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Identification of Mediterranean marine gobies (Actinopterygii: Gobiidae) of the continental shelf from photographs of in situ individuals

June 2022
Zootaxa 5144(1):1-103

DOI:10.11646/zootaxa.5144.1.1

Authors:

Marcelo Kovačić

Natural History Museum Rijeka

Roberto Pillon

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Numerous photographs of live fishes posted by anglers and divers on social media and citizen science databases are important sources of information for ichthyological research. However, validating records that extend the known ecology and bathymetric or geographic distribution of species should rely on a rigorous identification process. The family Gobiidae, with their small size, superficial resemblance among species and high species richness are particularly difficult to identify. Therefore, the identification from photographs of live individuals of Mediterranean marine gobies from the continental shelf was studied. A dichotomous identification key is provided based on photographs of live individuals, allowing positive identification of 41 out of the 66 species reviewed in this publication. Then, for all 66 species we provide a brief description of important characters, which can be used for provisional identification for those species that could not be positively identified using the key. Pending further progress in identification of live individuals, we suggest that records extending the known geographic and ecological species distribution be taken into account only if they could be validated using the dichotomous identification key.

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https://doi.org/10.11646/zootaxa.5144.1.1

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Accepted by B. Frable: 26 Apr. 2022; published: 1 Jun. 2022

5144

ZOOTAXA

Magnolia Press

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Zootaxa 5144 (1): 001–103

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Identification of Mediterranean marine gobies (Actinopterygii: Gobiidae)

of the continental shelf from photographs of in situ individuals

MARCELO KOVAČIĆ1*, JULIEN P. RENOULT2, ROBERTO PILLON3, RUDOLF SVENSEN4,

SERGEY V. BOGORODSKY5,6, SEMIH ENGIN7 & PATRICK LOUISY8,9

1Natural History Museum Rijeka, Lorenzov prolaz 1, HR-51000 Rijeka, Croatia

�

marcelo@prirodoslovni.com; https://orcid.org/0000-0002-4049-9366

2CEFE—Univ Montpellier, CNRS, EPHE, IRD, Montpellier, France.

�

jurenoult@gmail.com; https://orcid.org/0000-0001-6690-0085

3Ente Fauna Marina Mediterranea, Via M. Rapisardi trav. VIII, 2, 96012 Avola (SR), Italy

�

robertopillon@gmail.com; https://orcid.org/0000-0002-9360-1774

4Stavanger Museum, Department of Natural History, Musegata 16, 4010 Stavanger, Norway.

�

rudolf@uwphoto.no; https://orcid.org/0000-0002-0467-5393

5Senckenberg Research Institute and Natural History Museum Frankfurt, Senckenberganlage 25, 60325 Frankfurt a.M., Germany.

�

ic187196@yandex.ru; https://orcid.org/0000-0002-8679-9735

6Station of Naturalists, Omsk, Russia.

�

ic187196@yandex.ru

7Faculty of Fisheries, Izmir Katip Celebi University, HavaalanýSosesi Cd. Aosb 35620 Balatcýk/Cigli/Ýzmir, Turkey

�

engin.semih@gmail.com; https://orcid.org/0000-0002-2918-2552

8Association Peau-Bleue, 46 rue des Escais, 34300 Agde, France.

�

patrick.louisy@wanadoo.fr; https://orcid.org/0000-0001-6453-9636

9UMR BOREA (MNHN, CNRS, Sorbonne Universit, IRD, UniCaen, UGA), Station marine de Dinard, CRESCO, 38 rue du Port

Blanc, 35800 Dinard, France.

*Corresponding author

KOVAČIĆ ET AL.

MARCELO KOVAČIĆ, JULIEN P. RENOULT, ROBERTO PILLON, RUDOLF SVENSEN, SERGEY V.

BOGORODSKY, SEMIH ENGIN & PATRICK LOUISY

Identification of Mediterranean marine gobies (Actinopterygii: Gobiidae) of the continental shelf from

photographs of in situ individuals

(Zootaxa 5144)

103 pp.; 30 cm.

1 Jun. 2022

ISBN 978-1-77688-526-8 (paperback)

ISBN 978-1-77688-527-5 (Online edition)

FIRST PUBLISHED IN 2022 BY

Magnolia Press

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https://www.mapress.com/zt

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purpose other than private research use.

ISSN 1175-5326 (Print edition)

ISSN 1175-5334 (Online edition)

Table of Contents

Abstract ..................................................................................................4

Introduction ................................................................................................4

Material and methods ........................................................................................9

Results ..................................................................................................10

Family identification ........................................................................................10

Key for the identification of Mediterranean marine gobies from photographs of live individuals . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

Species accounts ..........................................................................................42

Aphia minuta (Risso, 1810) (Fig. 31)—Transparent Goby ..........................................................42

Aulopareia unicolor (Valenciennes, 1837) (Fig. 48)—Greenspot Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

Buenia affinis Iljin, 1930 (Fig. 41)—De Buen’s Goby .............................................................43

Buenia massutii Kovačić, Ordines & Schliewen, 2017 (Fig. 49)—Massuti’s Goby .......................................44

Chromogobius quadrivittatus (Steindachner, 1863) (Fig. 24)—Banded Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

Chromogobius zebratus (Kolombatović, 1891) (Fig. 25)—Kolombatovic’s Goby .......................................45

Corcyrogobius liechtensteini (Kolombatović, 1891) (Fig. 21)—Liechtenstein’s Goby ....................................46

Coryogalops ocheticus (Norman, 1927) (Fig. 50)—Canal Goby .....................................................46

Cryptocentrus caeruleopunctatus (Rüppell, 1830) (Fig. 3)—Red Sea Bluespotted Shrimpgoby . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

Cryptocentrus steinhardti Goren & Stern, 2021 (Fig. 46)—Steinhardt’s Shrimpgoby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Crystallogobius linearis (Düben, 1845) (Fig. 32)—Crystal Goby. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Deltentosteus collonianus (Risso, 1820) (Fig. 51)—Toothed Goby ...................................................49

Deltentosteus quadrimaculatus (Valenciennes, 1837) (Fig. 52)—Four-spotted Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

Didogobius schlieweni Miller, 1993 (Fig. 7)—Andromeda Goby .....................................................50

Didogobius splechtnai Ahnelt & Patzner, 1995 (Fig. 14)—Splechtna’s Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

Gammogobius steinitzi Bath, 1971 (Fig. 22)—Steinitz’s Goby. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

Gobius auratus Risso, 1810 (Figs. 36 and 40)—Golden Goby .......................................................53

Gobius bucchichi Steindachner, 1870 (Fig. 34)—Bucchich’s Goby ...................................................53

Gobius cobitis Pallas, 1814 (Fig. 53)—Giant Goby ................................................................54

Gobius couchi Miller & El-Tawil, 1974 (Fig. 43)—Couch’s Goby ....................................................55

Gobius cruentatus Gmelin, 1789 (Fig. 15)—Red-mouthed Goby. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

Gobius fallax Sarato, 1889 (Fig. 39)—Sarato’s Goby ..............................................................56

Gobius gasteveni Miller, 1974 (Fig. 54)—Steven’s Goby ...........................................................56

Gobius geniporus Valenciennes, 1837 (Fig. 55)—Slender Goby .....................................................57

Gobius incognitus Kovačić & Šanda, 2016 (Fig. 37)—Incognito Goby ................................................58

Gobius kolombatovici Kovačić & Miller, 2000 (Fig. 18)—Kolombatović’s goby ........................................59

Gobius niger Linnaeus, 1758 (Fig. 56)—Black Goby ..............................................................59

Gobius ophiocephalus Pallas, 1814 (Fig. 44)—Grass Goby .........................................................61

Gobius paganellus Linnaeus, 1758 (Fig. 57)—Rock Goby ..........................................................61

Gobius roulei de Buen, 1928 (Fig. 58)—Roule’s Goby .............................................................63

Gobius vittatus Vinciguerra, 1883 (Fig. 10)—Striped Goby .........................................................65

Gobius xanthocephalus Heymer & Zander, 1992 (Fig. 38)—Yellow-headed Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

Gobius xoriguer Iglésias, Vukić & Šanda 2021 (Fig. 59)—Kestrel Goby. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66

Hazeus ingressus Engin, Larson & Erhan, 2018 (Fig. 35)—Invaded Sand Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66

Hetereleotris vulgaris (Klunzinger, 1871) (Fig. 47)—Common Gogglegoby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67

Lebetus guilleti (Le Danois, 1913) (Fig. 8)—Guillet’s Goby ........................................................67

Lebetus patzneri Schliewen, Kovačić & Ordines, 2019 (Fig. 11)—Patzner’s Goby .......................................68

Lesueurigobius friesii (Malm, 1874) (Fig. 17)—Fries’s Goby .......................................................68

Lesueurigobius sanzi (de Buen, 1918) (Fig. 60)—Sanzo’s Goby .....................................................68

Lesueurigobius suerii (Risso, 1810) (Fig. 16)—Lesueur’s Goby .....................................................69

Millerigobius macrocephalus (Kolombatović, 1891) (Fig. 27)—Large-headed Goby .....................................70

Odondebuenia balearica (Pellegrin & Fage, 1907) (Fig. 26)—Coralline Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

Oxyurichthys petersii (Klunzinger, 1871) (Fig. 45)—Red Sea Tentacle-goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71

Papillogobius melanobranchus (Fowler, 1934) (Fig. 61)—Blackthroat Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

Pomatoschistus adriaticus Miller, 1973 (Fig. 62)—Mediterranean Painted Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

Pomatoschistus bathi Miller, 1982 (Fig. 63)—Bath’s Goby .........................................................74

Pomatoschistus knerii (Steindachner, 1861) (Fig. 64)—Kner’s Goby .................................................75

Pomatoschistus marmoratus (Risso, 1810) (Fig. 65)—Marbled Goby. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76

Pomatoschistus microps (Krøyer, 1838) (Fig. 66)—Common Goby ..................................................77

Pomatoschistus minutus (Pallas, 1770) (Fig. 67)—Sand Goby ......................................................79

Pomatoschistus nanus Engin & Seyhan, 2017 (Fig. 68)—Pygmaean Goby. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80

Pomatoschistus norvegicus (Collett, 1902) (Fig. 69)—Norway Goby .................................................81

Pomatoschistus quagga (Heckel, 1837) (Fig. 70)—Quagga Goby ....................................................82

Pomatoschistus tortonesei Miller, 1968 (Fig. 71)—Tortonese’s Goby .................................................83

Pseudaphya ferreri (de Buen & Fage, 1908) (Fig. 30)—Ferrer’s Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

KOVAČIĆ ET AL.

Silhouettea aegyptia (Chabanaud, 1933) (Fig. 72)—Egyptian Goby ..................................................84

Speleogobius llorisi Kovačić, Ordines & Schliewen, 2016 (Fig. 13)—Lloris’ Grotto Goby. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

Speleogobius trigloides Zander & Jelinek, 1976 (Fig. 12)—Grotto Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

Thorogobius ephippiatus (Lowe, 1839) (Fig. 5)—Leopard-spotted Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

Thorogobius macrolepis (Kolombatović, 1891) (Fig. 6)—Large-scaled Goby ..........................................86

Tridentiger trigonocephalus (Gill, 1859) (Fig. 9)—Chameleon Goby .................................................87

Trypauchen vagina (Bloch & Schneider, 1801) (Fig. 1)—Pink Worm Goby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87

Vanderhorstia mertensi Klausewitz, 1974 (Fig. 4)—Mertens’ Shrimpgoby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87

Vanneaugobius dollfusi Brownell, 1978 (Fig. 19)—Dollfus’ Goby ...................................................88

Zebrus pallaoroi Kovačić, Šanda & Vukić, 2021 (Fig. 29)—Pallaoro’s Goby ...........................................89

Zebrus zebrus (Risso, 1827) (Fig. 28)—Zebra Goby ...............................................................89

Discussion ................................................................................................90

Acknowledgements .........................................................................................91

References ................................................................................................91

Abstract

Numerous photographs of live fishes posted by anglers and divers on social media and citizen science databases are

important sources of information for ichthyological research. However, validating records that extend the known ecology

and bathymetric or geographic distribution of species should rely on a rigorous identification process. The family

Gobiidae, with their small size, superficial resemblance among species and high species richness are particularly difficult

to identify. Therefore, the identification from photographs of live individuals of Mediterranean marine gobies from the

continental shelf was studied. A dichotomous identification key is provided based on photographs of live individuals,

allowing positive identification of 41 out of the 66 species reviewed in this publication. Then, for all 66 species we

provide a brief description of important characters, which can be used for provisional identification for those species that

could not be positively identified using the key. Pending further progress in identification of live individuals, we suggest

that records extending the known geographic and ecological species distribution be taken into account only if they could

be validated using the dichotomous identification key.

Key words: identification protocol, visual identification criteria, citizen science, underwater visual census, small benthic

fishes

Introduction

The Gobiidae is the most species-rich fish family both in the Mediterranean Sea (Kovačić 2020) and worldwide

(Fricke et al. 2021; Parenti 2021). Knowledge of the diversity and taxonomy of Mediterranean gobiid species was

consolidated in the second half of the 20th century by a series of descriptions and redescriptions of species and

genera and by synthesis works clarifying synonymies, most of them by Peter Miller (Kovačić 2020). Since the

1980s, the taxonomy of the Mediterranean gobies has been relatively stable at the species level (Miller 1986); yet

new species are still regularly described (Kovačić 2020; Iglésias et al. 2021a; Kovačić et al. 2021; Goren & Stern

2021). Currently, the known Mediterranean gobiid fauna includes 64 native species and 13 alien species, thus

totaling 77 gobiid species (Table 1). However, the small size of gobies, the superficial resemblance among species

and high species richness can make their identification difficult, even in a laboratory with a stereomicroscope

and staining procedures (Kovačić 2020). Presently, laboratory identification of Mediterranean species of Gobiidae

can be achieved using Miller’s (1986) key and short descriptions of species of the Northeastern Atlantic and the

Mediterranean, Kovačić’s (2008a) key to species of the Adriatic Sea, and Kovačić’s (2020) key to Mediterranean

species.

Species identification is even more challenging for live individuals recorded from photographs. Many

characters based on morphological details in standard keys (Miller 1986; Kovačić 2008a, 2020) cannot be checked

on photographs. Photographic records of marine fishes in situ have dramatically increased over the last decade,

thanks to underwater cameras becoming increasingly affordable and to the development of citizen science databases

and data sharing through the social media. These photographic records can be an important source of information

on the geographic and ecological distribution of species, provided that identification is reliable (Kovačić & Svensen

2018). However, species misidentifications are not uncommon for these photographic records. While erroneous

TABLE 1. The checklist of the species of gobies (Gobiidae) in the Mediterranean Sea adapted from Kovačić (2020), updated by Iglésias et al. (2021), Kovačić et al. (2021), Goren

& Stern (2021), Özden et al. (2022). *alien species from Indo-West Pacific genera first described in Mediterranean as non-native area.

Species Species characteristics Identification status in present work

Aphia minuta (Risso, 1810) Pelagic above continental shelf, sometimes close to

the bottom, live coloration known

In the key, positive identification, with species description

Aulopareia unicolor (Valenciennes, 1837) Alien benthic on continental shelf, only freshly

dead coloration known

In the key, no positive identification, with species description

Buenia affinis Iljin, 1930 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Buenia lombartei Kovačić, Ordines & Schliewen, 2018 Bathyal on upper continental slope, only freshly

dead coloration known

Not in the key and with no description since bathyal

Buenia massutii Kovačić, Ordines & Schliewen, 2017 Benthic on continental shelf, only freshly dead

coloration known

In the key, no positive identification, with species description

Chromogobius quadrivittatus (Steindachner, 1863) Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Chromogobius zebratus (Kolombatović, 1891) Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Corcyrogobius liechtensteini (Kolombatović, 1891) Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Coryogalops ocheticus (Norman, 1927) Alien benthic on continental shelf, only freshly

dead coloration known

In the key, no positive identification, with species description

Cryptocentrus caeruleopunctatus (Rüppell, 1830) Alien benthic on continental shelf, live coloration

known

In the key, positive identification, with species description

Cryptocentrus steinhardti Goren & Stern, 2021 Alien benthic on continental shelf, live coloration

known*

In the key, positive identification, with species description

Crystallogobius linearis (Düben, 1845) Pelagic above continental shelf, sometimes close to

the bottom, extending to upper continental slope,

live coloration known

In the key, positive identification, with species description

Deltentosteus collonianus (Risso, 1820) Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Deltentosteus quadrimaculatus (Valenciennes, 1837) Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Didogobius bentuvii Miller, 1966 Benthic on continental shelf, known from a single

record with no live or freshly dead coloration

known

Not in the key and with no description since rare and with no

records of live or freshly dead specimens

Didogobius schlieweni Miller, 1993 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Didogobius splechtnai Ahnelt & Patzner, 1995 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Gammogobius steinitzi Bath, 1971 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

......continued on the next page

KOVAČIĆ ET AL.

TABLE 1. (Continued)

Species Species characteristics Identification status in present work

Gobius ater Bellotti, 1888 Benthic on continental shelf, very rarely recorded,

with no positive records of live or freshly dead

coloration

Not in the key and with no description since rare and with no

records of live or freshly dead specimens

Gobius auratus Risso, 1810 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Gobius bucchichi Steindachner, 1870 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Gobius cobitis Pallas, 1814 Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Gobius couchi Miller & El-Tawil, 1974 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Gobius cruentatus Gmelin, 1789 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Gobius fallax Sarato, 1889 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Gobius gasteveni Miller, 1974 Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Gobius geniporus Valenciennes, 1837 Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Gobius incognitus Kovačić & Šanda, 2016 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Gobius kolombatovici Kovačić & Miller, 2000 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Gobius niger Linnaeus, 1758 Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Gobius ophiocephalus Pallas, 1814 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Gobius paganellus Linnaeus, 1758 Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Gobius roulei de Buen, 1928 Benthic on continental shelf, extending to upper

continental slope, live coloration known

In the key, no positive identification, with species description

Gobius vittatus Vinciguerra, 1883 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Gobius xanthocephalus Heymer & Zander, 1992 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Gobius xoriguer Iglésias, Vukić & Šanda, 2021 Benthic on continental shelf, only freshly dead

coloration known

In the key, no positive identification, with species description

Gymnesigobius medits Kovačić, Ordines, Ramirez-Amaro &

Schliewen, 2019

Bathyal on upper continental slope, only freshly

dead coloration known

Not in the key and with no description since bathyal

Hazeus ingressus Engin, Larson & Erhan, 2018 Alien benthic on continental shelf, live coloration

known*

In the key, positive identification, with species description

Hetereleotris vulgaris (Klunzinger, 1871) Alien benthic on continental shelf, live coloration

known

In the key, positive identification, with species description

Knipowitschia caucasica (Berg, 1916) Euryhaline at river mouths and lakes Not in the key and with no description since euryhaline

......continued on the next page

TABLE 1. (Continued)

Species Species characteristics Identification status in present work

Knipowitschia panizzae (Verga, 1841) Euryhaline at river mouths and lakes Not in the key and with no description since euryhaline

Lebetus guilleti (Le Danois, 1913) Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Lebetus patzneri Schliewen, Kovačić & Ordines, 2019 Benthic on continental shelf, only freshly dead

coloration known

In the key, positive identification, with species description

Lesueurigobius friesii (Malm, 1874) Benthic on continental shelf, extending to upper

continental slope, live coloration known

In the key, positive identification, with species description

Lesueurigobius sanzi (De Buen, 1918) Benthic on continental shelf, only freshly dead

coloration known

In the key, no positive identification, with species description

Lesueurigobius suerii (Risso, 1810) Benthic on continental shelf, extending to upper

continental slope, live coloration known

In the key, positive identification, with species description

Millerigobius macrocephalus (Kolombatović, 1891) Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Neogobius melanostomus (Pallas, 1811) Euryhaline at river mouths and lakes Not in the key and with no description since euryhaline

Ninnigobius canestrinii (Ninni, 1883) Euryhaline at river mouths and lakes Not in the key and with no description since euryhaline

Odondebuenia balearica (Pellegrin & Fage, 1907) Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Oxyurichthys keiensis (Smith, 1938) Alien, with no positive records of live or freshly

dead coloration

Not in the key and with no description since rare and with no

records of live or freshly dead specimens

Oxyurichthys petersii (Klunzinger, 1871) Alien benthic on continental shelf, live coloration

known

In the key, positive identification, with species description

Papillogobius melanobranchus (Fowler, 1934) Alien benthic on continental shelf, only freshly

dead coloration known

In the key, no positive identification, with species description

Pomatoschistus adriaticus Miller, 1973 Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Pomatoschistus anatoliae Engin & Innal, 2017 Euryhaline at river mouths and lakes Not in the key and with no description since euryhaline

Pomatoschistus bathi Miller, 1982 Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Pomatoschistus knerii (Steindachner, 1861) Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Pomatoschistus marmoratus (Risso, 1810) Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Pomatoschistus microps (Krøyer, 1838) Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Pomatoschistus minutus (Pallas, 1770) Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

Pomatoschistus nanus Engin & Seyhan, 2017 Benthic on continental shelf, live coloration known In the key, no positive identification, with species description

......continued on the next page

KOVAČIĆ ET AL.

