ele12351-sup-0001-suppInfo

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Supplementary Information for:
“External morphology explains the success of biological invasions”
Ernesto Azzurro*, Victor M. Tuset, Antoni Lombarte, Francesc Maynou, Daniel Simberloff, Ana
Rodríguez-Pérez and Ricard V. Solé
*correspondence to: eazzurr@gmail.com
Supplementary Material Table S1: Eco-morphological guilds of species used for the morphological analyses. The
first two warps that explained the most variation of the group were used to project the morphological position of
each species/group in a two-dimensional plot.
Guild
Orders/Families
Eco-morphological charcteristics
1
Anguiliformes, Ophidiiformes, Lampridiformes, Siluriiformes
Benthic fishes with very enlongated body-shape living in caves
or buried
2
Gastereosteiformes
Benthic fishes, mouth small and tube-shaped snout
3
Tetraodontiformes, Zeiformes, Perciformes (Caproidae)
Fishes with specializated locomotion and/or capacity of inflation
4
Perciformes (Blenniidae, Callionymidae, Clinidae, Gobiidae,
Gobiesocidae, Tripterygiidae)
Small benthic fishes with gobiid-like body shape
5
Aulopiformes, Batrachoidiformes, Lophiiformes, Perciformes
(Trachinidae and Uranoscopidae), Scorpaeniformes
Benthic fishes mainly living on the sea bottom or buried in soft
bottoms
6
Pleuronectiormes
Benthic fishes with asymmetrical oval body-shape, living buried
in soft bottom
7
Perciformes (Moronidae, Serranidae, Sillaginidae)
Benthic species with serranid-like body shape, carnivorous, with
ability to swim fast over short distances
8
Percifomes (Centracanthidae, Chaetodontidae, Haemulidae,
Labridae, Leiognathidae
Lobotidae, Lutjanidae, Mullidae, Nemipteridae,
Pomacanthidae, Pomacentridae, Scaridae, Sciaenidae,
Siganidae, Sparidae, Terapontidae)
Benthic fishes living isolated or in school, low to medium trophic
level
9
Mugiliformes
Benthic fishes, many euryhaline with low trophic level
10
Warps
1 and 2
Atheriniformes, Clupeiformes
1 and 3
Small coastal pelagic fishes of low trophic level
11
Beloniformes, Perciformes (Echeniidae and Sphyraenidae)
Coastal pelagic fishes with enlongated body-shape
12
Perciformes (Carangidae, Coryphaenidae, Istiophoridae,
Luvaridae, Nomeidae, Pomatomidae, Scombridae,
Stromateidae, Xiphiidae)
Oceanic and coastal pelagic fishes, fast swimmers with high
2 and 3
trophic level
13
Beryciformes and Perciformes (Apogonidae, Priacanthidae,
Pempheridae and Holocentridae)
Benthic fishes with nocturnal activity, large eyes and usually in
caves during daytime
Supplementary Material Table S2: Native (Med) and introduced fishes of Lessepsian origin (Les) used in the
analysis according to the guilds defined in Table S1. Values of the first three warps are indicated.
Guild
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Origin
Med
Med
Med
Les
Med
Les
Med
Les
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Les
Order
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Anguilliformes
Lampriformes
Ophidiiformes
Ophidiiformes
Ophidiiformes
Perciformes
Siluriiformes
Family
Anguillidae
Congridae
Congridae
Congridae
Muraenesocidae
Muraenesocidae
Muraenidae
Muraenidae
Muraenidae
Muraenidae
Ophichthidae
Ophichthidae
Ophichthidae
Ophichthidae
Ophichthidae
Ophichthidae
Trachipteridae
Carapidae
Ophidiidae
Ophidiidae
Cepolidae
Plotosidae
Species
Anguilla anguilla
Ariosoma balearicum
Conger conger
Rhynchoconger trewavasae
Cynoponticus ferox
Muraenesox cinereus
Anarchias euryurus
Gymnothorax reticularis
Gymnothorax unicolor
Muraena helena
Apterichtus anguiformis
Apterichtus caecus
Dalophis imberbis
Echelus myrus
Ophichthus rufus
Ophisurus serpens
Zu cristatus
Carapus acus
Ophidion barbatum
Ophidion rochei
Cepola macrophthalma
Plotosus lineatus
Warp 1
0.079
-0.174
-0.113
-0.196
-0.088
-0.175
-0.019
-0.163
-0.125
-0.131
-0.152
-0.134
-0.077
-0.109
-0.073
-0.188
0.009
-0.244
-0.128
-0.138
-0.261
-0.122
Warp 2
-0.29
-0.28
-0.273
-0.278
-0.268
-0.251
-0.308
-0.296
-0.265
-0.255
-0.32
-0.327
-0.32
-0.299
-0.318
-0.303
-0.198
-0.238
-0.188
-0.195
-0.245
-0.132
Warp 3
0.068
0.023
0.127
0.054
-0.027
0.033
-0.041
-0.013
-0.054
-0.011
-0.035
-0.053
0.041
-0.015
-0.002
0.03
-0.247
0.18
0.124
0.106
0.105
0.029
Guild
2
2
2
2
2
2
2
2
2
2
2
2
2
2
Origin
Les
Med
Les
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Order
Gastereostiformes
Gastereostiformes
Gastereostiformes
Gastereostiformes
Gastereostiformes
Gastereostiformes
Gastereostiformes
Gastereostiformes
Gastereostiformes
Gastereostiformes
Gastereostiformes
Gastereostiformes
Gastereostiformes
Gastereostiformes
Family
Fistulariidae
Macroramphosidae
