ArticlePDF Available

Taxonomic review of the genus Trisopterus (Teleostei: Gadidae) with recognition of the capelan Trisopterus capelanus as a valid species

Journal of Fish Biology

November 2011
79(5):1236-60

DOI:10.1111/j.1095-8649.2011.03107.x

Source
PubMed

Authors:

Bo Delling

Swedish Museum of Natural History

Michael Norén

Swedish Museum of Natural History

Sven Kullander

Swedish Museum of Natural History

José A. González

University of Las Palmas de Gran Canaria

Trisopterus is demonstrated to be monophyletic, including four distinct species: T. capelanus, T. esmarkii, T. luscus and T. minutus. The capelan T. capelanus is resurrected from the synonymy of poor cod T. minutus, and is shown to be morphologically more similar to T. luscus, to which species it is also more closely related, indicated by a phylogenetic analysis presented here. A lectotype is designated for T. luscus. Trisopterus fasciatus, the type species of Trisopterus, is a junior synonym of T. luscus, and the lectotype of T. luscus is designated as the neotype of T. fasciatus. The lectotype of T. luscus is also designated as the neotype of Gadus barbatus. Gadus barbatus has priority over T. luscus but the name is suppressed by prevailing usage of T. luscus. A neotype is designated also for T. minutus. A phylogenetic analysis using mitochondrial cytochrome b, and a fragment of the nuclear rhodopsin gene, shows that T. capelanus and T. luscus are sister species, and in turn sister to a clade formed by T. minutus and T. esmarkii.

Neotypes of (a) Trisopterus minutus (NRM 60756) and (b) Trisopterus capelanus (TFMC BMVP/1262), and (c) lectotype of Trisopterus luscus (NRM 5678).

…

Figures - uploaded by Bo Delling

Content may be subject to copyright.

Content uploaded by Bo Delling

Content may be subject to copyright.

Journal of Fish Biology (2011) 79, 1236–1260

doi:10.1111/j.1095-8649.2011.03107.x, available online at wileyonlinelibrary.com

Taxonomic review of the genus Trisopterus (Teleostei:

Gadidae) with recognition of the capelan Trisopterus

capelanus as a valid species

B. Delling*†, M. Noren*, S. O. Kullander* and J. A. Gonz ´

alez‡

*Swedish Museum of Natural History, P.O. Box 50007, SE-104 05 Stockholm, Sweden and

‡Instituto Canario de Ciencias Marinas (Biología pesquera) and Universidad de Las Palmas

de Gran Canaria (Ecología marina aplicada y pesquerías), P.O. Box 56, Telde, E-35200 Las

Palmas, Spain

(Received 30 June 2010, Accepted 23 August 2011)

Trisopterus is demonstrated to be monophyletic, including four distinct species: T. capelanus,

T. esmarkii,T. luscus and T. minutus. The capelan T. capelanus is resurrected from the synonymy

of poor cod T. minutus, and is shown to be morphologically more similar to T. luscus, to which

species it is also more closely related, indicated by a phylogenetic analysis presented here. A lec-

totype is designated for T. luscus.Trisopterus fasciatus, the type species of Trisopterus, is a junior

synonym of T. luscus, and the lectotype of T. luscus is designated as the neotype of T. fasciatus.

The lectotype of T. luscus is also designated as the neotype of Gadus barbatus.Gadus barbatus

has priority over T. luscus but the name is suppressed by prevailing usage of T. luscus.Aneotype

is designated also for T. minutus. A phylogenetic analysis using mitochondrial cytochrome b,and

a fragment of the nuclear rhodopsin gene, shows that T. capelanus and T. luscus are sister species,

Key words: cytochrome b; morphology; phylogeny; rhodopsin; species.

INTRODUCTION

The gadid genus Trisopterus Raﬁnesque-Schmaltz 1814 includes three species, pout-

ing T. luscus (L. 1758), poor cod T. minutus (L. 1758) and Norway pout T. esmarkii

(Nilsson 1855), restricted in geographical distribution to the north-eastern Atlantic

Ocean and Mediterranean Sea (Svetovidov, 1986). Trisopterus minutus is generally

regarded as composed of two subspecies, T. m. minutus in the north-eastern Atlantic

Ocean, and capelan T. m. capelanus (Lac´

ep`

ede 1800) in the Mediterranean Sea,

following the revision by Svetovidov (1948). This taxonomy has been challenged

repeatedly, based on parasites (Tirard et al., 1992), allozymes (Tirard et al., 1992;

Mattiangeli et al., 2000) and otolith morphology (Gaemers, 1976) suggesting that

T. luscus and T. m. capelanus are more closely related to each other, than T. m.

minutus to T. m. capelanus, or even that they would represent the same species

(Tirard et al., 1992). No ﬁrm conclusion based on these observations has been

†Author to whom correspondence should be addressed. Tel.: +46 5195 4240; email: bo.delling@nrm.se

1236

TAXONOMY OF TRISOPTERUS CAPELANUS 1237

presented, however, and no taxonomic revision ever attempted. The collaborative

FishTrace project (Sevilla et al., 2007), aiming at barcoding European marine ﬁshes

using mitochondrial cytochrome band nuclear rhodopsin genes as markers, made

available specimens of all species of Trisopterus, enabling both a taxonomic and

a molecular reassessment of Trisopterus. The objective of this study was to reana-

lyze Trisopterus taxonomy, in particular with the aim of presenting a solution to the

prevailing taxonomic problem of the status of the subspecies of T. minutus.

MATERIALS AND METHODS

MATERIALS EXAMINED

Specimens are deposited in the Swedish Museum of Natural History, Stockholm (NRM)

and Museo de Ciencias Naturales de Tenerife (TFMC). Voucher specimens for molecular

analyses are listed in Table I.

Forty specimens were subjected to morphological analysis: Trisopterus capelanus NRM

5677, 47244, 53175, 54703, 5674, 57475, TFMC BMVP/1262 (neotype), TFMC BMVP/

1214; Trisopterus esmarkii NRM 46279, 46280, 49622, 55564; Trisopterus luscus NRM

5678 (lectotype), 40290, 46873, 54533, 54534, 57474; Trisopterus minutus NRM 21829,

54535, 55274, 55619, 60756 (neotype).

MOLECULAR ANALYSIS

Sequences from two genes were used in this study, the complete mitochondrial cytochrome

bgene (cytb) (1141 bp) and a 460 bp fragment of the nuclear rhodopsin gene (rho ). These were

selected because preliminary investigation (Sevilla et al., 2007) showed that these fragments

were easy to amplify, with low risk of paralogy, and with suitable variation and rate of

evolution.

Nuclear and mitochondrial DNA was extracted from specimens using the QiAmp DNA

Mini Kit (Qiagen; www.qiagen.com) with recommended protocol. For all PCR ampliﬁcation

a subset of the primers of Sevilla et al. (2007) was used, with the puReTaq Ready-To-Go

PCR kit (Amersham Biosciences; www.gelifesciences.com). PCR products were checked on

minigel, and then puriﬁed using the QIAquick PCR Puriﬁcation Kit (Qiagen).

The protocol used for ampliﬁcation of the cytochrome bgene was as follows. PCR cycling:

94◦C, 4 min; 35 cycles of 94◦C, 30 s; 52◦C, 30 s; 72◦C, 30 s; 72◦C, 8 min; PCR1: 2 μl

DNA extract was used to amplify the complete 1141 bp cytochrome bgene using the primers

Gluﬁsh-F with BacCyt b-R. PCR2: 0·5μl of PCR1 reaction product was used to amplify

overlapping 850 bp fragments, using the primers Gluﬁsh-F with CytbI-3R; and CytbI-6F

with BacCytb-R.

In a few cases where these primers failed to amplify a product, Gluﬁsh-F was replaced

with FishCytb-F and BacCytb-R replaced with TrucCytb-R, and CytbI-6F with CytbI-7F, in

both PCR1 and PCR2 and for sequencing.

The protocol used for ampliﬁcation of the rhodopsin gene fragment was as follows. PCR1:

3μl DNA extract was used to amplify a 460 bp fragment of the rhodopsin gene, using the

primers Rod-F2B with Rod-5R. PCR cycling: 94◦C, 4 min; 35 cycles of 94◦C, 30 s; 52◦

C, 30 s; 72◦C, 30 s; 72◦C, 8 min; PCR2: 0·5μl of PCR1 reaction product was reampliﬁed

using the Rod-F2W and Rod-R4N primers, producing a 411 bp fragment. PCR cycling: 94◦

C,4min;35cyclesof94

◦C, 30 s; 52◦C, 30 s; 72◦C, 30 s; 72◦C, 8 min.

Sequencing reactions were performed using BigDye 3.1 with recommended protocol (Qia-

gen) and the same primers as for respective PCR2 reactions, and then the reactions were

puriﬁed with the DyeEx 96 Kit (Qiagen). Both strands of the reactions were sequenced using

an ABI 3700 (Applied Biosystems; www.appliedbiosystem.com) automatic sequencer, and

the obtained fragments were proof-read and assembled using the computer software Geneious

version 4.7 (Drummond et al., 2009).

1238 B. DELLING ET AL.

Table I. Alphabetical list of the 34 sequences of 13 gadiform species included in this study

Species

Area of

origin

GenBank

accession

number

cytochrome b

Genbank

accession

number

rhodopsin Specimen voucher

Gadus morhua SB EU492304 EU492304 NRM 49416

Merlangius merlangus SB EU492299 EU492206 NRM 49463

Merluccius merluccius SB EU492330 EU492239 NRM 49447

Micromesistius poutassou WM EF439548 EF439402 TFMC BMVP/0877

Micromesistius poutassou WM EF439549 EF439403 TFMC BMVP/0878

Micromesistius poutassou CB EU224066 EU224165 MNHN 2004 – 0612

Micromesistius poutassou CB EU224067 EU224166 NRM 54503

Micromesistius poutassou EM EU264028 EU264055 MNHN 2005–1060

Micromesistius poutassou EM EU264029 EU264056 MNHN 2005–1061

Micromesistius poutassou NS EU492136 EU492040 MNHN 2005–1613

Micromesistius poutassou NS EU492137 EU492041 MNHN 2005–1614

Micromesistius poutassou SB EU492307 EU492214 NRM 49628

Micromesistius poutassou SB EU492308 EU492215 NRM 49629

Molva molva SB EU492328 EU492237 NRM 49627

Pollachius virens SB EU492302 EU492209 NRM 49452

Trisopterus capelanus WM EF439619 EF439486 TFMC BMVP/1262

Trisopterus capelanus WM EF439620 EF439487 TFMC BMVP/1214

Trisopterus capelanus EM EU036518 EU036622 MNHN 2005 – 1132

Trisopterus capelanus EM EU036519 EU036623 MNHN 2005 – 1133

Trisopterus esmarkii NS EU492144 EU492048 MNHN 2005 – 1673

Trisopterus esmarkii NS EU492145 EU492049 MNHN 2005 – 1674

Trisopterus esmarkii SB EU492305 EU492212 NRM 49622

Trisopterus esmarkii SB EU492306 EU492213 NRM 49623

Trisopterus luscus CB EU224041 EU224138 MNHN 2004 – 0602

Trisopterus luscus CB EU224042 EU224139 MNHN 2004 – 0603

Trisopterus luscus NS EU492067 EU491973 MNHN 2005–1677

Trisopterus luscus NS EU492068 EU491974 MNHN 2005–1678

Trisopterus minutus CB EU224043 EU224141 MNHN 2004 – 0607

Trisopterus minutus CB EU224044 EU224140 NRM 54535

Trisopterus minutus NS EU492138 EU492042 MNHN 2005–1681

Trisopterus minutus NS EU492139 EU492043 MNHN 2005–1682

Trisopterus minutus SB GU304597 GU304595 NRM 55274

Trisopterus minutus SB GU304596 GU304594 NRM 55619

SB, Skagerrak and Baltic Sea; WM, western Mediterranean Sea; CB, Bay of Biscay; EM, northern

Aegean Sea; NS, North Sea; NRM, Swedish Museum of Natural History; TFMC, Museo de Ciencias

Naturales de Tenerife; MNHN, Mus´

eum national d’Histoire naturelle.