TABLE 1. (Continued)

Species Species characteristics Identification status in present work

Pomatoschistus norvegicus (Collett, 1902) Benthic on continental shelf, extending to upper

continental slope, live coloration known

In the key, no positive identification, with species description

Pomatoschistus quagga (Heckel, 1837) Benthic on continental shelf, sometimes free-swim-

ming close to the bottom, live coloration known

In the key, no positive identification, with species description

Pomatoschistus tortonesei Miller, 1968 Benthic on continental shelf, only freshly dead

coloration known

In the key, no positive identification, with species description

Pseudaphya ferreri (de Buen & Fage, 1908) Pelagic above continental shelf, sometimes close to

the bottom, live coloration known

In the key, positive identification, with species description

Silhouettea aegyptia (Chabanaud, 1933) Alien benthic on continental shelf, only freshly

dead coloration known

In the key, no positive identification, with species description

Speleogobius llorisi Kovačić, Ordines & Schliewen, 2016 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Speleogobius trigloides Zander & Jelinek, 1976 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Thorogobius ephippiatus (Lowe, 1839) Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Thorogobius macrolepis (Kolombatović, 1891) Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Tridentiger trigonocephalus (Gill, 1859) Alien benthic on continental shelf, live coloration

known

In the key, positive identification, with species description

Trypauchen vagina (Bloch & Schneider, 1801) Alien benthic on continental shelf, only freshly

dead coloration known

In the key, positive identification, with species description

Vanderhorstia mertensi Klausewitz, 1974 Alien benthic on continental shelf, live coloration

known

In the key, positive identification, with species description

Vanneaugobius dollfusi Brownell, 1978 Benthic on continental shelf, live coloration known In the key, positive identification, with species description

Vanneaugobius pruvoti (Fage, 1907) Benthic on continental shelf, very rare with no con-

firmed live or freshly dead coloration known

Not in the key and with no description since rare and with no

records of live or freshly dead specimens

Zebrus pallaoroi Kovačić, Šanda & Vukić, 2021 Benthic on continental shelf, live coloration not

well known

In the key, positive identification, with species description

Zebrus zebrus (Risso, 1827) Benthic on continental shelf, live coloration known In the key, positive identification, with species description

identification has limited consequences when a specimen is recorded within the range of already well-known

distributions, extralimital misidentifications can lead to erroneous knowledge of gobiid species (Kovačić 2020).

Challenging in situ identification of gobies has another consequence for studies of fish assemblages based on

visual data. The most common method to study coastal marine fish assemblages is underwater visual census by

SCUBA diving (Watson et al. 2005), which usually rarely or roughly report gobiid species. In the Mediterranean,

gobies have generally remained unidentified during such censuses, being listed as Gobiidae (e.g. Guidetti & Bussotti

2002) or Gobius sp. (e.g. Relini et al. 1998), or identification was provided to the species level only for a few of the

largest species (Azzuro et al. 2013). Only when visual census involves an expert in Gobiidae, the true richness of

this group and its importance in fish assemblages is revealed (e.g. 14 gobiid species recorded in Kovačić 2002a).

It is expected that the citizen science data and in situ visual observations will continue to increase in the future.

A well-defined protocol that includes both the positive and provisional identifications of gobies from visual data

would help both purposes, i.e. reliably expanding our knowledge of the geographic and ecological distribution of

gobiid species, and increasing the proportion of identified gobiid individuals and recognized species in studies of

fish assemblages based on visual data, ultimately revealing their real significance in the benthic fish community.

The goal of this article is to propose such a protocol for the identification of Mediterranean marine gobies of

the continental shelf from photographs of live individuals. We first provide a dichotomous key allowing positive

identification to species level, for some but not all of the considered species. Then, for all species we provide a brief

description of important characters, which can help in identification for those species that could not be resolved

by the dichotomous key. With this work, we thus also aim to define the limits of positive species identification in

Mediterranean gobies, based on current knowledge.

Material and methods

Among 77 gobiid species known for the Mediterranean Sea (Table 1; Kovačić 2020; Iglésias et al. 2021a; Kovačić

et al. 2021; Özden et al. 2022), five species (Knipowitschia caucasica (Berg, 1916), Knipowitschia panizzae (Verga,

1841), Neogobius melanostomus (Pallas, 1811), Ninnigobius canestrinii (Ninni, 1883), Pomatoschistus anatoliae

Engin & Innal, 2017) are euryhaline and therefore do not occur in regular marine conditions, and two species are

exclusively bathyal (Buenia lombartei Kovačić, Ordines & Schliewen, 2018 and Gymnesigobius medits Kovačić,

Ordines, Ramirez-Amaro & Schliewen, 2019), not occurring on the continental shelf (Table 1). Moreover, three

(Didogobius bentuvii Miller, 1966, Gobius ater Bellotti, 1888, and Vanneaugobius pruvoti (Fage, 1907)) of the

remaining shelf species are very rare with no records of live or freshly dead specimens, as well as alien Oxyurichthys

keiensis (Smith, 1938) (Table 1). Hence 66 gobiid species were reviewed in this article, including 54 native and

12 alien species (Table 1). Five of the 54 native species have no published information on live coloration (Buenia

massutii Kovačić, Ordines & Schliewen, 2017, Gobius xoriguer Iglésias, Vukić & Šanda, 2021, Lebetus patzneri

Schliewen, Kovačić & Ordines, 2019, Lesueurigobius sanzi (De Buen, 1918), and Pomatoschistus tortonesei Miller,

1968), but their freshly dead coloration has been described. Among these five species, only L. patzneri has a unique

and distinct freshly dead coloration that is expected to match live coloration; therefore this was the only species

for which coloration allowed a positive identification among these species. Similarly, five of the 12 alien species

(Aulopareia unicolor (Valenciennes, 1837), Coryogalops ocheticus (Norman, 1927), Papillogobius melanobranchus

(Fowler, 1934), Silhouettea aegyptia (Chabanaud, 1933), and Trypauchen vagina (Bloch & Schneider, 1801)) lack

published information on live coloration but have known freshly dead coloration. Among them, only T. vagina can

be positively identified to species thanks to its very peculiar morphological and color characteristics. In addition,

for B. massutii and T. vagina, along with a photograph showing the species freshly dead coloration, we provide

photographs of live individuals which identity could not be confirmed with certainty.

This identification guide is organized in two parts: i) a key for identification of Mediterranean marine gobies of

the continental shelf from photographs of live individuals, and ii) brief accounts of all reviewed species, providing

a secondary check for species already included in the key, or information for provisional identification for the

remaining species.

The key is organized as a dichotomous bracket with the key characters provided as couplets with alternatives.

The present key considers all 66 species selected according to the criteria listed previously. However, it is incomplete

in a sense that it does not allow to identify some species to species-level. Those species are placed together at the end

of the key as an unresolved group within the second alternative of the last couplet. The key is restricted to adults of

KOVAČIĆ ET AL.

both sexes, and to good-quality underwater photographs of entire fish with clearly visible lateral and possibly dorsal

views. Furthermore, it should be used only for identification of live, unstressed individuals in their natural habitat,

and not for photographs of preserved or freshly dead specimens, or of live specimens kept in artificial containers

on land as their coloration could be altered in these unnatural conditions. Some photographs illustrating detailed

characters mentioned in the key were nevertheless made in portable aquaria; we included such photographs only

when we were certain that they illustrated characters as they are expressed in natural conditions. The characters used

in the key came from published data on live coloration (see the species accounts for the references), and from the

authors’ experience of underwater observations and analyses of underwater photographs, notably of specimens that

were subsequently collected and identified from their morphology.

For the species that cannot be positively identified from live photographs, the short account for each of these

species can be used to achieve provisional identification. The species accounts include important characters that

can give clues to identify species, including coloration, body shape, and also habitat preferences and geographic

distribution; they are accompanied with photographs. The benthic zonation and habitats for the Mediterranean

follows Bellan-Santini et al. (2002). Meristic abbreviations used in the species accounts: D - dorsal fins, A - anal

fin, P - pectoral fin(s).

Results

The key for identification of marine gobies of the Mediterranean continental shelf allows positive identification

for 41 species, leaving 25 species as an unresolved group in the second alternative of the last couplet. For these 25

species, we suggest that it is only possible to reach a provisional identification, based on the descriptions provided

in the species accounts. In addition, at the present state of knowledge, we are not sure if Zebrus zebrus (Risso, 1827)

and Zebrus pallaoroi Kovačić, Šanda & Vukić, 2021 can be positively distinguished on underwater photographs

considering general similarity and poor knowledge on Z. pallaoroi’s live aspect.

Family identification

Fishes belonging to the family Blenniidae (blennies) and Tripterygiidae (triplefins) can have similar size and

coloration to and be syntopic with gobies on rocky and mixed bottoms. However, individuals from these families

can be easily distinguished by their dorsal fins: blennies have a single dorsal fin, triplefins have three, while gobies

have two dorsal fins. One exception is the alien gobiid T. vagina (with a single dorsal fin, but whose unique shape

and proportions prevents any confusion with a blenny). Dragonets (family Callionymidae) can also superficially

resemble gobies and have two dorsal fins; sharing sandy bottom habitats with some gobies. However, they are

readily identified by their wide, largely depressed and triangular head shape, and pointed snout.

Key for the identification of Mediterranean marine gobies from photographs of live individuals

1a One dorsal fin; dorsal and anal fins confluent with caudal fin (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

.......................................................... Trypauchen vagina (Bloch & Schneider, 1801)

1b Two dorsal fins: if first dorsal fin reduced, then second dorsal fin and anal fin clearly separated from caudal fin .......2

2a (1) Head and predorsal area completely covered with oval to rounded, clearly defined, dark to orange or yellow-orange spots

or blotches (Fig. 2) .................................................................................3

2b (1) Head and predorsal area without oval to rounded, clearly defined, dark or orange spots ...........................6

3a (2) Side of head with blue-edged dark reddish spots; dorsal half of body with 2 large green-edged black ocellated spots (Fig.

3) ......................................................Cryptocentrus caeruleopunctatus (Rüppell, 1830)

3b (2) No pale to bluish edge around head spots; no ocellated spots on body .........................................4

4a (3) Head shorter than caudal fin; body slender; body with a mix of yellow and dark brown markings, first dorsal fin with yellow

spots (Fig. 4) ....................................................Vanderhorstia mertensi Klausewitz, 1974

4b (3) Head longer than caudal fin; body moderately deep; body with either orange or dark brown markings, first dorsal fin with

orange or dark brown longitudinal bands ...............................................................5

5a (4) Body pale bluish gray, with pattern of large black, brown-purple or orange-brown blotches and spots (Fig. 5) ..........

.................................................................Thorogobius ephippiatus (Lowe, 1839)

5b (4) Body pale, with orange to yellow-orange spots (Fig. 6) . . . . . . . . . . . . . . Thorogobius macrolepis (Kolombatović, 1891)

6a (2) Head and body are completely dark brown to purplish, with definite white markings, including 3–6 blotches at least as large

as eye diameter on dorsal half of body, a white transverse band in predorsal area, and dorsal and caudal fins with a broad

white outer edge (Fig. 7) ................................................Didogobius schlieweni Miller, 1993

6b (2) Head and body are not completely dark brown to purplish or if it is, then without conspicuous white blotches .........7

7a (6) A well delineated dark area on body between second dorsal-fin base and anal-fin base, contrasting with a plain pale caudal

peduncle (Fig. 8) .......................................................Lebetus guilleti (Le Danois, 1913)

7b (6) Color pattern without a dark area below second dorsal fin contrasting with pale caudal peduncle ...................8

8a (7) Two continuous or interrupted dark brown stripes on body, sometimes not well defined, the upper stripe running from above

eyes along back and ending at upper caudal-fin base, the lower stripe running from snout tip through eyes along mid-body

to caudal fin (Fig. 9) ................................................Tridentiger trigonocephalus (Gill, 1859)

8b (7) Color pattern with no or with a single longitudinal dark stripe ...............................................9

9a (8) Head and body pale to white, with one broad longitudinal dark stripe beginning from the upper lip, continuing over eye and

above body midline, to caudal-fin base (Fig. 10) ..............................Gobius vittatus Vinciguerra, 1883

9b (8) Color pattern without longitudinal dark stripe along body side, or if dark stripe is present, then body with other dark

markings ........................................................................................10

10a (9) Head and body red with irregular white markings, first dorsal fin entirely dark (Fig. 11) ...........................

....................................................Lebetus patzneri Schliewen, Kovačić & Ordines, 2019

10b (9) Head and body coloration not red, or if the primary pattern is reddish, then the first dorsal fin is not entirely dark (although

it can display a dark spot or blotch) ................................................................... 11

11a (10) Body red with 5 whitish saddles on back, the first anterior at the origin of the first dorsal fin, the last on caudal peduncle just

before caudal fin; 5–6 barely-defined white to yellowish blotches along ventral profile; in males, caudal fin and sometimes

posterior body yellow, and dorsal fins yellowish with at least one oblique pink to light purple stripe on each fin (Fig. 12) .

......................................................... Speleogobius trigloides Zander & Jelinek, 1976

11b (10) Head and body not red or if the primary pattern is reddish, then dorsal and ventral whitish marks are either absent or there

are more than 5 dorsal and 6 ventral whitish marks ......................................................12

12a (11) Body brown to reddish or beige, head reddish; 3 dark bars on dorsal half of body alternating with 4 pale areas (Fig. 13),

bars linked with 4 dark blotches on lower side, one more midlateral blotch on caudal peduncle (Fig. 13) . . . . . . . . . . . . . .

................................................. Speleogobius llorisi Kovačić, Ordines & Schliewen, 2016

12b (11) Head and body not red or brown-red, or if primary pattern is reddish, then there are more than 3 dark bars on dorsal half,

alternating with more than 4 pale areas, and without 4 dark blotches on lower side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

13a (12) Body mostly dark brown with 3 narrow pale bars extending onto ventral half of body (the first just before first dorsal fin

and extending onto pectoral-fin base), plus a very thin one on caudal-fin base (Fig. 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . .

.......................................................... Didogobius splechtnai Ahnelt & Patzner, 1995

13b (12) Body not mostly dark brown or, if mostly dark brown, then without well-defined transverse pattern of 3 pale bars extending

onto ventral half of body ...........................................................................14

14a (13) Body mostly mottled brown with midlateral row of broad rectangular ill-defined dark blotches; thin whitish saddles on back

(three below second dorsal fin); side of lips vermillion with prominent white bar on lips at angle of mouth (red color usually

not detectable under natural light) (Fig. 15) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gobius cruentatus Gmelin, 1789

14b (13) Head and body usually without mottled brown pattern and midlateral brown blotches, if mottled brown pattern present, then

lips not orange or red ..............................................................................15

15a (14) Head and body pale to grayish background often with a bluish hue, 3–4 slightly oblique yellow to yellow-orange stripes on

side of head, and one slanting yellow stripe above them behind eye (Fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

....................................................................Lesueurigobius suerii (Risso, 1810)

15b (14) Head without oblique yellow to yellow-orange stripes ....................................................16

16a (15) Body with two longitudinal rows of brownish spots (often with yellow to orange hue), one row along back and another

along lateral midline; second dorsal fin with a yellow to orange outer margin and whitish to blue submarginal band; both

dorsal fins and upper half of caudal fin with yellow to orange spots (Fig. 17) . . . . . . Lesueurigobius friesii (Malm, 1874)

16b (15) Body without longitudinal rows of gray-brown or orange-brown spots, or if such rows are present, then second dorsal fin

without a combination of yellow margin and whitish to blue submarginal band . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17a (16) Head and body with irregular orange blotches arranged in longitudinal series; two horizontal irregular orange stripes on

head, one behind eye, the other from below eye to upper pectoral-fin base; V-shaped orange stripe across snout; first dorsal

fin with blackish oval blotch between third and fifth spines (sometimes faint) (Fig. 18) . . . . . . . . . . . . . . . . . . . . . . . . . . . .

...........................................................Gobius kolombatovici Kovačić & Miller, 2000

17b (16) Head and body without orange blotches, or if orange blotches present, then no orange V-shaped line on snout, or if V-shaped

line is present, then no horizontal stripe present behind eye; dark blotch on first dorsal fin absent, or if present blotch is

elongate and extends along fin base, usually from first to fifth spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

18a (17) Head and body grayish with pale orange, reddish, rarely with pinkish blotches and speckles, or indistinct bars of these

colors on paler background; a series of short pearly white bars arranged in 3-5 pairs along lower body; anterior nostrils

contrastingly white (Fig. 19) ......................................... Vanneaugobius dollfusi Brownell, 1978

18b (17) No pure orange markings, although overall color can be brown-orange, reddish or with traces of orange pigmentation . . .

...............................................................................................19

19a (18) Body with pattern of transverse narrow pale bars on reddish to brownish background; 3 pale, uninterrupted bars radiating

downwards from eye to ventral part of head, two bars below eye and one behind (Figs. 20–22) . . . . . . . . . . . . . . . . . . . . 20

KOVAČIĆ ET AL.

19b (18) Body and head usually without transverse bars, or if pattern is transverse, then less than three complete pale bars radiating

downwards from eye to the ventral part of head .........................................................21

20a (19) Body reddish to orangish brown with 10–15 narrow bluish bars (Fig. 21) ......................................

