Main Article Content
Abstract
In this study the sagitta morphology of seven species of Gobiidae including three species of the genera Neogobius and four Ponticola, inhabiting the southern part of the Caspian Sea was described and analysed. The goal was to investigate the usefulness and efficiency of sagitta for species identification. The morphological analysis was based on four shape indices, roundness (4S/πOL2), rectangularity (S/OL×OW), ellipticity (OL-OW/OL+OW), and aspect ratio (OL/OW). The results indicated that otolith morphology and morphometry can be used as diagnostic characters distinguishing the Caspian Sea gobies. While the overall shape of otoliths differentiated the genera, the differences in the shape of the dorsal rim and development of posterior dorsal and anterior ventral lobes were particularly important for inter-specific distinction. The discriminant analysis could separate the specimens of genus Ponticola and Neogobius with 61% and 82% classification successes, respectively, proves that otolith shape indices were a more efficient tool for discrimination of otoliths in Neogobius than Ponticola species. Cluster analysis based on otoliths shape indices showed two main clusters, one cluster contains Neogobius species with higher roundness values, possessed more rounded shaped otoliths and the other includes Ponticola species with spindle and elongated shaped otoliths, except P. bathybius, recognized P. bathybius and Neogobius melanostomus as more similar taxa. These results were consistent with the phylogeny of these species proposed by recent molecular studies and indicated that otolith characteristics are partly genetically encoded and therefore have phylogenetic signals. The current research revealed a high degree of morphological variability of otolith shape in different species and confirmed its usefulness in Gobiidae taxonomy and phylogeny.
Keywords
Taxonomy
Otolith
Neogobius
Ponticola.
Article Details
License
The authors transfer the copyrights of their papers to the Iranian Society of Ichthyology. However, the information could be used in accordance with the Creative Commons licence (Attribution-NonCommercial 4.0 International (CC BY-NC 4.0)
How to Cite
DAVOODI, P. (2021). Characteristics of sagittae morphology in seven goby species from the Caspian Sea. Iranian Journal of Ichthyology, 8(3), 170–180. https://doi.org/10.22034/iji.v8i3.538
References
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Bani, A.; Poursaeid, S. & Tuset, V.M. 2013. Comparative morphology of the sagittal otolith in three species of south Caspian gobies. Journal of Fish Biology 82: 1321-1332.
Bell, J.D. & Harmelin-Vivien, M.L. 1983. Fish fauna of French Mediterranean Posidonia oceanica seagrass meadows. 2. Feeding habits. Tethys 11: 1-14.
Berg, L.S. 1949. Freshwater Fishes of the U.S.S.R. and Adjacent Countries. Izdatel'vesto Akademii Nauk SSSR, Moscow.
Campana, S.E. 2004. Photographic atlas of fish otoliths of the Northwest Atlantic Ocean. Canadian Special Publication of Fisheries and Aquatic, Canada.
Corkum, L.D.; Sapota, M.R. & Skora, K.E. 2004. The round goby, Neogobius melanostomus, a fish invader on both sides of the Atlantic Ocean. Biological Invasions 6: 173-181.
Davoodi, P. & Rahimian, H. 2016. Identifying Neogobius species from the southern Caspian Sea by otolith shape (Teleostei: Gobiidae). Zootaxa 4205: 081-086.
Eagderi, S.; Nikmehr, N. & Poorbagher, H. 2020. Ponticola patimari sp. nov. (Gobiiformes: Gobiidae) from the southern Caspian Sea basin, Iran. FishTaxa 17: 22-31.
Esmaeili H.R., Mehraban H., Abbasi K., Keivany Y., Coad B.W. 2017. Review and updated checklist of freshwater fishes of Iran: taxonomy, distribution and conservation status. Iranian Journal of Ichthyology 4(Suppl. 1): 1-114.
Esmaeili, H.R., Sayyadzadeh, G., Eagderi, S., & Abbasi, K. 2018. Checklist of freshwater fishes of Iran. FishTaxa, 3(3), 1-95.
Ghanbarifardi, M.; Gut, C.; Gholami, Z.; Esmaeili, H.R.; Gierl, C. & Reichenbacher, B. 2020. Possible link between the structure of otoliths and amphibious mode of life of three mudskipper species (Teleostei: Gobioidei) from the Persian Gulf. Zoology in the Middle East 66(4): 311-320.
