Economidichthys mornosensis / Mornos Goby / Μορνογωβιός (photo: S. Zogaris)
A new species of Goby!
Inland ichthyofaunas in Greece are more than just species lists; they are living indicators of biogeographic history. Fishes have guided us in the study of describing evolutionary regional units: freshwater ecoregions. Their distribution and evolutionary relationships help us reconstruct geological events that shaped aquatic biota at various scales in time and space.
Yet despite decades of research, some small fish species remain taxonomically unresolved. Among them are the tiny freshwater gobies: tiny morphologically subtle creatures with disproportionate biogeographic significance. They are good biogeographic indicators because they ''stay put'' - they don't migrate or disperse easily through space. One of them turned out to be a new species - we discribed it, published just a few days ago, we are calling it Economidichthys mornosensis - the Mornos Goby...endemic to the Mornos Delta in western Greece!
The Suspicion: Something Didn’t Fit
In the 1990s, Professor P.S. Economidis (1934-2022) noted that a very small goby lived in the Mornos Delta - on the coast of the huge and deep Gulf of Corinth. It resembled a species of the genus Knipowitschia, but he expressed doubts about this. Years later, genetic analyses we conducted with our Belgian partner Maarten P.M. Vanhove confirmed that the Mornos goby population was clearly distinct from known species. It does not belong in the genus Knipowitschia, but with the gobies of the genus Economidichthys — a group of small endemic fish of naturally restricted to western Greece, named in honor of Economidis. For many years, our fish team at HCMR, often collaborating with experts from abroad, has worked on these tiny goby fishes. My mentor Alcibiades Economou always thought something was worth the detective work at the Mornos Delta.
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| Fig. 2. The map in Professor Economidis's article in the popular periodical "Ι Physis" in 1992. It shows all the major freshwater gobies in Greece with approximate distributions known at the time. It provides an erroneous record of Economidichthys pygmaeus at the Mornos Delta (yellow highlight) and the question mark of a proposed new species which he called Knipowitschia stephanidisi - question mark (site 9? in this map). Conclusion: many decades later: Νο E. pygmaeus or Knipowitschia (or any other small goby) exist in freshwaters of the Mornos Delta - but a new, very tiny Economidichthys goby (obviously, the one he called K. stephanidisi), now named Economidichthys mornosensis. |
The Genetic Evidence
My friend, the naturalist and fish taxonomy celebrity, Jörg Freyhof, came in to help close the case of the Mornos fishes. We needed a complete review and survey of all populations of the related fish, the Western Greece Goby - Economidichthys pygmaeus. This included a search for the original population in the type locality - where it was first found (on Lefkada Island), specimens originally described there and published back in 1929. We collected Economidichthys widely and also confirmed a population extirpation (local extinction) at the type locality (on Lefkada, unfortunately). We now focused on resolving the problem at the Mornos once and for all.
First the genetics: This was already evident in our publication with Maarten P.M.Vanhove back in 2007. Using DNA barcoding (mitochondrial COI gene), our new chapter in this research confirmed a 5.7% genetic divergence from the Mornos population to its closest relative, Economidichthys pygmaeus. In fish taxonomy, a divergence of more than 2% in COI typically indicates the potential for a separate species rather than a local population variant (however, opinion on such % thresholds varies). The COI genetic divergence from Economidichthys pygmaeus is remarkably high despite the more widespread E. pygmaeus being found just 40 kilometers to the west of Mornos (in the river basin, next-door, the Acheloos - Evinos system). This divergance is well above typical intraspecific variation (~0.04% among the E. pygmaeus populations!). In other words, the Mornos goby represents an independent evolutionary lineage. Although related to E. pygmaeus, it evolved in a different world, within a state of a high degree of isolation.
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| Fig. 3. Genetic results shown through phylogenetic analyses (ML, NJ, MP trees); A dendrogram here summarizes the relationships among sampled populations: the Mornos population forms a distinct clade (highlighted yellow), apart from the widespread E. pygmaeus populations; an even more distinctive one is Economidichthys trichonis an endemic of the Lakes Trichonis and Lysimachia. From: the paper, Freyhof et al. (2026) |
What Makes It Different?