.......................................................Corcyrogobius liechtensteini (Kolombatović, 1891)

20b (19) Body brownish with 6–7 narrow pale bars (Fig. 22) ............................Gammogobius steinitzi Bath, 1971

21a (19) Head clearly depressed (distinctly dorsoventrally flattened) (Figs. 23, 24 and 25) ...............................22

21b (19) Head not clearly depressed (not distinctly dorsoventrally flattened) ..........................................23

22a (21) Body fawn to blackish with a broad pale to white predorsal bar extending ventrally on pectoral-fin base; no pale saddle at

origin of first dorsal fin; prominent to indistinct white saddle at origin of second dorsal fin, sometimes another short saddle

below posterior part of second dorsal fin; cheeks pigmented, densely dotted or with reticulate pattern (Fig. 24) . . . . . . . . .

.......................................................Chromogobius quadrivittatus (Steindachner, 1863)

22b (21) Broad, often faint pale bar on predorsal area, extending ventrally on pectoral-fin base; 5 or 6, often faint, pale saddles of

same intensity, extending or not as lateral bars, the first at origin of first dorsal fin, the last just behind second dorsal fin;

cheeks pale with two dark oblique bands radiating downwards from eye in the shape of an inverted V or a triangular mark

(Fig. 25) ....................................................Chromogobius zebratus (Kolombatović, 1891)

23a (21) Body pattern of pale transverse bars on reddish, brownish or greenish background, sometimes reduced to pale dorsal

saddles, side of body then more or less uniform or mottled reddish, brownish or greenish (Figs. 26–29) . . . . . . . . . . . . . 24

23b (21) No transverse bars on the body; if patterned with pale dorsal saddles, then pigmentation on body side not uniform or

mottled .........................................................................................26

24a (23) Head and body prevailing reddish or orangish with pale bars (Fig. 26a), or with a series of short white saddles along back

and side of body more or less uniform (Fig. 26b); no midlateral row of small black spots; whitish spot at the upper edge of

opercle ..................................................Odondebuenia balearica (Pellegrin & Fage, 1907)

24b (23) Head and body not reddish or orangish, usually with brownish or greenish background; 6−11 pale bars on body (Figs. 27a,

28a and 29a) or series of short white saddles along back, side of body being then more or less mottled, with midlateral row

of small black spots (Figs. 27b, 28b and 29b); often a pale, crescent-shaped, transverse band across anterior nape behind

eyes, reaching to pectoral-fin base (band may be absent); no whitish spot at the upper edge of opercle . . . . . . . . . . . . . . 25

25a (24) When visible, a broad pale crescent-shaped band across anterior nape behind eyes extends onto rear part of eyes; anterior

nostril tube without tentacle (Fig. 27) ........................Millerigobius macrocephalus (Kolombatović, 1891)

25b (24) When visible, a broad pale crescent-shaped band across anterior nape behind eyes does not extend onto eyes; anterior

nostril with a tentacle (Figs. 28 and 29) ..................................................................

.......................... Genus Zebrus: Z. zebrus (Risso, 1827) or Z. pallaoroi Kovačić, Šanda & Vukić, 2021

26a (23) Body translucent or semitranslucent and skin surface poorly pigmented, prominent melanophores form a series along lower

body or as caudal-fin mark; other pigments, if present, internal, on vertebrae and organs; fins translucent (Figs. 30, 31 and

32) ............................................................................................27

26b (23) Body not translucent or only partially translucent and skin with more prominent pigmentation than just a series of

melanophores along lower body or a caudal-fin mark; at least some pigment on fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

27a (26) Body semitranslucent with 4 elongate black markings along vertebral column and large, conspicuous, triangular black spot

at base of caudal fin (Fig. 30) .....................................Pseudaphya ferreri (de Buen & Fage, 1908)

27b (26) Body translucent with row of melanophores along anal-fin base, no triangular black spot at base of caudal fin, no black

markings along vertebral column .....................................................................28

28a (27) Second dorsal fin of moderate length, with I+11–13 rays; side of abdomen with no well defined black spots (Fig. 31) ....

.......................................................................... Aphia minuta (Risso, 1810)

28b (27) Second dorsal fin long, with I+18–20 rays; side of abdomen with longitudinal row of 3–4 well-defined tiny black spots (Fig.

32) ..............................................................Crystallogobius linearis (Düben, 1845)

29a (26) Body with longitudinal rows or series of dots or small dashes (Figs. 33–39) ...................................30

29b (26) Body without longitudinal rows of dots or small dashes ...................................................35

30a (29) No defined dark dot at posterior angle of mouth (Figs. 33a and 33b) .........................................31

30b (29) A well-defined dark dot at posterior angle of mouth (Fig. 33c, 33d, 33e and 33f) ...............................32

31a (30) Two longitudinal rows of elongate spots on cheek, one longitudinal mideye stripe (Fig. 34) ........................

..................................................................Gobius bucchichi Steindachner, 1870

31b (30) No rows of elongate spots on cheek and no longitudinal eye stripe (Fig. 35) .....................................

.........................................................Hazeus ingressus Engin, Larson & Erhan, 2018

32a (30) Ground coloration of body yellow, upper lip unmarked except dot at corner of mouth (Figs. 33c and 36) ..............

...................................................................Gobius auratus Risso, 1810 morph 2

32b (30) Ground coloration of body not yellow, or if body coloration yellow then upper lip with at least one brown or orange mark

in addition to dot on corner of mouth (Figs. 37b, 38b and 39b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

33a (32) On eye, upper iris with separate brown dots or radiating stripes, without longitudinal curved mark (Fig. 37b) ..........

..............................................................Gobius incognitus Kovačić & Šanda, 2016

33b (32) On eye, a dark curved mark on upper iris connects longitudinally other marks (Figs. 38b and 39b) . . . . . . . . . . . . . . . . . 34

34a (33) Dark red to light orange dots or tiny spots on body (Fig. 38) ........ Gobius xanthocephalus Heymer & Zander, 1992

34b (33) Brown and dark brown dots on body (Fig. 39) ......................................Gobius fallax Sarato, 1889

35a (29) Head and body uniformly yellow (Fig. 40) .................................Gobius auratus Risso, 1810 morph 1

35b (29) Head and body with irregularly scattered dots, midlateral dark marks, mottled brown pattern or any combination of these

characters .......................................................................................36

36a (35) 4–5 well-defined, usually dark-edged, pale dorsal saddles, each terminating with midlateral dark blotch; ventral half of

body contrasting white with 10–12 short and narrow brownish bars (Fig. 41) . . . . . . . . . . . . . . . Buenia affinis Iljin, 1930

36b (35) No pale dorsal saddles or, if such saddles or blotches present, no short narrow brownish bars on white lower side of body

...............................................................................................37

37a (36) One black spot on the corner of the mouth, usually followed by two elongate dark spots on ventral part of cheek (Figs. 42

and 43) .......................................................... Gobius couchi Miller & El-Tawil, 1974

37b (36) No black spot at corner of mouth .....................................................................38

38a (37) Body with irregular, zigzag vertical dark bars, 8 or more along lateral midline, sometimes coalescing in upper lateral stripe,

10 or more on lower side, alternating with pale or white interspaces about the same width (Fig. 44) . . . . . . . . . . . . . . . . . .

....................................................................Gobius ophiocephalus Pallas, 1814

38b (37) No or fewer than 8 dark markings along lateral midline, and/or no multiple alternating dark and pale areas along lower

body ...........................................................................................39

39a (38) Eyes high on head, top of eye protruding above dorsal profile; head length no more than 3/5 of caudal-fin length (Figs. 45

and 46) .........................................................................................40

39b (38) Eye not high on head, their top forming part of dorsal profile, and head length more than 3/5 of caudal-fin length .....41

40a (39) Body with 5 elongate mid-lateral brown blotches (excluding that on pectoral-fin base) and light blue metallic markings

(Fig. 45) ........................................................Oxyurichthys petersii (Klunzinger, 1871)

40b (39) Pale beige background with three slightly oblique brown bars, first bar below first dorsal fin, second and third bars below

anterior and posterior parts of second dorsal fin, and large irregular brown spot at caudal-fin base (Fig. 46) . . . . . . . . . . . .

..........................................................Cryptocentrus steinhardti Goren & Stern, 2021

41a (39) Semitranslucent scaleless body with 6 wavy, irregular gray-brown bars and no lateral midline markings (Fig. 47) .......

...............................................................Hetereleotris vulgaris (Klunzinger, 1871)

41b (39) Body not semitranslucent with gray-brown irregular bars or, if semitranslucent with dark bars and no scales visible, then

lateral midline marks present . . . . . . . . . . . . . . . .Remaining species: Aulopareia unicolor (Valenciennes, 1837), Buenia

massutii Kovačić, Ordines & Schliewen, 2017, Coryogalops ocheticus (Norman, 1927), Deltentosteus collonianus

(Risso, 1820), Deltentosteus quadrimaculatus (Valenciennes, 1837), Gobius cobitis Pallas, 1814, Gobius gasteveni

Miller, 1974, Gobius geniporus Valenciennes, 1837, Gobius niger Linnaeus, 1758, Gobius paganellus Linnaeus, 1758,

Gobius roulei de Buen, 1928, Gobius xoriguer Iglésias, Vukić & Šanda 2021, Lesueurigobius sanzi (De Buen, 1918),

Papillogobius melanobranchus (Fowler, 1934), Pomatoschistus adriaticus Miller, 1973, Pomatoschistus bathi Miller,

1982, Pomatoschistus knerii (Steindachner, 1861), Pomatoschistus marmoratus (Risso, 1810), Pomatoschistus microps

(Krøyer, 1838), Pomatoschistus minutus (Pallas, 1770), Pomatoschistus nanus Engin & Seyhan, 2017, Pomatoschistus

norvegicus (Collett, 1902), Pomatoschistus quagga (Heckel, 1837), Pomatoschistus tortonesei Miller, 1968, Silhouettea

aegyptia (Chabanaud, 1933) (males of Gobius gasteveni Miller, 1974, Gobius niger Linnaeus, 1758 and Gobius

paganellus Linnaeus, 1758 (can have almost entirely dark body and head, see Figs. 53b, 55b and 56b).

FIGURE 1. A) Freshly dead coloration of Trypauchen vagina. Photo by D. Darom, trawled between Hadera and Atlit, Israel. B)

Photo of Trypauchen sp. alive, photo by C. Telford, Thailand.

KOVAČIĆ ET AL.

FIGURE 2. Head and predorsal area completely covered with oval to rounded, clearly defined spots. Photos by M. Kovačić,

S.V. Bogorodsky, R. Pillon.

FIGURE 3. Live coloration in its natural habitat of Cryptocentrus caeruleopunctatus. Photo by S.V. Bogorodsky, Lahami Bay,

Egypt, Red Sea.

FIGURE 4. Live coloration in its natural habitat of Vanderhorstia mertensi. Photo by R. Pillon, Kondyli beach, Peloponnese,

Greece.

KOVAČIĆ ET AL.

FIGURE 5. Live coloration in its natural habitat of Thorogobius ephippiatus. Photo by S. Guerrieri, Tuscany, Italy.

FIGURE 6. Live coloration in its natural habitat of Thorogobius macrolepis. Photo by S. Guerrieri, Tuscany, Italy.

FIGURE 7. Live coloration in its natural habitat of Didogobius schlieweni. Photo by C. Balisky, Marseille, Southern France.

FIGURE 8. Live coloration in its natural habitat of Lebetus guilleti. Photo by J. Herler, Selce, Croatia, Northern Adriatic Sea.

KOVAČIĆ ET AL.

FIGURE 9. Live coloration in its natural habitat of Tridentiger trigonocephalus. Photo by Sascha Schulz/iNaturalist Australasian

Fishes Project, Victoria, Australia.

FIGURE 10. Live coloration in its natural habitat of Gobius vittatus. Photo by S.V. Bogorodsky, Turkey, Aegean Sea.

FIGURE 11. Freshly dead coloration of Lebetus patzneri. Photo by F. Ordines, Mallorca, Balearic Islands, Spain.

KOVAČIĆ ET AL.

FIGURE 12. Live coloration in its natural habitat of Speleogobius trigloides: A) male, B) female. Photos by R. Svensen, Selce,

Croatia, Northern Adriatic Sea.

FIGURE 13. Live coloration in its natural habitat of Speleogobius llorisi. Photo by R. Tournier-Broer, Saint-Raphaël, Southern

France.

FIGURE 14. Live coloration in its natural habitat of Didogobius splechtnai. Photo by A. Pagano, Sicily, Italy.

KOVAČIĆ ET AL.

FIGURE 15. Live coloration in its natural habitat of Gobius cruentatus. Photo by S. Guerrieri, Tuscany, Italy.

FIGURE 16. Live coloration in its natural habitat of Lesueurigobius suerii. Photo by S. Le Bris, Cagnes-sur-Mer, Southern

France.

FIGURE 17. Live coloration in its natural habitat of Lesueurigobius friesii. Photo by R. Svensen, Stavanger, Norway.

FIGURE 18. Live coloration in its natural habitat of Gobius kolombatovici. Photo by S. Le Bris, Marseille, Southern France.

KOVAČIĆ ET AL.

FIGURE 19. Live coloration in its natural habitat of Vanneaugobius dollfusi. Photo by S. Le Bris, Marseille, Southern France.

FIGURE 20. Three pale, uninterrupted bars (marked with numbers) extending downwards from eye radiating downwards from

eye to ventral side: A) C. liechtensteini, B) G. steinitzi. Photos by M. Kovačić (fishes photographed in a tank).

FIGURE 21. Live coloration in its natural habitat of Corcyrogobius liechtensteini (Kolombatović, 1891). Photo by S. Le Bris,

Marseille, Southern France.

FIGURE 22. Live coloration in its natural habitat of Gammogobius steinitzi Bath, 1971. Photo by S. Guerrieri, Tuscany, Italy.

KOVAČIĆ ET AL.

FIGURE 23. Head clearly depressed (dorsoventrally flattened). A) Chromogobius quadrivittatus, photo by M. Colombo. B)

Chromogobius zebratus, photo by M. Kovačić, photographed in a tank.

FIGURE 24. Live coloration in its natural habitat of Chromogobius quadrivittatus, photos by A) S. Engin, Aegean Sea, Turkey

and B) M. Colombo, Area Marina Protetta Capo Caccia-Isola Piana, Alghero, Sardinia, Italy.

KOVAČIĆ ET AL.

FIGURE 25. Live coloration in its natural habitat of Chromogobius zebratus. Photo by S. Guerrieri, Tuscany, Italy.

FIGURE 26. Live coloration of Odondebuenia balearica. A) Transverse coloration pattern with vertical pale bars, photo by

M. Kovačić, Rijeka, Croatia, northern Adriatic Sea (fish photographed in a tank). B) Coloration pattern reduced to pale dorsal

saddles while the rest of body lateral side is more or less uniform or mottled, photo by S. Guerrieri, Rijeka, Croatia, the Northern

Adriatic Sea.

KOVAČIĆ ET AL.

FIGURE 27. Live coloration of Millerigobius macrocephalus. Broad pale crescent-shaped band across anterior nape behind

eyes extending onto rear part of eyes. A) Transverse coloration pattern with pale vertical bars, photo by M. Kovačić, Karinsko

ždrilo, eastern Adriatic Sea (in aquarium). B) Coloration pattern reduced to pale dorsal saddles while the rest of body lateral side

is more or less uniformly colored or mottled brown, photo by J. Renoult, Corsica, France. C) Anterior nostril without tentacle,

photo by J. Renoult, Argelès-sur-Mer, Southern France.

FIGURE 28. Live coloration in its natural habitat of Zebrus zebrus. A) Transverse coloration pattern with pale vertical bars. B)

Coloration pattern reduced to pale dorsal saddles while the rest of body lateral side is more or less uniformly colored or mottled

brown; broad pale crescent-shaped band across anterior nape behind eyes, not extending onto posterior part of eyes. C) Anterior

nostril with tentacle. Photos by M. Kovačić, A) Karinsko ždrilo, Novigrad Sea, B) Rava island, Zadar Arcipelago, both central

Adriatic Sea.

KOVAČIĆ ET AL.

FIGURE 29. Zebrus pallaoroi. A) Freshly dead coloration, transverse coloration pattern with pale vertical bars. B) Live

coloration in its natural habitat, pattern reduced to pale dorsal saddles while the rest of body lateral side is more or less uniformly

colored or mottled dark. C) anterior nostril with tentacle. A) and B) photos of the same individual. Photos by J. Renoult, Palavas-

les-Flots, Southern France.

FIGURE 30. Live coloration in its natural habitat of Pseudaphya ferreri. Photo by S. Guerrieri, Rijeka, Croatia, Northern

Adriatic Sea.

FIGURE 31. Live coloration in its natural habitat of Aphia minuta. Photo by S. Guerrieri, Tuscany, Italy.

FIGURE 32. Live coloration in its natural habitat of Crystallogobius linearis. Photo by R. Svensen, Stavanger, Norway.

KOVAČIĆ ET AL.

FIGURE 33. The mouth unmarked at corner in A) G. bucchichi and B) H. ingressus. Dark dot on the corner of the mouth in C)

G. auratus, D) G. incognitus, E) G. xanthocephalus and F) G. fallax. Area marked with hollow circle. Photos by R. Svensen (A),

S. Engin (B), M. Kovačić (C, D, F), and P. Louisy (E). C and F photographed in a tank.

FIGURE 34. Live coloration in its natural habitat of Gobius bucchichi. Photo by R. Pillon, Rab Island, Croatia, Northern

Adriatic Sea.

FIGURE 35. Live coloration in its natural habitat of Hazeus ingressus. Photo by S. Engin, Fethiye, Turkey, Levant.

KOVAČIĆ ET AL.

FIGURE 36. Live coloration in its natural habitat of Gobius auratus morph 2. Photo by A. Falleni, Pag, Croatia, Northern

Adriatic Sea.

FIGURE 37. A) Live coloration in its natural habitat of Gobius incognitus. B) Upper lip with brown marks in addition to the

dot on the corner of the mouth; upper half of the eye with separate brown dots or radiating stripes without longitudinal curved

mark. Photos by S.V. Bogorodsky, Turkey, Aegean Sea.

FIGURE 38. A) Live coloration in its natural habitat of Gobius xanthocephalus. B) Upper lip with at least one orange mark in

addition to the dot on the corner of the mouth; dark curved mark on upper border of eye connects longitudinally other marks.

Photos by P. Louisy, Banyuls, Pyrénées Orientales, Southern France.

FIGURE 39. Live coloration in its natural habitat of Gobius fallax. B) Upper lip with brown mark in addition to the dot on the

corner of the mouth; dark curved mark on the upper border of the eye connects longitudinally other marks. Photo by R. Pillon,

Hvar Island, Croatia, Central Adriatic Sea.

KOVAČIĆ ET AL.

FIGURE 40. Live coloration in its natural habitat of Gobius auratus morph 1. Photo by S.V. Bogorodsky, Turkey, Aegean

Sea.

FIGURE 41. Live coloration in its natural habitat of Buenia affinis. Photos by R. Pillon, Rab Island, Croatia, Northern Adriatic

Sea.