Syngnathidae
Syngnathidae
Syngnathidae
Syngnathidae
Syngnathidae
Syngnathidae
Syngnathidae
Syngnathidae
Syngnathidae
Syngnathidae
Syngnathidae
Syngnathidae
Species
Fistularia commersonii
Macroramphosus scolopax
Hippocampus fuscus
Hippocampus guttulatus
Hippocampus hippocampus
Nerophis lumbriciformis
Nerophis maculatus
Nerophis ophidion
Syngnathus tenuirostris
Syngnathus abaster
Syngnathus acus
Syngnathus rostellatus
Syngnathus taeniotus
Syngnathus typhle
Warp 1
0.321
0.261
0.275
0.292
0.278
0.287
0.265
0.273
0.257
0.29
0.284
0.262
0.258
0.296
Warp 2
-0.184
0.101
-0.181
-0.193
-0.155
-0.305
-0.31
-0.302
-0.254
-0.271
-0.255
-0.253
-0.247
-0.241
Warp 3
0.182
0.199
-0.321
-0.236
-0.267
-0.245
-0.287
-0.229
-0.178
-0.182
-0.175
-0.105
-0.156
-0.11
Guild
Origin
Order
Family
Species
Warp 1
Warp 2
Warp 3
Med
Med
Les
Med
Les
Les
Les
Med
Les
Les
Les
Les
Med
Perciformes
Tetraodontiformes
Tetraodontiformes
Tetraodontiformes
Tetraodontiformes
Tetraodontiformes
Tetraodontiformes
Tetraodontiformes
Tetraodontiformes
Tetraodontiformes
Tetraodontiformes
Tetraodontiformes
Zeiformes
Caproidae
Balistidae
Diodontidae
Molidae
Monacanthidae
Ostraciidae
Ostraciidae
Tetraodontidae
Tetraodontidae
Tetraodontidae
Tetraodontidae
Tetraodontidae
Zeidae
Capros aper
Balistes capriscus
Cyclichthys spilostylus
Ranzania laevis
Stephanolepis diaspros
Ostracion cubicus
Tetrosomus gibbosus
Lagocephalus lagocephalus lagocephalus
Lagocephalus sceleratus
Lagocephalus spadiceus
Lagocephalus suezensis
Torquigener flavimaculosus
Zeus faber
-0.012
0.028
0.257
0.335
-0.047
0.217
0.224
0.285
0.26
0.238
0.247
0.263
0.05
0.242
0.152
0.104
0,000
0.154
0.004
0.038
-0.028
-0.01
0.07
-0.01
0.05
0.153
-0.011
-0.101
0.143
0.249
-0.075
0.06
0.033
0.121
0.138
0.096
0.129
0.13
0.101
3
3
3
3
3
3
3
3
3
3
3
3
3
Supplementary Material Table S2 continued
Guild
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
Origin
Order
Family
Species
Warp 1
Warp 2
Warp 3
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Les
Med
Med
Med
Med
Les
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Les
Med
Med
Med
Med
Med
Med
Les
Med
Med
Med
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Blenniidae
Callionymidae
Callionymidae
Callionymidae
Callionymidae
Callionymidae
Callionymidae
Callionymidae
Clinidae
Gobiesocidae
Gobiesocidae
Gobiesocidae
Gobiesocidae
Gobiesocidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Aidablennius sphynx
Blennius ocellaris
Coryphoblennius galerita
Hypleurochilus bananensis
Lipophrys adriaticus
Lipophrys canevae
Lipophrys dalmatinus
Lipophrys nigriceps
Lipophrys pholis
Parablennius gattorugine
Parablennius incognitus
Parablennius pilicornis
Parablennius rouxi
Parablennius sanguinolentus
Parablennius tentacularis
Parablennius zvonimiri
Paralipophrys trigloides
Petroscirtes ancylodon
Salaria basilisca
Salaria pavo
Scartella cristata
Callionymus fasciatus
Callionymus filamentosus
Callionymus lyra
Callionymus maculatus
Callionymus pusillus
Callionymus reticulatus
Callionymus risso
Clinitrachus argentatus
Apletodon dentatus dentatus
Apletodon incognitus
Diplecogaster bimaculata bimaculata
Lepadogaster candolii
Lepadogaster lepadogaster
Aphia minuta
Benthophilus stellatus
Buenia affinis
Chromogobius quadrivittatus
Chromogobius zebratus
Corcyrogobius liechtensteini
Coryogalops ochetica
Crystallogobius linearis
Deltentosteus collonianus
Deltentosteus quadrimaculatus
Didogobius bentuvii
Didogobius schlieweni
Didogobius splechtnai
Favonigobius melanobranchus
Gammogobius steinitzi
Gobius ater
Gobius auratus
-0.115
-0.098
-0.119
-0.147
-0.099
-0.09
-0.116
-0.087
-0.122
-0.123
-0.105
-0.165
-0.143
-0.125
-0.129
-0.113
-0.134
-0.124
-0.139
-0.128
-0.148
-0.094
-0.102
0.026
-0.048
-0.111
-0.066
-0.091
-0.189
0.21
0.253
0.247
0.134
0.144
0.049
-0.011
0.039
-0.002
-0.015
0.046
-0.03
-0.073
0.052
0.05
-0.004
0.014
0.041
-0.023
0.009
-0.024
-0.017
0.014
0.089
-0.041
0.057
-0.041
-0.038
-0.087
-0.014
-0.072
0.028
-0.02
0.008
-0.048
0.006
-0.011
-0.05
-0.043
-0.04
-0.019
0.008
-0.022
0.049
0.041
0.047
0.067
-0.007
0.013
0.026
-0.095
-0.002
-0.05
-0.019
-0.08
-0.068
-0.088
0.052
0.041
-0.017
-0.046
0.001
-0.013
-0.098
0.018
0.016
-0.07
-0.055
0.015
0.021
0.02
-0.032
-0.021
-0.031
-0.008
0.005
-0.001
-0.02
0.014
-0.002
0.022
-0.012
-0.014
0.008
0.018
0.007
-0.008
0.01
0.03
0.005
0.018
0.046
0.042
-0.012
0.02
0.007
0.059
0.002
0.01
0.01
-0.006
-0.089
0.174
0.176
0.179
0.16
0.172
-0.032
0.026
0.025
0.041
0.023
0.03
-0.002
0.03
0.005
0.004
0.016
0.024
0.037
0.02
0.008
0.005
0.036
Supplementary Material Table S2 continued
Guild
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
Origin