The sequences were manually aligned using the computer programme BioEdit (Hall, 1999).

All sequences have been deposited at GenBank. Alignments are available from the authors.

Table I lists all sequences and associated vouchers used for the molecular analysis.

The cytochrome band rhodopsin datasets were checked for saturation using the computer

programme Dambe (Xia & Xie, 2001), and no signiﬁcant saturation was found in any of the

datasets, in any of the three codon positions.

The cytochrome band the rhodopsin datasets were analysed individually and combined;

prior to the combined analysis they were tested for compatibility with the ILD test (Farris

TAXONOMY OF TRISOPTERUS CAPELANUS 1239

Table II. Observed uncorrected Pdistances (% dissimilarity) of cytochrome bfor

Trisopterus

T. capelanus T. minutus T. luscus T. esmarkii

T. capelanus 0·2–0·3 12·7–13·44·1–4·5 12·2–12·5

T. minutus 12·7–13·40·4–1·012·5–13·19·8–10·5

T. luscus 4·1–4·5 12·5–13·10·0–0·5 11·4–11·7

T. esmarkii 12·2–12·59·8–10·5 11·4–11·70·0–0·3

et al., 1994) using the computer programme PAUP (Swofford, 2002), and were found not to

be signiﬁcantly incongruent (P>0·05).

In all analyses, European hake Merluccius merluccius (L. 1758) (Merlucciidae) was des-

ignated as the outgroup.

Bayesian phylogenetic analysis was performed using MrBayes v3.2 (Huelsenbeck & Ron-

quist, 2001; Ronquist & Huelsenbeck, 2003), with the GTR ++I model as suggested by

MrModelTest (Nylander, 2004). The data were divided into partitions based on codon posi-

tion (ﬁrst, second and third), and in the combined analysis additionally partitioned by gene,

and all parameters except topology and branch length were allowed to vary independently for

each partition. The ‘temperature’ (level of perturbation) of the heated chains was 0·1. Two

parallel analyses with four chains each were run for three million generations, at which point

the average S. D. of split frequencies reported by MrBayes was lower than 0·01. Convergence

was checked using the computer programme Tracer (Rambaut & Drummond, 2007) version

1.4, and using the diagnostics of the SUMP and SUMT commands of MrBayes. The Bayesian

majority rule consensus tree was created from samples taken from both parallel analyses every

1000 generations, with the earliest 25% of samples discarded as burn-in, resulting in a total

of 4500 sampled trees.

Parsimony bootstrap analysis was performed using the computer programme TNT v1.1

(Goloboff et al., 2003), with the traditional search method, 2000 pseudoreplicates, 10 random

addition sequences per pseudoreplicate and Tree Bisection and Reconnection (TBR) branch

swapping. Genetic distances (uncorrected P-value) within and between putative species of

Trisopterus were calculated for both cytochrome b(Table II) and rhodopsin (Table III), using

the software MEGA4 (Tamura et al., 2007).

MORPHOLOGICAL ANALYSIS

A total of 40 specimens were analysed for morphology. Data were recorded for three

measurements and seven meristic characters, selected on the basis of previously reported

distinctions between species of Trisopterus (Fage, 1911; Svetovidov, 1986). Standard length

(LS), snout to anal-ﬁn base and anal-ﬁn base length were measured with a digital calliper

on the left side, rounded to the nearest 0·1 mm. Gill rakers were counted in situ on the right

side. Vertebrae and pterygiophore counts were obtained from X-rays (Fig. 1). Pterygiophore

Table III. Observed uncorrected Pdistances (% dissimilarity) of rhodopsin fragment for

Trisopterus

T. capelanus T. minutus T. luscus T. esmarkii

T. capelanus 0·0–0·32·4–3·31·8–2·03·1–3·3

T. minutus 2·4–3·30·0–0·72·4–2·70·9–1·6

T. luscus 1·8–2·02·4–2·70·03·1

T. esmarkii 3·1–3·30·9–1·63·10·0

1240 B. DELLING ET AL.

(b)

(c)

(d)

(a)

Fig. 1. Radiographs of (a) Trisopterus esmarkii, NRM 46280, 166·0 mm standard length (Ls); (b) T.minutus,

NRM 60756, 215·5mm LS, neotype; (c) T.capelanus, TFMC BMVP/1262, 207·0mm Ls, neotype

and (d) T.luscus, NRM 54533, 158·9mm Ls. , anteriormost anal ﬁn pterygiophore and tip of

postcleithrum.

counts largely correspond to ﬁn-ray counts and are easier to obtain with accuracy compared

to ﬁn-ray counts. The methodology described above also permitted the inclusion of material

in a poor state of preservation. For a single historical specimen of T. luscus (NRM 5678),

actual ﬁn rays were also counted using different methods and technical aids. Morphometric

data in Fage (1911) obtained from 50 specimens of Trisopterus were also analysed separately.

Fage (1911) used total length (LT), body depth at level of anus, vertical distance between

lateral line and (body midline) at level of anus, snout to anus, and anal-ﬁn base length.

Principal component analyses (PCA) on log10 transformed measurements and square root

of counts were used as an ordination method. Log10-transformation tends to make relations

TAXONOMY OF TRISOPTERUS CAPELANUS 1241

between measurements more linear and the factor loadings interpretable in terms of allometric

relationships (Bookstein et al., 1985). Meristic characters are summarised in frequency tables.

For the morphometric data set, principal component (PC) I represents size, whereas PC II

most often lacks correlation to size. For the meristic data set PC I usually provides most

information. Graphs contrasting the most informative PC of the morphometric and meristic

analyses visualize most effectively polarization of clusters representing different taxa which

overlap in separate analyses. The PCA were done using the SYSTAT 10 package (SPSS;

www.lbm.com/SPSS).

RESULTS

GENETIC DISTANCES

Intraspeciﬁc distance was <0·4% (rhodopsin) and 0·6% (cytochrome b) whereas

distances between putative congeneric species were >1·2% (rhodopsin) and 4·3%

(cytochrome b), respectively. The uncorrected mean ±s.d. distance between T.

minutus and T. capelanus was 2·9±0·4% for rhodopsin and 12·9±0·4% for

cytochrome b, respectively.

MOLECULAR PHYLOGENETIC ANALYSIS

In all analyses Trisopterus was recovered as a monophyletic group in which

T. esmarkii and T. minutus were sister taxa, and T. luscus and T. capelanus were

sister taxa. Trisopterus esmarkii,T. luscus and T. capelanus were strongly (100%

bootstrap frequency) supported as distinct taxa. Trisopterus minutus was strongly

(100% bootstrap frequency) supported as a distinct taxon in all analyses except when

analysing rhodopsin data alone. In the Bayesian analysis of rhodopsin (Fig. 2) the

paraphyly of T. minutus was moderately (87% bootstrap frequency) supported owing

to the position of T. esmarkii. In the parsimony analysis of rhodopsin T. minutus col-

lapsed to a polytomy which also included the monophyletic T. esmarkii. The Bayesian

(Fig. 3) and parsimony analyses of cytochrome bdata, as well as the Bayesian

(Fig. 4) and parsimony analyses of cytochrome band rhodopsin data combined,

recovered a monophyletic T. minutus.

MORPHOLOGY

Results from PCA are given in Figs 5 and 6 with corresponding loadings in

Tables IV– VI. Meristic characters are summarized in frequency tables (Tables VII –

XII). The sequence of taxa, i.e.,T.esmarkii,T.minutus,T.capelanus and T.lus-

cus in frequency tables is the same as in Fig. 1 and shows the overall decrease

in counts for meristic characters for the ﬁrst three species. Gill rakers (Table VII)

and pterygiophores supporting the second and third dorsal ﬁn (Table VIII) and sec-

ond anal ﬁn (Table IX) show no overlap for T.minutus and T.capelanus, whereas

vertebra counts (Table X) overlap slightly. The distinction of T. capelanus from

T. luscus is not strongly manifested in meristic data but morphometry, especially

the short preanal length and the long anal-ﬁn base length, distinguishes T. luscus. In

the multivariate analysis these two measurements almost distinguish the two species

along morphometric PC II in Fig. 5. If body depth is included as in data from Fage

(1911) the separation is more pronounced along PC II (Fig. 6) when specimens of

1242 B. DELLING ET AL.

Molva molva

Merluccius merluccius

Merlangius merlangus

Pollachius virens

Gadus morhua

Micromesistius poutassou WM

Micromesistius poutassou EM

Micromesistius poutassou NS

Micromesistius poutassou SB

Micromesistius poutassou CB

Trisopterus capelanus EM

Trisopterus capelanus WM

Trisopterus luscus NS

Trisopterus luscus CB

Trisopterus minutus SB

Trisopterus minutus CB

Trisopterus minutus SB

Trisopterus minutus CB

Trisopterus minutus NS

Trisopterus esmarkii SB

Trisopterus esmarkii NS

0·4

100

Fig. 2. Phylogenetic relationships of Trisopterus as inferred through Bayesian analysis of rhodopsin data.