FIGURE 42. One black spot on the corner of the mouth, usually followed by two elongated dots on ventral border of the cheek

(all marked with arrows) in G. couchi. Photo by M. Kovačić.

FIGURE 43. Live coloration in its natural habitat of Gobius couchi. Photo by R. Pillon, Rab Island, Croatia, Northern Adriatic

Sea.

KOVAČIĆ ET AL.

FIGURE 44. Live coloration in its natural habitat of Gobius ophiocephalus. Photo by S.V. Bogorodsky, Russia, Black Sea.

FIGURE 45. Live coloration in its natural habitat of Oxyurichthys petersii. Photo by J.E. Randall, Port Sudan, Sudan, Red

Sea.

FIGURE 46. Live coloration in its natural habitat of Cryptocentrus steinhardti Goren & Stern, 2021. Photo by P. Louisy, Kas,

Turkey, Eastern Mediterranean.

FIGURE 47. Live coloration in its natural habitat of Hetereleotris vulgaris. Photo by S.V. Bogorodsky, Mangrove Bay, El

Quseir, Egypt, Red Sea.

KOVAČIĆ ET AL.

Species accounts

Important identification characters in bold.

Aphia minuta (Risso, 1810) (Fig. 31)—Transparent Goby

Atherina minuta Risso, 1810: 340; type locality: northwestern Mediterranean Sea, France, Nice.

Size. Maximum known size 5.8 cm total length (La Mesa et al. 2005).

Morphology. D IV–VI + I,11–13; A I,11–15; P 15–19. Pedomorphic habitus. Body laterally compressed. Eyes

lateral, interorbital wide. Mouth superior, oblique, tip of lower jaw at the horizontal level of pupil. Caudal peduncle

slender in females, deeper in males, lower than body height. Dorsal fins distant from each other, with a broad

interdorsal space. The first dorsal fin small, roughly of trapezoid or parallelogram shape. Second dorsal fin high,

with moderately short base and ray lengths decreasing posteriorly giving the fin triangular appearance, at

least in females. Pelvic disc complete (Miller 1986; Patzner 2021). Caudal fin truncate to slightly emarginate.

Scales present, but not usually visible on photographs.

Live coloration. Body translucent, usually with a row of melanophores along anal-fin base, no triangular

black spot at the base of caudal fin and no well-defined black spots laterally on abdomen (Fig. 31).

Similar species. Crystallogobius linearis.

Habitat. Pelagic and neritic species, widely distributed in inshore and estuarine waters from the surface to

80–100 m depth over sand, mud, sea-weed (Cystoseira spp.) and over seagrasses (Zostera marina and Posidonia

oceanica) (La Mesa et al. 2005).

Geographic distribution. The northeastern Atlantic, from the Strait of Gibraltar to the western Baltic and the

coasts of Norway, as well as in the Mediterranean, including the Adriatic Sea, Black Sea and Azov Sea (La Mesa et

al. 2005; Boltachev & Karpova 2017).

Aulopareia unicolor (Valenciennes, 1837) (Fig. 48)—Greenspot Goby

Gobius unicolor Valenciennes in Cuvier & Valenciennes, 1837: 88; type locality: Indonesia, Jawa.

Size. Reaches 13 cm total length.

Morphology. D VI + I,10; A I,9; P 18–19; body stout, moderately elongate and somewhat laterally

compressed, body depth 5.2–5.6 in standard length. Head subcylindrical, snout short, obtuse. Caudal peduncle

deep, but lower than body depth. Dorsal fins of similar height, the first dorsal fin with more or less rounded margin

and first to fifth spines longest, not elongate. Pelvic fins united to form a disc. Caudal fin rounded, subequal to head

length. Body and nape covered with scales. Scales prominent, visible on body and predorsal area, head naked except

for dorsal part of operculum (Kimura et al. 2018).

Fresh coloration. Body dark grayish brown with many small bright green spots, small black spots on side

of abdomen and usually a diffuse blue-green mark above posterior part of opercle; a small black spot at upper

caudal-fin base (Fig. 48).

Similar species. Dark males of Gobius niger and G. paganellus.

Habitat. Inhabits brackish estuaries or shallow sandy areas.

Geographic distribution. In the Indian Ocean known only from Kerala, southwestern India (based on the

specimen CAS 29748 identified by H. Larson), not recorded in the Red Sea. Also reported from the Gulf of Thailand,

western Indonesia, Vietnam, and China in the western Pacific Ocean. Mediterranean records of the species are based

on specimens collected from Port Said and Damietta, Egypt (Akel & Rizkalla 2017).

FIGURE 48. Freshly dead coloration of Aulopareia unicolor. Photo by Z. Jaafar, Lake Songkla, Thailand.

Buenia affinis Iljin, 1930 (Fig. 41)—De Buen’s Goby

Buenia affinis Iljin, 1930: 51, Figs. 33–34; no type locality stated.

Size. Maximum known size 3.8 cm total length (Kovačić 2002b).

Morphology. D VI + I,8; A I,7; P 15–16 (Kovačić 2002b). Small goby with subcylindrical body, laterally

compressed towards caudal fin. Head slightly depressed and fairly long. Eyes large and close together, with narrow

interorbital space. Caudal peduncle slender, clearly lower than body depth. Dorsal fins of similar height, first dorsal

fin more or less with rounded edge with no elongate spines. Caudal fin rounded. Scales visible on upper body only

by reticulate pattern of pigmentation along scale edges.

Live coloration. Body dorsally mainly brown or beige with 4–5 well-defined, usually dark-edged, pale dorsal

saddles, each terminating with a midlateral dark blotch; ventral half of body abruptly white with 10–12 narrow

brownish bars (Fig. 41). The cheek below orbit with darkened rectangular area formed by many small close-set

dark dots. Typically 2 parallel dark dashes between eye and upper lip (Fig. 41a): lower one broad and always

visible, the upper one thin and sometimes hard to see (Fig. 41b). Iris and eye rim dark gray to dark brown,

often with well-defined light gray dots (Fig. 41). First dorsal fin very lightly pigmented, sometimes appearing

transparent. When visible, fin with 2 white longitudinal stripes alternating with 3 brown-yellow bands; sometimes

small blackish dot in the posterior fin corner. No specific sexual dichromatism reported (Kovačić 2002b).

Similar species. Buenia massutii, Pomatoschistus adriaticus, P. bathi, P. quagga.

Habitat. Infralittoral and circalittoral species, known from 1–48 m depth on coarse and medium grain sand

bottoms (Kovačić 2002b; Kovačić et al. 2018).

Geographic distribution. Mediterranean Sea. Recorded from the Balearic Islands (Kovačić et al. 2018), France

(Kovačić & Patzner 2009; Duhau et al. 2019; P. Louisy & P. Renoult, unpublished observations), Naples (Sanzo

1911), Croatia in the Kvarner area (Kovačić 2002b) and Mljet Island (Kovačić et al. 2012b), Turkey, from Yeni

Foça-Çakmaklý Island, Izmir in the Aegean Sea and Erdek-Horkos Island in the Sea of Marmara (Engin et al.

2014).

KOVAČIĆ ET AL.

Buenia massutii Kovačić, Ordines & Schliewen, 2017 (Fig. 49)—Massuti’s Goby

Buenia massutii Kovačić, Ordines & Schliewen, 2017a; type locality: western Mediterranean Sea, Spain, Balearic Islands,

Mallorca.

Size. Maximum size 4.1 cm total length (Ordines et al. 2019a).

Morphology. D VI + I,7; A I,6–7; P 16–17 (Kovačić et al. 2017a; Ordines et al. 2019a, 2019b). Small goby with

subcylindrical body, laterally compressed towards caudal fin. Head slightly depressed and fairly long. Eyes large

and close together, with narrow interorbital space. Caudal peduncle slender, clearly lower than body height. Second

spine of first dorsal fin elongate in males, reaching to or behind posterior end of the second dorsal fin when

folded down. Caudal fin rounded. Scales present, visibility on live specimens unknown.

Freshly dead coloration. Head and body yellowish white to dusky white with scattered yellow, orange and

dark brown markings. Body with 5 black marks, the first one over abdomen below first dorsal fin, three short

vertical black marks at and below lateral midline and fifth longer vertical mark on caudal-fin base (Fig. 49a).

Eyes grayish to brownish, occasionally with small pale gray dots. A rectangular area of small close-set dark dots

below eye.

FIGURE 49. A) Freshly dead coloration of Buenia massutii, photo by M. Kovačić, Mallorca, Balearic Islands, Spain. B) Buenia

cf. massutii live coloration in its natural habitat, photo by X. Rufray, Saint-Raphaël, Southern France.

Similar species. Buenia affinis.

Habitat. Circalittoral, on Peyssonnelia and maerl red algae beds, coarse sand and deep-sea oyster shell remains

bottoms from around 50 m to 116 m depth (Kovačić et al. 2017a; Ordines et al. 2019a, 2019b). A small individual

photographed at 41 m depth on detritic sand (Fig. 49B) might be this species (Rufray et al. 2021).

Geographic distribution. Mediterranean and adjacent Atlantic Ocean: Gulf of Cádiz, Atlantic; Alboran Island,

Águilas and off Balearic Islands, Spain (Kovačić et al. 2017a; Ordines et al. 2019a, 2019b); possibly southern

France (Rufray et al. 2021).

Chromogobius quadrivittatus (Steindachner, 1863) (Fig. 24)—Banded Goby

Gobius quadrivittatus Steindachner, 1863; type locality: Adriatic Sea, Croatia, Hvar.

Size. Maximum size 6.6 cm total length (Miller 1986).

Morphology. D VI + I,8–11; A I,7–9; P 16–18 (Miller 1971). Small goby with elongate body. Head clearly

depressed (distinctly dorsoventrally flattened), head and predorsal profile horizontal (Fig. 23a). Snout prominent

and mouth opening upwards (Fig. 23a). Caudal peduncle deep, almost as high as body. Dorsal fins of similar height,

the first dorsal fin may be slightly higher in appearance, with more or less rounded shape and no elongate spines.

Caudal fin rounded.

Live coloration. Body fawn, brown or blackish with a broad pale predorsal bar extending ventrally on

pectoral-fin base (Fig. 24), in some specimens indistinct. No pale saddle at origin of the first dorsal fin, prominent

to poorly visible white saddle at origin of the second dorsal fin (Fig. 24a), while another short saddle may be present

below posterior part of that fin (Fig. 24b). Tip of lower jaw, top of snout and interorbital space grayish or white.

Head usually with reticulate pattern, including cheeks (Fig. 24a), or cheeks densely dotted (Fig. 23a) (Colombo &

Langeneck 2013; Boltachev & Karpova 2017; Trkov et al. 2019). Scales present, small, at best visible on body as

pattern of pale dots.

Similar species. Chromogobius zebratus.

Habitat. Infralittoral species, 0–2 m depth on rocky shores, in clefts, below stones, hidden in biocover, in the

Black Sea also in brackish waters (Kovačić 1997; Patzner 2021).

Geographic distribution. Mediterranean and Black Sea (Miller 1986; Ahnelt 1990; Colombo & Langeneck

2013; Boltachev & Karpova 2017). In the Mediterranean Sea, there are records from northwestern Mediterranean,

Adriatic Sea, Aegean Sea and Levant (Miller 1986).

Chromogobius zebratus (Kolombatović, 1891) (Fig. 25)—Kolombatovic’s Goby

Gobius depressus var. zebratus Kolombatović, 1891: 27; type locality: Adriatic Sea, Dalmatia.

Size. Maximum size 6.2 cm total length (Alberto & Nieto 1993).

Morphology. D VI + I,11; A I,9–10; P 15–17 (Miller 1971, Alberto & Nieto 1993). Head clearly depressed

(distinctly dorsoventrally flattened), head and predorsal profile horizontal (Fig. 23a). Snout prominent and mouth

opening upwards (Fig. 23a). Caudal peduncle deep, almost as deep as body. Dorsal fins of similar height, the first

dorsal fin may be slightly higher in appearance, with more or less rounded shape and no elongate spines. Caudal fin

rounded. Scales present, small, at best visible on body as pattern of pale dots.

Live coloration. Body pale brown with a broad, pale to dusky bar on predorsal area extending ventrally

on pectoral-fin base, and 5 or 6 faint pale saddles, distinctly to poorly visible, from origin of the first dorsal fin

to just behind second dorsal fin, usually continuing to below side as faint bars half the width of dark interspaces

(Fig. 25); cheeks pale with 2 oblique dark bands radiating downwards from eye in the shape of an inverted V

or a triangular mark (Fig. 23b); base of pectoral fin with a narrow black bar preceded by a white or yellowish bar

across fin rays (Miller 1971).

Similar species. Chromogobius quadrivittatus.

Habitat. Infralittoral species, known from 0.5–20 m depth on rocky shores. Inside boulder fields, clefts, cavities,

also under individual stones and boulders of various size (Kovačić 1997; Engin & Dalgýç 2008; Patzner 2021), and

KOVAČIĆ ET AL.

in intertidal pools (Alberto & Nieto 1993).

Geographic distribution. Northeastern Atlantic, Mediterranean and Black Sea. Northeastern Atlantic near

Gibraltar (Alberto & Nieto 1993). In the Mediterranean Sea, there are records from western Mediterranean

(Bouchereau & Tomasini 1989; Iglésias et al. 2020), Sicily (Miller 1986), Malta (Kovačić & Schembri 2019),

Adriatic Sea, Aegean Sea, Levant (Miller 1986). The species is also known from the Black Sea (Engin & Dalgýç

2008; Boltachev & Karpova 2017).

Corcyrogobius liechtensteini (Kolombatović, 1891) (Fig. 21)—Liechtenstein’s Goby

Gobius liechtensteini Kolombatović, 1891: 25; type locality: Adriatic Sea, Croatia, Korčula.

Size. Reaches 2.5 cm total length (Miller 1986).

Morphology. D VI + I,9; A I,7–9; P 15–16. A very small species with a somewhat elongated body; head not

particularly flattened. Caudal peduncle deep, almost as high as body. Caudal fin rounded. First dorsal fin pointed

(first spine longest), at least in males (Miller 1972; Miller 1986; Ahnelt et al. 1994; Herler et al. 1999a; Louisy

2015).

Live coloration. Background coloration brick red (may vary from dark reddish brown to orangish brown,

rarely yellowish) with numerous narrow, transverse, bright blue to dark blue bars (usually 10 to 15 visible

on body), becoming more irregular on top of head (Fig. 21). Head sometimes darker, especially in males. Three

continuous pale to bluish, more or less radiating bars extend from lower eye to underside of head (Fig. 20a).

When present, dark branchiostegal spots located on lower head below each opercle are diagnostic (Fig. 21).

Anterior nostril tube not contrastingly white. Dorsal fins usually bear rows of dark reddish spots that may coalesce

in slightly oblique lines (Ahnelt et al. 1994; Ahnelt & Patzner 1996; Herler et al. 1999a; Louisy 2015). Scales visible

on body only by reticulate pattern of pigmentation along scale edges.

Similar species. Odondebuenia balearica, Vanneaugobius dollfusi.

Habitat. Infralittoral and circalittoral species known from 0.5 to about 40 m depth, being most common in 3–25

m depth. It is mostly found in sciaphilous areas of bedrock habitats, mainly on the walls and ceilings of caves, cavities

or overhangs. The species has also occasionally been observed or collected below stones, in coralligenous habitats

or on coralline grounds (Patzner 1999a; Francour et al. 2010; Kovacicì et al. 2012a; Louisy 2015; Bilecenoglu 2016;

Engin et al. 2018a; Ragkousis et al. 2021).

Geographic distribution. Northern Mediterranean, known from the Balearic Islands (Ahnelt et al. 1994; Ahnelt

& Patzner 1996) to the Aegean Sea (Kovačić et al. 2011; Gerovasileiou et al. 2015; Bilecenoglu 2016; Engin et

al. 2018; Ragkousis et al. 2021), including France at Marseilles (Scsepka & Ahnelt 1999), Corsica (Francour et al.

2010), French Riviera (Menut et al. 2019), Tyrrhenian Sea, Elba (Ahnelt et al. 1998) and Adriatic Sea (Kovačić et

al. 2012b).

Coryogalops ocheticus (Norman, 1927) (Fig. 50)—Canal Goby

Gobius ocheticus Norman, 1927: 381; type locality: Egypt, Suez Canal.

Size. Reaches 7 cm total length.

Morphology. D VI + I,10–11; A I,9–10; P 16–18, the uppermost 2 rays free of membrane. Body moderately

elongate, the depth 4.3–5.5 in standard length. Head subcylindrical, with a short rounded snout. The second spine

of first dorsal fin longest, with very short filament in males. Caudal peduncle deep, but lower than body depth.

Pelvic fins united to form a disc. Caudal fin rounded, shorter than head. Scales ctenoid to below origin of first dorsal

fin, cycloid ventrally on abdomen, no scales on head, chest, predorsal and prepectoral areas. Scales visible on dead

specimens by reticulate pattern of pigmentation along scale edges, visibility on live specimens unknown.

Fresh coloration. Greenish or brownish gray, scale edges dark brown, scales on lower body outlined with

orange (Fig. 50); a series of indistinct light brown blotches along back, the first on nape, ending at origin of first

dorsal fin; side of body with a series of 7 indistinct blackish blotches; head with three blackish spots or double

spots below eye, side of head with irregular small orange spots; first dorsal fin with 2 or 3 curved oblique

crossbands; pectoral fins with 2 blackish spots at base; pelvic fins blackish (visible also on Fig. 50).

Similar species. Millerigobius macrocephalus, Zebrus zebrus, Zebrus pallaoroi.

Habitat. Inhabits shallow water, isolated hard substrate but found occasionally on sand bottom (Goren 1985).

Geographic distribution. Reported from the Red Sea: Egypt (Suez Canal, Gulf of Suez, and Ras Muhammad),

Saudi Arabia (Ras Hatiba) and Eritrea (Dahlak Archipelago). A single record in the Mediterranean Sea from Port

Said, Egypt (Norman 1927), one specimen was collected from the Bardawil lagoon, Egypt (Kovačić & Golani

2007b).

FIGURE 50. Freshly dead coloration of Coryogalops ocheticus. Photo by S.V. Bogorodsky, Gulf of Suez, Egypt, Red Sea.

Cryptocentrus caeruleopunctatus (Rüppell, 1830) (Fig. 3)—Red Sea Bluespotted Shrimpgoby

Gobius caeruleopunctatus Rüppell, 1830: 134; type locality: Red Sea, Eritrea, Massawa.

Size. Reaches 13 cm total length.

Morphology. D VI + I,10; A I,9; P 17. Body moderately elongate, the depth about 5 times in standard length.

Head subcylindrical, snout short and blunt. Eyes elevated above dorsal profile of head (Fig. 3). Caudal peduncle

deep, but lower than body depth. Spines of first dorsal fin not elongate. Second dorsal fin higher than the first

(Fig. 3). Pelvic fins joined medially, reaching anus. Caudal fin rounded, its length about equal to head length (Goren

1979). Body covered with scales; no scales on head, prepelvic area, and pectoral-fin base, but scales poorly or not

visible on live specimen photographs (Fig. 3).