Gierl, C.; Liebl, D.; Šanda, R.; Vukić, J.; Esmaeili, H.R. & Reichenbacher, B. 2018. What can goby otolith morphology tell us? Cybium 42(4): 349-363.
Gut, C.; Vukić, J.; Šanda, R.; Moritz, T. & Reichenbacher, B. 2020. Identification of past and present gobies: distinguishing Gobius and Pomatoschistus (Teleostei: Gobioidei) species using characters of otoliths, meristics and body morphometry. Contributions to Zoology 89: 282-323.
Horvatić, S.; Cavraro, F.; Zanella, D. & Malavasi, S. 2015. Sound production in the Ponto-Caspian goby Neogobius fluviatilis and acoustic affinities within the Gobius lineage: implications for phylogeny. Biological Journal of the Linnaean Society 117:564-573.
Horvatić, S.; Lea, B.; Malavasi, S.; & Marčić, Z.; Buj, I.; Mustafić, P.; Ćaleta, M. & Zanella, D. 2019. Comparative analysis of sound production between the bighead goby Ponticola kessleri and the round goby Neogobius melanostomus: Implications for phylogeny and systematics. Environmental Biology of Fishes 102: 1-13.
Kovačić, M. & La Mesa, M. 2008. Feeding ecology of de Buen’s goby in the Adriatic Sea. Vie et Milieu 58: 249-256.
Kovačić, M.; Esmaeili, H.R.; Zarei, F.; Abbasi, K. & Schliewen, U.K. 2021. A new species of tadpole-goby, Benthophilus persicus sp. nov. (Teleostei: Gobiidae) from the southern Caspian Sea. Zootaxa 4980(1): 45-63.
Kovačić, M.; Ordines, F. & Schliewen, U.K. 2017. A new species of Buenia (Teleostei: Gobiidae) from the western Mediterranean Sea, with the description of this genus. Zootaxa 4250: 447-460.
Lombarte, A. & Castellon, A. 1991. Interespecific and intraspecific otolith variability in the genus Merluccius as determined by image analysis. Canadian Journal of Zoology 69: 2442-2449.
Lombarte, A.; Chic, O.; Parisi-Baradad, V.; Olivella, R.; Piera, J. & Garcia–Ladona, E. 2006. A web-based environment from shape analysis of fish otoliths. The AFORO database. Scientia Marina 70: 147-152.
Lombarte, A.; Miletić, M.; Kovačić, M.; Otero‐Ferrer, J.L. & Tuset, V.M. 2018. Identifying sagittal otoliths of Mediterranean Sea gobies: variability among phylogenetic lineages. Journal of Fish Biology 92: 1768-1787.
Lord, C.; Morat, F.; Lecomte-Finiger, R. & Keith, P. 2012. Otolith shape analysis for three Sicyo¬pterus (Teleostei: Gobioidei: Sicydiinae) species from New Caledonia and Vanuatu. Environmental Biology of Fishes 93: 209-222.
Medvedev, D.A.; Sorokin, P.A.; Vasil’ev V.P.; Chernova, V.P. & Vasil’eva, E.D. 2013. Reconstruction of phylogenetic relations of Ponto-Caspian gobies (Gobiidae, Perciformes) based on mitochondrial genome variation and some problems of their taxonomy. Journal of Ichthyology 53: 702-712.
Neilson, M.E. & Stepien, C.A. 2009. Escape from the Ponto–Caspian: Evolution and biogeography of an endemic goby species flock (Benthophilinae: Gobiidae: Teleostei). Molecular Phylogenetics and Evolution 52: 84-102.
Nolf, D. 1985. Handbook of Paleoichthyology, Otolithi piscium. Verlag Dr. Friedrich Pfeil, München.
Patzner, R.A.; Van Tassell, J.L.; Kovačić, M. & Kapoor, B.G. 2011. The Biology of Gobies. First Edition. CRC press, USA.
Platt, C. & Popper, A.N. 1981. Fine Structure and Function of the Ear. In: Tavolga, W.N.; Popper, A.N. & Fay, R.R. (eds), Hearing and Sound Communication in Fishes, Springer, New York. pp: 3-38.
Ponton, D. 2006. Is geometric morphometrics efficient for comparing otolith shape of different fish species? Journal of Morphology 267: 750-757.