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Les
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Les
Med
Med
Med
Les
Med
Med
Med
Med
Med
Med
Med
Med
Order
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Family
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobiidae
Gobioesocidae
Gobioesocidae
Tripterygiidae
Tripterygiidae
Tripterygiidae
Species
Gobius bucchichi
Gobius cobitis
Gobius couchi
Gobius cruentatus
Gobius fallax
Gobius gasteveni
Gobius geniporus
Gobius kolombatovici
Gobius niger
Gobius paganellus
Gobius roulei
Gobius vittatus
Gobius xanthocephalus
Gobiusculus flavescens
Knipowitschia caucasia
Knipowitschia panizzae
Lebetus guilleti
Lesueurigobius friesii
Lesueurigobius sanzi
Lesueurigobius suerii
Millerigobius macrocephalus
Odondebuenia balearica
Oxyurichthys petersi
Pomatoschistus bathi
Pomatoschistus canestrinii
Pomatoschistus knerii
Pomatoschistus marmoratus
Pomatoschistus microps
Pomatoschistus minutus
Pomatoschistus norvegicus
Pomatoschistus pictus
Pomatoschistus quagga
Pomatoschistus tortonesei
Pseudaphya ferreri
Silhouetta aegyptia
Speleogobius trigloides
Thorogobius ephippiatus
Thorogobius macrolepis
Trypauchen vagina
Vanneaugobius dollfusi
Zebrus zebrus
Zosterisessor ophiocephalus
Gouania willdenowi
Opeatogenys gracilis
Tripterygion delaisic
Tripterygion melanurus
Tripterygion tripteronotus
Warp 1
-0.042
0.023
0.003
-0.021
0.004
-0.022
-0.025
-0.011
0.028
-0.02
0.028
-0.023
-0.04
0.009
0.06
0.059
0.021
-0.025
0.045
-0.042
-0.033
0.031
-0.115
0.001
0.01
0.028
0.015
0.027
-0.033
0.013
-0.024
0.009
0.016
0.051
-0.055
0.04
-0.009
-0.011
-0.192
0.019
-0.021
-0.043
0.213
0.226
-0.08
-0.102
-0.114
Warp 2
-0.029
0,000
-0.044
-0.025
0.015
-0.021
-0.027
0.007
0.007
-0.023
0.024
-0.007
0.029
0.028
-0.036
0.017
-0.011
-0.003
0.059
-0.001
0.023
0.045
-0.041
-0.029
-0.057
-0.029
-0.035
-0.048
-0.028
-0.018
-0.023
0.001
0,000
-0.014
-0.049
-0.008
0.016
0.012
-0.187
0.002
-0.018
0.007
-0.149
-0.142
-0.012
0.006
0.038
Warp 3
0.002
0.016
0.037
0.013
0.013
0.021
0.004
0.009
0.054
0.037
0.07
-0.019
-0.022
-0.004
-0.017
0.001
0.006
0.044
0.11
0.03
0.034
0.032
0.064
-0.034
-0.022
-0.008
-0.006
-0.011
-0.013
0.001
-0.026
-0.04
0,000
-0.045
0.06
-0.014
-0.001
-0.003
0.076
0.036
-0.006
0.026
0.197
0.163
0.096
0.054
0.04
Supplementary Material Table S2 continued
Guild
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
Guild
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
Origin
Order
Family
Species
Warp 1
Warp 2
Warp 3
Les
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Les
Les
Les
Med
Les
Med
Med
Med
Med
Med
Med
Les
Med
Med
Med
Med
Med
Med
Med
Aulopiformes
Aulopiformes
Batrachoidiformes
Lophiiformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Scorpaeniformes
Synodontidae
Synodontidae
Batrachoidae
Lophiidae
Ammodytidae
Ammodytidae
Trachinidae
Trachinidae
Trachinidae
Trachinidae
Uranoscopidae
Cottidae
Dactylopteridae
Peristediidae
Platycephalidae
Platycephalidae
Platycephalidae
Scorpaenidae
Scorpaenidae
Scorpaenidae
Scorpaenidae
Scorpaenidae
Scorpaenidae
Scorpaenidae
Scorpaenidae
Synanceniidae
Triglidae
Triglidae
Triglidae
Triglidae
Triglidae
Triglidae
Triglidae
Saurida undosquamis
Synodus saurus
Halobatrachus didactylus
Lophius piscatorius
Ammodytes tobianus
Gymnammodytes cicerelus
Echiichthys vipera
Trachinus araneus
Trachinus draco
Trachinus radiatus
Uranoscopus scaber
Taurulus bubalis
Dactylopterus volitans
Peristedion cataphractum
Papilloculiceps longiceps
Platycephalus indicus
Sorsogona prionota
Helicolenus dactylopterus dactylopterus
Pterois miles
Scorpaena loppei
Scorpaena maderensis
Scorpaena notata
Scorpaena porcus
Scorpaena scrofa
Scorpaenodes arenai
Synanceia verrucosa
Aspitrigla cuculus
Aspitrigla obscura
Chelidonichthys lucerna
Eutrigla gurnardus
Lepidotrigla cavillone
Trigla lyra
Trigloporus lastoviza
0.138
0.197
-0.041
-0.1
-0.047
-0.034
-0.186
-0.187
-0.206
-0.161
-0.054
-0.051
0.025
-0.088
0.001
-0.027
0.02
-0.007
-0.03
-0.002
0.007
0.004
0.017
0.019
-0.019
0.028
-0.036
0.01
-0.037
-0.09
-0.104
-0.109
-0.074
-0.075
-0.086
-0.001
-0.031
-0.204
-0.196
0.012
-0.034
-0.06
-0.012
0.008
0.127
0.101
-0.048
-0.074
-0.103
-0.049
0.176
0.222
0.137
0.15
0.143
0.159
0.146
0.14
0.118
-0.005
0.023
-0.005
-0.033
0.045
0.05
0.036
-0.023
-0.111
0.009
-0.169
-0.012
0.006
0.021
0.016
0.017
0.021
0.057
-0.012
0.006
0.005
-0.006
0.029
0.009
-0.052
-0.049
-0.033
-0.04
-0.04
-0.036
-0.025
-0.048
0,000
-0.082
-0.058
-0.006
0.003
-0.05
-0.004
-0.036
Origin
Order
Family
Species
Warp 1
Warp 2
Warp 3
Med
Med
Med
Med
Med
Les
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Pleuronectiformes
Bothidae