Live coloration. Grayish brown, grading posteriorly to dark olive-brown with deep red blue-edged spots

subequal to pupil size on head, anterior body, and dorsal fins; dorsal half of body with 2 dark spots of about

eye size, circled with blue-green dots, one under middle of first dorsal fin and one below fourth or fifth dorsal-fin

soft ray; a series of 7 oblique pale lines with blue dots on mid and rear body; head with an irregular dark red bar,

KOVAČIĆ ET AL.

edged with blue-green, extending ventrally from eye (Fig. 3). The anal and paired fins olivaceous, with small pale

yellowish spots, those on anal fin mainly in 2 rows along base (Fig. 3).

Similar species. No similar species in the Mediterranean.

Habitat. Silty to coarse sand bottom, generally adjacent to sheltered reefs or seagrass beds, but also in calm

seaward areas from depths of 2–24 m. Shares a burrow with snapping shrimps.

Geographic distribution. Restricted to the Red Sea and Gulf of Tadjoura, Djibouti. Reported from the eastern

Mediterranean coast of Israel (Rothman & Goren 2015).

Cryptocentrus steinhardti Goren & Stern, 2021 (Fig. 46)—Steinhardt’s Shrimpgoby

Cryptocentrus steinhardti Goren & Stern, 2021: 1; type locality: eastern Mediterranean Sea, Israel, Ashdod.

Size. Reaches 8.2 cm total length.

Morphology. D VI + I,10; A I,9; P 15. Body elongate, the depth 6.5 in standard length. Head subcylindrical,

snout short and blunt. Eyes elevated above dorsal profile of head (Fig. 46). Caudal peduncle slender, clearly lower

than body height. Third and fourth spines of first dorsal fin longest, elongate. Second dorsal fin subequal in

height to first fin. Pelvic fins joined medially, reaching anus. Caudal fin rounded, clearly longer than head length

(Fig. 46). Body covered with scales; no scales on head and pectoral-fin base, but scales not visible or poorly visible

on live specimen photographs.

Live coloration. Body whitish or yellowish with three slightly oblique brown bars (Fig. 46), the first below

first dorsal fin, second and third below anterior and posterior part of second dorsal fin respectively, large irregular

brown spot or bar at caudal-fin base, and scattered brown spots about pupil size in pale interspaces; predorsal area,

nape and dorsal part of opercle with small irregular dark brown blotches.

Similar species. Oxyurichthys petersii.

Habitat. In the Mediterranean, inhabits soft substrata of open sand areas at depths of 11–80 m; observed to share

a burrow with snapping shrimps (Goren & Stern 2021; P. Louisy unpublished observations).

Geographic distribution. At present is known only from three specimens caught in the eastern Mediterranean off

the southern Israeli coast and underwater photographs taken in southern Turkey at Kas (Goren & Stern 2021; present

study), recently was photographed in the Red Sea at Eilat, Israel (Bogorodsky & Goren, in prep.).

Crystallogobius linearis (Düben, 1845) (Fig. 32)—Crystal Goby

Gobius linearis Düben, 1845: 111; type locality: eastern North Atlantic Sea, Norway, Bergen.

Size. Maximum known size of male 4.7 cm and of female 3.9 cm total length (Miller 1986).

Morphology. DII–III in male, 0 or rudimentary in female + I,18–20; A I,20–21; P 15–16 (Miller 1986).

Pedomorphic habitus. Body laterally compressed, slender compared to other gobies. Eyes lateral, interorbital wide.

Mouth superior, oblique, tip of lower jaw at the horizontal level of pupil. Caudal peduncle almost as deep as body.

First dorsal fin reduced or absent. If present, distant from the second fin with a broad interdorsal space. Second

dorsal fin low, long-based and with subequal ray lengths. Pelvic disc complete in male, reduced or lacking in

female. Caudal fin slightly emarginate. Scales absent.

Live coloration. Body transparent to translucent white; a row of melanophores along anal-fin base; side

of abdomen with the longitudinal row of 3–4 well defined tiny black spots; no triangular black spot at the base

of caudal fin (Fig. 32).

Similar species. Aphia minuta.

Habitat. Nectonic, coastal to offshore in 400 m, over shell, sand, mud (Miller 1986).

Geographic distribution. Northeastern Atlantic, from the Strait of Gibraltar to the coasts of Norway, as well as

in the Mediterranean, from the Alboran Sea to northeastern Aegean Sea including the Adriatic Sea (Miller 1986).

Deltentosteus collonianus (Risso, 1820) (Fig. 51)—Toothed Goby

Gobius collonianus Risso, 1820: 252; type locality: northwestern Mediterranean Sea, France, Nice.

Size. Maximum size 7 cm total length (Miller 1986).

Morphology. D VI + I,10–11; A I,10–11; P 17 (Miller 1986; Kovačić & Golani 2007b). Moderately small goby

with subcylindrical body, laterally compressed towards caudal fin, head slightly depressed and moderately large.

Snout large and pointed. Eyes lateral, interorbital moderately wide. Mouth superior, oblique, tip of lower jaw at

FIGURE 51. Live coloration in its natural habitat of Deltentosteus collonianus. A) photo by S. Guerrieri, Tuscany, Italy.

B) Breeding male (the rounded, notably enlarged first dorsal fin, has been injured), photo by S. Le Bris, Saména, Southern

France.

KOVAČIĆ ET AL.

horizontal level of pupil. Angle of jaws below rear half of orbit. Predorsal area and nape scaled. Caudal peduncle

slender, lower than body. The first dorsal fin higher than the second dorsal fin, no elongated spines, but breeding

males with notably enlarged, sail-shaped, first dorsal fin (Fig. 51b) (Louisy 2015; Le Bris et al. 2017; 2019a).

Caudal fin truncate to slightly emarginate.

Live coloration. Body dorsally mainly brown or beige with 5 pale dorsal saddles, each above a midlateral

dark blotch including one at the caudal-fin base (Fig. 51). The saddles on the back generally conspicuous and

usually with black anterior and posterior outlines. The midlateral dark blotches of irregular rhomboid to

rounded shape, anteriorly about 1/3 of body height. Black mark on caudal-fin base triangular and larger than

other midlateral marks. Ventral half of body whitish. The first dorsal fin without dark markings in females;

one or two black spots on rear margin in males (Le Bris et al. 2017; 2019a) (Fig. 51).

Similar species. Deltentosteus quadrimaculatus, Pomatoschistus adriaticus.

Habitat. Infralittoral to circalittoral species, known from 1.5–120 m on soft bottoms and seagrass meadows

(Miller 1986; Le Bris et al. 2017; Iglésias et al. 2021b; Patzner 2021).

Geographic distribution. Mediterranean and adjacent Atlantic Ocean. D. collonianus is known from the eastern

Atlantic coast of Portugal (Miller 1986). In the Mediterranean, it is known from the western basin to the Adriatic

Sea (Miller 1986). It was later reported from Levant (Kovačić & Golani 2007b).

Deltentosteus quadrimaculatus (Valenciennes, 1837) (Fig. 52)—Four-spotted Goby

Gobius quadrimaculatus Valenciennes in Cuvier & Valenciennes, 1837: 44; type locality: northwestern Mediterranean Sea,

France, Nice.

Size. Maximum size 8 cm total length (Miller 1986).

Morphology. D VI + I,8–9; A I,8–9; P 17–18 (Miller 1986). Moderately small goby with subcylindrical body,

laterally compressed towards caudal fin, head slightly depressed and moderately large. Snout large, gently sloping.

Eyes large. Mouth superior, oblique, tip of lower jaw at the horizontal level of eye. Angle of jaws below anterior

half of orbit. Predorsal area and nape scaled. Caudal peduncle slender, lower than body depth. The first dorsal fin

higher than the second, but with no elongated spines, except in adult males where the second spine is elongated and

filamentous (Miller 1986). Caudal fin rounded to truncate.

Live coloration. Body mainly brown or beige dorsally with 5 pale dorsal saddles, faint with no black outlines,

each terminating with a midlateral dark blotch. These midlateral dark blotches of irregular shape, less than 1/3

of body height (Fig. 52). Caudal-fin base mark T-shaped and higher than other midlateral marks, covering

about entire caudal peduncle height (Fig. 52). Ventral half of body whitish. Females may display a pale blue dot

on nape just behind eyes, partly circled in brown. The first dorsal fin with a black spot distally between fifth and

sixth spines (Fig. 52a), and with three pronounced dark streaks along first dorsal spine in males (Fig. 52b) (Miller

& Loates 1997; Louisy 2015; Pillon et al. 2016a).

Similar species. Deltentosteus collonianus, Pomatoschistus adriaticus, P. marmoratus, P. minutus.

Habitat. Infralittoral to circalittoral species, known from 2–150 m on soft bottoms, mostly sand, also on mud

(Louisy 2015; Pillon et al. 2016a; Patzner 2021).

Geographic distribution. Mediterranean and adjacent Atlantic Ocean: in the Mediterranean, from Gibraltar

along northern coast to the Levant; in the Atlantic, from Galicia to Morocco (Miller 1986).

Didogobius schlieweni Miller, 1993 (Fig. 7)—Andromeda Goby

Didogobius schlieweni Miller, 1993: 1413, type locality: Adriatic Sea, Croatia, Unije Island, near Cres Island, Croatia.

Size. Maximum size about 4.6 cm total length (Ballesta et al. 1998).

Morphology. D VI + I,12–13; A I,11; P 17 (Miller 1993; Ballesta et al. 1998). Small goby with elongate body,

head depressed and dorsally flattened. Interdorsal space membranous. Caudal peduncle deep, almost as deep

as body. Dorsal fins of similar height. The first dorsal fin with more or less rounded margin (second spine the longest

but not elongate). Caudal fin rounded. Scales present, small, at best visible on body as pattern of paler dots.

FIGURE 52. Live coloration in its natural habitat of Deltentosteus quadrimaculatus. A) Photo by R. Pillon, Maddalena, Sardinia,

Italy. B) Male with enlarged, pointed first dorsal fin, photo by L. Bérenger, Cagnes-sur-Mer, Southern France.

Live coloration. Head and body dark brown to purplish with a longitudinal series of 3–6 definite white

blotches as large as or larger than eye diameter on dorsal half of body and white transverse band in predorsal

area; scattered white dots in variable amount on head and sometimes on body; median fins black, outer margin

of dorsal fins and posterior margin of caudal fin with a broad white band (Fig. 7) (Ballesta et al. 1998; Louisy

2015; Le Bris et al. 2019b, 2021).

Similar species. Chromogobius quadrivittatus.

Habitat. Infralittoral species. Only observed at night among layers of pebbles in rocky gullies or in stony slopes

at depths of 1.5–14 m; also observed in Posidonia meadows (Miller 1993; Francour 2008; Kampouris et al. 2019;

Le Bris et al. 2019b, 2021; Patzner 2021).

KOVAČIĆ ET AL.

Geographic distribution. Mediterranean, with records from the northwestern Mediterranean (Ballesta et al.

1998; Francour 2008; Iglésias et al. 2021b), Adriatic Sea (Miller 1993; Kovačić 2005), and Greece, Aegean Sea

(Kampouris et al. 2019).

Didogobius splechtnai Ahnelt & Patzner, 1995 (Fig. 14)—Splechtna’s Goby

Didogobius splechtnai Ahnelt & Patzner, 1995: 97; type locality: western Mediterranean Sea, Spain, Balearic Islands, Ibiza

Island, Near Portinatx.

Size. Maximum published size 2.8 cm total length (Ahnelt & Patzner 1995).

Morphology. D VI + I,9–10; A I,9; P 13–16 (Ahnelt & Patzner 1995; Francour 2008). Small goby with stocky

body, large depressed head and a gently sloping snout. Caudal peduncle deep, almost as deep as body. The first

dorsal fin higher than the second, its first spine longest, giving the fin a triangular shape. Caudal fin rounded.

Scales visible on body from reticulate pigmentation pattern along scale edges.

Live coloration. Body mostly dark brown with 3 moderately broad whitish to pale grayish bars extending

onto ventral half of body, the first bar immediately in front of the first dorsal fin and extending onto ventral

half of pectoral fins, plus a very thin one at the base of caudal fin (Fig. 14). First dorsal fin colored like adjacent

body, with scattered white flecks and streaks, outer membrane between first two spines white (Louisy 2015, Patzner

2021).

Similar species. Gammogobius steinitzi.

Habitat. Infralittoral to circalittoral species, known from depth of 4–56 m. Present in small cavities in innermost

parts of large marine caves and in small bottom caves and cavities. Mostly found lying on the soft sediment in

recesses, ready to dive into a nearby hole or crevice (summarized by Francour 2008).

Geographic distribution. Mediterranean, there are records from the northwestern Mediterranean (Ahnelt &

Patzner 1995; Francour 2008; Iglésias et al. 2021b), Lampedusa (Stefanni 1999) and Adriatic Sea (Herler & Patzner

2002; Kovačić 2005). A record from Turkey, Aegean Sea (Francour et al. 2007), seems probable but was not

substantiated by specimens or photos, later confirmed by Ragkousis et al. (2021) for Aegean Sea.

Gammogobius steinitzi Bath, 1971 (Fig. 22)—Steinitz’s Goby

Gammogobius steinitzi Bath, 1971: 202, type locality: Mediterranean Sea, France, off Marseilles.

Size. Maximum size about 5.4 cm total length (Kovtun & Manilo 2013).

Morphology. D VI + I,8–9; A I,7–9; P 15–17 (summarized in Kovtun & Manilo 2013). Small goby with stocky

body, large depressed head and a gently sloping snout. Eyes large, with a narrow interorbital space (Fig. 22).

Caudal peduncle deep, almost as deep as the body. Dorsal fins of similar height, the first dorsal fin with more or less

rounded shape and no elongated spines, all but the sixth spines being subequal in length. Caudal fin rounded. Scales

visible on body from reticulate pigmentation pattern along scale edges.

Live coloration. Body with barred coloration pattern of 6–7 narrow pale bars on a mostly brownish

background (Fig. 22). Head brownish with 3 pale, continuous bars extending downwards from eye over cheek

and preopercle to ventral side (Fig. 20b). Fins mostly colorless; some black spots at the base of both dorsal fins,

extending as dots along rays or spines (Fig. 22).

Similar species. Corcyrogobius liechtensteini, Didogobius splechtnai.

Habitat. Infralittoral species, known from 5–25 m in marine caves, more common in deeper water. Mostly

inhabits the crevices and holes of cave walls and ceilings (summarized by Patzner 2021).

Geographic distribution. Mediterranean and Black Sea. Mediterranean records from the northwestern

Mediterranean (Bath 1971), Adriatic Sea (Kovačić 1999), Crete (Kovačić et al. 2011), and Turkey, Aegean Sea

(Engin et al. 2018a; Ragkousis et al. 2021). In the Black Sea, it was recorded in Crimea (Kovtun & Manilo 2013).

Gobius auratus Risso, 1810 (Figs. 36 and 40)—Golden Goby

Gobius auratus Risso, 1810: 160, type locality: Mediterranean Sea, France, La Fourmigue.

Size. Known adult size about 9 cm total length.

Morphology. D VI + I,13–15; A I,12–14; P 17–20. Medium-sized goby, with a moderately elongate body,

laterally compressed, large head, and a short, slightly pointed snout. Caudal peduncle deep, but lower than body

depth. Dorsal fins of similar height, the first dorsal-fin spines decreasing posteriorly, giving fin a triangular shape.

Caudal fin rounded. Scales present on body and on predorsal area, usually visible on photographs at least dorsally

(Herler et al. 2005; Renoult et al. 2022).

Live coloration. Body plain yellow (G. auratus morph 1) (Fig. 40), except in northern Adriatic (Herler et al.

2005) where the yellow body is covered with numerous orange or red-brown dots forming longitudinal rows,

including 3 or 4 rows below the lateral midline (G. auratus morph 2) (Fig. 36). Upper lip unmarked except the

dot on the corner of the mouth (Fig. 33c) (Renoult et al. 2022).

Similar species. Morph 1 with no similar species in the Mediterranean. For morph 2: Gobius fallax and G.

xanthocephalus.

Habitat. Infralittoral and circalittoral species known from 8–80 m depth on rocky and coralligenous habitats

(Herler & Patzner 2005; Patzner 2021).

Geographic distribution. Northern Mediterranean, presently known from the Provence basin in France westwards

(Herler et al. 2005) to west of Antalya, Turkey, Levantine Sea (Francour et al. 2007). In the Adriatic, a transition

form between the ‘dotted’ morph (morph 2) and the ‘plain colored’ morph (morph 1) can be found in the Zadar

region in Croatia (R. Pillon and M. Kovačić, unpublished observation).

Gobius bucchichi Steindachner, 1870 (Fig. 34)—Bucchich’s Goby

Gobius bucchichi Steindachner, 1870: 627, type locality: Adriatic Sea, Croatia, Hvar.

Size. Known adult size about 9 cm total length.

Morphology. D VI + I,14; A I,13–14; P 17–18. Medium-sized goby with moderately long body, and large head

with short, slightly acute and oblique snout. Caudal peduncle deep, but lower than body depth. Dorsal fins of similar

height, the first dorsal fin may be slightly taller, with more or less rounded margin and no elongate spines. Caudal

fin rounded. Scales present on body and on predorsal area, usually visible on photographs, at least dorsally (Kovačić

& Šanda 2016).

Live coloration. Body pale gray, pale yellowish, or yellowish brown with pattern of brown dots forming

longitudinal lines. Dots are most marked along the lateral midline but always remain separated and generally

form a continuous row with no marked spaces (Fig. 34). Eyes with a conspicuous horizontal dark brown stripe

through the center, narrowly continuing on postorbital part of head as series of small close-set spots, an irregular

longitudinal marking along the upper iris, a variable number of additional markings in orbit’s upper half but

unmarked in ventral half, except sometimes with a small dot in its anterior portion. Snout with dark dashes

forming M-shaped line. Usually it has only 2 longitudinal rows of dots on the cheek with a large unmarked

area in the middle (Fig. 33a). Rarely, a few dots are visible also in the middle cheek but usually limited to the

anterior part of the cheek, very close together, well aligned and not parallel to the lower row. Posterior angle of

mouth without a well-defined dark dot (Fig. 33a), so the lower row of dots, which runs along the ventral edge of

the cheek, starts with 2 dashes never preceded by a marked dot on the corner of the mouth (Kovačić & Šanda 2016;

Renoult et al. 2022).

Similar species. Gobius incognitus, G. fallax.

Habitat. Infralittoral species, known from 1–8 m depth from all kinds of rocky substrata: gravel, cobbles,

boulders and bedrock, but on sand only when it is mixed with a hard substratum, never on pure sand. The substratum

was bare or covered with short thallus algae (Kovačić & Šanda 2016).

Geographic distribution. Presently known from the eastern coast of the Adriatic Sea, it is common from the Gulf

of Trieste (Tiralongo & Pillon 2020) to southern Croatia (Kovačić & Šanda 2016); there is only one record in the

Ionian Sea from Butrint, southern Albania (Kovačić & Šanda 2016), and a record in the Aegean Sea from Kondyli

KOVAČIĆ ET AL.

beach, eastern Peloponnese (Tiralongo & Pillon 2020). Population from the Black Sea more similar in pattern

of coloration to G. bucchichi than to G. incognitus in absence of black dot at the corner of mouth and any mark

ventrally on iris (Vasil’eva 2007; Boltachev & Karpova 2017), and requires additional study.