Popper, A.N.; Ramcharitar, J. & Campana, S.E. 2005. Why otoliths? Insights from inner ear physiology and fisheries biology. Marine and Freshwater Research 56: 497-504.
Purrafee Dizaj, L.; Esmaeili, H.R.; & Teimori, A. 2020. Comparative otolith morphology of clupeids from the Iranian brackish and marine resources (Teleostei: Clupeiformes). Acta Zoologica. DOI: 10.1111/azo.12353
Reichenbacher, B. & Reichard, M. 2014. Otoliths of five extant species of the annual killifish Nothobranchius from the East African savan¬nah. PLoS ONE 9: e112459.
Rohlf, F.J. 2006. TpsDig2, digitize landmarks and outlines, version 2.10. Department of Ecology and Evolution, State University of New York, Stony Brook.
Sadeghi, R.; Esmaeili, H.R.; Zarei, F. & Reichenbacher, B. 2020. Population structure of the ornate goby, Istigobius ornatus (Teleostei: Gobiidae), in the Persian Gulf and Oman Sea as determined by otolith shape variation using ShapeR. Environmental Biology of Fishes, 103(10): 1217-1230.
Sanjarani Vahed, N.; Esmaeili, H.R.; Masoudi, M. & Reichenbacher, B. 2019. Ontogenetic otolith development in an endemic tooth-carp, Aphanius vladykovi (Teleostei: Aphaniidae). Journal of Ichthyology 59(3): 336-343.
Schulz-Mirbach, T.; Friedrich, L.; Plath, M. & Heß, M. 2018. Enigmatic ear stones: what we know about the functional role and evolution of fish otoliths: The role of fish otoliths in inner ear function. Biological Reviews 94: 1-26.
Schwarzhans, W. 2014. Otoliths from the middle Miocene (Serravallian) of the Karaman basin, Turkey. Cainozoic Research 14: 35-69.
Teimori, A.; Khajooei, A.; Motamedi, M. & Askari Hesni, M. 2019. Characteristics of sagittae morphology in sixteen marine fish species collected from the Persian Gulf: Demonstration of the phylogenetic influence on otolith shape. Regional Studies in Marine Science 29: 1-12.
Thacker, C. & Roje, D. 2011. Phylogeny of Gobiidae and identification of gobiid lineages. Systematics and Biodiversity 9: 1-19.
Tuset, V.M.; Lombarte, A. & Assis, C. 2008. Otolith atlas for the western Mediterranean, north and central eastern Atlantic. Scientia Marina 72: 1-198.
Tuset, V.M.; Lozano, I.J.; Gonzalez, J.A.; Pertusa, J.F. & Garcia-Diaz, M.M. 2003. Shape indices to identify regional differences in otolith morphology of comber, Serranus cabrilla. Journal of Applied Ichthyology 19: 88-93.
Vasil’eva, E.D.; Schwarzhans, W.W.; Medvedev, D.A. & Vasil’ev, V.P. 2016. Cryptic Species of Ponto-Caspian Bighead Goby of the Genus Ponticola (Gobiidae). Journal of Ichthyology 56: 3-21.
Xin, Y.; Liang, C.; Jinhu, L.; Bo, Z.; Xiujuan, S. & Shuozeng, D. 2014. Application of otolith shape analysis for stock discrimination and species identification of five goby species (Perciformes: Gobiidae) in the northern Chinese coastal waters. Chinese Journal of Oceanology and Limnology 32: 1060-1073.
Yu, X.; Cao, L.; Liu, J.; Zhao, B.; Shan, X. & Dou, S. 2014. Application of otolith shape analysis for stock discrimination and species identification of five goby species (Perciformes: Gobiidae) in the northern Chinese coastal waters. Chinese Journal of Oceanology and Limnology 32: 1060-1073.
Zander, C.D. 2011. Gobies as predator and prey. In: Patzner, R.A.; Van Tassell, J.L.; Kovačić, M. & Kapoor, B.G. (eds), The Biology of Gobies, CRC press, USA. pp. 291–344.
Zarei, F.; Esmaeili, H.R.; Schliewen, U.K.; Abbasi, K. & Sayyadzadeh, G. 2021. Mitochondrial phylogeny, diversity, and ichthyogeography of gobies (Teleostei: Gobiidae) from the oldest and deepest Caspian sub-basin and tracing source and spread pattern of an introduced Rhinogobius species at the tricontinental crossroad. Hydrobiologia 848(6): 1267-1293.