Bothidae
Bothidae
Bothidae
Citharidae
Cynoglossidae
Pleuronectidae
Scophthalmidae
Scophthalmidae
Scophthalmidae
Scophthalmidae
Soleidae
Soleidae
Soleidae
Soleidae
Soleidae
Soleidae
Soleidae
Soleidae
Soleidae
Soleidae
Soleidae
Soleidae
Arnoglossus kessleri
Arnoglossus laterna
Arnoglossus thori
Bothus podas
Citharus linguatula
Cynoglossus sinusarabici
Platichthys flesus
Lepidorhombus boscii
Psetta maxima
Scophthalmus rhombus
Zeugopterus regius
Buglossidium luteum
Dicologlossa cuneata
Dicologlossa hexophtalma
Microchirus azevia
Microchirus ocellatus
Microchirus variegatus
Monochirus hispidus
Pegusa impar
Pegusa lascaris
Pegusa nasuta
Solea solea
Synapturichthys kleinii
-0.335
-0.315
-0.33
-0.306
-0.264
-0.354
-0.252
-0.32
-0.313
-0.278
-0.354
-0.344
-0.333
-0.335
-0.337
-0.333
-0.327
-0.349
-0.363
-0.332
-0.347
-0.328
-0.327
0.002
0.026
0.007
0.036
-0.005
-0.141
0.062
0.001
0.098
0.134
0.043
-0.076
-0.082
-0.029
-0.082
-0.038
-0.066
-0.002
-0.072
-0.075
-0.027
-0.103
-0.122
0.036
0.02
0.015
0.024
0.016
0.014
-0.026
0.022
0.014
0.026
0.046
0.023
0.041
0.025
0.026
0.036
0.015
0.044
0.061
0.04
0.007
0.032
0.003
Supplementary Material Table S2 continued
Guild
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
Origin
Med
Med
Med
Med
Med
Les
Med
Les
Les
Med
Med
Med
Med
Med
Med
Les
Order
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Family
Moronidae
Moronidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Serranidae
Sillaginidae
Species
Dicentrarchus labrax
Dicentrarchus punctatus
Anthias anthias
Epinephelus aeneus
Epinephelus caninus
Epinephelus coioides
Epinephelus costae
Epinephelus fasciatus
Epinephelus malabaricus
Epinephelus marginatus
Mycteroperca rubra
Serranus atricauda
Serranus cabrilla
Serranus hepatus
Serranus scriba
Sillago suezensis
Warp 1
0.075
0.094
0.013
0.027
0.047
0.017
-0.005
0,000
0.023
0.004
0.014
-0.02
0.012
-0.031
0.032
-0.044
Warp 2
0.023
0.04
0.229
0.079
0.078
0.111
0.09
0.124
0.078
0.133
0.127
0.094
0.066
0.145
0.111
-0.055
Warp 3
-0.074
-0.043
-0.064
-0.026
-0.029
-0.02
0.003
-0.014
-0.025
-0.013
-0.046
-0.002
-0.033
-0.01
-0.01
0.024
Guild
Origin
Order
Family
Species
Warp 1
Warp 2
Warp 3
Med
Med
Med
Les
Les
Les
Med
Med
Med
Les
Med
Med
Med
Med
Med
Med
Med
Med
Med
Les
Med
Med
Med
Med
Med
Med
Med
Med
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Centracanthidae
Centracanthidae
Centracanthidae
Chaetodontidae
Chaetodontidae
Chaetodontidae
Haemulidae
Haemulidae
Haemulidae
Haemulidae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Labridae
Centracanthus cirrus
Spicara maena
Spicara smaris
Chaetodon austriacus
Chaetodon larvatus
Heniochus intermedium
Parapristipoma octolineatum
Plectorhinchus mediterraneus
Pomadasys incisus
Pomadasys stridens
Acantholabrus palloni
Centrolabrus trutta
Coris julis
Ctenolabrus rupestris
Labrus bergylta
Labrus merula
Labrus mixtus
Labrus viridis
Lappanella fasciata
Pteragogus pelycus
Symphodus bailloni
Symphodus cinereus
Symphodus doderleini
Symphodus mediterraneus
Symphodus melanocercus
Symphodus melops
Symphodus ocellatus
Symphodus roissali
0.046
0.004
-0.04
-0.046
-0.074
0.115
0.015
-0.007
0.05
0.025
-0.02
-0.022
-0.049
0.004
0.027
0.029
0.007
0.004
0.008
-0.073
-0.037
-0.024
-0.011
-0.034
-0.001
-0.01
-0.032
0.026
0.007
0.056
0.026
0.188
0.273
0.294
0.157
0.159
0.143
0.111
0.041
0.12
-0.032
0.073
0.059
0.047
0.054
0.06
0.027
0.16
0.078
0.133
0.044
0.093
0.039
0.123
0.084
0.083
0.002
0.033
-0.017
-0.002
-0.049
0.078
-0.034
-0.07
-0.036
-0.036
0,000
0.006
0.005
0.014
-0.016
-0.013
-0.01
-0.019
-0.01
0.013
-0.024
-0.02
-0.008
-0.01
0.006
0.006
-0.008
-0.024
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
Supplementary Material Table S2 continued
Guild
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
Guild
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
Origin
Order
Family
Species
Warp 1
Warp 2
Warp 3
Med
Med
Med
Med
Les
Les
Med
Les
Med
Med
Les
Les
Les
Les
Les
Les
Med
Les
Med
Med
Med
Med
Med
Med
Les
Les
Les
Med
Les
Med
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Labridae
Labridae
Labridae
Labridae
Leiognathidae
Leiognathidae
Lobotidae
Lutjanidae
Mullidae
Mullidae
Mullidae
Mullidae
Mullidae
Nemipteridae
Pomacanthidae
Pomacanthidae
Pomacentridae
Scaridae
Scaridae
Sciaenidae
Sciaenidae
Sciaenidae
Sciaenidae
Sciaenidae
Siganidae
Siganidae
Sparidae
Sparidae
Sparidae
Sparidae
Symphodus rostratus
Symphodus tinca
Thalassoma pavo
Xyrichtys novacula
Equulites elongatus
Equulites klunzingeri
Lobotes surinamensis
Lutjanus argentimaculatus
Mullus barbatus barbatus
Mullus surmuletus
Parupeneus forsskali
Upeneus moluccensis
Upeneus pori
Nemipterus randalli