Majority rule consensus tree, numbers at nodes indicate bootstrap frequency, nodes with <50% bootstrap

frequency collapsed. Branch lengths are proportional to number of expected substitutions per site. Scale

bar indicates number of expected substitutions per site. SB, Skagerrak and Baltic Sea; WM, western

Mediterranean Sea; CB, Bay of Biscay; EM, northern Aegean Sea; NS, North Sea.

similar size, i.e. PCI, are compared. Vertical distance between the lateral line and

body midline at the level of the anus was excluded from PCA on data from Fage

(1911). The distinction between Trisopterus species is further manifested in the inter-

nal anatomy comparing positions of the ventrally directed tip of the postcleithrum

TAXONOMY OF TRISOPTERUS CAPELANUS 1243

Molva molva

Merluccius merluccius

Merlangius merlangus

Pollachius virens

Gadus morhua

Micromesistius poutassou CB

Micromesistius poutassou NS

Micromesistius poutassou SB

Micromesistius poutassou NS

Micromesistius poutassou SB

Micromesistius poutassou WM

Micromesistius poutassou EM

Trisopterus capelanus WM

Trisopterus capelanus EM

Trisopterus luscus CB

Trisopterus luscus NS

Trisopterus esmarkii NS

Trisopterus esmarkii SB

Trisopterus minutus CB

Trisopterus minutus NS

Trisopterus minutus SB

100

100 100

95 100

100

0·4

Fig. 3. Phylogenetic relationships of Trisopterus as inferred through Bayesian analysis of cytochrome bdata.

Majority rule consensus tree, numbers at nodes indicate bootstrap frequency, nodes with <50% bootstrap

frequency collapsed. Branch lengths are proportional to number of expected substitutions per site. Scale

bar indicates number of expected substitutions per site. SB, Skagerrak and Baltic Sea; WM, western

Mediterranean Sea; CB, Bay of Biscay; EM, northern Aegean Sea; NS, North Sea.

and the anteriormost anal-ﬁn pterygiophore. Irrespective of size and condition the

ventrally directed tip of the postcleithrum points towards the anteriormost anal-ﬁn

pterygiophore in T. luscus, whereas T. capelanus possesses a distinct displacement

along the vertebral axis (Fig. 1). This displacement along the vertebral axis is most

pronounced in T. esmarkii.

1244 B. DELLING ET AL.

Molva molva

Merluccius merluccius

Merlangius merlangus

Gadus morhua

Pollachius virens

Micromesistius poutassou WM

Micromesistius poutassou EM

100

0·3

Micromesistius poutassou EM

Micromesistius poutassou CB

Micromesistius poutassou NS

Micromesistius poutassou SB

Micromesistius poutassou CB

Micromesistius poutassou NS

Trisopterus capelanus WM

Trisopterus capelanus EM

Trisopterus luscus CB

Trisopterus luscus NS

Trisopterus esmarkii SB

Trisopterus esmarkii NS

Trisopterus esmarkii SB

Trisopterus esmarkii NS

Trisopterus minutus NS

Trisopterus minutus CB

Trisopterus minutus SB

Fig. 4. Phylogenetic relationships of Trisopterus as inferred through Bayesian analysis of combined

cytochrome band rhodopsin data. Majority rule consensus tree, numbers at nodes indicate bootstrap

frequency, nodes with <50% bootstrap frequency collapsed. Branch lengths are proportional to number

of expected substitutions per site. Scale bar indicates number of expected substitutions per site. SB,

Skagerrak and Baltic Sea; WM, western Mediterranean Sea; CB, Bay of Biscay; EM, northern Aegean

Sea; NS, North Sea.

TAXONOMY OF TRISOPTERUS CAPELANUS 1245

Morphometric PC II

Meristic PC I

Fig. 5. Plot of scores of ﬁrst meristic principal component (PC) on second morphometric principal component

for Trisopterus:T.esmarkii (), T. minutus (), T. capelanus ()andT. luscus ( ). Neotypes and

lectotype indicated as ﬁlled symbols.

Morphometric PC I

Morphometric PC II

Fig. 6. Plot of scores of second on ﬁrst morphometric principal component (P) for Trisopterus based on data

from Fage (1911): T. minutus ()T. capelanus ()andT. luscus ().

1246 B. DELLING ET AL.

Table IV. Character loadings on principal components (PC) I – III for seven meristic char-

acters taken on Trisopterus

PCI PCII PCIII

First dorsal ﬁn pterygiophores 0·903 −0·128 −0·360

Second dorsal ﬁn pterygiophores 0·918 −0·008 0·344

Third dorsal ﬁn pterygiophores 0·973 0·116 −0·088

First anal ﬁn pterygiophores 0·785 −0·601 0·075

Second anal ﬁn pterygiophores 0·941 0·286 0·051

Vertebrae 0·981 0·016 0·033

Gill rakers 0·942 0·193 −0·046

Variance explained 5·959 0·511 0·268

Per cent of total variance 85·127 7·259 3·822

Table V. Character loadings on principal components (PC) I and II for three measurements

taken on Trisopterus

PC I PC II

Standard length 0·222 0·011

Preanal length 0·214 0·086

Anal ﬁn length 0·248 −0·084

Variance explained 0·156 0·014

Per cent of total variance 90·111 8·417

Table VI. Character loadings on principal components (PC) I – III for four measurements

taken on Trisopterus by Fage (1911)

PCI PCII PCIII

Total length 0·251 −0·023 0·007

Preanal length 0·220 −0·094 0·000

Body depth 0·314 0·035 −0·038

Anal ﬁn length 0·326 0·048 0·031

Variance explained 0·317 0·013 0·002

Per cent of total variance 95·441 3·919 0·729

Table VII. Frequency distribution of gill raker counts in Trisopterus

Gill rakers

15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37

T. esmarkii 1221

T. minutus 2231 1

T. capelanus 33931

T. luscus 21 12

Counts for neotypes and lectotype in bold.

TAXONOMY OF TRISOPTERUS CAPELANUS 1247

Table VIII. Frequency distribution of dorsal ﬁn pterygiophore counts in Trisopterus

Dorsal ﬁn pterygiophores

First dorsal ﬁn Second dorsal ﬁn Third dorsal ﬁn

12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 17 18 19 20 21 22 23 24 25 26 27 28 29 30

T. esmarkii 113 113 122

T. minutus 243 1521 1521

T. capelanus 79312293118621

T. luscus 1511311221

Counts for neotypes and lectotype in bold.

1248 B. DELLING ET AL.

Table IX. Frequency distribution of anal ﬁn pterygiophore counts in Trisopterus

Anal ﬁn pterygiophores

First anal ﬁn Second anal ﬁn

26 27 28 29 30 31 32 33 34 35 36 17 18 19 20 21 22 23 24 25 26 27 28 29 30

T. esmarkii 1 22 1121

T. minutus 114111 144

T. capelanus 135721 14392

T. luscus 13 11141

Counts for neotypes and lectotype in bold.

DISCUSSION

PHYLOGENETIC RELATIONSHIP OF TRISOPTERUS

The problem of distinguishing the species of Trisopterus has a long history, com-

plicated by the fact that T. luscus and T. capelanus are sympatric in the Mediterranean

Sea. Steindachner (1868) considered T. minutus (including T. capelanus) and T. lus-

cus to be the same species. Lilljeborg (1886: 70) noted that Linnaeus (1758) gave the

Mediterranean as type locality of T. minutus, but based on literature and examina-

tion of specimens of T. luscus from Nice (received as Morhua capelanus) Lilljeborg

somehow came to the conclusion that T. minutus did not exist in the Mediterranean

Sea, but used the name in the sense of M¨

uller (1776) for the North Atlantic species.

Fage (1911) reviewed earlier analyses and presented a detailed morphological com-

parison of T. luscus,T. capelanus and T. minutus. Fage’s (1911) data show that

T. capelanus and T. luscus are indeed more similar to each other, but he neverthe-

less came to the conclusion that T. capelanus is evolutionarily in between the more

primitive T. luscus, and the more advanced T. minutus. Also in Fage’s review (1911)

there is no clear taxonomic statement although it is obvious that he considers and

diagnoses three species.

The present molecular study is the ﬁrst to include all species of Trisopterus, and

strongly supports T.capelanus as a distinct taxon separate from T.minutus. This

is corroborated by the uncorrected molecular distance for cytochrome bbetween the

putative species (13%), which is incompatible with published intraspeciﬁc genetic

distances for the cytochrome bgene, but compatible with genetic distances for

Table X. Frequency distribution of total vertebra counts in Trisopterus

Vertebrae

45 46 47 48 49 50 51 52 53

T. esmarkii 14

T. minutus 1341

T. capelanus 7911

T. luscus 141

Counts for neotypes and lectotype in bold.

TAXONOMY OF TRISOPTERUS CAPELANUS 1249

Table XI. Fin ray and pterygiophore counts in NRM 5678, lectotype of Gadus luscus

Dorsal ﬁns Anal ﬁns

Method/source First Second Third First Second

Linnaeus (1764), ﬁn rays 13 23 19 31 18

X-ray, pterygiophores 14 25 20 34 20

X-ray, ﬁn rays 13 23 19 35 20

Microscope, ﬁn rays 13 23 18 33 19

No technical aids, ﬁn rays 13 23 18 32 19

Table XII. Selected characters of Trisopterus species, useful for species identiﬁcation

T. esmarkii T. minutus T. capelanus T. luscus

Mouth Superior Sub-terminal Sub-terminal Sub-terminal

Colour pattern

with bars

Absent Absent Absent Usually present

Gill rakers ≥33 24–32 15–21 18–22

Fin-ray counts

Second dorsal ﬁn 25–27 23–27 17–22 21–24

Third dorsal ﬁn 27–29 21–25 16–20 18–22

Second anal ﬁn 26–29 22–24 16–21 18–21

Preanal: anal ﬁn base

length ratio

>1>1>1<1

Tip of postcleithrum Points anterior

of anal ﬁn

insertion

Points anterior

of anal ﬁn

insertion

Points anterior

of anal ﬁn

insertion

Points to anal

ﬁn insertion

congeneric species. Bradley & Baker (2001) studied a fragment of cytochrome b

of bats and mice and concluded that distances of <2% were indicative of intraspe-

ciﬁc variation, and Hsieh et al. (2001) who studied a fragment of cytochrome bof a

range of organisms concluded that distances of <2·7% were indicative of intraspe-

ciﬁc variation. Farias et al. (2001) studied complete cytochrome bfrom a wide range

of cichlids, and found no species-pair with a genetic distance of <2%. For compari-

son, the uncorrected intraspeciﬁc genetic distance for the cytochrome bgene within

Trisopterus species is 0 to 1%, and the intraspeciﬁc genetic distance within the related

species Micromesistius poutassou (Risso 1827) which has a similar distribution to

the studied species of Trisopterus,is0·5%.

The phylogenetic analyses reveal that the closest relative of T. capelanus is not

T. minutus, but T. luscus.Micromesistius potaussou as sister species to Trisopterus

is also in agreement with the gadiform phylogeny based on mitochondrial 12S and

16S and nuclear recombination activating gene 1 (RAG1) presented by Roa-Var´

& Ortí (2009).