Of course, to most people, these tiny gobies really look alike. But, beyond the genetics, morphology supports the Mornos species status. Males show a prominent black blotch on the first dorsal fin (unlide the E. pygmaeus!). The perianal organ is smaller and structurally different from that of E. pygmaeus. The posterior back lacks scales at the base of the second dorsal fin a final distinguishing character.
The mornos fish is also extremely small — not exceeding 4 cm in total length (TL). Even when reared in large tanks under captive conditions, individuals remained tiny, suggesting that small body size is an evolutionary trait, not just plasticity due to environmental constraints. The species is very close to the size of the smallest adult fish in Europe, the Trichonis Dwarf Goby - E. trichonis (which has an adult size of about 3 cm TL at its maximum). The closer relative, the Western Greece Goby - E. pygmaeus- is by contrast a 'giant'; we have caught several adult specimens at 6 cm TL max. (as published in our fish checklist - Barbieri et al. 2015).
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| Fig. 4. Snapshots from our field aquarium photography-all from the September 2025 - comparison of the three species of the genus Economidichthys (not shown exactly to scale): The Mornos Goby Economidichthys mornosensis - A,B,C (three adult males shown). The Western Greece Goby Economidichthys pygmaeus - D. The Trichonis Dwarf Goby Economidichthys trichonis - E. (Photos by S. Zogaris). |
So, based on all the above data, we decided to name and publish the new species as:
Economidichthys mornosensis
The suggested common names are: Mornos Goby and Μορνογωβιός (Mornogovios).
Standardized ''common names'' for animals are important in order to maintain consistancy and public interest. I see no reason not to call it ''Mornos Goby'' (as you will read below, we consider it extremely range-restricted). In Greek, I propose a “genus-based compound name” following the naming style used by Antonis Ch. Kanellis and P. S. Economidis. As a side note, Professor A. Ch. Kanellis (1908–1992) was instrumental in compiling the first Greek lists of bird and mammal names in the '60s and '70s. He also had German colleagues who assisted him and he adopted the genus-based compound naming method (although, unfortunately, this was sometimes applied inconsistently and with various errors- long story...). But note, “genus-based compound name” i.e., where the generic identity is embedded in the common name is common practice in zoological naming (e.g., “sand goby,” “marsh harrier,” “stonechat”). I think this is practical and easy to remember. Thus, I proposed Μορνογωβιός - a compound name, instead of Γωβιός του Μόρνου (Goby of the Mornos).
A species with a tiny geographical range - part of an important biogeographical unit!
The Mornos goby is a stenoendemic (i.e. a very range restricted, narrow endemic species). Based on our studies and our inventory so far, it survives only in small spring-fed wetlands within the Mornos Delta, nowhere else. Its entire distribution covers less than 5 km² and we have found it in only three water bodies in that system (actually just two in the survey of spring and fall 2025). And, there may be a biogeographical reason for this extreme range-restricted state. We did not find it anywhere else in the Gulf of Corinth basin. This is where the biogeography comes in - read on please.
The geological history of the Mornos River is critical to understand. The river has and always had drained into the Gulf of Corinth, which once was a lake, just before the start of the Holocene. This likely explains the species isolation and the context for its evolution. In contrast, major river basins flowing westward into the Ionian Sea host the related Economydichthys pygmaeus - a much more widespread species because population exchanges (as did river confluences) existed among these systems - all facing sea-ward, westward.
How many people know the Gulf of Cornith was not connected to the Ionian sea just 13 K years ago?
So the Gulf of Corinth freshwaters may make up a biogeographical unit of extreme conservation interest - but only its freshwater wetlands, not the marine gulf it is today....Today, most freshwater coastal wetlands around the Gulf of Corinth have been degraded or totally lost. To the east of the Gulf is a strong rain-shadow area - see Galaxidi and Corinth: dry lands and intermittent/ephemeral streams, almost seasonally semi-arid (no long-term freshwater wetlands). And humans have been activley stealing waters and wastefully destryong wetlands for a long time here. So, the spring-fed flowing waters on level ground (or near the coast), are extremely rare. Most are found solely in the Mornos Delta. The map below shows the gulf and the major river basins flowing within it; the Mornos is the largest, with the largest wetland remnants at its Delta.