References
Bani, A.; Poursaeid, S. & Tuset, V.M. 2013. Comparative morphology of the sagittal otolith in three species of south Caspian gobies. Journal of Fish Biology 82: 1321-1332.
Bell, J.D. & Harmelin-Vivien, M.L. 1983. Fish fauna of French Mediterranean Posidonia oceanica seagrass meadows. 2. Feeding habits. Tethys 11: 1-14.
Berg, L.S. 1949. Freshwater Fishes of the U.S.S.R. and Adjacent Countries. Izdatel'vesto Akademii Nauk SSSR, Moscow.
Campana, S.E. 2004. Photographic atlas of fish otoliths of the Northwest Atlantic Ocean. Canadian Special Publication of Fisheries and Aquatic, Canada.
Corkum, L.D.; Sapota, M.R. & Skora, K.E. 2004. The round goby, Neogobius melanostomus, a fish invader on both sides of the Atlantic Ocean. Biological Invasions 6: 173-181.
Davoodi, P. & Rahimian, H. 2016. Identifying Neogobius species from the southern Caspian Sea by otolith shape (Teleostei: Gobiidae). Zootaxa 4205: 081-086.
Eagderi, S.; Nikmehr, N. & Poorbagher, H. 2020. Ponticola patimari sp. nov. (Gobiiformes: Gobiidae) from the southern Caspian Sea basin, Iran. FishTaxa 17: 22-31.
Esmaeili H.R., Mehraban H., Abbasi K., Keivany Y., Coad B.W. 2017. Review and updated checklist of freshwater fishes of Iran: taxonomy, distribution and conservation status. Iranian Journal of Ichthyology 4(Suppl. 1): 1-114.
Esmaeili, H.R., Sayyadzadeh, G., Eagderi, S., & Abbasi, K. 2018. Checklist of freshwater fishes of Iran. FishTaxa, 3(3), 1-95.
Ghanbarifardi, M.; Gut, C.; Gholami, Z.; Esmaeili, H.R.; Gierl, C. & Reichenbacher, B. 2020. Possible link between the structure of otoliths and amphibious mode of life of three mudskipper species (Teleostei: Gobioidei) from the Persian Gulf. Zoology in the Middle East 66(4): 311-320.
Gierl, C.; Liebl, D.; Šanda, R.; Vukić, J.; Esmaeili, H.R. & Reichenbacher, B. 2018. What can goby otolith morphology tell us? Cybium 42(4): 349-363.
Gut, C.; Vukić, J.; Šanda, R.; Moritz, T. & Reichenbacher, B. 2020. Identification of past and present gobies: distinguishing Gobius and Pomatoschistus (Teleostei: Gobioidei) species using characters of otoliths, meristics and body morphometry. Contributions to Zoology 89: 282-323.
Horvatić, S.; Cavraro, F.; Zanella, D. & Malavasi, S. 2015. Sound production in the Ponto-Caspian goby Neogobius fluviatilis and acoustic affinities within the Gobius lineage: implications for phylogeny. Biological Journal of the Linnaean Society 117:564-573.
Horvatić, S.; Lea, B.; Malavasi, S.; & Marčić, Z.; Buj, I.; Mustafić, P.; Ćaleta, M. & Zanella, D. 2019. Comparative analysis of sound production between the bighead goby Ponticola kessleri and the round goby Neogobius melanostomus: Implications for phylogeny and systematics. Environmental Biology of Fishes 102: 1-13.
Kovačić, M. & La Mesa, M. 2008. Feeding ecology of de Buen’s goby in the Adriatic Sea. Vie et Milieu 58: 249-256.
Kovačić, M.; Esmaeili, H.R.; Zarei, F.; Abbasi, K. & Schliewen, U.K. 2021. A new species of tadpole-goby, Benthophilus persicus sp. nov. (Teleostei: Gobiidae) from the southern Caspian Sea. Zootaxa 4980(1): 45-63.
Kovačić, M.; Ordines, F. & Schliewen, U.K. 2017. A new species of Buenia (Teleostei: Gobiidae) from the western Mediterranean Sea, with the description of this genus. Zootaxa 4250: 447-460.