MORPHOLOGY OF TRISOPTERUS

Among the four species of Trisopterus,T. esmarkii and T. minutus are easily

distinguished both from each other and from T. capelanus and T. luscus by counts

1250 B. DELLING ET AL.

of gill-rakers, vertebrae and pterygiophores in the third dorsal ﬁn and second anal

ﬁn (Tables VII– X). The common pattern is for T. capelanus and T. luscus to have

overlapping lower counts, and T. minutus to have slightly higher counts but lower

than those of T. esmarkii. Consequently, T. capelanus and T. luscus end up closer in

multivariate statistics (Fig. 5). Adult T. luscus may appear very distinct, e.g. deep-

bodied and often strikingly colourful with its barred colour pattern. Juvenile and

preserved specimens without the typical barred colour-pattern, however, are not eas-

ily distinguished from T. capelanus. Trisopterus luscus is indeed more deep-bodied

than T. capelanus as revealed by data from Fage (1911), however, the preanal length

compared to anal-ﬁn base length is probably the most efﬁcient external morphologi-

cal distinguishing character; shorter in T. luscus, longer in T. capelanus. For material

studied here preanal:anal ﬁn base length ratios range from 0·73 to 0·96 in T. luscus

compared to 1·02 to 1·37 in T. capelanus. The position of the postcleithrum in rela-

tion to the anteriormost anal-ﬁn pterygiophore (Fig. 1) also seems to be a reliable

character distinguishing the two species from each other. Counts in Tables VIII and

IX may differ slightly from published ﬁn-ray counts, because pterygiophores rather

than the actual ﬁn rays were counted. The thick skin over the ﬁn bases makes it

difﬁcult to count ﬁn rays directly. X-radiography shows the occurrence of supranu-

merary rays in anteriormost ﬁns, and the presence of pterygiophores without ﬁn rays

between the second and third dorsal ﬁns, and the ﬁrst and second anal ﬁns (Fage,

1911). Consequently, counts reported here are often slightly higher than actual ﬁn-ray

counts. The allocation of pterygiophores without rays to the second or third dorsal

ﬁn and the ﬁrst or second anal ﬁn, respectively, in Tables VIII and IX also includes

an element of arbitrariness, however, not affecting the overall distinction between

studied species.

Fin bases in Trisopterus are more or less embedded in the skin. This character is

most strongly developed in the anal ﬁns of T. luscus. This is probably the explanation

for the small differences that remain between Linnaeus’ (1764) counts and counts

presented here (Table XI) of the lectotype of T. luscus without using technical aids,

i.e. it is easier to detect embedded rays with a needle after 250 years in ethanol, than

in a fresh and ﬂeshy specimen.

Distinctive characters for all four Trisopterus species are summarized in Table XII.

Ranges for meristic characters are slightly adjusted in accordance with Svetovidov

(1973). Pterygiophore counts (Tables VIII and IX) are also reduced by one when

converted to ﬁn-ray counts (Table XII).

TAXONOMIC REVIEW OF TRISOPTERUS

Trisopterus was described by Raﬁnesque-Schmaltz (1814) with Trisopterus fas-

ciatus Raﬁnesque-Schmaltz 1814 as the only included species, and therewith type

species of Trisopterus by monotypy. Trisopterus fasciatus is currently regarded as

a junior synonym of Gadus capelanus Lac ´

ep`

ede (1800. Morua Risso 1827, with

Gadus capelanus as type species by monotypy, is a junior synonym, objective or

subjective depending on the identity of T. fasciatus.Brachygadus Gill 1862 (type

species Gadus minutus L. 1758, by monotypy), and Gadulus Malm 1877 (type

species Gadus luscus L. 1758, by original designation), are considered as junior

subjective synonyms, by inclusion of the respective type species in Trisopterus by

Svetovidov (1973).

TAXONOMY OF TRISOPTERUS CAPELANUS 1251

Trisopterus was diagnosed by Raﬁnesque-Schmaltz (1814) by compressed body,

scaled head, and three dorsal and anal ﬁns placed opposite each other, the middle ones

the largest (‘V. G. TRISOPTERUS. Corps comprim´

e, tˆ

ete ´

ecailleuse; 3. nageoires

dorsales et anales oppos´

ees, les interm´

ediaires les plus grandes. – Obs. Il appartient

a la familles [sic] des Gadiens ou Gadinia.’). The description of the ﬁns is open to

alternative interpretation because there is no gadid species known to have three anal

ﬁns, but may have been inspired by the drawing in Willoughby (1685: pl. I, 1, ﬁg. 2:

Asellus mollis minor), where it looks like there are three anal ﬁns. The unexpected

period following the ﬁgure ‘3’ may be a typographical mistake. An unexplained

period appears also on pages 18 (‘10. rayons’), 19 (‘8. rayons’) and 20 (‘5. lobes’).

Current understanding of the taxonomy of Trisopterus derives essentially from

Svetovidov (1948, 1962, 1986), who resurrected the genus from the synonymy

under Gadus. Svetovidov diagnosed Trisopterus by the long anterior anal ﬁn with

its ﬁrst ray below the anterior dorsal ﬁn, dorsal ﬁns closely apposed or in contact

basally, a dark blotch at the base of the pectoral ﬁn, presence of a chin barbel,

presence of a trigeminofacial foramen for the facial nerve, and the median frontal

mucous cavity almost closed anteriorly. None of these character states is unique to

Trisopterus within the Gadidae. In Dunn’s (1989) phylogenetic analysis of the Gadi-

dae, Trisopterus was recovered as sister group of Eleginus, and supported by four

homoplasies: dorsal ﬂange of mesethmoid sharply pointed and high (shared with

Gadiculus); posterior margin of maxilla initially developing in a bifurcate manner

which persists during ontogeny (shared with Merlangius and Pollachius); number

of precaudal vertebrae low (13–19) (shared with Gadiculus); reduced number of

caudal-ﬁn rays (<45) (shared with Gadiculus). Teletchea et al. (2006) did not include

Eleginus in their phylogenetic analysis based on mitochondrial cytochrome coxidase

subunit I (CO I), and cytochrome b, but found Trisopterus (T. minutus and T. luscus

were analysed) as sister group to Gadiculus +Micromesistius. Their morphologi-

cal analysis did not provide resolution within the Gadidae. Roa-Var´

on & Ortí (2009)

included Eleginus and their phylogenetic analysis places Eleginus inside Microgadus

and not as sister to Trisopterus (T. minutus and T. esmarkii were analysed). Although

no autapomorphy seems to be available for Trisopterus, DNA data strongly supports

the genus, and data presented here also shows strong support for two internal clades.

Fifteen nominal species have been referred to Trisopterus (Eschmeyer, 2008),

of which ﬁve are currently synonymized under Brosme brosme (Ascanius 1772) in

the family Lotidae. As noted, Svetovidov (1948, 1962, 1986) distinguished three

species of Trisopterus diagnosed by differences in mouth shape, relative length of

the anal ﬁn, body depth, and interorbital width, viz.T. esmarkii (Nilsson 1855),

T. minutus (L. 1758), and T. luscus (L. 1758). He further distinguished two sub-

species of T. minutus,viz., T. m. minutus in the north-eastern Atlantic Ocean, and

T. m. capelanus (Lac´

ep`

ede 1800) in the Mediterranean Sea. Molecular and morpho-

logical analysis corroborates this species composition, with the only major difference

that T. capelanus is demonstrated to be a distinct species, more closely related to,

and morphologically more similar to T. luscus than to T. minutus (Figs 1 – 6). While

researching the nomenclatural history of T. capelanus, it also became apparent that

there are several other nomenclatural and taxonomic problems within the genus, and

which are relevant for assessing the taxonomic status of T. capelanus.

1252 B. DELLING ET AL.

Taxonomy of Trisopterus esmarkii

Trisopterus esmarkii is recognized by a projecting lower jaw, minute chin barbel,

and a countershaded colouration with dark dorsum, silvery sides, and white abdomen.

It has been reported from a very wide area in the northern Atlantic Ocean, including

Iceland, the Bay of Biscay, the British Isles, and the North Sea, and the coast north

to Svalbard and Jan Mayen, and Barents Sea (Svetovidov, 1986). There are no

synonyms, and identiﬁcation of this species has never been considered problematic.

Syntypes are preserved in the Lund University Museum of Zoology, LZM L849/3700

and LZM L849/3014.

Taxonomy of Trisopterus luscus

Trisopterus luscus is recognized by a projecting upper jaw, chin barbel about as

long as the eye diameter, deep body, and long ﬁrst anal ﬁn (longer than preanal

distance). It occurs along the north-eastern Atlantic Ocean coast from Norway to

Morocco, and in the western Mediterranean Sea (Svetovidov, 1986). It has four syn-

onyms, viz.Gadus barbatus (L. 1758), Gadus bibus Lac´

ep`

ede 1800, Gadus tacaud

Lac´

ep`

ede 1800 and Gadus colias Gray 1854. Trisopterus fasciatus is a potential

synonym, as discussed below.

Gadus luscus was based on a specimen for which Linnaeus (1758) gave ﬁn counts,

and a literature reference to Artedi’s (1738) Gadus No. 5, in its turn based on

descriptions of Asellus luscus in Raius (1713) and Willughbeij (1686). Trisopterus

luscus was also listed in Linnaeus (1764), with counts taken from a specimen in the

collection of King Adolf Fredrik. Because the ﬁn counts given in Linnaeus (1764)

agree perfectly with the ﬁn counts in Linnaeus (1758), and because Linnaeus had

already written the text for Linnaeus (1764) before the publication of Linnaeus (1758)

(Fernholm & Wheeler, 1983), it is assumed that the specimen of T. luscus presently in

the NRM collection (NRM 5678) and coming from King Adolf Fredrik’s collection,

is a syntype of Gadus luscus. No specimens are known to have survived from

Artedi (1738), and consequently NRM 5678 is the only surviving syntype. Owing

to earlier problems with the distinction of the sympatric T. luscus and T. capelanus

(Steindachner, 1868; Lilljeborg, 1886; Fage, 1911), and uncertainty about the identity

of the lost specimens of Artedi (1738), NRM 5678 is ﬁxed as the lectotype of

T. luscus (Fig. 7), thereby removing any uncertainty concerning the identity of this

species in relation to T. capelanus.

Thus, NRM 5678, 181 mm Ls, 199 mm LT, lectotype of T. luscus is identiﬁed as

the specimen described above despite slight differences in ﬁn ray counts depending

on methodology (see Table XI). It has 25 gill rakers and pterygiophore counts in

dorsal and anal ﬁns: D1 14, D2 25, D3 20, A1 34 and A2 20. Short preanal dis-

tance, 66 mm in comparison to anal ﬁn base length of 73 mm, as well as position

of postcleithrum in relation to anteriormost anal ﬁn pterygiophore also clearly dis-

tinguish NRM 5678 as T. luscus. There is no locality information associated with

the specimen, but in Linnaeus (1764) the locality is given as the Mediterranean Sea.

The Spanish and Portuguese local name (Cohen et al., 1990; Corbera et al., 1998),

Faneca, is printed next to the locality, suggesting probably a Spanish origin.

