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Supplementary Materials: Ellouet et al. A trait-based approach for assessing and mapping niche overlaps between native
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and exotic species: the Mediterranean coastal fish fauna as a case study.
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Appendix 1. List of ecological and biological traits
Modalities
Variables
Vertical
distribution
Habitats
Migration
Reproduction
Semelparity
Diet
Behavior
Sex reversal
Larvae
Length
A
Demersal
Soft
Catadromous
Oviparous
Yes
Carnivorous
Solitary
Protogynous
Planktotrophic
—
B
Benthic
Rocky
Amphidromous
Viviparous
No
Omnivorous
Gregarious
Hermaphrodism
Leicitotrophic
—
C
Pelagic
Posidonia
Oceanodromous
Ovoviviparous
—
Herbivorous
Both
Protandrous
—
—
D
—
—
Anadromous
—
—
—
—
—
—
—
Nature of
traits
Qualitative
ordinal
Qualitative
Qualitative
Qualitative
Qualitative
Qualitative
Qualitative
Qualitative
Qualitative
Quantitative
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Appendix 2. List of ecomorphological measures and associated functional traits
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Morphological traits were measured on side view of fish using the ImageJ software. Bi is the body
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standard length; Hd is the head depth along the vertical axis of the eye; Mo is the vertical distance
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between the tip of upper jaw and the head bottom on the same axis as Hd; Cfd is the caudal fin
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depth; CPd: caudal peduncle; Ed: eye diameter; Eh: distance between the bottom of the; head and
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the eye center along the head depth axis; PFb: body depth at the level of the pectoral fin insertion;
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PFi: distance between the insertion of pectoral fin and the bottom of the body; Cfs caudal fine
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surface. The list of 7 functional traits derived from 10 morphological measures with abbreviations
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and formulas is included in the following table (see also the associated figure for definitions):
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Functional trait
Abbreviation Formula
Ecological meaning
Oral gape position
Ops
Mo/Hd
Feeding position in the water column
(adapted from Sibbing & Nagelkerke 2001)
Eye size
Edst
Ed/Hd
Prey detection (adapted from Boyle & Horn
2006)
Eye position
Eps
Eh/Hd
Vertical position in the water column
(Gatz1979)
Pectoral fin position
Pfps
Pfi/PFb
Pectoral fin use for maneuverability (Dumay
et al. 2004)
Caudal propulsion efficiency through
reduction of drag (Webb 1984)
Caudal fin use for propulsion and/or direction
Aspect ratio of the caudal fin
Cfar
CFd²/CFs
(Webb 1984)
Vertical position in the water column (Gatz
Body shape
Bsh
PFB/Bi
1979)
For Flatfishes PFps was fixed to 0 for these species, by default Ops was fixed to 0 and Eyes position was
fixed to 1
Caudal peduncle throttling
Cpt
CFd/CPd
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Appendix S3. List of scientific names of the 81 Mediterranean endemic fish species. Species were
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ranked according to the FGO index, (Functional and Geographic Overlap). IUCN status: CR:
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Critically Endangered, EN: Endangered, VU: Vulnerable, NT: Near Threatened, LC: Least Concern,
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DD: Data Deficient, NL: Non Listed.