Pomacanthus imperator
Pomacanthus maculosus
Chromis chromis
Scarus ghobban
Sparisoma cretense
Argyrosomus regius
Sciaena umbra
Umbrina canariensis
Umbrina cirrosa
Umbrina ronchus
Siganus luridus
Siganus rivulatus
Acanthopagrus bifasciatus
Boops boops
Crenidens crenidens
Dentex dentex
-0.053
-0.018
-0.038
-0.129
-0.032
-0.034
-0.025
0.027
0.042
0.027
0.053
0.067
0.068
-0.008
-0.057
0.014
0.043
-0.019
-0.038
0.054
0.014
0.036
0.03
0.023
-0.107
-0.084
0.038
0.034
0.036
0.028
0.1
0.073
0.077
0.038
-0.002
0.141
0.139
0.137
0.084
0.096
0.09
0.075
0.043
0.133
0.178
0.153
0.143
0.088
0.079
-0.019
0.076
0.081
-0.001
0.049
0.068
0.067
0.206
-0.007
0.129
0.124
-0.017
-0.031
-0.017
-0.033
0.038
0.023
-0.028
-0.034
-0.066
-0.054
-0.018
-0.064
-0.061
-0.046
-0.029
-0.026
-0.03
-0.018
-0.045
-0.052
-0.105
-0.074
-0.08
-0.073
-0.002
0.006
-0.035
-0.049
-0.042
0.002
Origin
Order
Family
Species
Warp 1
Warp 2
Warp 3
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Les
Med
Med
Med
Les
Les
Les
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Sparidae
Terapontidae
Terapontidae
Terapontidae
Dentex gibbosus
Dentex macrophthalmus
Dentex maroccanus
Diplodus annularis
Diplodus cervinus cervinus
Diplodus puntazzo
Diplodus sargus sargus
Diplodus vulgaris
Lithognathus mormyrus
Oblada melanura
Pagellus acarne
Pagellus bogaraveo
Pagellus erythrinus
Pagrus auriga
Pagrus caeruleostictus
Pagrus pagrus
Rhabdosargus haffara
Sarpa salpa
Sparus aurata
Spondyliosoma cantharus
Pelates quadrilineatus
Terapon jarbua
Terapon puta
0.067
0.035
0.027
0.022
0.023
0.028
0.035
0.013
0.044
0.015
0.026
0.026
0.028
0.035
0.009
0.013
0.013
0.045
0.011
0.009
0.021
0.067
0.025
0.1
0.162
0.162
0.184
0.189
0.096
0.176
0.173
0.098
0.08
0.081
0.119
0.117
0.269
0.256
0.149
0.14
0.07
0.121
0.143
0.074
0.099
0.039
-0.001
-0.013
-0.009
-0.022
-0.017
-0.022
-0.025
0,000
-0.018
-0.01
-0.023
-0.03
-0.003
-0.033
-0.044
-0.015
-0.018
-0.01
-0.025
-0.031
-0.02
-0.039
-0.018
Supplementary Material Table S2 continued
Guild
9
9
9
9
9
9
9
Guild
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
Guild
11
11
11
11
11
11
11
11
11
11
11
11
11
11
11
11
11
11
11
11
11
11
Origin
Order
Family
Species
Warp 1
Warp 2
Warp 3
Med
Med
Les
Med
Med
Med
Med
Mugiliformes
Mugiliformes
Mugiliformes
Mugiliformes
Mugiliformes
Mugiliformes
Mugiliformes
Mugilidae
Mugilidae
Mugilidae
Mugilidae
Mugilidae
Mugilidae
Mugilidae
Chelon labrosus
Liza aurata
Liza carinata
Liza ramada
Liza saliens
Mugil cephalus
Oedalechilus labeo
0.14
0.129
0.135
0.151
0.136
0.129
0.148
-0.033
-0.026
-0.001
-0.011
-0.051
-0.027
-0.015
0,000
-0.006
0,000
0.018
-0.013
0.008
-0.003
Origin
Order
Family
Species
Warp 1
Warp 2
Warp 3
Med
Med
Med
Les
Med
Med
Les
Les
Les
Med
Med
Med
Les
Med
Med
Les
Atheriniformes
Atheriniformes
Atheriniformes
Atheriniformes
Clupeiformes
Clupeiformes
Clupeiformes
Clupeiformes
Clupeiformes
Clupeiformes
Clupeiformes
Clupeiformes
Clupeiformes
Clupeiformes
Clupeiformes
Clupeiformes
Atherinidae
Atherinidae
Atherinidae
Atherinidae
Clupeidae
Clupeidae
Clupeidae
Clupeidae
Clupeidae
Clupeidae
Clupeidae
Clupeidae
Clupeidae
Clupeidae
Engraulidae
Engraulidae
Atherina boyeri
Atherina hepsetus
Atherina presbyter
Atherinomorus forskali
Alosa alosa
Alosa fallax
Dussumieria elopsoides
Etrumeus golanii
Herklotsichthys punctatus
Sardina pilchardus
Sardinella aurita
Sardinella maderensis
Spratelloides delicatulus
Sprattus sprattus sprattus
Engraulis encrasicolus
Stolephorus insularis
0.115
0.142
0.114
0.141
0.179
0.182
0.253
0.258
0.217
0.204
0.219
0.22
0.229
0.241
0.208
0.141
-0.028
-0.086
-0.055
-0.04
0.049
0.021
-0.048
-0.056
-0.003
-0.021
-0.022
0.023
-0.001
-0.044
-0.048
0.028
-0.001
0.011
0.001
0.052
-0.077
-0.102
-0.052
-0.168
-0.106
-0.125
-0.154
-0.1
-0.133
-0.043
-0.048
0.003
Origin
Order
Family
Species
Warp 1
Warp 2
Warp 3
Med
Med
Med
Med
Med
Med
Les
Les
Med
Les
Med
Med
Med
Med
Med
Med
Med
Med
Les
Les
Med
Med
Beloniformes
Beloniformes
Beloniformes
Beloniformes
Beloniformes
Beloniformes
Beloniformes
Beloniformes
Beloniformes
Beloniformes
Beloniformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Belonidae
Belonidae
Belonidae
Exocoetidae
Exocoetidae
Exocoetidae
Exocoetidae
Hemiramphidae
Hemiramphidae
Hemiramphidae
Scomberesocidae
Echeneidae
Echeneidae
Echeneidae
Echeneidae
Echeneidae
Exocoetidae
Exocoetidae
Sphyraenidae
Sphyraenidae
Sphyraenidae
Sphyraenidae
Belone belone
Belone svetovidovi
Tylosurus acus imperialis
Exocoetus obtusirostris
Exocoetus volitans
Hirundichthys rondeletii
Parexocoetus mento
Hemiramphus far
Hemiramphus picarti
Hyporhamphus affinis
Scomberesox saurus saurus
Echeneis naucrates
Remora australis