Lombarte, A. & Castellon, A. 1991. Interespecific and intraspecific otolith variability in the genus Merluccius as determined by image analysis. Canadian Journal of Zoology 69: 2442-2449.
Lombarte, A.; Chic, O.; Parisi-Baradad, V.; Olivella, R.; Piera, J. & Garcia–Ladona, E. 2006. A web-based environment from shape analysis of fish otoliths. The AFORO database. Scientia Marina 70: 147-152.
Lombarte, A.; Miletić, M.; Kovačić, M.; Otero‐Ferrer, J.L. & Tuset, V.M. 2018. Identifying sagittal otoliths of Mediterranean Sea gobies: variability among phylogenetic lineages. Journal of Fish Biology 92: 1768-1787.
Lord, C.; Morat, F.; Lecomte-Finiger, R. & Keith, P. 2012. Otolith shape analysis for three Sicyo¬pterus (Teleostei: Gobioidei: Sicydiinae) species from New Caledonia and Vanuatu. Environmental Biology of Fishes 93: 209-222.
Medvedev, D.A.; Sorokin, P.A.; Vasil’ev V.P.; Chernova, V.P. & Vasil’eva, E.D. 2013. Reconstruction of phylogenetic relations of Ponto-Caspian gobies (Gobiidae, Perciformes) based on mitochondrial genome variation and some problems of their taxonomy. Journal of Ichthyology 53: 702-712.
Neilson, M.E. & Stepien, C.A. 2009. Escape from the Ponto–Caspian: Evolution and biogeography of an endemic goby species flock (Benthophilinae: Gobiidae: Teleostei). Molecular Phylogenetics and Evolution 52: 84-102.
Nolf, D. 1985. Handbook of Paleoichthyology, Otolithi piscium. Verlag Dr. Friedrich Pfeil, München.
Patzner, R.A.; Van Tassell, J.L.; Kovačić, M. & Kapoor, B.G. 2011. The Biology of Gobies. First Edition. CRC press, USA.
Platt, C. & Popper, A.N. 1981. Fine Structure and Function of the Ear. In: Tavolga, W.N.; Popper, A.N. & Fay, R.R. (eds), Hearing and Sound Communication in Fishes, Springer, New York. pp: 3-38.
Ponton, D. 2006. Is geometric morphometrics efficient for comparing otolith shape of different fish species? Journal of Morphology 267: 750-757.
Popper, A.N.; Ramcharitar, J. & Campana, S.E. 2005. Why otoliths? Insights from inner ear physiology and fisheries biology. Marine and Freshwater Research 56: 497-504.
Purrafee Dizaj, L.; Esmaeili, H.R.; & Teimori, A. 2020. Comparative otolith morphology of clupeids from the Iranian brackish and marine resources (Teleostei: Clupeiformes). Acta Zoologica. DOI: 10.1111/azo.12353
Reichenbacher, B. & Reichard, M. 2014. Otoliths of five extant species of the annual killifish Nothobranchius from the East African savan¬nah. PLoS ONE 9: e112459.
Rohlf, F.J. 2006. TpsDig2, digitize landmarks and outlines, version 2.10. Department of Ecology and Evolution, State University of New York, Stony Brook.
Sadeghi, R.; Esmaeili, H.R.; Zarei, F. & Reichenbacher, B. 2020. Population structure of the ornate goby, Istigobius ornatus (Teleostei: Gobiidae), in the Persian Gulf and Oman Sea as determined by otolith shape variation using ShapeR. Environmental Biology of Fishes, 103(10): 1217-1230.
Sanjarani Vahed, N.; Esmaeili, H.R.; Masoudi, M. & Reichenbacher, B. 2019. Ontogenetic otolith development in an endemic tooth-carp, Aphanius vladykovi (Teleostei: Aphaniidae). Journal of Ichthyology 59(3): 336-343.
Schulz-Mirbach, T.; Friedrich, L.; Plath, M. & Heß, M. 2018. Enigmatic ear stones: what we know about the functional role and evolution of fish otoliths: The role of fish otoliths in inner ear function. Biological Reviews 94: 1-26.
Schwarzhans, W. 2014. Otoliths from the middle Miocene (Serravallian) of the Karaman basin, Turkey. Cainozoic Research 14: 35-69.