Species
Didogobius bentuvii
Sphyraena viridensis
Panturichthys fowleri
Millerigobius macrocephalus
Tripterygion delaisi
Chromogobius zebratus
Gobius vittatus
Symphodus melanocercus
Solea aegyptiaca
Pomatoschistus tortonesei
Symphodus doderleini
Symphodus cinereus
Aphanius iberus
Aphanius fasciatus
Tripterygion tripteronotus
Gobius strictus
Diplodus sargus
Raja melitensis
Psetta maxima
Parablennius zvonimiri
Scorpaenodes arenai
Microichthys sanzoi
Gouania willdenowi
Didogobius splechtnai
Chromogobius quadrivittatus
Eutrigla gurnardus
Engraulis encrasicolus
Tetrapturus belone
Cyclothone pygmaea
Lepadogaster lepadogaster
Sprattus sprattus
Gammogobius steinitzi
Pomatoschistus bathi
Ophichthus rufus
Arnoglossus kessleri
Lesueurigobius suerii
Tripterygion melanurus
Notoscopelus elongatus
Ophidion rochei
Zosterisessor ophiocephalus
Lipophrys nigriceps
FGO IUCN status
100.50
DD
58.90
LC
40.71
LC
31.88
NL
22.80
DD
22.38
NL
21.71
NL
20.93
DD
18.87
LC
18.71
NL
18.68
DD
18.21
NL
17.90
NL
17.61
NL
17.60
DD
17.47
NL
17.35
DD
16.33
NL
16.08
NL
16.08
NL
15.87
LC
15.49
NL
15.47
LC
15.37
NL
14.48
NL
14.43
NT
13.96
NL
13.82
LC
13.72
NL
13.44
NL
13.40
NL
13.28
NL
13.09
NL
12.94
NL
12.86
DD
12.47
DD
12.22
DD
12.12
NL
11.93
NL
11.84
DD
11.61
DD
Species
Odondebuenia balearica
Bathypterois mediterraneus
Buenia affinis
Microichthys coccoi
Paralepis coregonoides
Dasyatis tortonesei
Gobius geniporus
Gobius xanthocephalus
Raja rondeleti
Gobius ater
Paraliparis murieli
Pomatoschistus quagga
Paralepis speciosa
Platichthys luscus
Raja polystigma
Gymnammodytes cicerelus
Pseudaphya ferreri
Syngnathus tenuirostris
Trisopterus luscus
Lipophrys basiliscus
Knipowitschia panizzae
Gobius fallax
Parophidion vassali
Lepidion lepidion
Corcyrogobius liechtensteini
Huso huso
Didogobius schlieweni
Oligopus ater
Paraliparis leptochirus
Opeatogenys gracilis
Speleogobius trigloides
Thorogobius macrolepis
Acipenser naccarii
Syngnathus taenionotus
Lipophrys adriaticus
Acipenser stellatus
Pomatoschistus knerii
Pomatoschistus pictus
Pomatoschistus canestrinii
Nansenia iberica
FGO
11.54
11.42
11.17
11.14
10.85
10.83
10.77
10.59
9.92
9.76
9.29
9.28
9.18
8.57
8.53
8.35
8.24
7.95
7.75
7.36
7.19
7.03
7.03
6.25
6.22
5.70
5.41
4.99
4.95
4.83
4.22
4.19
3.88
2.13
1.90
1.85
1.71
1.67
1.21
0.00
IUCN status
NL
DD
DD
CR
NL
NL
NL
LC
LC
NL
DD
NL
VU
NL
LC
NL
NL
LC
LC
NL
NL
DD
NL
LC
DD
NL
NL
NL
NL
NL
DD
NL
NL
DD
NL
NL
DD
NL
NL
LC
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Appendix S4 Scientific names of the 57 Atlantic fish species.
Species
Species
Acanthurus monroviae
Microchirus hexophthalmus
Aluterus monocerus
Pagellus bellottii bellottii
Arius parki
Parablennius pilicornis
Beryx splendens
Parapristipoma octolineatum
Carcharhinus altimus
Pinguipes brasilianus
Carcharhinus brachyurus
Pisodonophis semicinctus
Carcharhinus falciformis
Pontinus kuhlii
Centrolabrus exoletus
Pristis pectinata
Cephalopholis taeniops
Psenes pellucidus
Chaunax pictus
Pseudupeneus prayensis
Cheilopogon furcatus
Rhizoprionodon acutus
Cyclopterus lumpus
Scorpaena stephanica
Diodon hystrix
Selene dorsali
Diplodus bellotii
Seriola carpenteri
Enchelycore anatina
Seriola fasciata
Entelurus aequoraeus
Seriola rivoliana
Ephippion guttiferum
Serranus atricauda
Fistularia petimba
Solea senegalensis
Galeocerdo cuvieri
Sphoeroides marmoratus
Galeoides decadactylus
Sphoeroides pachygaster
Gephyroberyx drawini
Sphoeroides spengleri
Gobius couchi
Sphyrna mokarran
Gymnammodytes semisquamatus
Synaptura lusitanica
Halosaurus ovenii
Syngnathus rostellatus
Laemonema latifrons
Tetrapturus georgei
Lagocephalus lagocephalus
Trachyscorpia cristulata
Lepidion guentheri
Umbrina canariensis
Lesueurigobius sanzoi
Zenopsis conchifer
Lipophrys pholis
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Appendix S5 Scientific names of the 74 Lessepsian fish species.