Gobius cobitis Pallas, 1814 (Fig. 53)—Giant Goby

Gobius cobitis Pallas, 1814: 160, type locality: Black Sea, Ukraine, Crimea, Feodosiya [Feodosia].

Size. Known adult size about 20–25 cm total length.

Morphology. D VI + I,13–14; A I,10–12; P 19–22. Large goby with a robust body, a large head, long snout and

thick lips. Caudal peduncle deep, but lower than body depth. Dorsal fins of similar height, the first dorsal fin with

more or less rounded edge and no elongated spines. Pectoral fins with well-developed, short free rays. Caudal fin

rounded. Predorsal area and nape scaled. Scales small, visible on body and predorsal area as pattern of pale dots.

Live coloration. Head and body variegated, finely mottled greenish to beige, with fairly invariable pattern

of large and irregular dark midlateral blotches (Fig. 53). Dorsally three dark blotches at the posterior base of

the dorsal fins and a smaller blotch just before the caudal peduncle on the back, and three fainter (sometimes not

visible) blotches located in the center of the nape, over the pectorals, and just behind origin of the second dorsal fin,

respectively. Along midline: 5 medium-sized dark blotches. Below midline: 9-10 smaller, aligned dark blotches.

Often a series of roundish white spots on lower side of head (Fig. 53): one on lower lip, one at the angle of mouth,

3–5 along lower head reaching posterior border of opercle, and sometimes some smaller ones along opercular edge

(Renoult et al. 2022).

Similar species. Gobius ophiocephalus, G. paganellus.

Habitat. Infralittoral and intertidal species, known from 0.2–10 m depth on hard or mixed bottoms and in

lagoons with limited variation in salinity, except for the ocassional record from Israel (Kovačić & Golani 2007b,

Patzner 2021).

FIGURE 53. Live coloration in its natural habitat of Gobius cobitis. Photo by T. Menut, Thau lagoon, Southern France.

Geographic distribution. Mediterranean, Eastern Atlantic, and Black Sea (Vasil’eva 2007). This species has

migrated to the Gulf of Suez, in the Red Sea (Goren & Klausewitz 1978).

Gobius couchi Miller & El-Tawil, 1974 (Fig. 43)—Couch’s Goby

Gobius couchi Miller & El-Tawil, 1974: 546, type locality: England, English Channel, Cornwall, Helford.

Size. Known adult size about 7–9 cm total length.

Morphology. D V–VI + I,12–14; A I,11–13; P 15–18. Relatively small goby. Snout short and with a steep

profile. Caudal peduncle deep, but lower than body depth. Dorsal fins of similar height; first dorsal fin with a

relatively rounded shape, the dorsal spines often slightly prolonged (membranes slightly incised between the

spines); in breeding males, both dorsal fins have prolonged free tips on spines and rays, and deeply incised

interradial membranes (Fig. 42). Caudal fin rounded. Scales present on body and on predorsal area, usually visible

on photographs, at least on body (Renoult et al. 2022).

Live coloration. Body gray, light brown or olive with mottled or reticulated pattern (Fig. 43). Lateral

midline is always darker, formed by a longitudinal series of small black spots. One black spot on the corner of the

mouth, usually followed by 2 more or less rectangular dark spots on lower cheek (Fig. 42).

Similar species. Gobius niger.

Habitat. Infralittoral species, known from 1–20 m depth on soft bottoms with algae or marine plants and

shelters such as rocks and holes. In the Mediterranean, it is generally found in sheltered bays, often associated with

Cymodocea nodosa (Pillon et al. 2016b, 2019; Patzner 2021).

Geographic distribution. Atlantic, Mediterranean and Black Sea. In the Atlantic, known from Ireland, the south

of Great Britain and northern Brittany, France (Miller & El-Tawil 1974; Baldock & Kay 2012; Iglésias et al. 2020).

In the Mediterranean Sea, recorded from Cadaquès, Spain (Renoult et al. 2021a), Port-Vendres and Saint-Mandrier,

France (Louisy 2005; Iglésias et al. 2021b), Liguria, Italy (Liu et al. 2009a), Ischia Island, Italy (Stefanni & Mazzoldi

1999), Malta (Kovačić et al. 2013), Croatia (Kovačić 2001; Kovačić et al. 2012b), Corfu Island, Greece (Šanda &

Kovačić 2009), Crete (Kovačić et al. 2011), Cyprus (Kovačić & Golani 2007a) and southern part of the Dardanelles

Strait, Turkey (Özen et al. 2009). In the Black Sea, from Crimea (Karpova & Boltachev 2018).

Gobius cruentatus Gmelin, 1789 (Fig. 15)—Red-mouthed Goby

Gobius cruentatus Gmelin, 1789: 30, type locality: western Mediterranean Sea, France, Marseilles.

Size. Known adult size about 13–15 cm total length.

Morphology. D VI + I,14; A I,12–13; P 20–21. Large goby with stout body, moderately long snout with

moderately steep profile, thick lips. Caudal peduncle deep, but lower than body depth. First dorsal fin higher than

the second dorsal fin; first dorsal fin spines decreasing posteriorly, shaping a curved margin. Caudal fin rounded.

Predorsal area and nape scaled. Scales small, visible on body and predorsal area as dense pattern.

Live coloration. Body mostly mottled brown, with large darker, squarish midlateral blotches, each one

preceded and followed by a white patch (Fig. 15). Back with 4 broad dark transverse bands, the second and

fourth the darkest, separated by thin white saddles (Fig. 15). Lips mostly vermilion, with a white bar below eye,

at angle of mouth (Fig. 15) (red color usually not detectable under natural light only). Membranes of both dorsal

fins are highly variegated, mixing irregular patches of red, dark brown and white coloration. Rows of head sensory

papillae well-visible and bicolored black and white.

Similar species. No similar species in the Mediterranean. However, with no artificial light, may be confused

with Gobius paganellus or G. niger.

Habitat. Infralittoral species, known from 1–40 m depth on inshore rocky habitats, sand with stones and boulders,

and in seagrass meadows (Miller 1986; Louisy 2015; Patzner 2021).

Geographic distribution. Northeastern Atlantic, Mediterranean and Black Sea (Engin et al. 2007; Vasil’eva

2007). In the Atlantic, occurs from the counties of Cork and Kerry in Ireland to Western Sahara (Miller 1986). In the

Mediterranean, all along the northern coast, and along the southern coast in Morocco, Algeria and Tunisia (Miller

1986), as well as in Benghazi in Libya (Al-Hassan & El-Silini 1999).

KOVAČIĆ ET AL.

Gobius fallax Sarato, 1889 (Fig. 39)—Sarato’s Goby

Gobius fallax Sarato, 1889: 506, type locality: northwestern Mediterranean Sea, France, Nice.

Size. Known adult size about 9 cm total length.

Morphology. D VI + I,14–16; A I,13–15; P 18. Medium-sized goby with moderately elongate body, large head,

snout moderately steep and shorter than eye diameter. Body often held slightly arched. Caudal peduncle deep,

but lower than body depth. Dorsal fins of similar height, first dorsal fin may be slightly higher, with more or less

rounded margin and no elongate spines. Caudal fin rounded. Scales present on body and on predorsal area, usually

visible on photographs on body.

Live coloration. Body light gray or whitish, usually with blue or yellow reflections on the back (Fig. 39).

Yellow hues sometimes more visible on the head. Head and body covered with longitudinal lines of well-defined

brown or dark brown dots, those along midline slightly paler (Fig. 39). Dots are broadest along the lateral

midline but always remain separated. The eyes have brown stripes radiating from the pupil to the orbital rim, 2 of

them forming a broad horizontal stripe across the pupil. Dark curved mark on the upper iris on the eye connects

longitudinally other marks (Fig. 39b). Lower lip with a median W-shaped mark. A well-defined dark dot on

posterior angle of mouth (Fig. 39b). Upper lip with brown marks in addition to the dot on the corner of the mouth

(Fig. 39b). Snout with M-shaped line and an oblique preorbital bar (Herler et al. 2005; Renoult et al. 2022).

Similar species. Gobius bucchichi, G. incognitus, G. xanthocephalus, G. auratus morph 2.

Habitat. Infralittoral species, known from 1–32 m depth on mixed sandy-rocky habitats, rock crevices or stones

nearby soft bottom, sometimes covered with seagrass (Herler & Patzner 2005; Louisy 2015).

Geographic distribution. Mediterranean. Recorded from many places from the Balearic Islands to Turkey (Fricke

et al. 2007), including Croatia (Kovačić & Patzner 2011), Crete (Kovačić et al. 2011), Cyprus (Kovačić & Golani

2007a) and Libya (Al-Hassan & El-Silini 1999). Also recorded in Gran Canaria, Atlantic (Dooley et al. 1985), but

this needs to be confirmed because of taxonomic progress on the Gobius auratus complex since this record.

Gobius gasteveni Miller, 1974 (Fig. 54)—Steven’s Goby

Gobius gasteveni Miller, 1974: 468, type locality: England, English Channel, Plymouth.

Size. Known adult size about 10–12 cm total length.

Morphology. D V–VI + I,12–15; A I,11–14; P 18–22. Medium-sized goby with a moderately long body, and

a short snout with a steep profile. Caudal peduncle deep, but lower than body. Dorsal fins of similar height, the

first may be slightly higher, with a more or less trapezoid shape, its first spine no shorter than the others. Caudal

fin rounded. Scales present on body and on predorsal area, usually visible on photographs at least on body (Miller

1974, 1986).

Live coloration. Body light gray to light pink with gold and greenish reflections on the back. Lateral

midline with 6 to 11 rectangular, orange to dark brown blotches, some of them usually separated by isolated dark

dots surrounded with white (Fig. 54). Lips white, or light gray or gold with white spots. Snout with a well-defined

V-shaped line. Cheeks pale or shaded, and then often with a gold sheen and round white spots. An oculoscapular

orange stripe starts behind eye and continues above upper pectoral-fin base into an irregular dorsolateral

body stripe until top of caudal peduncle (Fig. 54a). Dorsal fins with white spots and white margin in some males,

possibly only in Atlantic population. Breeding males dark (Fig. 54b) (Alberto et al. 1999; Renoult et al. 2022).

Similar species. Gobius roulei, G. geniporus.

Habitat. Infralittoral and circalittoral species, known from 5–120 m depth on soft bottoms with mud, sand, silt,

shell or small-stone deposits (Miller 1986; Patzner 2021).

Geographic distribution. Eastern Atlantic from Great Britain (Miller 1974) to Madeira and Canary Islands

(Miller 1984) and in the western Mediterranean: Alboran and Balearic Islands (Ahnelt & Dorda 2004), southern

France (J. Renoult unpublished observation), Gulf of Genoa in Italy (Ahnelt et al. 2011), Malta (Kovačić & Schembri

2019) and Lebanon (Bitar & Badreddine 2021).

FIGURE 54. Live coloration in its natural habitat of Gobius gasteveni. A) common coloration, photo by B. Guichard, Brest,

northwestern France. B) dark coloration, photo by L.S. Tocino, Granada, Southern Spain.

Gobius geniporus Valenciennes, 1837 (Fig. 55)—Slender Goby

Gobius geniporus Valenciennes in Cuvier & Valenciennes, 1837: 32, type locality: Mediterranean Sea, Italy.

Size. Known adult size about 14–16 cm total length.

Morphology. D VI + I,12–14; A I,11–12; P 17–19. Large goby with a long body, moderately long snout with

a shallow profile. Caudal peduncle deep, but lower than body depth. Dorsal fins of similar height, the first dorsal

KOVAČIĆ ET AL.

fin with more or less rounded edge and no elongated spines. Caudal fin rounded. Scales small, visible on body and

predorsal area as dense pattern. Predorsal area and nape scaled (Miller 1986).

Live coloration. Body ground coloration whitish with beige to pale brown speckling with a longitudinal

series of 5 rectangular dark blotches alternating with smaller, paler and less rectangular brown markings

along lateral midline (Fig. 55). Body below the lateral midline white with diffuse brown speckles, or with a

longitudinal series of small brown blotches (Fig. 55). Usually a large dark or shaded area below eye, often

containing 2 dark bars. Some specimens with dorsal fins intensively patterned with large brown to reddish spots,

sometimes coalescing to form longitudinal bands, and terminated by a white margin. Caudal fin with brown spots.

Head sensory papillae black and visible (Renoult et al. 2022).

Similar species. Gobius gasteveni, G. roulei.

Habitat. Infralittoral species, known from 1–40 m depth on sandy or gravel bottoms nearby rock beds or seagrass

(Louisy 2015; Patzner 2021).

Geographic distribution. Mediterranean, including the Sea of Marmara and Cyprus (Miller 1986; Kovačić &

Golani 2007a).

FIGURE 55. Live coloration in its natural habitat of Gobius geniporus. Photo by S.V. Bogorodsky, Turkey, Aegean Sea.

Gobius incognitus Kovačić & Šanda, 2016 (Fig. 37)—Incognito Goby

Gobius incognitus Kovačić & Šanda, 2016: 1106, type locality: northwestern Mediterranean, France, Gulf of Lion, Banyuls-

sur-Mer, Paulliles beach.

Size. Known adult size about 9–10 cm total length.

Morphology. D VI + I,13–14; A I,12–13; P 19–20. Medium-sized goby with moderately long body, large head,

short snout. Caudal peduncle deep, but lower than body. Dorsal fins of similar height, the first dorsal fin with more

or less rounded edge and no elongated spines.

Caudal fin rounded. Scales small, visible on body and usually on predorsal area as dense pattern. Predorsal area

and nape scaled (Kovačić & Šanda 2016).

Live coloration. Body uniformly pale gray, greenish gray or light brown, covered with longitudinal lines

of brown dots (Fig. 37A). The dots are broadest, darkest and most numerous along the lateral midline, where they

coalesce in well-aligned midlateral dashes or blotches separated by white spaces. Eyes with a conspicuous

horizontal stripe through the center, upper iris with separate brown dots or radiating stripes without a

longitudinal mark uniting them (Fig. 37b). V-shaped line in the center of the snout. Three longitudinal rows

of dots recognizable on cheek or in irregular pattern (Fig. 33d); the median row more or less regularly crosses

the center of the cheek, parallel to the lower row. There is a well-defined dark dot at posterior angle of mouth, so

the lower row begins with an isolated dot at the corner of the mouth, usually followed by 2 short horizontal dashes.

Upper lip with brown marks in addition to the dot on the corner of the mouth (Fig. 37b) (Kovačić & Šanda

2016; Renoult et al. 2022).

Similar species. Gobius bucchichi, G. fallax.

Habitat. Infralittoral species, known from 0.5–12 m depth mostly on shallow and sub-horizontal bottoms with

sand and stones or rocks (Kovačić & Šanda 2016). At low depths it is typically associated with the sea anemone

Anemonia viridis (Tiralongo et al. 2020).

Geographic distribution. Mediterranean. Confirmed presence from the northwestern Mediterranean to the

Aegean Sea and Israel (Kovačić & Šanda 2016).

Gobius kolombatovici Kovačić & Miller, 2000 (Fig. 18)—Kolombatović’s goby

Gobius kolombatovici Kovačić & Miller, 2000: 232, type locality: Adriatic Sea, Croatia, Kamenjak, the Island of Krk.

Size. Known adult size about 11 cm total length.

Morphology. D VI + I,13–14; A I,13; P 17–19. Medium-sized goby, snout moderately short but longer than

the eye diameter, with a moderately steep profile. Caudal peduncle deep, but lower than body depth. Dorsal fins

of similar height, the first dorsal fin may be slightly higher, with more or less rounded distal edge and no elongate

spines. Caudal fin rounded. Scales present on body and on predorsal area, usually visible on photographs at least

on body (Kovačić & Miller 2000).

Live coloration. Head and body on back pale to fawn, pearly white ventrally, with irregular orange blotches

arranged in longitudinal series (Fig. 18). Snout with a conspicuous, V-shaped orange line. A broad horizontal

stripe across eye and a continuous and well-marked orange oculoscapular line from behind eye to above pectoral-fin

base. First dorsal fin with a distinct oval blackish blotch between the third and fourth (or fifth) interspinous

membranes (Fig. 18) (Kovačić & Miller 2000; Francour & Mangialajo 2007; Renoult et al. 2022).

Similar species. Thorogobius macrolepis.

Habitat. Infralittoral and circalittoral species, known from 12–90 m depth. Typically found on patches of soft

bottoms in steep rocky areas, often on coarse shell sand at the bottom of rocky or coralligenous walls, close to

shelter (Kovačić & Miller 2000; Francour & Mangialajo 2007).

Geographic distribution. Scattered records in the Mediterranean Sea: western Mediterranean (e.g., Columbretes

and Balearic Islands in Spain: Kersting & Ballesteros 2010); Ligurian (Francour & Mangialajo 2007), Adriatic

(Kovačić & Miller 2000), Ionian (Tiralongo & Pagano 2015) and Aegean seas (Bilecenoglu 2013), and Lebanon

(Bitar & Badreddine 2021).

Gobius niger Linnaeus, 1758 (Fig. 56)—Black Goby

Gobius niger Linnaeus, 1758: 262; no type locality.

Size. Known adult size about 15–18 cm total length.

Morphology. D V–VII + I,11–13; A I,10–13; P 15–20. Large goby with a relatively stocky body, proportionately

large head with a short and steep snout. Caudal peduncle deep, almost as deep as body. The first dorsal fin higher or

about equal to the second, mature males with long and pointed fin with elongated spines (third and fourth spines

longest). Pectoral-fin free rays moderately developed. Caudal fin rounded. Rows of sensory papillae well developed

and clearly visible on the nape. Scales present on body and on predorsal area, usually visible in photographs at least

on body (Miller 1986).

KOVAČIĆ ET AL.

FIGURE 56. Live coloration in its natural habitat of Gobius niger. A) Usual coloration, photo by S.V. Bogorodsky, Turkey,

Aegean Sea. B) Male with erected enlarged first dorsal fin, photo by P. Louisy, Thau Lagoon, Southern France. C) Dark male

during reproduction, photo by R. Svensen, Stavanger, Norway.

Live coloration. Body ground coloration variable, usually light cream or gray, sometimes greenish (Fig. 56a &

56b), to black in nesting males (Fig. 56c). In light-colored individuals, the body has dark mottling on the back, 5 or

6 black midlateral blotches and becomes white ventrally with few scattered black dots. Head with a white preorbital

bar and a black suborbital bar, lips mottled dark gray or brown. Oculoscapular line limited to three inconspicuous

black or brown dashes. Membrane of first dorsal fin light blue-green, or light gray, or light reddish (blackish in

nesting males) in the distal half, reddish with 3 irregular white horizontal bands in the proximal half. One black

blotch often visible distally on the first membrane of each dorsal fin or just on the first dorsal fin (Fig. 56a);

rarely a black spot is visible at the lower posterior corner of the first dorsal fin. Sensory papillae black and well

visible on the nape (Renoult et al. 2022).

