The Mornos Goby - A new species restricted to the Mornos Delta, Greece

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. 

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.

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). 

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. 

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).

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). 

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.... 

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).  

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. 

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:

https://www.researchgate.net/publication/400759296_Economidichthys_mornosensis_a_new_freshwater_sand-goby_from_Greece_Teleostei_Oxudercidae

Our man in Lake Baringo: Joseph Aengwo

 Focus on Ecoguides: Joseph Aengwo

February 2026 - Kenya (our third Fam tour visit)

I’ve always wanted to talk more about the eco-guides — about the people who quietly dedicate their lives to it.

We often celebrate destinations. We photograph landscapes. Tick species off our lists. But we rarely pause to acknowledge the individuals whose deep knowledge, lived experience and interpretation abilities make wildlife tourism and nature promotion possible.

This is one of those stories.

On a recent FAM trip to Lake Baringo in Kenya, I had the privilege of meeting someone who embodies what true "eco-guiding-with-a-local" is all about: Joseph Aengwo, one of the famous bird guides who began at Baringo but has developed a company, now guiding nearly everywhere in East Africa.

Joseph is a prominent and highly experienced ornithological guide from Lake Baringo. And when I say experienced, I don’t just mean he knows bird names. I mean he understands birds really deeply and their environment too— how the Rift Valley breathes, how the seasons shift, how bird calls change with the light.

Born and raised on the shores of Lake Baringo — a true birding haven in the heart of the Rift Valley — this landscape is not just his workplace. It’s his boyhood wonderland, memory, culture, heritage.

This was one of our top birding targets at Baringo, the Northern Carmine Bee-Eater. Tick at the Baringo Lake causeway to the Tumbili Cliffs! 

Professionally, Joseph is formally trained in tourism and wildlife management from Moi University. He is also a bronze badge member of the Kenya Professional Safari Guides Association — a credential that reflects both skill and commitment to guiding standards in Kenya. But what truly sets him apart isn’t the certification. It’s the way he guides.

There’s a quiet confidence in how he organizes guiding. A patience in how he adjusts to the clients/guests. A generosity in how he shares stories — not just about species, but about conservation, community, and the life-forces that keeps Lake Baringo thriving. 

Joe also has been active in conservation and he knows the situation of the lake ecosystems very well. Also, he has a great sense of humour...our man in Baringo!  

He told us there are now about 20 boys (and a one or two girls) involved locally in nature conservation actions, and some of them are active in in ecoguiding as well;  most of them beginners. We met two of them who had staked-out owls and other cryptic birds for Joe. Joe is still associated with the Lake Baringo Biodiversity Conservation Group (LBBCG) formed in 2008, a non-profit local environmental group that plays a role striving to protect this place. The group aims to promote community awareness on environmental conservation and protection of key species sites and habitats, also encouraging community participation. Its all very low-key, very informal. We saw very few tourists. 

Now all these pics may bring to mind a wonderful beautifully managed and manocured National Park. But this is not to be in Baringo!  The situation here is not protected, its not a fortress park or any such park. And there are problems. Things are rapidly changing and there are also issues of serious poverty. The lake waters have risen substantially in the last few years, displacing people, flooding wildlife habitat and ruining agricultural areas, drowning villages. Between before 2009 and late 2020, Lake Baringo’s water level rose by approximately 9.5 meters (from about 972 m to around 981.5 m above sea level) — a dramatic rise over roughly a decade.

This situation is unusual, but it is also seen in Lake's Bogoria and Naivasha, also to a lesser extent at Lake Nakuru. 

The main hypothesized causes based on current understanding point to:

🌧️ Chagnes in Rainfall patterns (more intense rain) and catchment changes (vegetation change; increased run-off). These are currently the most strongly supported explanations for the main driver of  the recent water level rise.

🌍 Geological forces - unknown. The lake’s tectonic context matters for its hydrology, but there is no definitive evidence that plate movement is the main cause of the contemporary lake level rise.

🧠 Some discussions about underground fault-seepage changes exist, but they remain hypotheses rather than confirmed causes.

Also, despite this serious and unresolved problem of the lake waters, the area has a strong wildlife - human population conflict too. Large mammals are scarce. Poaching is rampant and displaced peoples have settled in difficult conditons. Overgrazing is also a serious problem. 

So, in this case, organized ecotourism may support and promote solutions to these big problems, big landscape scale measures are needed for conservation. Of course, since we work in science of this sort, we also would promote science-led initiatives to deal with such serious issues. This place is too important to neglect!