Gadus barbatus was based on literature sources (Artedi, 1738), and Linnaeus’s

own observations from the Swedish west coast in Linnaeus (1747). The variation in

ﬁn-ray counts given in Linnaeus (1758) suggests that more than one species may be

TAXONOMY OF TRISOPTERUS CAPELANUS 1253

(a)

(b)

(c)

Fig. 7. Neotypes of (a) Trisopterus minutus (NRM 60756) and (b) Trisopterus capelanus (TFMC

BMVP/1262), and (c) lectotype of Trisopterus luscus (NRM 5678).

involved, and it is only the relatively deep body (‘longitudine ad latitudinem tripla’)

that points to G. luscus.

Gadus barbatus was described simultaneously with G. luscus. Relative priority

of these names must thus be decided with reference to ﬁrst reviser action. Bloch

(1786) seems to be the ﬁrst author to synonymize the two species. He did not use

a scientiﬁc name in the text, but on plate 166 the name is Gadus barbatus. In the

text there is a reference to Gmelin’s (1789) edition of Systema Naturae, but it is

nevertheless clear that Bloch selected G. barbatus over G. luscus.Gadus barbatus

was never used as the valid name after 1899, whereas G. luscus appears in numerous

publications after 1899. In accordance with the International Code of Zoological

1254 B. DELLING ET AL.

Nomenclature (International Commission on Zoological Nomenclature, 1999) this

means that G. luscus takes priority over G. barbatus under article 23.9.1, as herewith

established as speciﬁed in article 23.9.2., i.e. that the junior synonym has been used

in at least 25 works, published by at least 10 authors in the immediately preceding

50 years and encompassing a span of not <10 years. The works are Svetovidov

(1962, 1973, 1986), Gaemers (1976), Fernholm & Wheeler (1983), Cohen et al.

(1990), Tirard et al. (1992), Corbera et al. (1998), Mattiangeli et al. (2000) and 17

additional references given in the Appendix.

There are no known preserved type specimens of G. barbatus. Because the type

series may be a composite, contrasting with the already established synonymy with

G. luscus, it is useful to designate a neotype for G. barbatus. This action will establish

a deﬁnite identity for the name G. barbatus. NRM 5678, the lectotype of G. luscus

is selected as the neotype of Gadus barbatus. Distinguishing characters are detailed

above. This act makes G. barbatus an objective junior synonym of G. luscus.

The status of T. fasciatus, type species of the genus, has never been fully assessed.

Jordan & Evermann (1917) considered G. capelanus to be the type species, with

no mention of T. fasciatus. Jordan & Evermann were probably guided by Risso

(1827) because they refer in passing to the synonymy of Morua and Trisopterus as

having been established by Risso. Risso (1827) listed Trisopterus in the bibliography

under Morua capelanus Risso 1827, indicating that T.fasciatus would be a junior

synonym but did not explicitly mention T. fasciatus. Risso did, however, mention

that T. minutus of Bloch appeared to be the same species. The name T. fasciatus

has never been used as valid after the original description. It is not even listed

as a synonym under any species of Trisopterus in the bibliographies provided by

Svetovidov (1948, 1962, 1973).

The original description of Trisopterus and the only included species, T. fasciatus,

is very brief, and does not contain any information permitting deﬁnite recognition

of the species: ‘15. Trisopterus fasciatus. Jaune dor´

eray

e transversalement de bleu,

ligne lat´

erale droite et brune, queue fourchue’ (Raﬁnesque-Schmaltz, 1814); (golden

yellow, transversally striped with blue, lateral line straight and brown, caudal forked).

Because vertical bars are more prominent in T. luscus, and Raﬁnesque-Schmaltz

(1814) had already included T. minutus (as G. minutus) in his ﬁrst list of Sicilian

ﬁshes, it seems more likely that T. fasciatus is a junior synonym of T. luscus.

Because there is no extant type material of T. fasciatus, and it seems important to

establish the correct type species for the genus in light of the nomenclatural problems,

the lectotype of T. luscus is selected as neotype of T. fasciatus.

Morrhua sycodes (Cocco 1884) was listed by Eschmeyer (2008) as a synonym of

T. minutus, but is probably a synonym of T. luscus.ForM. sycodes Cocco (1884)

mentions ﬁve foramina close to the suborbital and on the side of the lower jaw,

and the colour as ash-colored on the back, silvery on the side and belly. Cocco

(1884) included in M. sycodes ‘Var. I’ of M. capelanus from Risso (1827), said to

be silvery, with larger head and better developed pelvic ﬁn than T. capelanus. The

only other species in the area that is very similar to T. capelanus is T. luscus, and

it is possible that M. sycodes is actually a synonym of T. luscus, especially as that

species is not mentioned by Risso (1810, 1827) or Cocco (1884–1885). It is less

likely that their T. capelanus is actually T. luscus, and consequently M. sycodes a

synonym of T. capelanus, because the meristics given by Risso (1827) are more

similar to T. capelanus.

TAXONOMY OF TRISOPTERUS CAPELANUS 1255

Taxonomy of Trisopterus minutus

Trisopterus minutus is recognized by a projecting upper jaw, chin barbel about

as long as the eye diameter, slender body and short anal ﬁn (shorter than preanal

distance). The only proposed synonyms are G. capelanus and M. sycodes, the latter

by Eschmeyer (2008).

Linnaeus (1758: 253) based his description of G. minutus on Artedi’s (1738:

Genera: 21; Synonymia: 36) account of Gadus dorso tripterygio, ore cirrato, corpore

sescunicali, ano in medio corporis, and apparently based only on literature accounts

in Gesner (1558), Rondeletius (1559), Willughbeij (1686) and Raius (1713), who

refer to a species from the coast of southern France near Montpellier and Mar-

seille, and Venice at the northern extreme of the Adriatic Sea, as well as to Raius’s

(1713) description of Asellus mollis minimus from Cornwall at the south-western

tip of Great Britain. Linnaeus (1758) listed a number of ﬁn counts after the refer-

ence to Artedi (1738), which are from Willughbeij (1686) rather than from Artedi

(1738). Linnaeus (1758) gave the locality as the Mediterranean Sea (M. Mediter-

raneo). Neither Linnaeus (1758) nor Artedi (1738) had specimens of T. minutus,

but specimens of Rondeletius, Raius and Willughbeij for which illustrations or data

are given are syntypes. None of these syntypes have been preserved. If Linnaeus

(1758) is strictly followed in regarding T. minutus as a Mediterranean Sea species,

and recognizing that T. capelanus was based on Atlantic Ocean material, prevailing

usage of the names T. minutus (for an Atlantic Ocean species) and T. capelanus

(for a Mediterranean Sea species) should be reversed. To preserve current usage, a

specimen (Fig. 7) collected in Kattegat is herewith selected as neotype of T. minutus,

thus ﬁxing the name for an Atlantic Ocean species. The neotype has the following

data: NRM 60756, 219 mm LS, 240 mm LT; Denmark: Kattegat near Laes ¨

o Island;

K. Frohlund, 5 February 2007. It is identiﬁed as T. minutus based on meristic char-

acters. It has 25 gill rakers and pterygiophore counts in the dorsal and anal ﬁns

are D1 13, D2 25, D3 23, A1 31 and A2 25. Gill raker counts and pterygiophores

supporting the second anal ﬁn are outside the range of variation for material of the

three other species examined.

Taxonomy of Trisopterus capelanus

Gadus capelanus Lac´

ep`

ede (1800: 411) is based on 12 literature references, of

which all except two (Bloch, 1783; Gmelin, 1789) are non-binominal, and make

references to local names Mollo at Venice and Poor and Power in Cornwall. It was

synonymized with G. minutus L. 1758 by Risso (1810). No specimens are pre-

served that were deﬁnitely examined by Lac´

ep`

ede. It is quite clear from Lac´

ep`

ede’s

description that he was simply redescribing G. minutus, under a different name, e.g.

by explicitly referring to G. minutus in Gmelin (1789), and considering a species

having a wide Atlanto-Mediterranean distribution: ‘Le capelan vit dans les m`

emes

mers [North European Seas, Baltic Sea]; que le tacaud et le callarias; mais il habite

aussi dans la M´

editerran´

ee’. The description may very well be exclusively adapted

from earlier literature.

Among the references cited by Lac´

ep`

ede, a specimen is preserved only from Bloch

(1783: ﬁg. 1), viz. ZMB 2291 (cf. Paepke, 1999), with locality Atlantic Ocean. It

was identiﬁed by Bloch (1783) as G. minutus, and determined by Paepke (1999) as

T. minutus. As explained above Linnaeus (1768) based his description of G. minutus

entirely on literature data, including both Mediterranean Sea and North Atlantic

1256 B. DELLING ET AL.

Ocean populations and gave Mediterranean Sea as the locality. The count of 17 rays

in the posterior anal ﬁn is relatively low, and more similar to counts reported by

Svetovidov (1948, 1962) for T. capelanus (19 – 20) than for T. minutus (23–25).

The only surviving syntype of G. capelanus is probably of North Atlantic Ocean

origin, and if selected as lectotype would ﬁx the name G. capelanus as a synonym

of T. minutus from the north-east Atlantic Ocean. It may be desirable, however, to

preserve the epithet capelanus for the Mediterranean species. The specimen illus-

trated by Rondeletius (1559) for his description of Anthiae secunda specie, cited by

Lac´

ep`

ede (1800), is thus designated as lectotype of G. capelanus. The description

makes explicit mention of the use of capelan as the local name. Rondelet lived and

worked in Montpellier, on the Mediterranean Sea coast, and it may be assumed

that the Anthia secunda specie was founded on Mediterranean Sea specimens. The

lectotype is illustrated with a large eye, three dorsal ﬁns of which the ﬁrst is trian-

gular, a long chin barbel, a concave caudal ﬁn, and a single anal ﬁn opposite the

posteriormost dorsal ﬁn. Except for the number of anal ﬁns, the illustration is com-

patible with current understanding of the morphology of T. capelanus. The lectotype

is no longer extant. No specimens are known to have been preserved from Rondelet.

TFMC BMVP/1262 is thus designated here as neotype of G. capelanus Lac´

ep`

ede

(1800) (Fig. 7). A neotype is justiﬁed with reference to the confused nomenclatural

situation in Trisopterus where authors have apparently assumed that G. capelanus

is a Mediterranean Sea species and G. minutus a north-east Atlantic Ocean species.