Similar species. Gobius roulei, G. couchi.

Habitat. Infralittoral and circalittoral species known from 1–96 m depth, mostly on sheltered or muddy soft

bottoms and coastal lagoons (Miller 1986; Patzner 2021).

Geographic distribution. Northeastern Atlantic, from the Baltic Sea to Canary Islands and in the whole

Mediterranean (Miller 1986), the Black Sea (Tserkova et al. 2016) and the Sea of Azov (Miller 1986).

Gobius ophiocephalus Pallas, 1814 (Fig. 44)—Grass Goby

Gobius ophiocephalus Pallas, 1814: 153, type locality: Black Sea, coasts of Crimea.

Size. Known adult size about 15–20 cm, reaching 25 cm total length.

Morphology. D V–VII + I,13–16; A I,12–16; P 17–20. Body stout and distinctly compressed laterally (for

a Gobius species), especially on caudal peduncle. Proportionally small eyes. Snout moderately long and with a

moderately steep profile. Caudal peduncle deep, but lower than body depth. The first dorsal fin about equal to,

or shorter than the second dorsal fin, with more or less rounded distal edge. Dorsal-fin ray tips become free from

membrane in reproductive males (Toricelli et al. 2000). Caudal fin rounded. Scales small, visible on body and

usually also on predorsal area as pattern of pale dots. Predorsal area and nape scaled (Miller 1986).

Live coloration. Body coloration gray-green to pale fawn dorsally, whitish to yellow-green ventrally (Fig. 44).

Eight to fourteen irregular, vertical dark bars along the lateral midline, forming dark lateral zigzag markings

in combination with a series of 10 or more dark areas alternating with pale interspaces on lower side. A

diffuse, light to golden dorsolateral stripe runs from predorsal area to caudal-fin base, not always visible

(Fig. 44). It is bordered ventrally by an olive-brown stripe, and dorsally by a dark vertebral stripe. In the Adriatic,

these stripes can be interrupted by dark transverse saddles. One oblique dark preorbital bar, bordered ventrally by a

white suborbital bar. Cheek and preopercle mottled brown-olive with white spots of various sizes and irregular

shapes (Fig. 44). First dorsal fin reddish with 4 gray-blue transverse bands. Second dorsal fin with the same color

but with gray-blue irregular markings not forming clearly defined bands, and with many small white dots (Renoult

et al. 2022).

Similar species. Gobius cobitis.

Habitat. Known from 0.1–3 m depth, eurythermal and euryhyaline, on soft bottoms with marine phanerogams

in estuaries and coastal lagoons (Kara & Quignard 2019).

Geographic distribution. Mediterranean, Adriatic, Black Sea and Sea of Azov. Along the African coasts,

confirmed records from Libya and Tunisia (Hajji et al. 2013).

Remark. Often classified in its own genus Zosterisessor Whitley, 1935, but its position within the genus Gobius

was confirmed by a phylogenetic analysis (Iglésias et al. 2021a).

Gobius paganellus Linnaeus, 1758 (Fig. 57)—Rock Goby

Gobius paganellus Linnaeus, 1758: 263, type locality: Mediterranean Sea.

Size. Known adult size about 12 cm total length.

Morphology. D VI + I,12–15; A I,10–13; P 15–20. Medium-sized goby with a proportionally short and stocky

body. Head wide and moderately depressed. Caudal peduncle deep, but lower than body. Dorsal fins of similar

KOVAČIĆ ET AL.

height, the first dorsal fin may be slightly higher, with a more or less rounded distal edge and no elongate spines.

Pectoral-fin free rays very well developed. Caudal fin rounded. Predorsal area and nape entirely covered with

large scales. Scales visible on body and usually also on predorsal area (Miller 1986; Louisy 2015).

FIGURE 57. Live coloration in its natural habitat of Gobius paganellus. A) Light colored individual, photo by M. Kovačić,

Bol, island of Brač, Croatia, central Adriatic Sea. B) Dark-colored male, photo by G. Kunz, Donji Kraj, Istria, Croatia, northern

Adriatic Sea.

Live coloration. Background coloration highly variable, ranging from light beige to dark brown (Fig. 57a).

Nesting males occasionally entirely black, with just the first dorsal fin margin colored (Fig. 57b). Most frequent

pattern: broad dark bars and thin alternate white saddles on the back, medium-sized dark blotches along lateral

midline, and small dark markings on a light background below. Usually a pale to white bar between eye and

corner of mouth; it is often preceded by a dark preorbital bar. First dorsal fin usually dark with an orange to

yellow marginal band (Renoult et al. 2022).

Similar species. Gobius ater, G. cobitis, Millerigobius macrocephalus, Zebrus zebrus, Z. pallaoroi.

Habitat. Infralittoral species known from 0.5–20 m depth on very diverse hard or mixed bottoms (Patzner 2021;

P. Louisy unpublished observation).

Geographic distribution. The whole of the Mediterranean (Miller 1986), Black Sea (Engin & Seyhan 2009) and

Eastern Atlantic from Great Britain to Senegal (Miller 1986), including the Azores (Santos et al. 1997), the Canary

Islands (Wirtz 1994) and Madeira (Wirtz et al. 2008).

Gobius roulei de Buen, 1928 (Fig. 58)—Roule’s Goby

Gobius roulei de Buen, 1928: 1, figs. 1–2, type locality: western Mediterranean Sea, Spain, Baleares Islands, Mallorca, Palma

Bay, Porto Pi.

Size. Known adult size about 8–9 cm total length.

Morphology. D VI + I,12–14; A I,12; P 17–19. Medium-sized goby with a relatively slender body, proportionately

large head with a short and steep snout. Caudal peduncle deep, but lower than body. First dorsal fin high and

pointed with elongate spines in males, more often with a triangular rounded shape in females (and about height of

second dorsal fin). Pectoral-fin free rays moderately developed. Caudal fin rounded. Scales present on body, usually

visible on photographs, at least dorsally (Fig. 58). Predorsal area naked (Miller 1986). Rows of dark sensory papillae

sometimes visible on nape.

Live coloration. Background coloration whitish, grayish or pale orange. Lateral midline covered with an almost

continuous line of brown to black dots between pectoral and caudal fins; this line is slightly broadened at regular

intervals to form 5 to 6 midlateral blotches (Fig. 58). Back mottled orange to dark brown, with white or gray patches

sometimes forming narrow saddles, ventral side almost uniformly light cream, with only faint grayish markings

below the midlateral blotches (Fig. 58). Lips white, unmarked. Oculoscapular dark line well visible, beginning

with a short and broad orange to pink dash just behind eye. Dorsal-fin membranes pale, with a yellow shade on

distal half, and 2 to 3 faint, darker transverse bands on proximal half. Basal half of first spine of each dorsal fin with

alternating white and black dashes along leading edge (Fig. 58) (Renoult et al. 2022).

Similar species. Gobus niger, G. gasteveni, G. geniporus.

Habitat. Infralittoral and circalittoral to bathyal species known from 1 to 385 m depth (Maul 1976) on sand or

coarse sand, among seagrass or bedrocks (Kovačić 1995).

Geographic distribution Mediterranean, in the Atlantic recorded only in southern Portugal (Maul 1976). It was

described from the Balearic Islands (de Buen 1928), and since has been recorded throughout northern Mediterranean

from Gibraltar to Malta (Kovačić & Schembri 2019) and Cyprus (Kovačić & Golani 2006), including southern

France (Le Bris & Louisy 2015), the Ligurian Sea (Liu et al. 2009b), Adriatic Sea (Kovačić 1995; Lipej et al. 2005),

Aegean Sea and Levant (Bilecenoglu et al. 2014).

KOVAČIĆ ET AL.

FIGURE 58. Live coloration in its natural habitat of Gobius roulei. A) Pale male, photo by R. Pillon, Maddalena, Sardinia, Italy.

B) Male in nocturnal dark coloration, photo by S. Le Bris, Marseille, Southern France.

Gobius vittatus Vinciguerra, 1883 (Fig. 10)—Striped Goby

Gobius vittatus Vinciguerra, 1883: 527, type locality: western Mediterranean Sea, Italy, Sardinia, Isola Tavolara.

Size. Known adult size about 4–5 cm total length.

Morphology. D VI + I,11–13; A I,11–13; P 15–18. Short and stout body, nearly cylindrical, except for laterally

compressed caudal peduncle (Kovačić 2006). Caudal peduncle slender, lower than body. Dorsal fins of similar

height, the first with a more or less rounded edge. Caudal fin truncate to slightly emarginate. Body and predorsal

area covered with scales (Miller 1986), usually poorly visible on live specimen photographs (Fig. 10).

Live coloration. Background coloration pale to white, light yellow or beige, with a broad, dark-chestnut to

black stripe between the upper lip and snout and the caudal fin, passing through the eyes (Fig. 10). The lateral

stripes of both sides meet above the upper lip, forming a V-shaped snout line that slightly extends onto mid upper

lip (Fig. 10). Fins mostly clear and unmarked, yet often with a light pink hue on the rays and membranes (Fig. 10)

(Heymer & Zander 1978).

Similar species. Tridentriger trigonocephalus, also the blenny Parablennius rouxi.

Habitat. Infralittoral and circalittoral species, known from 2–85 m depth (Francour et al. 2010), most frequent

between 15–40 m depth, on steep to moderate rocky slopes made of boulders or coralligenous beds (Heymer &

Zander 1978), more rarely on sand (Kovačić 2004; Kovačić & Arko-Pijevac 2008).

Geographic distribution. Northern Mediterranean, presently known from the Strait of Gibraltar in Spain to

Leghorn in Italy, in the Gulf of Taranto and the Gargano Peninsula (Italy), Eastern Adriatic, the Aegean Sea (Miller

1986). Also recorded from Crete (Kovačić et al. 2011).

Gobius xanthocephalus Heymer & Zander, 1992 (Fig. 38)—Yellow-headed Goby

Gobius xanthocephalus Heymer & Zander, 1992: 305, type locality: western Mediterranean Sea, France, Banyuls-sur-Mer, Ile

Grosse.

Size. Known adult size about 9 cm total length.

Morphology. D VI + I,13–16; A I,13–15; P 18–20. Medium-sized goby with moderately elongate body, large

head, and a short, slightly acute and oblique snout. Caudal peduncle deep, but lower than body. Dorsal fins of

similar height, the first dorsal fin may be slightly higher in appearance, with more or less rounded distal edge and

no elongate spines. Caudal fin rounded.

Scales present on body and on predorsal area, usually visible on photographs at least on body.

Live coloration. Head yellowish, more intensely on the snout and in the interorbital space, contrasting

with a light gray or whitish body (Fig. 38). Head and body covered with longitudinal lines of well-defined orange

to dark red dots. Eyes with brown stripes radiating from the pupil over iris to the orbital rim, 2 of them constituting a

broad horizontal stripe across the pupil; the upper eye radial stripes are connected by an irregular longitudinal

curved line (Fig. 38b). Upper lip with orange markings. Lower lip with a median W-shaped mark. Snout with M-

shaped line and an oblique preorbital bar (Fig. 38b). One isolated brown to dark red dot at the corner of the

mouth, followed by 2 short horizontal dashes (Fig. 33e).

Similar species. Gobius auratus morph 2, G. fallax.

Habitat. Infralittoral species, known from 1–36 m depth (Heymer & Zander 1992; Wirtz & Herrera 1995) on

hard (e.g., bedrock, coralligenous) or mixed bottoms (e.g., sandy patches with pebbles, boulders, and seagrass).

Geographic distribution. Northeastern Atlantic, Mediterranean and Black Sea. In the Mediterranean, recorded in

the northwestern Mediterranean (Miller 1986), including the Balearic Island of Ibiza (Fisher et al. 2007) and Corsica

(Francour et al. 2010). Isolated populations confirmed in Crimea (Boltachev et al. 2009) and in the eastern part of

the Black Sea (Vasil’eva & Bogorodsky 2004). In the Atlantic Ocean: from Galicia, Northern Spain (Villegas-Riìos

& BanÞoìn 2010) to the Canary Islands (Wirtz & Herrera 1995).

KOVAČIĆ ET AL.

Gobius xoriguer Iglésias, Vukić & Šanda 2021 (Fig. 59)—Kestrel Goby

Gobius xoriguer Iglésias, Vukić & Šanda 2021a: 3, type locality: western Mediterranean, France, Corsica, Off Ghisonaccia,

Tyrrhenian Sea.

Size. Known adult size reaching 6.5 cm total length.

Morphology. D VI + I,14; A I,13; P 18–19. Moderately long body and distinctly large eyes. Snout short (shorter

than eye diameter), with moderately steep and rounded profile. Caudal peduncle deep, but lower than body depth.

First dorsal fin higher than second dorsal fin (maybe in mature males only), third spine longest and elongate,

interspinous membranes deeply incised. No free pectoral-fin rays detected. Caudal fin rounded. Scales visible on

dead specimens from reticulate pattern of pigmentation along scale edges, visibility on live specimens unknown.

Freshly dead coloration. Background coloration orange-beige, dark-brown or whitish, with orange markings

dorsally, and 7 rectangular, orange midlateral blotches (Fig. 59). Eyes with stripes radiating from the pupil to the

orbital rim, including a broad horizontal stripe across the eye. One indistinct orange preorbital bar. Lips unmarked.

Cheek and opercle orange, possibly smoked gray in some individuals, with numerous rounded white spots

(Fig. 59). On the anterior half of the first dorsal fin, an oblique, smokey gray blotch extending from the distal half of

first spine to the base of fourth or fifth. Upper base of pectoral fins with an orange blotch (Iglésias et al. 2021a).

Similar species. Gobius gasteveni, G. kolombatovici, Vanneaugobius dollfusi.

Habitat. Circalittoral species, recorded from deep coralline beds composed of fixed or unattached red algae,

between 62 and 102 m depth (Iglésias et al. 2021a).

Geographic distribution. Western Mediterranean, known from only three individuals collected in Corsica, off

the French-Spanish continental border, and in the Balearic Island of Menorca (Iglésias et al. 2021a).

FIGURE 59. Freshly dead coloration of Gobius xoriguer. Photo by S. Iglésias, off eastern Corsica, Southern France.

Hazeus ingressus Engin, Larson & Erhan, 2018 (Fig. 35)—Invaded Sand Goby

Hazeus ingressus Engin, Larson & Irmak, 2018: 318; eastern mediterranean, Turkey, Fethiye.

Size. Largest specimen examined 3.7 cm total length.

Morphology. D VI + I,8; A I,8–9; P 17–18. Body moderately elongate, the depth 5.0–6.3 in standard length.

Stout head with slightly rounded snout and large eyes extending slightly above dorsal profile. Caudal peduncle deep,

but lower than body depth. First spine of first and second dorsal fins stout and sharp-tipped; third spine of the

first dorsal fin slightly longest but not elongate. Caudal fin rounded, shorter than head length. Body, nape, cheek

and opercle scaled, but scales not visible or poorly visible on live specimen photographs (Engin et al. 2018).

Live coloration. Pale gray with numerous small irregular brown spots on body and dorsally on head, and

a row of 5 irregular blackish blotches along midside (Fig. 35), first below first dorsal fin, fifth on caudal-fin base.

Head with brown marking below orbit. Second dorsal fin with brown streaks along rays. No rows of elongate spots

on cheek and no longitudinal eye stripe (Fig. 33b).

Similar species. Gobius incognitus, G. bucchichi.

Habitat. Inhabits sand bottom in sheltered bays and lagoons at depth of 5–37 m (Engin et al. 2018).

Geographic distribution. Only known from the type specimens collected at Fethiye on the Levantine coast of

Turkey, eastern Mediterranean Sea. It was later observed in Abu Dabab lagoon, Marsa Alam, Egypt on silty sand

bottom (based on an unpublished photograph provided by B. Hazes).

Hetereleotris vulgaris (Klunzinger, 1871) (Fig. 47)—Common Gogglegoby

Gobiosoma vulgare Klunzinger, 1871: 484; type locality: Red Sea, Egypt, Al Quseir.

Size. Reaches 4 cm total length.

Morphology. D VI + I,11; A I,10; P 15–17 (rarely 17). Body moderately elongate, its depth 4.2–5.3 in standard

length. Head depressed, snout short, rounded. Caudal peduncle deep, but lower than body depth. Dorsal fins of

similar height, the first dorsal fin with more or less rounded margin having spines subequal in length, none elongate.

Caudal fin rounded, longer than head. Head and body scaleless (Hoese 1986).

Live coloration. Semitranslucent gray, with 6 irregular, sometimes indistinct, dark yellowish brown bars

on body, containing irregular pale spots, and extending into basal part of dorsal fins (Fig. 47); head with 2

irregular dark brown bands radiating anteroventrally and posteroventrally from eye with a whitish area between

them; a narrow dark brown bar at base of caudal fin (Fig. 47), this fin crossed by irregular vertical rows of dark

brown spots.

Similar species. Millerigobius macrocephalus, Zebrus zebrus, Z. pallaoroi.

Habitat. No data on habitat for the Mediterranean. In the Red Sea, inhabits crevices among stones on reef flat

or rocky substrata at coral reefs at depths of 1–24 m.

Geographic distribution. Reported from the Red Sea, Gulf of Aden and Socotra Archipelago, south to

Mozambique, east to southern Oman and Pakistan. A single specimen is known from the coast of Israel, eastern

Mediterranean Sea (Hoese 1986).

Lebetus guilleti (Le Danois, 1913) (Fig. 8)—Guillet’s Goby

Gobius (Lebetus) scorpioides var. guilleti Le Danois, 1913: 91, type locality: western Mediterranean Sea, northwestern France,

on English Channel, Morlaix Bay.

Size. Maximum known size 2.0 cm total length (Schliewen et al. 2019).

Morphology. D VI + I,7–9; A I,4-6; P 14-15 (Herler & Kovačić 2002). Small goby with subcylindrical body,

laterally compressed towards caudal fin. Head slightly depressed and fairly long, snout short, eyes large and close

together. Caudal peduncle deep, but lower than body depth. First dorsal fin higher than the second dorsal fin; no

elongated spines, but breeding males with notably enlarged, sail-like first dorsal fin. Caudal fin rounded. Body

covered with scales (Herler & Kovačić 2002), which are poorly visible on live specimen photographs (Fig. 8).

Live coloration. Head and most predorsal area grayish in males, brownish in females. Body with a pattern of

alternating dark and pale, slightly oblique broad bars. A white area below the first dorsal fin, followed by a well

obliquely-delineated dark area between bases of second dorsal and anal fins (which contains an oblong white

spot ventrally), contrasting with a plain white caudal peduncle (Fig. 8). Rear caudal peduncle and caudal-fin

base dark brown. White body area extending onto most of pectoral fin. Additional narrow rusty red bars can be

present over pale and dark areas below dorsal fins. Males’ enlarged first dorsal fin grayish to white with 2-3, often

faint, oblique yellow to orange bands, second dorsal fin with black margin, 4 oblique yellow to orange bands and

sometimes a large blue-edged black blotch on its anterior corner. Females’ triangular first dorsal fin white to gray-

brown with a small green round spot between fifth and sixth spines, second dorsal fin somewhat transparent, both

dorsal fins with visible or indistinct oblique orange bands (Herler & Kovačić 2002; Hope & Shucksmith 2010;

Schliewen et al. 2019).