Rhinos and Giraffes and other mega-mammals once roamed below these hills onthe shores of the lake, now there is nothing like that. Poaching is a problem. This beautiful and huge landscape is not protected and there are serious problems. 

Joe with one of the best Birders in Bulgaria, my good friend Tihomir Stefanov. This is the causeway to the Tumbili Cliffs- amazing place for birding and fish-watching.

The endemic Baringo Tilapia - Oreochromis (niloticus) baringoensis. 

This amazing semi-desert, arid land beauty. And full of birds.

Verreaux's eagle (Aquila verreauxii), used be known simply as Black Eagle when I was a kid. Rare bird indeed.

One of the most captivating places for birds are the famous Baringo cliffs. 

Looking accros the lake to the islands; a short rain-shower blessing the landscapes. 

Rising waters of Lake Baringo, looking north, from the causeway near Tumbili Cliff Lodge. 

The lake as viewed from Tumbili Cliff.

The location of Lake Baringo in the East African Rift Valley. A critical problem is the water level rise. Maps from: Herrnegger M et al. Paleohydrology repeating? Regional hydrological change may lead to an overflow and cross-mixing of an alkaline and a freshwater lake in East Africa. Journal of Hydrology: Regional Studies. 2024 Oct 1;55:101951.

Joseph Aengwo

Have a look at Joe's website at: https://kenya-birding.com/

Cell Phone: +254 727 856 048
email: joe.aengwo@gmail.com

Also, we thank our good friend DERRICK MURIUKI GICHIMU who rents out the best cars for any Safari. Derrick is one of our best friends in Nairobi and he knows Kenya. Find Derrick at: muriukidgichimu@gmail.com or at: Kompact safaris Kenya car hire | budget friendly Kenya car hire/ self drive 4x4 Nairobi

Electrofishing the large rivers of Western Greece - the Ionian Freshwater Ecoregion

I’m posting a handful of snapshots boat electrofishing in the rivers that spill into the Ionian Sea. Western Greece is often described with postcard words—gorges, plane-tree galleries, turquoise springs—but for a freshwater biologist the real poetry is in the fish.

These river and lake basins sit in an ecoregion of unusually high endemism. In plain terms: many of the species we net here exist nowhere else on Earth, restricted to a single basin or to a tight cluster of neighbouring basins. Each river is its own little evolutionary archive, shaped by histroical isolation, ancient geographical barriers, limestone karst and local ecological dynamics. You can feel that history in the bucket: a familiar “Mediterranean” fish assemblage, yes—but also odd local forms, subtle variations, and those small, secretive fishes that only make sense once you’ve seen the ecosystems that produced them.

Boat electrofishing looks dramatic (and, inevitably, people ask if it’s “fishing”....). It isn’t sport; it’s scientific sampling—carefully standardised, short pulses, tight safety protocols, capture technique and quick handling so fish can recover and swim away. The routine is half choreography: the skiff sliding along a run, the anode poles probing the margins, dip-nets ready, data sheets getting damp, GPS logging each transect. Meanwhile, we’re also reading the river: flow, substrate, cover, riparian conditiosn, the scars of water abstraction, the shadow of barriers, the die-back of the platanus trees due to an alien fungus invasion; the occasional surprise of an alien species where it shouldn’t be.

And still—despite multiple pressures and degradation—these Ionian rivers can feel astonishingly alive and wild. Every survey reminds me that conservation here isn’t an abstract idea. It’s local, basin by basin, decision by decision. Protect the habitat, and you protect an entire, irreplaceable lineage.

 

Fish Pictured here include (in order of appearance): Economidichthys pygmaeus - Goby, Knipowitschia sp. - Goby; Squalius peloponensis - Chub; Luciobarbus graecus (with D. Kommatas) - Barbel; Dicentrarchus labrax - Sea Bass; Luciobarbus albanicus - Barbel; Knipowitschia sp. - Goby; Liza (Chelon) ramada (in net) - Mullet; Salaria fluviatilis (juv.) - Freshwater Blenny; Alosa fallax - Twaite Shad. 

My colleagues: Alexandros Ntakis, Dimitris Kommatas, Vassilis Pappas, Anthi Economou and Petros Chronopoulos. Some pics taken by Vassiliki Vlami who was also on one of the trips. And again my thanks to HCMR, we are so grateful to work there...

