29.705 31.1 South Africa KwaZulu-Natal
TZMSA430-
04
ADC222.3-3 South African
Institute for
Aquatic Biodi-
versity
Chelon
dumerili
BOLD:
AAD7493
Chelon sp. B -29.705 31.1 South Africa KwaZulu-Natal
TZMSA449-
04
ADC222.3-2 South African
Institute for
Aquatic Biodi-
versity
Chelon
dumerili
BOLD:
AAD7493
Chelon sp. B -29.705 31.1 South Africa KwaZulu-Natal
TZSAL173-
04
L-III-A7-DB- Research Col-
lection of Allan
Connell
Chelon
dumerili
BOLD:
AAD7493
Chelon sp. B -30.431 30.851 South Africa KwaZulu-Natal
ANGBF5285-
12
JQ060495 Mined from
GenBank, NCBI
Chelon tri-
cuspidens
BOLD:
AAF6205
Chelon tri-
cuspidens
-33.705 25.84 South Africa
......continued on the next page
MAJOR REVIEW OF THE STATUS OF CHELON PERSICUS Zootaxa 5188 (1) © 2022 Magnolia Press · 41
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
DSLAF119-
07
LIIB9 #1 Research Col-
lection of Allan
Connell
Chelon tri-
cuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-30.333 30.75 South Africa KwaZulu-Natal
DSLAF120-
07
LIIB9 #2 Research Col-
lection of Allan
Connell
Chelon tri-
cuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-30.333 30.75 South Africa KwaZulu-Natal
DSLAF121-
07
LIIB9 #3 Research Col-
lection of Allan
Connell
Chelon tri-
cuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-30.333 30.75 South Africa KwaZulu-Natal
DSLAF122-
07
LIIB9 #4 Research Col-
lection of Allan
Connell
Chelon tri-
cuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-30.333 30.75 South Africa KwaZulu-Natal
DSLAG640-
10
ADC10-939 Research Col-
lection of Allan
Connell
Chelon tri-
cuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-30.431 30.851 South Africa KwaZulu-Natal
DSLAG904-
10
ADC10-1203 Research Col-
lection of Allan
Connell
Chelon tri-
cuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-30.431 30.851 South Africa KwaZulu-Natal
DSLAR176-
08
ADC08-L II
B9.1
Research Col-
lection of Allan
Connell
Chelon tri-
cuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-30.431 30.851 South Africa KwaZulu-Natal
DSLAR228-
08
ADC08-L II
B9.2
Research Col-
lection of Allan
Connell
Chelon tri-
cuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-30.333 30.75 South Africa KwaZulu-Natal
SAIAB236-06 SAIAB
78143-T223
78143 South African
Institute for
Aquatic Biodi-
versity
Chelon tri-
cuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-31.597 29.607 South Africa Eastern Cape
......continued on the next page
THIEME ET AL.
42 · Zootaxa 5188 (1) © 2022 Magnolia Press
APPENDIX 1. (Continued)
Process ID Sample ID Catalog number Institution
storing
Species
name
BIN BIN Species
name
lat lon Country Province/state
SAIAB237-06 SAIAB
78143-T436
78143 South African
Institute for
Aquatic Biodi-
versity
Chelon
tricuspidens
BOLD:
AAF6205
Chelon tri-
cuspidens
-31.597 29.607 South Africa Eastern Cape
SAIAE020-14 Smith 222.8
#2
SAIAB:FO: South African
Institute for
Aquatic Biodi-
versity
BOLD:
AAF6205
Chelon tri-
cuspidens
-32.135 28.526 South Africa Eastern Cape
TZMSB223-
04
ADC 222.8-1 South African
Institute for
Aquatic Biodi-
versity
Chelon
tricuspidens
BOLD:
AAF6205
Chelon tri-
cuspidens
-30.05 30.892 South Africa KwaZulu-Natal
TZSAL663-
13
CCDB-
17646_D07
Research Col-
lection of Allan
Connell
Chelon
tricuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-30.33 30.75 South Africa KwaZulu-Natal
TZSAL702-
13
CCDB-
17646_G10
Research Col-
lection of Allan
Connell
Chelon
tricuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-30.33 30.75 South Africa KwaZulu-Natal
TZSAL703-
13
CCDB-
17646_G11
Research Col-
lection of Allan
Connell
Chelon
tricuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-30.33 30.75 South Africa KwaZulu-Natal
TZSAL704-
13
CCDB-
17646_G12
Research Col-
lection of Allan
Connell
Chelon
tricuspidens
BOLD:
AAF6205
Chelon
tricuspidens
-30.33 30.75 South Africa KwaZulu-Natal