Remora brachyptera
Remora osteochir
Remora remora
Cheilopogon exsiliens
Cheilopogon heterurus
Sphyraena chrysotaenia
Sphyraena flavicauda
Sphyraena sphyraena
Sphyraena viridensis
0.317
0.325
0.28
0.198
0.193
0.268
0.286
0.323
0.294
0.323
0.236
-0.139
-0.062
-0.091
-0.05
-0.073
0.292
0.284
0.112
0.117
0.131
0.127
-0.189
-0.184
-0.172
0.054
0.038
0.103
0.012
-0.115
-0.138
-0.141
-0.184
-0.175
-0.108
-0.12
-0.148
-0.095
0.018
-0.008
-0.014
-0.075
-0.097
-0.104
0.172
0.177
0.191
0.186
0.18
0.176
0.225
0.15
0.174
0.134
0.182
-0.046
-0.045
-0.068
-0.055
-0.065
0.14
0.157
-0.012
-0.04
-0.035
-0.043
Supplementary Material Table S2 continued
Guild
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
Origin
Les
Med
Med
Med
Med
Les
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Order
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Family
Carangidae
Carangidae
Carangidae
Carangidae
Carangidae
Carangidae
Carangidae
Carangidae
Carangidae
Carangidae
Carangidae
Carangidae
Carangidae
Coryphaenidae
Coryphaenidae
Istiophoridae
Istiophoridae
Species
Alepes djedaba
Campogramma glaycos
Caranx crysos
Caranx hippos
Caranx rhonchus
Decapterus russelli
Lichia amia
Naucrates ductor
Pseudocaranx dentex
Seriola dumerili
Trachinotus ovatus
Trachurus mediterraneus
Trachurus trachurus
Coryphaena equiselis
Coryphaena hippurus
Istiophorus albicans
Tetrapturus albidus
Warp 1
-0.055
-0.059
-0.017
-0.017
-0.036
-0.016
-0.08
-0.004
-0.002
0.012
-0.035
-0.026
-0.055
-0.098
-0.115
0.003
-0.009
Warp 2
0.062
0.009
0.081
0.129
0.007
0.037
0.041
0.034
0.105
0.003
-0.002
-0.012
0.034
-0.065
-0.061
-0.02
0.016
Warp 3
-0.004
-0.045
0.016
0.016
0.006
0.018
-0.026
-0.05
-0.002
-0.057
-0.006
0.015
0.012
-0.056
-0.036
-0.046
0,000
Guild
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
Origin
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Med
Les
Med
Med
Med
Les
Med
Med
Med
Med
Med
Order
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Family
Istiophoridae
Istiophoridae
Luvaridae
Nomeidae
Pomatomidae
Scombridae
Scombridae
Scombridae
Scombridae
Scombridae
Scombridae
Scombridae
Scombridae
Scombridae
Scombridae
Scombridae
Scombridae
Scombridae
Scombridae
Stromateidae
Xiphiidae
Species
Tetrapturus belone
Tetrapturus georgii
Luvarus imperialis
Cubiceps gracilis
Pomatomus saltatrix
Acanthocybium solandri
Auxis rochei rochei
Auxis thazard thazard
Euthynnus alletteratus
Katsuwonus pelamis
Orcynopsis unicolor
Rastrelliger kanagurta
Sarda sarda
Scomber colias
Scomber scombrus
Scomberomorus commerson
Scomberomorus tritor
Thunnus alalunga
Thunnus thynnus
Stromateus fiatola
Xiphias gladius
Warp 1
-0.048
-0.025
-0.002
-0.003
0.019
-0.001
0.024
-0.024
-0.011
0.002
-0.085
-0.005
-0.012
0.02
-0.015
-0.12
-0.078
0.001
-0.006
-0.033
0.155
Warp 2
-0.05
-0.045
-0.002
0.052
-0.014
-0.15
-0.087
-0.067
-0.083
-0.029
-0.012
-0.016
-0.078
-0.093
-0.116
-0.102
-0.077
0.041
0.008
0.185
-0.012
Warp 3
-0.011
0.018
0.154
0.01
0.011
0.026
-0.037
-0.029
-0.037
-0.001
-0.037
-0.004
-0.035
-0.017
-0.024
-0.041
-0.005
0.022
-0.028
0.025
-0.076
Guild
Origin
Order
Family
Species
Warp 1
Warp 2
Warp 3
Les
Les
Med
Les
Les
Les
Les
Les
Les
Beryciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Perciformes
Holocentridae
Apogonidae
Apogonidae
Apogonidae
Apogonidae
Apogonidae
Apogonidae
Pempheridae
Priacanthidae
Sargocentron rubrum
Ostorhinchus fasciatus
Apogon imberbis
Apogonichthyoides pharaonis
Cheilodipterus novemstriatus
Jaydia queketti
Jaydia smithi
Pempheris rhomboidea
Priacanthus sagittarius
0.051
0.034
0.031
0.045
0.073
-0.013
0.036
0.065
-0.062
0.165
0.158
0.2
0.24
0.122
0.191
0.102
0.191
0.164
-0.075
-0.036
-0.05
-0.024
-0.051
-0.041
-0.051
-0.03
0.009
13
13
13
13
13
13
13
13
13
Supplementary Figure S1: Results of morphological analyses of the thirteen groups of species (Guilds 1-13)
(Supplementary Tables S1, S2). Native species are indicated with black triangles and Lessepsian species with
circles. The a priori known ranks of abundance were indicated as follows: (1) green circles = not established or rare;
(2) yellow circles = occasional to common but never very abundant; (3) red circles = very abundant. Note that
abundant invaders are generally added outside of or at the margins of the original convex hull, whereas other
introduced species are mostly found within the original morphospace and are morphologically similar to resident
species. Fish images from FAO. Analyses were computed sequentially, according to the date of first record of each
Lessepsian species: Colors indicating the four temporal windows are reported in the plot representing the first group
(Guild 1).