You are also welcome to download one of our recent papers on the fish assemblages within this Ecoregion: 

Fish_assemblage_taxonomy_highlights_spatial_patterns_of_b-diversity_within_a_freshwater_ecoregion

And a map by my good colleague Theocharis Vavalidis from the above paper. The major river basins where HCMR does electrofishing using a boat are numbered as so: 1) Kalamas, 2) Acheron, 3) Louros, 4) Arachthos, 5) Acheloos, 6) Evinos, 7) Mornos, 8) Pineios, 9) Alpheios, 10) Pamissos, 11) Evrotas.

Greece as a birding destination

Greece as a birding destination

Birding is big in nature-based tourism. However, it is not an easy 'industry' to develop. Greece is an interesting country to discuss such development. 

The interest in international wildlife watching tourism in Greece is probably old but probably rather slow growing. See multiple references to: ''wildflowers and archeological sites'', Spring birding on Lesvos (Lesbos), botanical excursion to Crete etc  (ever since the late 1970s...) and this "hidden Greece" approaches away from the beaches have been well promoted. Ever since people like Marc Dubin (Rough Guides) helped inland areas famous (at least since the early 1980s). "Wildlife" stands as a nice complement to cultural tours, i.e., many of the culutural monuments are in marvelous open landscapes, sometime close to wild lands too. Also, I should mention Oliver Rackham and his important contribution through the book 'The Making of the Creten Landscape' (1996). In the '80s and '90s, when there were magazines and tiny guide books sold at kiosks, people like George Sfikas provided an important impetus for promoting natural history tourism here. And there were many writers at the time (now they are bloggers, or vloggers...). Ok.

There were different stages of this promotion. And some of it really worked but was it sustainable? 

Ok, since the early 1980s there has been an active interest in some kind of quasi-ecotourism development in this country but nothing never really clicked, despite the glossy photos, nature protection areas (on paper), and good new accomodation facilities, etc. 

Lets focus on birds. Not a lot of the country was well known for birding before 2000 (and before Google Earth; it was not easy to explore). Since, 1997, when 'Birds of Greece' was published by Handrinos and Akriotis (this was a land-mark for birding developement as where some early "site-guides" (Crete, Lesbos, Northern Greece)).  

Times have changes a generation later (2015-2025 period). We know where the birds are and this has really been helped by Ornithologiki, Greece's birdlife partner, with many field trips and information and leBird, of course. One would expect Greece to be good/thriving/big for birding tourism. Simple criteria point to this and  support this claim. There are some amazing hotspot concentrations - with specialty birds, big wetlands, migration traps and an amazing combo with beach and cultural tourism. Also, Greece is so easy to drive around in, and you can combine driving in Greece with visiting four bordering Balkan countries next door. Also Greece is a beautiful. 

Is Greece a top birding country in Europe and the Med: no (i.e., compared to Spain, Isreal, Turkey Cyprus, France).

I generally feel a lot more could have been done here. And there are many lost opportunities and degraded sites (i.e, trashed infrastructure, wasted efforts by government, and poor conservation interest).  Basically Greece is a top and global nature tourism opportunity which we Greeks have trashed and neglected, for 4 decades straight! I know there is a lot of greenwashing paperwork saying otherwise. 

Yes, I think we have failed. And I say this from experience. I first led tours with my brother and then my wife in the late 1980s and 1990s; then I did a lot of education tourism in this century. So I have been doing this a lot (despite my other interests and varied activities).  I still lead educational tours. I know Greece. Why am is so negative?

Well, maybe I should not be so negative (leave that for another time)...here I will concentrate on the successes, the good places. Where a perfect combination of location, government support and private initiatives really made a difference. Where local people really benefited and birds were effectively used a as an incentive and tool for ecotourism development.  

Some areas, about 30 or so hotspots did become 'world famous' for birders in Greece. And the specific attractions have recived a steady development (ecodevelopment, ecotourism). This is looking at the glass half-full. Lets do this. 

Where are the top birding hotspots where we have good ecotourism experiance in Greece?

It depends on your choices, your criteria: lets use these: Richness (bird spp.), Distinctness (ambiguous I know...) and accessibility as categoreis. 

I would wager there are 13 top areas, and I know I am missing out a lot but lets see these together here...

