Abstract
Economidichthys mornosensis, a new species, is described from the Mornos River delta in southwestern Greece. It is related to Economidichthys pygmaeus, from which it is distinguished by having a large black blotch on the first dorsal fin in males, a short perianal organ, and a scaleless back along the complete base of the second dorsal fin. Molecular data suggest that the new species can be distinguished from E. pygmaeus by a minimum K2P distance of 5.7 % in the mtDNA COI barcode region. Economidichthys mornosensis is endemic to a very small area, and potentially Critically Endangered.
References
- Agorreta, A., San Mauro, D., Schliewen, U., Van Tassell, J.L., Kovačić, M., Zardoya, R. & Rüber, L. (2013) Molecular phylogenetics of Gobioidei and phylogenetic placement of European gobies. Molecular Phylogenetics and Evolution, 69 (3), 619–633. https://doi.org/10.1016/j.ympev.2013.07.017
- Barbieri, R., Zogaris, S., Kalogianni, E., Stoumboudi, M.T., Chatzinikolaou, Y., Giakoumi, S., Kapakos, Y., Kommatas, D., Koutsikos, N., Tachos, V., Vardakas, L. & Economou, A.N. (2015) Freshwater fishes and lampreys of Greece. An annotated checklist. Monographs on Marine Sciences 8. Hellenic Centre for Marine Research, Athens, 129 pp.
- Bianco, P.G., Bullock, A.M., Miller, P.J. & Roubal, F.R. (1987) A unique teleost dermal organ in a new European genus of fishes (Teleostei: Gobioidei). Journal of Fish Biology, 31 (6), 797–803. https://doi.org/10.1111/j.1095-8649.1987.tb05280.x
- Biomatters (2013) Geneious Pro. Available from: http://www.geneious.com (accessed 15 August 2025)
- Buj, I., Šanda, R., Zogaris, S., Freyhof, J., Geiger, M.F. & Vukić, J. (2019) Cryptic diversity in Telestes pleurobipunctatus (Actinopterygii; Leuciscidae) as a consequence of historical biogeography in the Ionian Freshwater Ecoregion (Greece, Albania). Hydrobiologia, 835, 147–163. https://doi.org/10.1007/s10750-019-3935-6
- Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9 (8), 772. https://doi.org/10.1038/nmeth.2109
- Economidis, P.S. (1992) Ο νανογωβιός και η παρέα του (The dwarf-goby and its friends): Part A. I Physi, 56 (Winter), 8–12.
- Economidis, P.S. (1995) Endangered freshwater fishes of Greece. Biological Conservation, 72 (2), 201–211. https://doi.org/10.1016/0006-3207(94)00083-3
- Economidis, P.S. & Miller, P.J. (1990) Systematics of freshwater gobies from Greece (Teleostei: Gobiidae). Journal of Zoology, 221, 125–170. https://doi.org/10.1111/j.1469-7998.1990.tb03781.x
- Economou, A.N., Giakoumi, S., Vardakas, L., Barbieri, R., Stoumboudi, M.T. & Zogaris, S. (2007) The freshwater ichthyofauna of Greece: an update based on a hydrographic basin survey. Mediterranean Marine Science, 8 (1), 91–166. https://doi.org/10.12681/mms.164
- Edgar, R.C. (2004) MUSCLE: Multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32, 1792–1797. https://doi.org/10.1093/nar/gkh340
- Freyhof, J., Kärst, H. & Geiger, M.F. (2014) Valencia robertae, a new killifish from southern Greece (Cyprinodontiformes: Valenciidae). Ichthyological Exploration of Freshwaters, 24 (4), 289–298.
- Geiger, M.F., Herder, F., Monaghan, M.T., Almada, V., Barbieri, R., Bariche, M., Berrebi, P., Bohlen, J., Casal-Lopez, M., Delmastro, G.B., Denys, G.P.J., Dettai, A., Doadrio, I., Kalogianni, E., Kärst, H., Kottelat, M., Kovacic, M., Laporte, M., Lorenzoni, M., Marcic, Z., Özuluð, M., Perdices, A., Perea, S., Persat, H., Porcelotti, S., Puzzi, C., Robalo, J., Šanda, R., Schneider, M., Šlechtová, V., Stoumboudi, M. & Freyhof, J. (2014) Spatial heterogeneity in the Mediterranean biodiversity hotspot affects barcoding accuracy of its freshwater fishes. Molecular Ecology Resources, 14, 1210–1221. https://doi.org/10.1111/1755-0998.12257
- Giakoumi, S., Zimmerman, B., Zogaris, S., Economou, A.N. & Kalogianni, E. (2006) Current population status of Valencia letourneuxi (Sauvage, 1880) in the Mornos River Delta. In: 8th Pan-Hellenic Congress on Oceanography and Fisheries, Thessaloniki, Greece, 4–8 June, 2006, Book of Abstracts, pp. 199.