In fact, the type locality of G. minutus was stated as the Mediterranean Sea by Lin-

naeus (1758), and the only surviving syntype of G. capelanus is apparently from

the north-east Atlantic Ocean. The neotype has the following collection data: TFMC

BMVP/1262, 207 mm Ls, 231 mm LT; eastern coast of Spain, El Campello, Ali-

cante: J. A. Gonz´

ales, 30 September 2009. It has 18 gill rakers and the pterygiophore

counts in the dorsal and anal ﬁns are D1 14, D2 22, D3 19, A1 30 and A2 20. It is

distinguished from T. luscus based on the long preanal distance, 80 mm in compari-

son to the anal ﬁn base length of 73 mm, as well as the position of the postcleithrum

in relation to the anteriormost anal ﬁn pterygiophore (Fig.1). Gill raker and ptery-

giophore counts further distinguish it from T. minutus. The neotype is included in

the molecular analyses (Figs 2–4).

Molecular and morphological analyses show that T. minutus,T. capelanus, and

T. luscus are three diagnosable, distinct species, which together with T. esmarkii

form a monophyletic group representing the genus Trisopterus. It has also been

demonstrated that T. luscus and T. capelanus form a monophyletic group. These

two species are sympatric and morphologically distinct from each other. By selecting

type specimens to ﬁx the usage of the prevailing names to agree with the taxonomy,

the nomenclature of the species of Trisopterus has been stabilized.

The European Commission funded the FishTrace project. The FishTrace Consortium (www.

ﬁshtrace.org) comprises 53 members from the following institutions: Complutense University,

Madrid; Joint Research Centre of the European Commission; Swedish Museum of Natural

History; Canarian Institute of Marine Sciences; French Research Institute for the Exploitation

of the Sea; Netherlands Institute for Fisheries Research; Natural History Museum of Funchal;

Natural History Museum of Tenerife; Fisheries Research Institute of Kavala; and National

Museum of Natural History, Paris. The Willi Hennig Society made TNT available for use

in phylogenetic analyses. M. F. Hern´

andez Martín and A. de Vera Hern´

andez of the Museo

de Ciencias Naturales de Tenerife kindly loaned the type specimen of Trisopterus capelanus.

J. M. Bautista, FishTrace project coordinator, and all the members of the FishTrace team

TAXONOMY OF TRISOPTERUS CAPELANUS 1257

contributed to this study. Special thanks go to G. Krey, E. G. Gonz´

alez, H. van Pelt-Heerschap

and P. Pruvost for providing specimens or data.

References

Artedi, P. (1738). Ichthyologia sive opera omnia de Piscibus scilicet: Biblioteca Ichthyolog-

ica. Philosophia Ichthyologica. Genera piscium. Synonymia specierum. Descriptiones

specierum. Omnia in hoc genere perfectiora, quam antea ulla. Lugduni Batavorum:

Conrad Wishoff.

Bloch, M. E. (1783). Oeconomische Naturgeschichte der Fische Deutschlands. Zweeter Theil.

Berlin: Published by the author.

Bloch, M. E. (1786). Naturgeschichte ausl¨andischer Fische. Zweiter Theil. Berlin: Published

by the author.

Bookstein, F., Chernoff, B., Elder, R., Humphries, J., Smith, G. & Strauss, R. (1985). Mor-

phometrics in evolutionary biology.Academy of Natural Sciences of Philadelphia,

Special Publication 15.

Bradley, R. D. & Baker, R. J. (2001). A test of the genetic species concept: cytochrome-b

sequences and mammals. Journal of Mammalogy 82, 960–973.

Cocco, A. (1884–1885). Indice ittiologico del mare di Messina. Il Naturalista Siciliano,

Giornale di scienze naturali 4, 25–29, 68–72, 85–88, 113–116, 177–180, 191–194,

228–232, 238–240, 291–294.

Cohen, D. M., Inada, T., Iwamoto, T. & Scialabba, N. (1990). FAO Species Catalogue. Vo l .

10. Gadiform ﬁshes of the world (Order Gadiformes). An annotated and illustrated

catalogue of cods, hakes, grenadiers and other gadiform ﬁshes known to date. Rome:

FAO.

Corbera, J., Sabat´

es, A. & García-Rubies, A. (1998). Peces de mar de la península Ib´erica.

Barcelona: Planeta.

Dunn, J. R. (1989). A provisional phylogeny on gadid ﬁshes based on adult and early life-

history characters. In Papers on the Systematics of Gadiform Fishes, Natural His-

tory Museum of Los Angeles County, Los Angeles, California (Cohen, D. M., ed),

pp. 209–235. Los Angeles, CA: Natural History Museum of Los Angeles County.

Fage, L. (1911). Le capelan de la M´

editerran´

ee: Gadus capelanus (Risso) et ses rapports

avec les esp`

eces voisines:G.luscus Linn ´

eetG. minutus O. Fr. M¨

uller. Archives de

Zoologie exp´erimentale et g´en´erale, 46, s ´erie 5 6, 257 – 282.

Farias, I. P., Ortí, G., Sampaio, I., Schneider, H. & Meyer, A. (2001). The cytochrome bgene

as a phylogenetic marker: the limits of resolution for analyzing relationships among

cichlid ﬁshes. Journal of Molecular Evolution 53, 89 – 103.

Farris, J. S., K¨

allersj¨

o, M., Kluge, A. G. & Bult, C. (1994). Testing signiﬁcance of incongru-

ence. Cladistics 10, 315–319.

Fernholm, B. & Wheeler, A. (1983). Linnaean ﬁsh specimens in the Swedish Museum of

Natural History, Stockholm. Zoological Journal of the Linnean Society 78, 199–286.

Gaemers, P. A. M. (1976). New concepts in the evolution of the Gadidae (Vertebrata, Pisces)

based on their otoliths. Mededelingen van de Werkgroep voor Tertiaire en Kwartaire

Geologie 13, 3–32

Gesner, C. (1558). Historiæ animalium liber IIII. qui est de piscium & aquatilium animantium

natura. Froschoverum: Tiguri.

Gmelin, J. F. (1789). Caroli a Linn´e Systema Naturae per regna tria naturae, secundum

classes, ordines, genera, species; cum characteribus, differentiis, synonymis, locis. Edi-

tio decimo tertia, aucta, reformata, Tomus I, Pars III. Lipsiae: G. E. Beer.

Goloboff, P., Farris, S. & Nixon, K. (2003). TNT: Tree Analysis using New Technology.

Tucum´

an: Published by the authors.

Hall, T. A. (1999). BioEdit: a user-friendly biological sequence alignment editor and analysis

program for Windows 95/98/NT. Nucleic Acids Symposium Series 41, 95 – 98.

Hsieh, H. M., Chiang, H. L., Tsai, L. C., Lai, S. Y., Huang, N. E., Linacre, A. & Lee, J. C.

(2001). Cytochrome b gene for species identiﬁcation of the conservation animals.

Forensic Science International 122, 7 – 18.

1258 B. DELLING ET AL.

Huelsenbeck, J. P. & Ronquist, F. (2001). MRBAYES: Bayesian inference of phylogeny.

Bioinformatics 17, 754–755.

International Commission on Zoological Nomenclature. (1999). International Code of Zoo-

logical Nomenclature, 4th edn. London: The International Trust for Zoological Nomen-

clature.

Jordan, D. S. & Evermann, B. W. (1917). The Genera of Fishes. From Linnaeus to Cuvier,

1758–1833, Seventy-ﬁve Years, with the Accepted Type of Each. Stanford, CA: Leland

Stanford Junior University Publications.

Lac´

ep`

ede, B. G. (1800). Histoire naturelle des poissons, Tome II. Paris: Plassan.

Lilljeborg, W. (1886). Sveriges och Norges ﬁskar [ Also as Sveriges och Norges Fauna,

ﬁskarne]. Uppsala: W. Schultz.

Linnaeus, C. (1747). W¨astg¨ota-Resa, p ˚ariksens h¨ogloﬂige st¨anders befallning f¨orr ¨attad ˚ar

1746. Med anm¨arkningar uti oeconomien, naturkunnogheten, antiquiteter, inv˚anarnes

seder och lefnads-s¨att. Stockholm: Lars Salvius.

Linnaeus, C. (1758). Systema naturae per regna tria naturae, secundum classes, ordines, gen-

era, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima,

reformata. Holmia: Laurentius Salvius.

Linnaeus, C. (1764). Museum S:ae R:ae M:tis Adolphi Friderici regis Svecorum, Gotho-

rum, Vanalorumque &c. &c. &c. In quo Animalia rariora Imprimis & Exorica: Aves,

Amphibia, Pisces Describuntur. Tomi Secundi Prodromus.Holmia: Laurentius Salvius.

Mattiangeli, V., Bourke, E. A., Ryan, A. W., Mork, J. & Cross, T. F. (2000). Allozyme anal-

yses of the genus Trisopterus: taxonomic status and population structure of the poor

cod. Journal of Fish Biology 56, 474–494.

M¨

uller, O. F. (1776). Zoologiae Danicae prodromus, seu Animalium Daniae et Norvegiae

indigenarum characteres, nomina et synonyma imprimis popularium. Havnia: Published

by the author.

Nylander, J. A. A. (2004). MrModeltest v2. Evolutionary Biology Centre, Uppsala University

Uppsala: Program distributed by the author.

Paepke, H.-J. (1999). Bloch’s ﬁsh collection in the Museum f ¨ur Naturkunde der Humboldt

Universit¨at zu Berlin. Ruggell: Theses Zoologicae, A.R.G. Gantner Verlag.

Raﬁnesque-Schmaltz, C. S. (1814). Pr´ecis des d´ecouvertes et travaux somiologiques de Mr.

C.S. Raﬁnesque-Schmaltz, entre 1800 et 1814. Palerme: Published by the author.

Raius, J. (1713). Synopsis methodica piscium. London: W. Innys.

Risso, A. (1810). Ichthyologie de Nice, ou Histoire naturelle des Poissons du D´epartement

des Alpes maritimes. Paris: F. Schoell.

Risso, A. (1827). Histoire naturelle des principales productions de l’Europe m´eridionale et

particuli`erement de celles des environs de Nice et des Alpes maritimes, 3. Paris &

Strasbourg: F.-G. Levrault.

Roa-Var´

on, A. & Ortí, G. (2009). Phylogenetic relationship among families of Gadiformes

(Teleostei, Paracanthopterygii) based on nuclear and mitochondrial data. Molecular

Phylogenetics and Evolution 52, 688 – 704.

Rondeletius, G. (1559). Libri de piscibus marinis, in quibus verae piscium essigies expressae

sunt. Lugduni: Matthias Bonhomme.

Ronquist, F. & Huelsenbeck, J. P. (2003). MRBAYES 3: Bayesian phylogenetic inference

under mixed models. Bioinformatics 19, 1572–1574.

Sevilla, R. G., Diez, A., Nor ´

en, M., Mouchel, O., J ´

erˆ

ome, M., Verrez-Bagnis, V., van Pelt,

H., Favre-Krey, L., Krey, G., the FishTrace Consortium & Bautista, J. M. (2007).

Primers and polymerase chain reaction conditions for DNA barcoding teleost ﬁsh based

on the mitochondrial cytochrome band nuclear rhodopsin genes. Molecular Ecology

Notes 7, 730–734.