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Fig. 5. Copy-paste of a map of the Gulf of Corinth, with the rivers entering the Gulf - and the Mornos - the largest (blue spot highlighted; where the Delta is today). The Mornos therefore did not drain into the open Ionian Sea like the other westward flowing rivers of Western Greece (i.e. Acheloos, Pineios), it entered a former ancient lake. The Rion Sill is only 60 m. deep; anytime global sea levels were below this depth, the Gulf was transformed back into a lake. This has been going on for at least 5 million years. Map from: McNeill, et al. 2019. High-resolution record reveals climate-driven environmental and sedimentary changes in an active rift. Scientific Reports, 9(1), p.3116.
Specifically what and when was the Gulf of Corinth a Lake?
Here I include references (as cited in Freyhof et al. 2026):
Unlike the other large rivers along the Ionian coast, the Mornos River flows into the semi-enclosed Gulf of Corinth. This gulf experienced very long periods of isolation and freshwater (lake) conditions starting at least in the Late Miocene (~5 Ma). Yes, at least 5 million years ago! Even more extensive lake conditions occurred during the Pliocene to early Pleistocene (~2–1 Ma) (Gawthorpe et al. 2018). Around ~1 Ma, increased subsidence and/or a lowering of the western basin threshold (the Rion Sill) allowed seawater from the Ionian Sea to enter the basin. As a result, the basin changed from a lake into a marine gulf (Lykousis et al. 2004; McNeill et al. 2019); freshwater life probably found refuge in the surrouning riparian wetlands - river mouths, springs. Later, during the Pleistocene glacial periods, global sea levels dropped repeatedly (down to ~120 m below present levels). When sea levels were low, the shallow Rion Sill became exposed, cutting off the marine connection with the Ionian Sea. During these times, the Gulf of Corinth once again turned into a very large lake basin (Perissoratis et al. 2000). After the Last Glacial Maximum, sea levels rose again and overtopped the Rion sill in the late Pleistocene (~13.5 ka before present), restoring marine conditions in the Gulf of Corinth (Mazzini et al. 2023). Because the Gulf of Corinth has had this unusual history of alternating between lake and marine conditions, and because it has long been connected to the Mornos River, freshwater environments in the area likely remained isolated for long periods (i.e. no real or ''easy'' connection to the adjacent Acheloos). This long-term isolation probably promoted vicariant separation of freshwater organisms. Such geographic isolation of freshwater ecosystems from the wider Ionian coastal region may have encouraged the local differentiation of fish species (especially poor dispersers, such as tiny gobies); maybe other aquatic species as well. 
| | Fig. 6. This is a freshwater mussel in the Unionidae family that we found in the Mornos Delta, it is a river mussel! It survived in a spring-fed former distributary channel with the Mornos Gobies! Usually you have mussels only in perrrenial river waters, usually bigger rivers. Our Portuguese colleagues are helping us explore the biogeography of these species - this one is a new location discovery, we never new that they existed in the Mornos Delta. So, due to the biogeographic idiosynchrasies of the river Delta we are now researching other aquatic species, there may be new stories here! (Photo: S. Zogaris). |
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Under Pressure
Somehow, the wetland wonders that were a refuge to this fish and other freshwater biota survived the ups and downs of a large inland lake, then a sea (back-and-forth). But, they could not easily have survived human-induced wetland ecosystem degradation (this has been going on for a long time in this 'heart of civilization' area....see Ancient Corinth etc).
Although the Mornos Delta is part of the Natura 2000 network, the remaining habitat faces serious pressures:
- Water abstraction and diversion from springs
- Wetland drainage and fragmentation
- Infrastructure development and scattered construction
- Increasing drought frequency
We saw and photographed a lot of degradation in 2025: Illegal dumping, trash heaps and garbage tipping, agro-chemical pollution, embankment and drainage channel digging and clearing, wetland in-filling, scattered housing being built, and over-abstraction (or so it seems) from a key hotspot wetland site at the fish species stronghold (during a drought period - see photos in Figs. 7 and 10 below).
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| Fig. 7. This is a pump-house immediately above the spring-fed wetland hotspot which is a critical refuge for the Mornos Goby (and other species). Please see Fig. 10 below for conditions at the wetland immediately below this pumphouse in Sept. 2025. For years we know of the pump-house, however, we know nothing of its impact on the water levels or spring flow. This area is within the heart of the Natura 2000 protected area - and area zones as highly important. No action on the ground. (Photo: J. Freyhof). |
At the Mornos Delta, the fish and freshwater biota populations are small and fragmented. Our study suggests the Mornos Goby likely qualifies as a Critically Endangered species. The delta is also home to the globally threatened killifish Valencia robertae, and prorably other rare freshwater animals. We found a species of mussel (Unionidae)... If research continues, it could be that this area is exceptionally important for aquatic biodiversity in general (Anyway, it is important, even with the two endangered species of fish).