- Gkenas, C., Malavasi, S. & Leonardos, I. (2012) Diet and feeding habits of Economidichthys pygmaeus (Perciformes: Gobiidae) in Lake Pamvotis, NW Greece. Journal of Applied Ichthyology, 28, 75–81. https://doi.org/10.1111/j.1439-0426.2011.01912.x
- Gawthorpe, R.L., Leeder, M.R., Kranis, H., Skourtsos, E., Andrews, J.E., Henstra, G.A., Mack, G.H., Muravchik, M., Turner, J.A. & Stamatakis, M. (2018) Tectono-sedimentary evolution of the Plio-Pleistocene Corinth rift, Greece. Basin Research, 30 (3), 448–479. https://doi.org/10.1111/bre.12260
- Holly, M. (1929) Zoologische Forschungsreise nach den Jonischen Inseln und dem Peloponnes von Max Beier. VI. T. Pisces. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, 138, 487–489.
- Huyse, T., Van Houdt, J. & Volckaert, F.A.M. (2004) Paleoclimatic history and vicariant speciation in the "sand goby" group (Gobiidae, Teleostei). Molecular Phylogenetics and Evolution, 32, 324–336. https://doi.org/10.1016/j.ympev.2003.11.007
- Ivanova, N.V., Zemlak, T.S., Hanner, R.H. & Hebert, P.D.N. (2007) Universal primer cocktails for fish DNA barcoding. Molecular Ecology Notes, 7, 544–548. https://doi.org/10.1111/j.1471-8286.2007.01748.x
- Kapakos, Y., Leris, I., Kalaitzakis, N., Fytilis, K., Kouraklis, P. & Kalogianni, E. (2025) First record of the western Greece goby Economidichthys pygmaeus (Gobiiformes: Gobiidae) in the Pinios River (Greece). Is it the first freshwater Greek endemic that will spread to eastern Europe? Acta Zoologica Bulgarica, 77 (1), 49–54. https://doi.org/10.71424/azb77.1.002810
- Karymbalis, E., Gaki-Papanastassiou, K. & Maroukian, H. (2007) Recent geomorphic evolution of the fan delta of the Mornos River, Greece: natural processes and human impacts. Bulletin of the Geological Society of Greece, 40 (4), 1538–1551. https://doi.org/10.12681/bgsg.17057
- Kimura, M. (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution, 16 (2), 111–120. https://doi.org/10.1007/BF01731581
- Kottelat, M. & Freyhof, J. (2007) Handbook of European Freshwater Fishes. Kottelat, Cornol and Freyhof, Berlin, xiv + 646 pp.
- Kumar, S., Stecher, G., Suleski, M., Sanderford, M., Sharma, S. & Tamura, K. (2024) MEGA12: Molecular Evolutionary Genetics Analysis version 12 for adaptive and green computing. Molecular Biology and Evolution, 41, 1–9. [publication PDF: https://www.megasoftware.net/citations] https://doi.org/10.1093/molbev/msae263
- Lykousis, V., Sakellariou, D., Moretti, I. & Kaberi, H. (2007) Late Quaternary basin evolution of the Gulf of Corinth: Sequence stratigraphy, sedimentation, fault slip and subsidence rates. Tectonophysics, 440 (1–4), 29–51. https://doi.org/10.1016/j.tecto.2006.11.007
- Masters, B.C., Fan, V. & Ross, H.A. (2011) Species delimitation – a geneious plugin for the exploration of species boundaries. Molecular Ecology Resources, 11, 154–157. https://doi.org/10.1111/j.1755-0998.2010.02896.x
- Mazzini, I., Cronin, T.M., McNeill, L.C., Shillington, D.J., Gawthorpe, R.L., Ll. Collier, R.E., de Gelder, G., Golub, A.R., Toomey, M.R., Poirier, R.K., Huang, H.-M.M. & Phillips, M.P. (2023) A new deglacial climate and sea-level record from 20 to 8 ka from IODP Site M0080, Alkyonides Gulf, eastern Mediterranean. Quaternary Science Reviews, 313, 108192. https://doi.org/10.1016/j.quascirev.2023.108192
- McKay, S.I. & Miller, P.J. (1997) The affinities of European sand gobies. Journal of Natural History, 31 (10), 1457–1482. https://doi.org/10.1080/00222939700770791
- McNeill, L.C., Shillington, D.J., Carter, G.D.O., Everest, J.D., Gawthorpe, R.L., Miller, C., Phillips, M.P., Collier, R.E.L., Cvetkoska, A., De Gelder, G., Diz, P., Doan, M.-L., Ford, M., Geraga, M., Gillespie, J., Hemelsdaël, R., Herrero-Bervera, E., Ismaiel, M., Janikian, L., Kouli, K., Le Ber, E., Li, S., Maffione, M., Mahoney, C., Machlus, M.L., Michas, G., Nixon, C.W., Oflaz, S.A., Omale, A.P., Panagiotopoulos, K., Pechlivanidou, S., Sauer, S., Seguin, J., Sergiou, S., Zakharova, N.V. & Green, S. (2019) High-resolution record reveals climate-driven environmental and sedimentary changes in an active rift. Scientific Reports, 9, 3116. https://doi.org/10.1038/s41598-019-40022-w
- Miller, P.J. (1990) The endurance of endemism – the Mediterranean freshwater gobies and their prospects for survival. Journal of Fish Biology, 37, 145–156. https://doi.org/10.1111/j.1095-8649.1990.tb05030.x
- Miller, P.J. (2004) The freshwater fishes of Europe, Vol 8/II Gobiidae. AULA-Verlag, Wiebelsheim, 478 pp.