Steindachner, F. (1868). Ichthyologischer Bericht ¨

uber eine nach Spanien und Portugal unter-

nommene Reise. (Sechste Fortsetzung.). Sitzungsberichte der Kaiserlichen Akademie

der Wissenschaften. Mathematisch-Naturwissenschaﬂiche Classe 57, 667 – 739.

Svetovidov, A. N. (1948). Fauna SSSR, Ryby, Treskoobraznye. Zoologischkij Institut

Akademia Nauk SSSR, Novaya Seriya 34, 1 – 221.

Svetovidov, A. N. (1962). Fauna of the U.S.S.R. (Fauna SSSR). Fishes (Ryby). Gadiformes

(Treskoobraznye). Jerusalem: Israel Program for Scientiﬁc Translations.

TAXONOMY OF TRISOPTERUS CAPELANUS 1259

Svetovidov, A. N. (1973). Gadidae. In Check-list of the Fishes of the North-eastern Atlantic

and of the Mediterranean, Vol. 1 (Hureau, J. C. & Monod, T., eds), pp. 303–320.

Paris: Unesco.

Svetovidov, A. N. (1986). Gadidae. In Fishes of the North-eastern Atlantic and the Mediter-

ranean, Vol. 2. (Whitehead, P. J. P., Bauchot, M.-L., Hureau, J.-C., Nielsen, J. & Tor-

tonese, E., eds), pp. 680–1001. Paris: Unesco.

Swofford, D. L. (2002). PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Meth-

ods). Version 4. Sunderland, MA: Sinauer Associates.

Tamura, K., Dudley, J., Nei, M. & Kumar, S. (2007). MEGA4: Molecular Evolutionary

Genetics Analysis (MEGA) software version 4.0. Molecular Biology and Evolution

24, 1596–1599.

Teletchea, F., Laudet, V. & H¨

anni, C. (2006). Phylogeny of the Gadidae (sensu Svetovidov,

1948) based on their morphology and two mitochondrial genes. Molecular Phyloge-

netics and Evolution 38, 189 – 199.

Tirard, C., Berrebi, P., Raibaut, A. & Frenaud, F. (1992). Parasites as biological markers: evo-

lutionary relationships in the heterospeciﬁc combination of helminths (monogeneans)

and teleosts (Gadidae). Biological Journal of the Linnean Society 47, 173–182.

Willoughby, F. (1685). Icthyographia. Londinum: Societatis Regalis Londinensis.

Willughbeij, F. (1686). De Historia Piscium Libri quatuor. Oxonium: Societatis Regiae Londi-

nensis.

Xia, X. & Xie, Z. (2001). DAMBE: data analysis in molecular biology and evolution. Journal

of Heredity 92, 371 – 373.

Electronic References

Drummond, A. J., Ashton, B., Cheung, M., Heled, J., Kearse. M., Moir, R., Stones-Havas, S.,

Thierer, T. & Wilson, A. (2009). Geneious v. 4.7. Available at http://www.geneious.

com/

Eschmeyer, W. N. (2008). Catalog of Fishes electronic version (updated 29 August 2008).

Available at http://www.calacademy.org/research/ichthyology/catalog/ﬁshcatsearch.

html/

Rambaut, A. & Drummond, A. J. (2007). Tracer v1.4. Available at http://beast.bio.ed.ac.uk/

Tracer/

Appendix: Additional references to fulﬁl requirements speciﬁed in Article

23.9.2. of the International Code of Zoological Nomenclature (1999)

Alonso-Fern´

andez, A., Domínguez-Petit, R., Bao, M., Rivas, C. & Saborido-Rey, F. (2008).

Spawning pattern and reproductive strategy of female pouting Trisopterus luscus (Gadi-

dae) on the Galician shelf of north-western Spain. Aquatic Living Resources 21,

383–393.

Armstrong, M. J. (1982). The predator-prey relationships of Irish Sea poor-cod (Trisopterus

minutus L.), pouting (Trisopterus luscus L.) and cod (Gadus morhua L.). ICES Journal

of Marine Science 40, 135 – 152.

Esteves, A., Seixas, F., Carvalho, S., Naz´

ario, N., Mendes, M. & Martins, C. (2009). Huff-

manela sp. (Nematoda: Trichosomoididae) muscular parasite from Trisopterus luscus

captured off the Portuguese coast. Diseases of Aquatic Organisms 84, 251 – 255.

Fernandes, D., Andreu-S´

anchez, O., Bebianno, M. J. & Porte, C. (2008). Assessment of pollu-

tion along the Northern Iberian shelf by the combined use of chemical and biochemical

markers in two representative ﬁsh species. Environmental Pollution 155, 327 – 335.

Fernandes, D., Bebianno, M. J. & Porte, C. (2008). Hepatic levels of metal and metalloth-

ioneins in two commercial ﬁsh species of the Northern Iberian shelf. The Science of

the Total Environment 391, 159–167.

Gestal, C. & Azevedo, C. (2006). Ultrastructural aspects of hepatic coccidiosis caused by

Goussia lusca n. sp. (Apicomplexa: Coccidia) infecting Trisopterus luscus (Gadidae)

from the NE Atlantic Ocean. Diseases of Aquatic Organisms 11, 25 – 31.

1260 B. DELLING ET AL.

Hamerlynck, O. & Hostens, K. (1993). Growth, feeding, production, and consumption in 0-

group bib (Trisopterus luscus L.) and whiting (Merlangius merlangus L.) in a shallow

coastal area of the south-west Netherlands. ICES Journal of Marine Science 50, 81– 91.

Henk, J., Heessen, L. & Daan, N. (1996). Long-term trends in ten non-target North Sea ﬁsh

species. ICES Journal of Marine Science 53, 1063 – 1078.

Hoff, P. T., Van de Vijver, K., Van Dongen, W., Esmans, E. L., Blust, R. & De Coen, W. M.

(2003). Perﬂuorooctane sulfonic acid in bib (Trisopterus luscus) and plaice (Pleu-

ronectes platessa) from the Western Scheldt and the Belgian North Sea: distribution

and biochemical effects. Environmental Toxicology and Chemistry 22, 608 – 614.

Hostens, K. & Mees, J. (1999). The mysid-feeding guild of demersal ﬁshes in the brackish

zone of the Westerschelde estuary. Journal of Fish Biology 55, 704 – 719.

Kastelein, R. A., van der Heul, S., van der Veen, J., Verboom, W. C., Jennings, N., de Haan,

D. & Reijnders, P. J. (2007). Effects of acoustic alarms, designed to reduce small

cetacean bycatch in gillnet ﬁsheries, on the behaviour of North Sea ﬁsh species in a

large tank. Marine Environmental Research 64, 160 – 180.

Last, J. M. (1978). The food of three species of gadoid larvae in the Eastern English Channel

and Southern North Sea. Marine Biology 48, 377–386.

Merayo, C. R. & Villegas, M. L. (1984). Age and growth of Trisopterus luscus (Linnaeus,

1758) (Pisces, Gadidae) off the coast of Asturias. Hydrobiologia 281, 115– 122.

Rello, F. J., Valero, A. & Adroher, F. J. (2008). Anisakid parasites of the pouting (Trisopterus

luscus) from the Cantabrian Sea coast, Bay of Biscay, Spain. Journal of Helminthology

82, 287–291.

Tanner, S. E., Fonseca, V. F. & Cabral, H. N. (2009). Condition of 0-group and adult pouting,

Trisopterus luscus L., along the Portuguese coast: evidence of habitat quality and

latitudinal trends. Journal of Applied Ichthyology 25, 387–393.

Tirard, C., Thomas, F., Raibaut, A. & Renaud, F. (1996). The distribution and abundance of

Lernaeocera lusci (Copepoda) on hake (Merluccius merluccius ) and bib (Trisopterus

luscus) (Teleostei). International Journal for Parasitology 26, 1387–1392.

Van Damme, P. A. & Ollevier, F. (1995). Morphological and morphometric study of crus-

tacean parasites within the genus Lernaeocera.International Journal for Parasitology

25, 1401–1411.

Trisopterus minutus voucher NRM 55619 rhodopsin (RHO) gene, partial cds

Nucleotide Sequence

December 2009

Sven Kullander · Michael Norén · B. Delling

Trisopterus minutus voucher NRM 55274 cytochrome b (CYTB) gene, complete cds; mitochondrial

Nucleotide Sequence

December 2009

Sven Kullander · Michael Norén · B. Delling

Trisopterus minutus voucher NRM 55274 rhodopsin (RHO) gene, partial cds

Nucleotide Sequence

December 2009

Sven Kullander · Michael Norén · B. Delling

Trisopterus minutus voucher NRM 55619 cytochrome b (CYTB) gene, complete cds; mitochondrial

Nucleotide Sequence

December 2009

Sven Kullander · Michael Norén · B. Delling

Pieter A. M. Gaemers. Taxonomy, Distribution and Evolution of Trisopterine Gadidae by Means of Otoliths and Other Characteristics. Fishes 2016, 1, 18–51.

Article

Full-text available

Jul 2017

Pieter Gaemers

The author has made the following corrections to his paper [1], which has been republished in order to comply with the rules of the International Code of Zoological Nomenclature (I.C.Z.N.). [...]

Taxonomy, Distribution and Evolution of Trisopterine Gadidae by Means of Otoliths and Other Characteristics

Article

Full-text available

Jul 2017

Pieter Gaemers

In a greater study of the recent fossil Gadidae, the object of this paper is to better define the trisopterine species and their relationships. The taxonomy of the four recent species usually included in the genus Trisopterus is further elaborated by means of published and new data on their otoliths, by published data on general external features and meristics of the fishes, and their genetics. Fossil otoliths, from the beginning of the Oligocene up to the present, reveal much of their evolution and throw more light on their relationships. Several succeeding and partly overlapping lineages representing different genera are recognized during this time interval. The genus Neocolliolus Gaemers, 1976, for Trisopterus esmarkii (Nilsson, 1855), is more firmly based. A new genus, Allotrisopterus, is introduced for Trisopterus minutus (Linnaeus, 1758). The similarity with Trisopterus capelanus (Lacepède, 1800) is an example of convergent evolution. The tribe Trisopterini Endo (2002) should only contain Trisopterus, Allotrisopterus and Neocolliolus as recent genera. Correct identification of otoliths from fisheries research and from sea bottom samples extends the knowledge of the present day geographical distribution of T. capelanus and T. luscus (Linnaeus, 1758). T. capelanus is also living along the Atlantic coast of Portugal and at least up to and including the Ría de Arosa, Galicia, Spain. There it can easily be mistaken for A. minutus that is also living there. Otoliths of T. luscus have been identified from the Evvoïkós Channel between Euboia and the mainland of Greece, thus it must live also in the Aegean Sea. Otoliths prove to be a powerful tool in taxonomy, biogeography and evolution of teleosts.

