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. 55. 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. 56. Navarro, N., Neige, P. & Marchand, D. (2005). Faunal invasions as a source of morphological constraints and innovations? The diversification of the early Cardioceratidae (Ammonoidea; Middle Jurassic). Paleobiology, 31, 98-116. 57. Patzner, R.A. (2008) Mediterranean gobies. available at http://www.patzner.sbg.ac.at/Gobiidae.htm. 58. Pausas, J. G. & Verdú, M. (2008). Fire reduces morphospace occupation in plant communities. Ecology, 89, 2181-2186. 59. Quignard, J.P & Tomasini J.A. (2000). Mediterranean fish biodiversity. Biol. Mar. Med. 7, 1-66. 60. 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. 61. Shen, B., Dong, L., Xiao, S., & Kowalewski, M. (2008). The Avalon explosion: evolution of Ediacara morphospace. Science, 319(5859), 81-84.