I divided GR in North and South based on a line across the country. This is a bit abstract but worthy of consideration since 8 of the sites are north of the line (only 5 south of the line). Note that all areas have wetlands and no non-wetland dominated area is included except say Crete which has some minor wetland sites as well. The north is exceptional in having many more 'lesser' areas close to and attached to the ones selected here - and some of these are uniquely rich near the wetlands. So in the North you may have the wetlands and the temperate mountain landscapes (this includes the spectacular Central Rhodope mountains near Nestos or the Kaimaktsalan Mountains with the 'lake district' of Kastoria-Ptolemaida for example). The south is interesting in having some specialty birds not easily found in the north (Bonelli's Eagle, Ruepell's Warbler, huge colonies of Eleanora's Falcon, etc). The scale of cover in the south and the distinctiveness is hard to express. There are many smaller areas that are not in the 13 top sites I picked here: some areas that probably should be included are: Naxos Island (wetlands and mountains with Griffin Vulture colonies), Kos (wetlands), Samos (Eastern specialties, small wetlands).

This is a quick description of the top sites I picked in this rapid review:

1. Evros Delta National Park (Eastern Macedonia and Thrace)

Why: Legendary. One of Europe’s great deltas (≈320 spp. most easily seen each year), guided tours/boat rides via the Visitor Centre (Loutros). Extensive seasonal checklists and reports going back to late 1960s. Best seasons: peak spring and autumn migrations and definatly winter (geese, raptors, swans, pelicans, rarities all the time). Facilities & access:easy; track network (4×4 helpful in wet months), boat trips (more so in the past); good access from Alexandroupolis and close to other hotspots along the border (and on the other side of the border as well).

2. Dadia–Lefkimi–Soufli Forest NP (Eastern Macedonia and Thrace)

Why: Raptor stronghold with Europe’s vulture trio; raptor observatory & good information centre; long-term scientific monitoring shows since early 1990s strong - outstanding conservation impact. Famous place. Good network of trials. Best seasons: All year round. Late spring–summer for migrants, breeders and soaring birds; autumn migration wonderful with storks and eagle passage and lots of passerines; forest birds. Winter: Wintering raptors and passerines. Facilities & access: Dadia visitor centre & hide network; one private hide for photography (Chris Vlachos). Area regenerating after a massive forest fire a couple of years back. Still a beautiful area, within a culturally interesting part of Greece; easy access to Turkey and Bulgaria increase the ecotourism potential.

3. Nestos Delta, Porto Lagos & Lakes Vistonida–Ismarida (Eastern Macedonia and Thrace)

Why: One of Greece’s largest national parks by extent although much of it under cultivation. Lots to see: mosaic of lagoons, reedbeds, ponds, lakes, & coastal steppe, saltmarshes, and the largest lowland riparian forest in the country (much of it regenerating after protection and restoration progremmes). Multiple vantage points and tracks (but area not well organized; poorly promoted).  Best seasons: spring for herons/warblers/raptors; winter for geese and ducks. Facilities & access: spread-out network of drivable embankments; visitor nodes at Porto Lagos area but still poorly developed. Nearby uplands also of interest, especially Kompsatos Gorge and Nestos Gorge. Site very close to spectacular mountaion areas of the Central and Eastern Rhodope massif. But generally, this area is underplayed, poorly organized (a transit area moslty). 

4. Lake Kerkini National Park (Central Macedonia)

Why: Without a doubt Greece's best all-round international birding destination! Year-round abundance and high diversity (≈300+ spp.). Look-outs, embankment roads & a staffed info-centre, easy boat trips for pelicans/waterfowl, shorebirds; easy road access from Thessaloniki. Conservation & monitoring are long-standing (Ramsar/Natura 2000) and work of local scientists and good management team on the ground.  Best seasons: All year: winter (pelicans, waterfowl), spring (pelicans in breeding plumage, herons, breeding forest and countryside birds), late spring for breeding waterbirds, forest birds in nearby mountains; Autumn is very good as well (low water, shorebirds, migrants, flamigos, etc). But I think late spring is best (May-late June) and combined with forest birding on Mount Belles and Mount Krousia - it is fantastic. Easy access to Bulgaria and North Macedonia as well. Also, the site is close to other lesser hotspots (Lake Doirani, Central Rhodope). If you are from the Americas and want to go to one site, go to Kerkini.

5. Axios–Loudias–Aliakmonas Delta (near Thessaloniki)(Central Macedonia)

Why: This place is totally underated. Vast coastal complex with easy urban access, active management unit & information centre, routes for independent birders but no over-all development for birding. Its important to note that the area is next door to Thessaloniki, Greece's 'second capital city'. In fact parts of it are within the city limits: This includes the Dendropotamos Estuary (Ekvoli Dedropotamou) and the Loudias Estuary at Kalochori. These places are full of birds at all seasons.  Best seasons: all year; spring/autumn for shorebirds, herons, storks and migrants; winter for a varierty of raptors, flamingos & wildfowl. Facilities & access: Info Centre; signed routes; near-urban walking paths (e.g. Kalochori Lagoon). Note that from this place you can look across the bay to see the towering Mount Olympus - often snow-capped. Its less than 60 minutes away and offers good forest birding potential (among other attractions). 