Teimori, A.; Khajooei, A.; Motamedi, M. & Askari Hesni, M. 2019. Characteristics of sagittae morphology in sixteen marine fish species collected from the Persian Gulf: Demonstration of the phylogenetic influence on otolith shape. Regional Studies in Marine Science 29: 1-12.
Thacker, C. & Roje, D. 2011. Phylogeny of Gobiidae and identification of gobiid lineages. Systematics and Biodiversity 9: 1-19.
Tuset, V.M.; Lombarte, A. & Assis, C. 2008. Otolith atlas for the western Mediterranean, north and central eastern Atlantic. Scientia Marina 72: 1-198.
Tuset, V.M.; Lozano, I.J.; Gonzalez, J.A.; Pertusa, J.F. & Garcia-Diaz, M.M. 2003. Shape indices to identify regional differences in otolith morphology of comber, Serranus cabrilla. Journal of Applied Ichthyology 19: 88-93.
Vasil’eva, E.D.; Schwarzhans, W.W.; Medvedev, D.A. & Vasil’ev, V.P. 2016. Cryptic Species of Ponto-Caspian Bighead Goby of the Genus Ponticola (Gobiidae). Journal of Ichthyology 56: 3-21.
Xin, Y.; Liang, C.; Jinhu, L.; Bo, Z.; Xiujuan, S. & Shuozeng, D. 2014. Application of otolith shape analysis for stock discrimination and species identification of five goby species (Perciformes: Gobiidae) in the northern Chinese coastal waters. Chinese Journal of Oceanology and Limnology 32: 1060-1073.
Yu, X.; Cao, L.; Liu, J.; Zhao, B.; Shan, X. & Dou, S. 2014. Application of otolith shape analysis for stock discrimination and species identification of five goby species (Perciformes: Gobiidae) in the northern Chinese coastal waters. Chinese Journal of Oceanology and Limnology 32: 1060-1073.
Zander, C.D. 2011. Gobies as predator and prey. In: Patzner, R.A.; Van Tassell, J.L.; Kovačić, M. & Kapoor, B.G. (eds), The Biology of Gobies, CRC press, USA. pp. 291–344.
Zarei, F.; Esmaeili, H.R.; Schliewen, U.K.; Abbasi, K. & Sayyadzadeh, G. 2021. Mitochondrial phylogeny, diversity, and ichthyogeography of gobies (Teleostei: Gobiidae) from the oldest and deepest Caspian sub-basin and tracing source and spread pattern of an introduced Rhinogobius species at the tricontinental crossroad. Hydrobiologia 848(6): 1267-1293.
Bell, J.D. & Harmelin-Vivien, M.L. 1983. Fish fauna of French Mediterranean Posidonia oceanica seagrass meadows. 2. Feeding habits. Tethys 11: 1-14.
Berg, L.S. 1949. Freshwater Fishes of the U.S.S.R. and Adjacent Countries. Izdatel'vesto Akademii Nauk SSSR, Moscow.
Campana, S.E. 2004. Photographic atlas of fish otoliths of the Northwest Atlantic Ocean. Canadian Special Publication of Fisheries and Aquatic, Canada.
Corkum, L.D.; Sapota, M.R. & Skora, K.E. 2004. The round goby, Neogobius melanostomus, a fish invader on both sides of the Atlantic Ocean. Biological Invasions 6: 173-181.
Davoodi, P. & Rahimian, H. 2016. Identifying Neogobius species from the southern Caspian Sea by otolith shape (Teleostei: Gobiidae). Zootaxa 4205: 081-086.
Eagderi, S.; Nikmehr, N. & Poorbagher, H. 2020. Ponticola patimari sp. nov. (Gobiiformes: Gobiidae) from the southern Caspian Sea basin, Iran. FishTaxa 17: 22-31.
Esmaeili H.R., Mehraban H., Abbasi K., Keivany Y., Coad B.W. 2017. Review and updated checklist of freshwater fishes of Iran: taxonomy, distribution and conservation status. Iranian Journal of Ichthyology 4(Suppl. 1): 1-114.
Esmaeili, H.R., Sayyadzadeh, G., Eagderi, S., & Abbasi, K. 2018. Checklist of freshwater fishes of Iran. FishTaxa, 3(3), 1-95.