A Small Fish, A Larger Message
The story of the discovery of the Mornos Goby reminds us that Greece’s freshwater biodiversity may still provide amazing surprises. Small and inconspicuous aquatic species are often the most vulnerable and there is no real concern for them. Scientists and naturalists must speak for them. These biota depend on fragile habitats that are easily altered or destroyed. Legal so-called protection alone is not enough. Without scientific work, active management and habitat restoration, conservation remains only “on paper.” Protecting this tiny fish means protecting an entire wetland ecosystem at the landscape scale — and the natural heritage it represents. This means an action plan and careful conservation politics - on the ground. We need to save the landscapes of the Mornos Delta and specifically help protect the spring-fed wetlands within these landscapes. Can we do it? Below I provide some important snap-shots of local conditions. First of all, please keep in mind, this area is a Protected Area....
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| Fig. 8. Within the terrestrial protected areas of the Gulf of Corinth (marine areas not shown) we have the Delta (top left) inside the SCI called PARALIAKI ZONI APO NAFPAKTO EOS ITEA - PERIOCHI PIGON CHILIADOU (GR2450004). In fact, back in 2017, we at HCMR (and with the support of a consortium surveying fish for the Natura project) expanded the protected area to include a critial spring area, the 'Periochi Pigon Chiliadou'. Our ''success'' in this really means ''nothing'' (I was responsible for delineating the new extenstion of the Natura on behalf of the fish - that, nearly 10 years ago). If you cannot promote measures on the ground you are doing practically nothing (see photos below). |
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| Fig. 9. Landscape scenes from the Mornos Delta within the Natura 2000 protected area (GR2450004). A typical agricultural-domainted, second-home building area and beach tourism hot-spot. Rapid land-use change are taking place. These photos are from Google Maps; I am not making a statement here or propagandizing, there is no conservation action for freswaters on the ground - none at all. |
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| Fig. 10. One of the most important habitats for the Mornos Goby, a spring area which I intentionally leave un-named since I a fear it may be vandalized or disturbed since, really, there is no real protection: Left: the hotspot area in Sept. 2014 and Right in Sept. 2025. (Photos: S. Zogaris). |
Last thoughts
Looking back, I have been to the Mornos many times in the last 20 years - Alcibiades Economou introduced me to the area. I have been there with nearly a dozen people and I can name them all and cherish memories with them. Our investigations, include great experiences with friends from Belgium, Croatia, and the Czech republic. I was fortunate to visit in 2014 with P.S. Economidis who told us the story of how the late Professor Alexandros Stephanidis had first collected the tiny mystery goby at the Mornos Delta. I have been there with nearly all my co-workers from HCMR and with wonderful naturalists, such as Nikos Petrou from the Hellenic Society for the Protection of Nature and our brilliant collaborator, Brian Zimmerman from Zoological Society London. Later on we helped expand the Natura 2000 boundaries there. All this reminds me of what we do - rarely talked about - Natural History and conservation science. For this you need inventory and taxonomy.
I would like to express my gratitude to everyone who has worked to collect aquatic animals and study their taxonomy in this country. Taxonomic research requires collaboration and decades of careful fieldwork, in-depth study, data-management, collection management and long-term dedication. This type of work rarely receives substantial funding or recognition. It is sustained mainly by the curiosity, commitment, and perseverance of naturalists who are driven by a genuine desire to understand biodiversity. Such combined efforts provide the foundation for conservation-relevant evolutionary research, biodiversity promotion in our cultures, conservation policy development, and conservation actions. Mornos Delta is microcosm of this. May we continue to collaborate, support one another, and advance this work with dedication and scientific integrity.
The paper:
FREYHOF, J., GEIGER, M. F., & ZOGARIS, S. (2026). Economidichthys mornosensis, a new freshwater sand-goby from Greece (Teleostei: Oxudercidae). Zootaxa, 5759(1), 26-40.
Our paper with the full details is available from the Authors through ResearchGate:
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