6. Prespa Lakes National Park (Western Macedonia)

Why: Globally important Dalmatian Pelican colony; strong NGO-led conservation (SPP), visitor info hubs, trails & viewpoints; excellent documentation and monitoring and good community spirit supporting conservation (both at the national and municipality level). Both upland wild land and lake-side cultural landscapes. Note this is a really unique place globally, but a lot of it looks like upland areas in some other Balkan countries or even in parts of Asia. Best seasons: Mar–Oct for pelicans/waterfowl; winter for waterfowl. Migration season and especially spring for both migrants and breeders. Not so good as a winter destination (continental cold!). Facilities & access: Agios Germanos Info Center; multiple park info points- well developed and with a proud local nature-positive vibe. And still authentic.

7. Amvrakikos Gulf National Park (Epirus)

Why: One of the Mediterranean’s most productive semi-enclosed gulfs and the most extensive complex of lagoons, vast saltmarshes and mudflats in Greece (comprable in size to Messolonghi). Hosts Dalmatian Pelican colonies, Greater Flamingo flocks, varied herons, raptors and many other attractions (dolphins) but all is low key and poorly organized. Ramsar/Natura 2000 protection with NGO/park support but poorly managed, poor coordination on the ground; ramshackle conditions. However, it is a great place to explore (you need a guide - it is huge!).  Best seasons: Year-round interest; peak spring/autumn passage; winter for wildfowl, hugh flocks. Facilities & access: Visitor centres at Salaora and Strongyli no longer functioning; observation tower at Louros (overgrown road access). The area is easily reached from Preveza airport or Arta but the best place to stay is at Koronissia. The problem with this area is that it is plainly too vast. Access is difficult in some areas (roads overgrown with vegetation), and poorly signed or promoted. However, any Greek ornithologists will tell you this area is within Greece's top five birding-nature attractions. 

8. Messolonghi–Aitoliko Lagoons (Western Greece)

Why: This area is the southern half of Greece's most important wetland complex and probably the most birdy area in the south in general. It is huge. It is also a culturally and aesthetically amazing place. A Ramsar wetland with excellent, diverse birding (salinas, lagoons, mudflats; rock hills, canyon, woods, coasts and more). Active management unit, small visitor information centre, and some ecotourism promotion (low-key). Amazing and easy place to see good wildlife and have large day-lists (with spectacular species, incl. vultures, flamingos etc. all together). Combines well with nearby areas (big lakes, mountain forests, rivers, etc - Acheloos-Evinos uplands). Best seasons: spring & autumn for shorebirds/terns; winter for wildfowl and raptors. Facilities & access: Info Centre at Aitoliko; guided tours upon request or from Greece's top birding guides (based in Athens). Messolonghi is the closest big wetlnad experiance near Athens. 

9. Strofylia National Park (Peloponnese)

Why: Greece’s largest coastal umbrella pine forest adjoining lagoons, dunes and wetlands plus rock hills and beatiful oak-sage savannas. A wonderful landscape with good birds (~270 bird species recorded including flamingos, spoonbills, forest birds, raptors). Active management but big poaching probs; popular for low-key nature tourism. Best seasons: Autumn/winter for waterbirds; spring for migrants and breeding herons/terns. Too hot in summer and not that birdy then (nearby Messolonghi is perhaps better in Summer). Facilities & access: Network of tracks and watchpoints; visitor information at Kotychi–Strofylia Management Body; within 1.5 hours drive from Patras; also close to Athens.

10. Gialova Lagoon / Pylos (Peloponnese)

Why: Key stopover at the south end of the Balkans (~250+ spp.), good trail access near major tourism infrastructure (Pylos/Costa Navarino) and beautiful landscapes.  Best seasons: Apr–May & Sep–Oct migration; winter holds some ducks, flamingos, etc. Facilities & access: graded tracks, local accommodation/food options in Gialova/Pylos. Note that in the Peloponese we have only two wetland areas as important hotspots; this is unfortunate because a lot of other areas are poorly developed (not well organized/poor access/ poor conservation intiatives or plainly too small - of localized or regional interest). There is an important migration in the southern part of the peninsula at other points besides Gialova/Pylos - and perhaps the best and best studied place is the 'pelagic' island of Antikythira, due south of the southernmost three pronged peninsula. But I think Antikythira, and its world-famous bird observatory, are mostly for an ornithological trip, not general birding (that's why its not included here). 