Ghanbarifardi, M.; Gut, C.; Gholami, Z.; Esmaeili, H.R.; Gierl, C. & Reichenbacher, B. 2020. Possible link between the structure of otoliths and amphibious mode of life of three mudskipper species (Teleostei: Gobioidei) from the Persian Gulf. Zoology in the Middle East 66(4): 311-320.
Gierl, C.; Liebl, D.; Šanda, R.; Vukić, J.; Esmaeili, H.R. & Reichenbacher, B. 2018. What can goby otolith morphology tell us? Cybium 42(4): 349-363.
Gut, C.; Vukić, J.; Šanda, R.; Moritz, T. & Reichenbacher, B. 2020. Identification of past and present gobies: distinguishing Gobius and Pomatoschistus (Teleostei: Gobioidei) species using characters of otoliths, meristics and body morphometry. Contributions to Zoology 89: 282-323.
Horvatić, S.; Cavraro, F.; Zanella, D. & Malavasi, S. 2015. Sound production in the Ponto-Caspian goby Neogobius fluviatilis and acoustic affinities within the Gobius lineage: implications for phylogeny. Biological Journal of the Linnaean Society 117:564-573.
Horvatić, S.; Lea, B.; Malavasi, S.; & Marčić, Z.; Buj, I.; Mustafić, P.; Ćaleta, M. & Zanella, D. 2019. Comparative analysis of sound production between the bighead goby Ponticola kessleri and the round goby Neogobius melanostomus: Implications for phylogeny and systematics. Environmental Biology of Fishes 102: 1-13.
Kovačić, M. & La Mesa, M. 2008. Feeding ecology of de Buen’s goby in the Adriatic Sea. Vie et Milieu 58: 249-256.
Kovačić, M.; Esmaeili, H.R.; Zarei, F.; Abbasi, K. & Schliewen, U.K. 2021. A new species of tadpole-goby, Benthophilus persicus sp. nov. (Teleostei: Gobiidae) from the southern Caspian Sea. Zootaxa 4980(1): 45-63.
Kovačić, M.; Ordines, F. & Schliewen, U.K. 2017. A new species of Buenia (Teleostei: Gobiidae) from the western Mediterranean Sea, with the description of this genus. Zootaxa 4250: 447-460.
Lombarte, A. & Castellon, A. 1991. Interespecific and intraspecific otolith variability in the genus Merluccius as determined by image analysis. Canadian Journal of Zoology 69: 2442-2449.
Lombarte, A.; Chic, O.; Parisi-Baradad, V.; Olivella, R.; Piera, J. & Garcia–Ladona, E. 2006. A web-based environment from shape analysis of fish otoliths. The AFORO database. Scientia Marina 70: 147-152.
Lombarte, A.; Miletić, M.; Kovačić, M.; Otero‐Ferrer, J.L. & Tuset, V.M. 2018. Identifying sagittal otoliths of Mediterranean Sea gobies: variability among phylogenetic lineages. Journal of Fish Biology 92: 1768-1787.
Lord, C.; Morat, F.; Lecomte-Finiger, R. & Keith, P. 2012. Otolith shape analysis for three Sicyo¬pterus (Teleostei: Gobioidei: Sicydiinae) species from New Caledonia and Vanuatu. Environmental Biology of Fishes 93: 209-222.
Medvedev, D.A.; Sorokin, P.A.; Vasil’ev V.P.; Chernova, V.P. & Vasil’eva, E.D. 2013. Reconstruction of phylogenetic relations of Ponto-Caspian gobies (Gobiidae, Perciformes) based on mitochondrial genome variation and some problems of their taxonomy. Journal of Ichthyology 53: 702-712.
Neilson, M.E. & Stepien, C.A. 2009. Escape from the Ponto–Caspian: Evolution and biogeography of an endemic goby species flock (Benthophilinae: Gobiidae: Teleostei). Molecular Phylogenetics and Evolution 52: 84-102.
Nolf, D. 1985. Handbook of Paleoichthyology, Otolithi piscium. Verlag Dr. Friedrich Pfeil, München.
Patzner, R.A.; Van Tassell, J.L.; Kovačić, M. & Kapoor, B.G. 2011. The Biology of Gobies. First Edition. CRC press, USA.
Platt, C. & Popper, A.N. 1981. Fine Structure and Function of the Ear. In: Tavolga, W.N.; Popper, A.N. & Fay, R.R. (eds), Hearing and Sound Communication in Fishes, Springer, New York. pp: 3-38.
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