- Perissoratis, C., Piper, D.J.W. & Lykousis, V. (2000) Alternating marine and lacustrine sedimentation during late Quaternary in the Gulf of Corinth rift basin, central Greece. Marine Geology, 167 (3–4), 391–411. https://doi.org/10.1016/S0025-3227(00)00038-4
- Rodríguez, N.V., Šanda, R., Zogaris, S. & Vukić, J. (2020) Distribution and genetic diversity of two species of Pelasgus minnows (Leuciscidae) in southern Greece. Knowledge and Management of Aquatic Ecosystems, 421, 27. https://doi.org/10.1051/kmae/2020019
- Rodríguez, N.V., Šanda, R., Zogaris, S. & Vukić, J. (2021) Evolutionary history of the Pelasgus minnows (Teleostei: Leuciscidae), an ancient endemic genus from the Balkan Peninsula. Molecular Phylogenetics and Evolution, 164, 107274. https://doi.org/10.1016/j.ympev.2021.107274
- Saitou, N. & Nei, M. (1987) The Neighbor-Joining Method—a new method for reconstructing phylogenetic trees. Molecular Biology and Evolution, 4, 406–425. https://doi.org/10.1093/oxfordjournals.molbev.a040454
- Shumka, S., Shumka, L., Trajce, K. & Ceci, S. (2020) First record of the Western Greece goby—Economidichthys pygmaeus (Holly, 1929), in Greater Prespa Lake (Albania). Ecologica Montenegrina, 35, 78–81. https://doi.org/10.37828/em.2020.35.6
- Stephanidis, A. (1939) [The freshwater fishes of northern Greece and the island of Corfu]. University of Athens, Athens, 163 pp. [in Greek]
- Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. [program] https://doi.org/10.1111/j.0014-3820.2002.tb00191.x
- Tamura, K. & Nei, M. (1993) Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees. Molecular Biology and Evolution, 10 (3), 512–26. https://doi.org/10.1093/oxfordjournals.molbev.a040023
- Thacker, C.E., Gkenas, C., Triantafyllidis, A., Malavasi, S. & Leonardos, I. (2018) Phylogeny, systematics and biogeography of the European sand gobies (Gobiiformes: Gobionellidae). Zoological Journal of the Linnean Society, 185 (1–14), 212–225. https://doi.org/10.1093/zoolinnean/zly026
- Tsaparis, D., Konstantinidis, I., Palandacic, A., Kalogianni, E., Stoumboudi, M.T., Barbieri, R., Vardakas, L., Koutsikos, N. & Tsigenopoulos, C.S. (2021) DNA barcoding provides new insights on the distribution, systematics and conservation of the freshwater genus Pelasgus (Cypriniformes: Cyprinidae) in Greece. Hydrobiologia, 848 (5), 1163–1176. https://doi.org/10.1007/s10750-021-04526-9
- van Oosterhout, F., Maliaka, V., van Bochove, K. & Lürling, M. (2022) First report of Economidichthys pygmaeus (Holly, 1929) (Gobiidae) in Lake Lesser Prespa, Greece. Check List, 18 (6), 1249–1253. https://doi.org/10.15560/18.6.1249
- Vanhove, M.P., Economou, A.N., Zogaris, S., Larmuseau, M.H., Giakoumi, S., Kalogianni, E., Volckaert, F.A.M. & Huyse, T. (2012) Phylogenetics and biogeography of the Balkan ‘sand gobies’ (Teleostei: Gobiidae): vulnerable species in need of taxonomic revision. Biological Journal of the Linnean Society, 105, 73–91. https://doi.org/10.1111/j.1095-8312.2011.01781.x
- Vavalidis, T., Zogaris, S., Kallimanis, A.S. & Bobori, D.C. (2025) Fish assemblage taxonomy highlights spatial patterns of β-diversity within a freshwater ecoregion. Aquatic Sciences, 87 (4), 95. https://doi.org/10.1007/s00027-025-01217-8
- Vogiatzi, E., Kalogianni, E., Zimmerman, B., Giakoumi, S., Barbieri, R., Paschou, P., Magoulas, A., Tsaparis, D., Poulakakis, N. & Tsigenopoulos, C.S. (2014) Reduced genetic variation and strong genetic population structure in the freshwater killifish Valencia letourneuxi (Valenciidae) based on nuclear and mitochondrial markers. Biological Journal of the Linnean Society, 111, 334–349. https://doi.org/10.1111/bij.12206
- Zogaris, D., Gkafas, G.A., Sarantopoulou, J., Koutsikos, N., Vardakas, L. & Exadactylos, A. (2024) Quietly invading the Balkans: Economidichthys pygmaeus (Holly, 1929) (Gobiidae) established in Northeastern Greece. BioInvasions Records, 13 (1), 215–224. https://doi.org/10.3391/bir.2024.13.1.19