Rôles des contraintes génomiques et des traits d'histoire de vie dans la spéciation : une approche de génomique comparative

Thesis

Full-text available

Apr 2022

Pierre Barry

La spéciation est le processus évolutif au cours duquel une espèce se scinde en deux lignées qui divergent en accumulant des barrières reproductives, jusqu'à l’acquisition d’un isolement reproductif total. Durant ce processus, les lignées divergentes peuvent toujours s’échanger des gènes par hybridation, mais le flux génique est progressivement limité par l’accumulation des barrières. Il en résulte une semi-perméabilité des génomes, où certains locus s’échangent librement entre lignées et restent indifférenciés tandis que d’autres n’introgressent pas, contribuant ainsi à l’établissement de régions génomiques divergentes, appelées îlots génomiques de spéciation. L'étude de l’établissement, l’accumulation, l’érosion et la maintenance de ces barrières et de leurs effets sur la semiperméabilité des génomes de lignées en cours de spéciation permet de comprendre comment de nouvelles espèces se forment. L'avènement des techniques de séquençage à haut débit a permis de caractériser le paysage génomique de divergence chez de multiples lignées en cours de spéciation à travers l’arbre du vivant. Ces études ont permis de mesurer l’influence de l’histoire démographique et de l’architecture génomique comme déterminants majeurs du paysage génomique de divergence. Toutefois, d'autres facteurs pourraient intervenir et expliquer la diversité des trajectoires évolutives pouvant conduire ou non à la spéciation. Le principal objectif de cette thèse est d'évaluer l'impact des traits d'histoire de vie des espèces sur la spéciation. Nous avons choisi d’étudier 20 espèces de poissons marins subdivisées en deux lignées (Atlantique et Méditerranéenne), et présentant une large diversité de niveaux de divergence et de traits d’histoire de vie. Dans le premier chapitre, nous avons étudié les déterminants de la diversité génétique, substrat sur lequel s’établit la divergence lors de la séparation initiale des lignées. Nous avons observé que la longévité adulte des po issons marins est corrélée négativement à la diversité génétique, et nous avons démontré que cette relation pouvait s’expliquer par une plus grande variance du succès reproducteur chez les espèces longévives à cause de stratégies reproductives particulières aux poissons marins (forte mortalité juvénile, faible mortalité adulte et augmentation de la fécondité avec l’âge). Puis, dans un second chapitre, nous avons détecté une grande diversité d’histoires évolutives entre espèces, caractérisée par un fort gradient de divergence génétique entre lignées atlantiques et méditerranéennes. Ce gradient reflète en partie le niveau de semi-perméabilité des génomes. Les espèces à faible différentiation présentent un isolement reproductif faible, alors que les espèces les plus fortement différenciées montrent un isolement reproductif quasi-complet. Les traits d’histoire de vie des espèces expliquent en partie cette diversité de niveaux d’isolement via différents mécanismes. La durée de vie larvai re influence négativement la différenciation génétique en modulant les capacités de dispersion, l’effet de la taille du corps indique un effet négatif de l’abondance long-terme sur la divergence, et la longévité semble impacter le nombre de générations écoulées depuis la séparation ancestrale. En conclusion, les 20 espèces étudiées présentent une variabilité surprenante d’histoires évolutives au regard des similitudes de leur histoire biogéographique et leur architecture génomique. Les relations entre traits d’histoire de vie et histoire évolutive des espèces sont complexes, mais nous avons pu éclairer certaines d’entre elles en décomposant l’implication des traits dans les différentes étapes de la spéciation. L’application de l’approche de génomique comparative développée au cours de cette thèse dans d’autres zones de suture permettra d’étendre nos connaissances des déterminants du tempo et du mode de la spéciation.

First length–weight relationships of 11 fish species in the Aegean Sea

Article

Apr 2015

Weight-length relationships were established for eleven marine fish species caught in the SE Aegean Sea, Turkey. Additionally, a bibliographic review of such relationships for these species was conducted. Based on the results, the values of b parameter varied between 2.477 and 3.496, with one species having isometric growth, five negative and six positive allometric growth. Furthermore, for Aulopus filamentosus there exist no information in the literature, whilst for Callanthias ruber and Gnathophis mystax, there are no such information available from the Mediterranean.

Diclidophora luscae (Monogenea: Diclidophoridae) in pouting, Trisopterus luscus (Linnaeus, 1758) from the northeast Atlantic; epidemiology, morphology, molecular and phylogenetic analysis

Article

Full-text available

Sep 2022
Parasitol Res

Diclidophora (Monogenea) species are gill parasites with a stenoxenic specificity occurring only in Gadiformes. Epidemiological, morphological, molecular and phylogenetic studies were performed on 594 Diclidophora specimens collected from 213 Trisopterus luscus captured in the northeast Atlantic off the Portuguese coast during 2012, 2013 and 2020. Prevalence, parasite abundance and infection intensity were determined. Positive correlation between fish weight and length and infection intensity was observed. The effects of preservation on the parasite morphological features were studied, highlighting that specimen's identification should be reinforced by molecular studies. A sequence of D. luscae capelanii from T. capelanus captured in the Mediterranean Sea included in the 28S rDNA molecular analysis was nested within a robust D. luscae clade. Data analysis suggested that this species is in fact D. luscae, which is compatible with T. luscus and T. capelanus. The identity of fish hosts was confirmed by barcoding. For the first time, data on the infection parameters is shown, highlighting the importance of including this parasite in the monitoring plans for a holistic approach with possible effects for the management of pouting resources aiming of attaining sustainable development and biodiversity conservation measures, according to the 14th objective of the 2030 agenda.

Manuale per la classificazione dell’Elemento di Qualità Biologica “Fauna Ittica” nelle lagune costiere italiane. “Fish Fauna” Biological quality Element classification manual for Italian coastal lagoons

Technical Report

Full-text available

Jan 2018

Under the Water Framework Directive 2000/60/ EC, ichthyofauna is considered an expression of health and biotic integrity of the transitional aquatic ecosystems, together with other Biological Quality Elements and physico-chemical variables. This Manual, describes the principal fish species occurring within transitional waters as well as the national classification index, Habitat Fish Bio-Indicator (HFBI), for coastal lagoons. The HFBI has been validated at European level and represents the official method for the classification of ecological status of Italian transitional water bodies.

Habilitation "Apports de la systématique à la domestication de nouvelles espèces en aquaculture : applications aux premiers stades vie

Thesis

Full-text available

Dec 2016

Fabrice Teletchea

Domestication is the process by which animals become progressively adapted, generations after generations, to both captive conditions and humans. In land, this process has started about 12 000 years ago, and has allowed modifying profoundly animals that are today domesticated and display numerous breeds. Contrary to land animals, domestication of aquatic animals is much more recent, thus reared animals have only slightly changed from their wild congeners. After a discussion of the main consequences for aquaculture, a new typology, based on the methods and concepts of systematics, will be described to promote the diversification of new fish species, particularly for European inland aquaculture. From this new typology, this research project aims at developing an approach of evolutionary developmental biology focused on European freshwater fish species, by realizing phylogenetic analyses and comparative biology in silico, as well as experimental studies. In the end, this project could allow a better understanding of the diversity of life strategies of fish and proposing “standard” protocols for the rearing of early life stages of new species in aquaculture, and thus enhance the diversification of fish production.

Systematics and Aquaculture: What Could They Bring to Each Other?

Article

Full-text available

Oct 2016

Fabrice Teletchea

Systematics is traditionally based on morphological characters to both define species and establish classifications. In the past decades, partly due to the advent of molecular biology, traditional systematics has declined while molecular systematics has tremendously increased. This results in that fewer funding are generally provided to traditional systematics, particularly for searching new morphological characters. Aquaculture, the farming of aquatic animals, has increased exponentially in the past decades, providing today more than half of fish consumed worldwide, and is expected to continue to rise. Aquaculture requires controlling the life cycle of the farmed fish in captivity, including the rearing of early life stages. Therefore, by coupling systematics and aquaculture, it could be possible to bring new funds and facilities to the former to study the early life stages of numerous fish species and to the latter it would offer a conceptual framework to perform comparative ontogeny. Together, this could help improving our knowledge on the early life stages that could be useful for both taxonomists and zootechnicians.

2016 : Bilan d’une année d’inventaires ichtyologiques subaquatiques en Méditerranée française.

Article

Full-text available

Dec 2018

Depuis quelques années, un petit groupe de plongeurs passionnés de poissons marins réalisent le long de la côte méditerranéenne française une quarantaine de plongées par an. L'objectif est d’inventorier la diversité spécifique et de dresser les listes les plus complètes possibles des poissons qu’ils détectent par observation directe, de jour ou de nuit, et ce dans le maximum d’habitats différents. Ce compte-rendu présente les données les plus intéressantes recueillies en 2016.

Ethnobiology and etymology. Greek καλλαρίας (a Mediterranean cod)

Article

Mar 2017

Andrea Guasparri

New concepts in the evolution of the Gadidae (Vertebrata, Pisces), based on their otoliths

Article

Full-text available

Jan 1976

Pieter Gaemers

MEGA4, a molecular evolutionary genetic analysis MEGA software version 4.0

Article

Full-text available

Jan 2007
MOL BIOL EVOL

International code of zoological nomenclature = Code international de nomenclature zoologique

Book

Jan 1999

W. D. L. Ride

Morphometrics in Evolutionary Biology

Article

Dec 1988
COPEIA

Ichthyologie de Nice, ou Histoire naturelle des poissons du département des Alpes Maritimes

Article

A. Risso

Perfluorooctane sulfonic acid in bib (Trisopterus luscus) and plaice (Pleuronectes platessa) from the Western Scheldt and the Belgian North Sea: Distribution and biochemical effects

Article

Mar 2003

A biomonitoring campaign was conducted in the Belgian North Sea and in the Western Scheldt (The Netherlands) with the primary goal to assess perfluorooctane sulfonic acid (PFOS) contamination and distribution in different biota. This study covers the results obtained for bib (Trisopterus luscus) and plaice (Pleuronectes platessa) and includes the assessment of some stress-related biochemical endpoints. Analysis of liver and muscle PFOS concentrations of both species provided evidence for the existence of a PFOS pollution gradient along the Western Scheldt with higher levels at the upstream locations and a lower degree of PFOS pollution at the marine locations. Cellular necrosis was studied by measuring aspartate aminotransferase (AST) and alanine aminotransferase (ALT) levels in the serum. Serum ALT but not serum AST was shown to correlate positively with the PFOS liver concentration in bib (r = 0.44, p < 0.05), indicating that PFOS might contribute to the induction of hepatic damage in bib in the area of study. Analysis of total carbohydrate, lipid, and protein content of bib liver tissue revealed a positive correlation between the protein content and the PFOS liver concentration (r = 0.55, p < 0.01). Whether this is due to induction of compensatory mechanisms, detoxification, or repair processes remains unclear.

Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis

Article