11. Crete (island-wide; key wetlands, mountains & gorges)

Why: Greece’s southernmost major island is a migration crossroads, with raptors funnelling through its mountain ridges and numbers of passerines and waterbirds using wetlands as stopovers. Resident raptors include amazing populations of vultures and several rarities (both breeding and wintering). Key sites include Lake Agia near Chania (with a hide and info boards), artificial reservoirs, and gorges such as Samaria and the Asterousia mountains for raptors. Several species show distinctive Cretan forms (but there are no endemics), adding extra interest for birders. Best seasons: Spring (Apr–May) for mass migration and breeding raptors; autumn passage is also rewarding. Facilities & access: International airports at Heraklion and Chania provide easy access. Lake Agia has basic birding infrastructure; other sites are reachable by car with nearby accommodation options.

12. Lesvos (island-wide, esp. Kalloni Gulf wetlands and areas around this central location)

Why: First Lesvos is in Asia, not Europe. It is one of the big islands along the coast of Anatolia. It has famous spring migration (falls of passerines, shorebirds, bee-eaters, shrikes) and some local Anatolian specialties (cinereous bunting, Kruper's nuthatch and others); superb documentation (decades of trip reports) and an Environmental Information Centre at Kalloni and Natural History museum (Sigri). Best seasons: late Apr–May (spring peak); also autumn passage. Facilities & access: Kalloni Centre programs; dense network of drivable tracks and easy accommodation near hotspots. At least two good local guides regularly lead tours on the island. Hands-down, Lesbos is Greece's best destination for visitors from Western and Northern Europe (and may be good also for North Americans who have seen Central European habitats). Lesvos does not really compete with the wetlands of Northeastern Greece or the mountains of that area; both areas are big and special for birding and natural history. 

13. Schinias–Marathon National Park (Attica)

Why: Greece's capital’s most important birding wetland (>210 spp. recorded), ease of accessibility (45–60 min from central Athens); some trails and observation towers; strong seasonality but great for quick visits or wetland targets in season (i.e., needs to be a bit 'wet'; not a good place in prolonged droughts...). Best seasons: peak passage is spring, also for breeders but during drought years things can be underwelming. Winter is interesting if there are flood waters and colder conditions (cold snaps up north); autumn brings rarities. Facilities & access:some "marked paths' (Schinias Reservoir); easy public/private transport from Athens; however its best to have a vehicle since the area is rather big and there are no circuit trails; not easy to walk it on a hot day. Schinias-Marathon is close to some other areas in Attika, notably Mount Parnitha (spring forest birds, winter); Artemis Lagoon, Oropos Lagoon and even the wonderful Tritsis Park within Athens metro area (see map below showing the top 15 birding wetland sites near Athens). 

15 important wetland sites for birding in Attika (circled in red). Since Athens attracts a great number of international tourists, it would be in the economy's great interest to promote ecotourism in these sites. At this point in time only one of them has some kind of infrastructure (observation towers, etc), and that is Schinias-Marathon National Park. Some of them are threatened and extremely degraded. 

A typical itinerary for spring birding in Greece; as marketed by one international tour company (FieldGuides.com). My wife and I led this kind of tour back since the early 1990s for Canadian visitors and it was really a fantastic experience for all. The trip starts with a flight from Athens to Alexandroupolis and then you are led down to Athens.

Greeks promoting birding within the National Parks of Northern Greece at the UK Bird Fair in 2025. The NECCA our authority for conservation areas is doing its part in promotion of biodiversity and awareness. Below are some campaign posters. 

Schinias-Marathon National Park in Attika (very near Athens) makes a wonderful airphoto at certain angles and there has been some effort made to suppress wildfires and save the coastal pine wood at the beach. The fantastic aspect here is the amazing wetland behind the wood.

One of the most unGreek landscapes in Greece is the remarkable wetland complex of the Lakes Vistonis, Ismarida and Thracian lagoons, also part of a National Park. This incredible area is one of the top 5 places for birding but it is poorly developed relative to its potential. 

Greece has no endemic bird species but several are range-restricted specialties and rarities which are not seen easily in other parts of Europe. Rueppel's Warbler is one such bird distributed in the south and a few of th larger Aegean Islands.