Species
Species
Abudefduf vaigiensis
Papilloculiceps longiceps
Alepes djedaba
Parexocoetus mento
Apogon pharaonis
Pelates quadrilineatus
Apogon queketti
Pempheris vanicolensis
Apogon smithi
Petroscirtes ancylodon
Atherinomorus forskali
Platycephalus indicus
Atherinomorus lacunosus
Plotosus lineatus
Callionymus filamentosus
Pomadasys stridens
Chilomycterus spilostylus
Priacanthus hamrur
Crenidens crenidens
Pteragogus pelycus
Cynoglossus sinusarabici
Pterois miles
Decapturus russelli
Rachycentron canadum
Dussumieria elapsoides
Rastrelliger kanagurta
Epinephelus coioides
Rhabdosargus haffara
Epinephelus malabaricus
Rhynchonger trewavasae
Etrumeus teres
Sargocentron rubrum
Fistularia commersonii
Saurida undosquamis
Glaucostegus halavi
Scarus ghobban
Hemiramphus far
Scomberomorus commerson
Heniochus intermedium
Siganus luridus
Herklotsichthys punctatus
Siganus rivulatus
Himantura uarnak
Silhouettea aegyptia
Hippocampus fuscus
Sillago sihama
Hyporhamphus affinis
Sorsogona prionota
Iniistius pavo
Sphyraena chrysotaenia
Lagocephalus sceleratus
Sphyraena flavicauda
Lagocephalus spadiceus
Spratelloides delicatulus
Lagocephalus suezensis
Stephanolepis diaspros
Leiognathus klunzingeri
Synagrops japonicus
Liza carinata
Terapon puta
Lutjanus argentimaculatus
Terapon therapes
Monishia ochetica
Tetrosomus gibbosus
Muraenesox cinereus
Torquigener flavimaculosus
Nemipterus randalli
Tylerius spinosissimus
Omobranchus punctatus
Tylosorus choran
Oxyurichthys petersi
Upeneus moluccensis
Pampus argentus
Upeneus pori
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Appendix S6: Influence of the weighting of traits on the FNNr and FGO indices.
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In the first approach (a, b) we classified the traits in two categories, the ecomorphological traits and
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the biological/ecological traits. To do so, we attributed a weight of 1/7 for each ecomorphological
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trait and a weight of 1/10 for each biological/ecological trait. In the second approach (c, d) we
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classified each trait into four major categories : foraging ecology, habitat use, life history, behaviour.
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Each category has a weight of 1 and the weighting of each trait depends on the number of traits by
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category. For example, we considered 8 traits in the habitat use category and each trait had therefore
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a weight of 1/8 for this category. Overall, the results showed that the FNNr and FGO indices are
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robust to the considered weighting schemes. We indeed found high spearman’s correlation values (ρ
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>0.90 for each pairwise comparison) between weighted and unweighted indices (*P<0.0001).
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Appendix S7: Influence of the introduction date on the FNNr and FGO indices
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The geographical range size of an exotic species is expected to be closely related to the date
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of introduction, which may influence the FNNr and FGO values. In this complementary analysis,
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introduction dates were compiled from Ben Rais Lasram & Mouillot (2009) and the CIESM
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(Golani et al., 2002) and grouped into four main periods (1: <1960, 2:1960-1980, 3:1980-2000, 4:
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>2000). We first tested whether the spatial distribution of FNNr values changed according to the
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date of introduction (see the figure below). To do so, we first calculated the FNNr by only
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considering exotic species introduced before 1960 (a). We then recalculated the FNNr index for
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each grid cell by further considering species introduced between 1960-1980 (b). We repeated this
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operation for the other time periods (c: 1980-2000 and d:>2000). Overall, this analysis showed that
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the spatial pattern of FNNr was consistent across the four time periods, with the Levantine basin
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showing the grestest values of FNNr.
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Then, we performed a similar analysis by considering the FGO index. The ranking of the 81
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endemic species, according to their FGO values, was highly comparable across the four time
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periods defined by the date of introduction of exotic species (Spearman’s correlation test: ρ>0.93
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for each pairwise comparison, P<0.0001).
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References
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Ben Rais Lasram, F. & Mouillot, D. (2009) Increasing southern invasion enhances congruence
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69
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between endemic and exotic Mediterranean fish fauna. Biological Invasions, 11, 697-711.
Golani, D., Massuti, E., Orsi Relini, L. & Quignard, J.P. (2002) The CIESM atlas of exotic species
in the Mediterranean Fishes edn. F.Briand CIESM, Monaco.
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