Fig S1, guilds 1 and 2
Guild 1
Entered(<(1951(
O.#barbatum#
Z.#cristatus#
Entered(in(1951%1975(
Entered(in(1976%2002(
C.#acus#
C.#macrophthlama#
Entered(>(2002(
Original(assemblage(
A.#anguilla#
O.#serpens#
A.#euryurus#
A.#caecus#
D.#imberbis#
P.#lineatus#
M.#cinereus#
R.#trewavasae#
G.#re3cularis#
>(2002(
1976%2002(
Guild 2
Original(assemblage(
1976%2002(
M.#scolopax#
H.#guE ulatus#
S.#taenionotus#
S.#typhle#
S.#tenuirostris#
N.#maculatus#
N.#lumbriciformis#
H.#fuscus##
F.#commersonii##
Fig S1, guilds 3 and 4
Guild 3
Original(assemblage(
<(1951(
C.#aper#
B.#capriscus#
S.#diaspros#
L.#spadiceus#
R.#laevis#
L.#lagocephalus#
C.#spilostylus#
T.#gibbosus#
T.#flavimaculosus#
O.#cubicus#
L.#suezensis#
1976%2002(
L.#sceleratus#
>(2002(
<"1951(
Original(assemblage(
Guild 4
B.#ocellaris#
L.#sanzoi#
H.#bananensis#
P.#pillicornis#
A.#dentatus#
D.#bimaculata#
C.#oche3ca*#
A.#incognitus#
C.#argentatus#
O.#gracilis#
G.#willdenowi#
C.#filamentosus##
F.#melanobranchus#
O.#petersi*#
S.#aegyp3a#
P.#ancylodon#
T.#vagina#
1951%1975(
1976%2002(
>(2002(
Fig S1, guilds 5 and 6
<(1951(
Original(assemblage(
Guild 5
U.#scaber#
S.#prionota#
D.#volitans#
T.#draco#
S.#saurus#
L.#piscatorius#
P.#indicus#
P.#longiceps#
S.#undosquamis#
S.#verrucosa#
P.#miles#
1951%1975(
>(2002(
1976%2002(
Guild 6
Original(assemblage(
P.#impar#
C.#linguatula#
P.#nasuta#
1951%1975(
1950'1075"
C.#sinusarabici#
P.#flesus#
Fig S1, guilds 7 and 8
E.#costae#
Guild 7
1951%1975(
Original(assemblage(
S.#atricauda#
S.#scriba#
E.#coioides##
S.#hepatus#
E.#malabaricus##
D.#punctatus#
A.#anthias#
D.#labrax#
S.#suezensis#
E.#fasciatus##
Groupr 9
>2002(
1976%2002(
Guild 8
Original(assemblage(
S.#maena#
<1951(
E.#klunzingeri#
D.#gibbosus#
S.#rivulatus#
X.#novacula#
A.#regius#
U.#pori#
U.#moluccensis#
S.#umbra#
H.#intermedium#
E.#elongatus#
P.#pelycus#
S.#luridus#
M.#grandoculis#
C.#austriacus#
S.#ghobban*#
P.#quadrilineatus*#
P.#stridens#
1951%1975(
T.#puta#
R.#haffaa #
P.#forsskali#
L.#argen3maculatus#
C.#crenidens#
1976%2002(
P.#imperator*# P.#maculosus#
C.#larvatus*#
>2002(
T.#theraps#
A.#bifasciatus*# T.#jarbua#
N.#randalli#
Fig S1, guilds 9 and 10
Guild 9
Original(assemblage(
<1951((
L.#ramada#
M.#cephalus#
L.#carinata#
O.#labeo#
L.#aurata#
L.#saliens#
Original(assemblage(
1951%1975(
<1951(
>2002(
Guild 10
Fig S1, guilds 11 and 12
T.#acus#
Guild 11
<(1951(
Original(assemblage(
P.#mento#
E.#obtrusirostris#
S.#saurus#
H.#rondele3#
B.#belone#
H.#far#
S.#chrysotaenia#
E.#naucrates#
R.#osteochir#
S.#viridensis#
R.#brachyptera#
H.#affinis#
S.#flavicauda#
1951%1975(
Original(assemblage(
1976%2002(
<1951(
Guild 12
L.#imperialis#
L.#imperialis#
S.#flatola#
S.#scrombrus#
A.#djedaba#
A.#solandrii#
S.#commerson#
C.#equiselis#
S.#gladius#
D.#russelli#
R.#kanagurta#
1951%1975(
>2002(
Fig S1, guilds 13
Guild 13
< 1951
A. pharaonis
A. imberbis
S. rubrum
P. sagi arius
P. rhomboidea
O. fasciatus
J. queke
J. smithi
C. novemstriatus
1976-2002
> 2002
Supplementary text
Introduction:
The idea of “unsaturated”, “empty” or “vacant” niches” has often been advanced to explain the
success of non-native species in the Mediterranean literature (e.g. Por 1978; Ben-Tuvia 1985).
Methods - the conceptual approach:
Using a landmark-based procedure (Bookstein 1991), we calculated the convex hull of
existing morphologies (recipient community), and we examined the relative position of a newly
introduced species in relation to its abundance in its invaded range.
Size is an important trait to be considered in ecological studies, since it is related to many
facets of fish niche such as diet, metabolism, mobility, and longevity. The reason why we did not
consider this attribute in our model is because its inclusion would have masked other traits
associated with shape. Thus, we considered body shape to be less variable than size within
species so that it may better predict interspecific differences in fish niche. If we take Fistularia
commersonii or other abundant invaders, the uniqueness of their external morphology seems to
be robust enough to inform us about other unmeasured dimensions of species strategy in
comparison to the other components of the receiving community. Nevertheless, we acknowledge
that one of the most important challenges in the field of geometric morphology is to deal with the
effect of size and allometry, and it remains challenging to test whether morphologically close
species tend to compete more than species with different morphologies (e.g., among native
species at a local scale).
We did not consider variables such as intraspecific variability, real species cooccurrences, and species abundances of natives in the receiving community. We worked at the
regional scale and used the entire set of Mediterranean species when testing the position of the
newcomers. Similar simplifications have been used to explore the morphological space of fossil
assemblages (e.g. Shen et al. 2008), where co-occurrence at the community level is very difficult
to infer. Reasons why we did not consider commonly used indices such as nearest-neighbour
distances or mean distance to other species are related to the fact that these metrics do not
provide information about the relative position of species within the morphospace. As an
example, Scomberomorus commerson is found near other scomberiformes, but at the same time
this invader also is located outside the original range of morphologies, the convex hull.
According to our model, this position identifies the maximum rank of morphological (ecological)
opportunities. On the contrary, Liza carinata shows a good morphological distance from other
components of its guild, but it remains enclosed within the convex hull.
Methods - the definition of the guilds:
Guilds are useful in the comparative study of communities, since it is usually impossible
to study all species living in an ecosystem at once. The (N=13) guilds represent proposed sets of
species defined on the basis of commonly accepted taxonomic, ecological, and morphological
principles, but other criteria could be used as well. Certainly, given the scale of our study (the
entire Mediterranean region), it is clear that in some cases the guilds used may include species
that do not really coexist. Indeed, guilds represent the entire morphospace occupation and
variation of species that share similar ways of exploiting environmental resources. A species like
Fistularia commersonii is very different from any other species of the Mediterranean Sea.
Nothing comparable to this fish exists in this basin and such morphological uniqueness is
confirmed also in other groups of species.
Methods - species lists and ranks of abundance:
The native species list was obtained from available bibliographic sources (Quignard &
Tomasini 2000) updated with the most recent literature (e.g. for gobid species Patzner 2008).
Depth ranges were obtained from the FAO guides (Fischer & Bianchi 1984; Fischer et al. 1987),
plus specific bibliographic searches. In the absence of density data on all Lessepsian fish species,
abundance levels (in the form of an ordered qualitative variable) were determined from the
available atlas (Golani et al. 2013) and guides (Bariche 2012). We took into account some recent
updates regarding the names of Lessepsian species and considered Pempheris rhomboidea
instead of P. vanicolensis, according to (Randall et al. 2013).
Results and Discussion:
Our approach can help elucidate mechanisms underlying invasions (e.g., biotic vs.
environmental filtering), and it could be extended to other systems - e.g. plants (Pausas & Verdú
2008), molluscs (Navarro et al. 2005), and birds (Moulton et al. 2001).
About one-third (36.7%) of Lessepsian fish had no confamilials among native species
and more than a half (54.4%) had fewer than two. The number of native confamilials was not
significantly related to the abundance level of Lessepsians (X2 =10.13, df=6, P=0.119).
Nevertheless it is interesting to highlight that Lessepsian migrants added two new orders
(Beryciformes and Siluriformes) to our list of Mediterranean species, and both are represented by
fishes (Sargocentron rubrum and Plotosus lineatus, respectively) that developed abundant
populations in their invaded range. Future studies might explore in detail the importance of
phylogenetic relationships within the framework of ecological theory.
Supplementary references
51.
Bariche, M. (2012). Field identification guide to the living marine resources of the Eastern and
Southern Mediterranean. FAO Species Identification Guide for Fishery Purposes, FAO,
Rome.
52.
Bookstein, F.L. (1991). Morphometric tools for landmark data. Geometry and Biology.
Cambridge University Press, New York.
53.
Fischer, W. & Bianchi, G. (1984). FAO Species Identification Sheet for Fishery Purposes.
Western Indian Ocean, (Fishing Area 51), Vol II, FAO, Rome.
54.
Fischer, W., Bauchot M.L., Schneider M. (1987). Fiches FAO d’Identificacion des Espèces pour
les Besoins de la Peche. Méditerranée et Mer Noire. Zone de Pêche 37. Volume II, Vertébrés,
FAO, Rome.
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Moulton, M.P., Sanderson, J.G. & Labisky, R.F. (2001). Patterns of success in game bird (Aves:
Galliformes) introductions to the Hawaiian islands and New Zealand. Evol. Ecol. Res., 3, 507519.
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Navarro, N., Neige, P. & Marchand, D. (2005). Faunal invasions as a source of morphological
constraints and innovations? The diversification of the early Cardioceratidae (Ammonoidea;
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Patzner, R.A. (2008) Mediterranean gobies. available at
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Pausas, J. G. & Verdú, M. (2008). Fire reduces morphospace occupation in plant communities.
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Quignard, J.P & Tomasini J.A. (2000). Mediterranean fish biodiversity. Biol. Mar. Med. 7, 1-66.
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Randall, J.E., Bogorodsky, S.V., Alpermann, T.J., Satapoomin, U., Mooi, R.D. & Mal, A.O.
(2013) Pempheris flavicycla, a new pempherid fish from the Indian Ocean, previously
identified as P. vanicolensis Cuvier. Journal of the Ocean Science Foundation, 9, 1–23.
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