Similar species. Lebetus patzneri, Speleogobius trigloides.

Habitat. Infralittoral and circalittoral species, known from 2–67 m depth on stone, shell, gravel, coralligenous

or red algae beds bottoms (Engin et al. 2015; Schliewen et al. 2019).

Geographic distribution. The northeastern Atlantic, from Normandy to Norway, as well as in the Mediterranean,

including the northwestern Mediterranean, Adriatic Sea and the Sea of Marmara (Miller 1986; Engin et al. 2015;

Schliewen et al. 2019).

KOVAČIĆ ET AL.

Lebetus patzneri Schliewen, Kovačić & Ordines, 2019 (Fig. 11)—Patzner’s Goby

Lebetus patzneri Schliewen, Kovačić & Ordines, 2019: 236; type locality: western Mediterranean Sea, Spain, Balearic Islands,

Menorca Channel.

Size. Maximum size 2.1 cm total length (Schliewen et al. 2019).

Morphology. D VI–VII+I,7–8; A I,6; P 15–16. Small goby with subcylindrical body, laterally compressed towards

caudal fin. Head slightly depressed and fairly long. Eyes moderately large and close together. Snout moderately long

and pointed. Lips thick, giving mouth a broad, almost rectangular shape in dorsal and lateral view. Caudal

peduncle deep, but lower than body depth. The first dorsal fin small, more or less with rounded distal edge. Caudal

fin rounded. Body covered with scales, but scales poorly visible on freshly dead specimen photographs, visibility

on live specimens unknown (Schliewen et al. 2019).

Freshly dead coloration. Head and body red, gently black dotted (peppered) and with irregular white

markings; first dorsal fin entirely dark (Fig. 11). Three red bars radiating from lower margin of eye, with white

in between. Caudal peduncle mostly red as the rest of body, with only a triangular white blotch or saddle (Fig. 11)

(Schliewen et al. 2019).

Similar species. Lebetus guilleti, Speleogobius trigloides, Vanneaugobius dollfusi.

Habitat. Circalittoral, on Peyssonnelia and rodolith red algae beds, from 60–72 m depth (Schliewen et al.

2019).

Geographic distribution. Mediterranean, only known from Balearic Islands, Spain (Schliewen et al. 2019).

Lesueurigobius friesii (Malm, 1874) (Fig. 17)—Fries’s Goby

Gobius friesii Malm, 1874: 383; type locality: Sweden, Gullmarfjord, Lysekil.

Size. Maximum size 10 cm total length (Miller 1986).

Morphology. D VI + I,13–16; A I,12–15; P 18–19 (Miller 1986). Moderately large goby with subcylindrical

body, laterally compressed towards caudal fin, head moderately large. Snout large and blunt. Eyes dorsolateral,

interorbital space narrow. Caudal peduncle deep, but lower than body depth. Dorsal fins of similar height, the

first dorsal fin may be slightly higher, with more or less rounded edge and no elongate spines. Caudal fin slightly

pointed, almost rounded. Predorsal area and nape scaled (Miller 1986). Scales large, visible on body and usually

also on predorsal area.

Live coloration. Body with 2 longitudinal rows of gray-brown to orange-brown spots, more or less overlaid

with yellow or orange, one row along back and another along lateral midline; smaller spots of the same color (and

not lines) on head (Fig. 17). Both dorsal fins and upper half of caudal fin with yellow to orange spots; second

dorsal-fin edge yellow to orange with a blue to whitish submarginal band (Moen & Svensen 2004; Louisy

2015).

Similar species. Lesueurigobius sanzi, L. suerii, Thorogobius macrolepis.

Habitat. Infralittoral and circalittoral to bathyal species, known from 10–440 m on soft bottoms and seagrass

meadows (Goren et al. 2019; Patzner 2021).

Geographic distribution. Mediterranean and Atlantic Ocean: present in the Atlantic from Mauritania to Norway

and along north Mediterranean coast from Gibraltar to the Sea of Marmara, also along Turkish coast south to Levant

(Miller 1986; Goren et al. 2019).

Lesueurigobius sanzi (de Buen, 1918) (Fig. 60)—Sanzo’s Goby

Gobius sanzi de Buen, 1918: 296; type locality: western Mediterranean Sea, Spain, Malaga coast, Calatoros.

Size. Maximum size 9.5 cm total length (Miller 1986).

Morphology. D VI + I,15; A I,16–17; P 22 (Miller 1986). Moderately large goby with high, laterally compressed

and robust body, head high and moderately large. Snout large and blunt. Eyes dorsolateral, interorbital space narrow.

Caudal peduncle deep, almost same depth as body. The first dorsal fin with elongated spines, third and fourth

spine longest. Caudal fin lanceolate. Predorsal area and nape scaled. Scales large, visible on freshly dead specimens

on body and usually also on predorsal area, visibility on live specimens unknown.

Freshly dead coloration. Body fawn, with diffuse vertical yellow and dark brown bands across body.

Margins of anal and caudal fins with a blackish shade (Fig. 60); live coloration unknown (Miller 1986).

Similar species. Lesueurigobius friesii, L. suerii.

Habitat. Circalittoral species, known from 47–100 m on soft bottoms, sand and mud (Miller 1986).

Geographic distribution. Atlantic Ocean and adjacent Mediterranean: known from the eastern Atlantic coast

from Portugal to Mauritania (Miller 1986) and also off northern Namibia. In the Mediterranean it is known from the

Alboran Sea (Miller 1986).

FIGURE 60. Freshly dead coloration of Lesueurigobius sanzi. Photo by A. Kassar, Beni Saf, Alboran Sea, Algeria.

Lesueurigobius suerii (Risso, 1810) (Fig. 16)—Lesueur’s Goby

Gobius suerii Risso, 1810: 387; type locality: northwestern Mediterranean Sea, France, Nice.

Size. Maximum size 7.6 cm total length (Lamprakis et al. 2003).

Morphology. D VI + I,13–14; A I,13–14; P 17–20 (Miller 1986). Moderately small goby with laterally

compressed body, head high and moderately large. Snout blunt, short. Eyes dorsolateral, high on head with top

forming part of dorsal profile. Caudal peduncle deep, but lower than body depth. The first dorsal fin slightly

higher than the second, spines not elongated. Caudal fin lanceolate. Predorsal area and nape naked (Miller

1986). Scales large, visible on body (Fig. 60).

Live coloration. Body with a midlateral series of poorly defined dark spots; more irregular, often oblique

markings along back. Background color grayish beige with light blue to reddish hue, sometimes with ill-defined

yellow bars (Fig. 16). Head with 3–4 oblique yellow to orangish stripes over cheek, preopercle and opercle, and

another slanting upper stripe on side of nape. Usually, second dorsal, caudal and anal fins with an orange outer

margin and a blue to whitish submarginal band, sometimes faint.

Similar species. Lesueurigobius friesii, L. suerii, Oxyurichthys petersii.

Habitat. Infralittoral and circalittoral to bathyal species, known from infralittoral to 440 m on soft bottoms,

mostly mud, muddy sand and possibly also sand (Miller 1986; Bérenger et al. 2015; Rufray et al. 2016; Goren et

al. 2019).

KOVAČIĆ ET AL.

Geographic distribution. Mediterranean and Atlantic Ocean. In the Mediterranean, known from the northern

coast from Alboran Sea to Levant, and also on the south coast from Algeria (Miller 1986). In the Atlantic Ocean, it

was recorded from Morocco and Canaries (Miller 1986).

Millerigobius macrocephalus (Kolombatović, 1891) (Fig. 27)—Large-headed Goby

Gobius macrocephalus Kolombatović, 1891: 22; type locality: Adriatic Sea, Croatia, Brač Island, Milna.

Size. Maximum size about 5 cm total length (Vanhove et al. 2011).

Morphology. D VI + I,9–11; A I,9–10; P 14–16. Small goby with stocky body, large head and a short steep

snout. Anterior nostril tube long, without tentacle. Caudal peduncle deep, almost as deep as body. Dorsal fins

of similar height, the first with a more or less rounded edge. Only the tip of upper pectoral fin rays is free from

membrane. Caudal fin rounded. Body scaled, predorsal area naked (Miller 1986; Vanhove et al. 2011). Scales more

or less visible on body from the reticulate pattern of pigmentation along scale edges (Fig. 27).

Live coloration. Body greenish, brownish or dark brown with 6–11 narrow, usually indistinct, pale bars

across the sides (Fig. 27a) (Vanhove et al. 2011; Patzner 2021). In unstressed animals in the wild, side of body

often more or less uniformly colored or mottled brown (Fig. 27b). When pale markings are visible on the back, they

most often coalesce into a highly irregular mediodorsal pale band (most dark markings on the body not reaching

to base of dorsal fins). Many small pale dots irregularly arranged, especially on snout, preopercle and cheek.

When present, the pale crescent band on head (broad whitish transverse band across anterior nape extending on

the sides down to pectoral-fin bases) extends onto the rear part of eyes; it is most often ill-defined (consists of

overlapping whitish spots). Often a distinctive and well-defined longitudinal median narrow line on nape and

predorsal area, usually formed by a dark dash followed by a white dash, another dark dash, and then a pale

line reaching to origin of first dorsal fin (Fig. 27c). First dorsal fin more or less dark, sometimes reddish, with a

whitish horizontal band and often a pale distal edge.

Similar species. Zebrus zebrus, Zebrus pallaoroi.

Habitat. Infralittoral species, known from 1–25 m depth on stones and pebbles near soft bottoms (summarized

by Patzner 2021). Also recorded in transition waters, hypersaline or brackish (Vanhove et al. 2011).

Geographic distribution. Mediterranean and Black Sea. In the Mediterranean Sea, there are records from Ibiza

(Ramos-Espla & Perez-Ruzafa 1987), Cerbère, Gulf of Lion (Duhau et al. 2021), southern Corsica (Bouchereau &

Tomasini 1989, Renoult et al. 2021b), Sicily (Giacobbe et al. 2016), Malta (Kovačić et al. 2013), Croatia (Kovačić

2005), Slovenia (Trkov et al. 2019), Greece, Lake Vouliagmeni (Vanhove et al. 2011), Crete (Kovačić et al. 2011),

Euboia Island (Kovačić et al. 2021), Turkey, Aegean Sea (Bogorodsky et al. 2010), Cyprus (Kovačić et al. 2021),

Israel and Lebanon (Miller 1977). Large-headed goby was also found in Crimea, in the Black Sea (Boltachev et al.

2010).

Odondebuenia balearica (Pellegrin & Fage, 1907) (Fig. 26)—Coralline Goby

Eleotris balearicus Pellegrin & Fage, 1907: 11; type locality: western Mediterranean Sea, Spain, Balearic Islands, north of

Cabrera Island.

Size. Maximum size about 3.6 cm total length.

Morphology. D VI + I,9–10; A I,9–10; P 14–16. Small species with a moderately elongated body. Head not

prominently dorsoventrally flattened. Deep caudal peduncle, nearly same as body depth below mid second dorsal

fin. First dorsal fin triangular, with the first spine longest, more elongated and pointed in males (Fig. 26).

Caudal fin rounded. Anterior nostril tubular without process from rear rim. Uppermost and lowermost scales on

caudal-fin base conspicuously enlarged and with elongated ctenii although this is rarely visible in photographs.

The pelvic fins are separated, which is also hardly noticeable from lateral view (Miller & Tortonese 1968; Miller

1986; Ahnelt et al. 1994; Kovačić & Golani 2007a; Louisy 2015). Scales visible on body from reticulate pattern of

pigmentation along scale edges.

Live coloration. Head and body usually reddish or orangish overall, with 5–8 indistinct transverse pale

bars (Fig. 26a) or, more often, with just a series of about 9 small whitish saddles or markings along back and

side of body more or less uniformly colored (Fig. 26b). The first 2 of these pale dorsal markings, located on nape

and predorsal area, may be faint. No midlateral row of small black spots. The barred pattern on the body mostly

appears when the fish is stressed after capture (or when anesthetized). Tubular anterior nostrils usually whitish. A

whitish spot at the dorsal edge of opercle. Possibly 2 small white spots aligned vertically on upper pectoral-fin

base with a darker patch in between (Louisy 2002, 2015). Outer margin of first dorsal fin broadly white or pale

gray.

Similar species. Vanneaugobius dollfusi, Corcyrogobius liechtensteini.

Habitat. Infralittoral and circalittoral species, known from a few meters to about 45 m depth on rocky or

coralligenous bottoms in small cavities and crevices, under stones or in multi-layered scree, sometimes among

macroalgae or on mixed sandy grounds (Louisy 2002; Kovačić et al. 2012a & b; Glavičić et al. 2016). Specimens

have also been caught from about 25–103 m depth, on coralline and stony grounds, but also on Cladophora beds or

on muddy substrates (Miller 1986; Ahnelt et al. 1994; Ahnelt & Dorda 2004; Stern & Rilov 2019).

Geographic distribution. Mediterranean. Northern and eastern Mediterranean, from the Gulf of Valencia

(Ordines et al. 2019a) and Balearic Islands (Ahnelt et al. 1994) to the Levant, Cyprus (Kovačić & Golani 2007a)

and Israel (Stern & Rilov 2019), including southern France, at Six-Fours (Escoubet et al. 1981) and Banyuls (Louisy

2002), southern Italy, Gulf of Taranto (Parenzan 1973), Strait of Sicily (Consoli et al. 2019), Malta (Kovačić &

Schembri 2019), the Adriatic Sea (Ahnelt et al. 1994; Kovačić et al. 2012b) and the Aegean Sea (Koukouras et al.

1985; Miller 1986).

Oxyurichthys petersii (Klunzinger, 1871) (Fig. 45)—Red Sea Tentacle-goby

Apocryptes (Gobiichthys) petersii Klunzinger, 1871: 480; type locality: Red Sea, Egypt, Al Quseir.

Size. Attains about 17 cm total length.

Morphology. D VI + I,12–13; A I,12–13; P 22–23. Body elongate, the depth 5.6–6.5 in standard length. Eyes

protruding, their upper part well above dorsal profile of head (Fig. 45), each topped by a short fleshy knob.

A narrow dermal crest present on nape. Caudal peduncle deep, almost as deep as body. Spines of first dorsal fin

elongate, each ending in a short filament. Dorsal and anal fins long-based. Pelvic fins united to form a disc.

Caudal fin lanceolate, nearly twice as long as head (Fig. 45). Scales on body ctenoid posteriorly, becoming cycloid

anteriorly below middle of first dorsal fin, no scales on cheek, opercle, and prepectoral area (Pezold & Larson

2015). Scales small, at best visible on body as pattern of pale dots.

Live coloration. Yellowish to greenish gray dorsally, abdomen whitish, with a midlateral row 6 slightly

elongate, blackish blotches, the first blotch on pectoral-fin base, the last on mid caudal peduncle at caudal-fin base

(Fig. 45), sometimes alternating with 5 smaller dark spots. Head with a broad dark bar below eye, indistinctly

edged with a pale blue line, followed by a row of dark blotches across operculum. Males with a series of oblique

iridescent blue streaks on body, the first below origin of first dorsal fin; membranes of both dorsal fins with yellowish

marks, forming longitudinal stripes, separated by short blue stripes.

Similar species. Cryptocentrus steinhardti.

Habitat. Inhabits silty sand or mud substrata of sheltered bays or estuaries, from the shallows to depths of 50 m;

takes refuge in a burrow, but is able to dive quickly into mud if threatened when away from shelter.

Geographic distribution. In the native area restricted to the Red Sea (Pezold & Larson 2015). The most common

Lessepsian goby migrant in the Mediterranean, where reported from Greece at Kastellorizo Island (Apostolopoulos

& Karachle 2016), Israel (Ben-Tuvia 1983; Golani 2021), Lebanon (Bariche & Fricke 2020), Syria (Ali 2018),

Tunisia (Golani et al. 2002), and Turkey from Aegean Sea and Mediterranean Sea coasts (Bilecenoglu et al. 2002;

Akyol et al. 2006).

KOVAČIĆ ET AL.

Papillogobius melanobranchus (Fowler, 1934) (Fig. 61)—Blackthroat Goby

Rhinogobius melanobranchus Fowler, 1934: 82; type locality: Indonesia, Bali.

Size. Reaches 5.5 cm total length.

Morphology. D VI + I,8–9 (usually 8); A I,8; P 15–16. Body depth 4.7–5.3 in standard length. Head slightly

laterally compressed, with relatively short snout. The second and third spines of first dorsal fin longest but not

elongate. Caudal peduncle deep, but lower than body depth. Pelvic fins joined to form a disc, reaching to or extending

slightly beyond anus. Caudal fin rounded, shorter than head. No scales on head and predorsal area. Body covered

with scales (Kovačić & Golani 2007b), but scales poorly visible on freshly dead specimen photographs, visibility

on live specimens unknown (Fig. 61).

Fresh dead coloration. Whitish or pale greenish gray, the upper 2/3 of body with small brown spots of variable

size, mainly on scale edges; body with a midlateral row of 6 larger and darker markings, mainly as double

spots (Fig. 61), the first beneath pectoral fin, the last at base of caudal fin. A large indistinct orangish brown blotch on

base of opercle. In males, ventral third of body with vertical orange lines, some linked to the midlateral blackish

spots. In males, outer margin of first dorsal fin deep purple, with an irregular pale yellow submarginal band;

caudal fin with a dark spot on upper edge (Fig. 61) (Kovačić & Bogorodsky 2013).

Similar species. Pomatoschistus adriaticus, P. marmoratus, P. microps.

Habitat. Inhabits silty sand and sparse seagrass, at depths of 0.3–5 m (Kovačić & Bogorodsky 2013).

Geographic distribution. Reported from the northern Red Sea and the Arabian Gulf south to Mozambique, east

to Singapore, northern Australia, Indonesia and Papua New Guinea. In the Mediterranean Sea one specimen was

collected from the Bardawil Lagoon, Egypt (Kovačić & Golani 2007b).

Remarks: Often classified in the genus Favonigobius Whitley, 1930; however, tentatively placed in Papillogobius

for the unique pattern of lateral line system papillae; see Kovačić & Bogorodsky (2013).

FIGURE 61. Freshly dead coloration of Papillogobius melanobranchus. Photo by S.V. Bogorodsky, Gulf of Suez, Egypt, Red

Sea.

Pomatoschistus adriaticus Miller, 1973 (Fig. 62)—Mediterranean Painted Goby

Pomatoschistus pictus adriaticus Miller, 1973: 72; type locality: Adriatic Sea, Croatia, Split.

Size. Maximum size 5.3 cm total length (Miller 1973).

Morphology. D VI + I,8–9; A I,8–9; P 17–19 (Miller 1973). Moderately small goby with subcylindrical body,

laterally compressed towards caudal fin, head slightly depressed and moderately large. Snout pointed, moderately

large. Eyes dorsolateral, interorbital space narrow. Caudal peduncle deep, but lower than body depth. Dorsal fins of

similar height, the first dorsal fin with more or less rounded margin and no elongated spines. Caudal fin rounded.

Scales present on body, usually visible on photographs at least dorsally. Predorsal area naked (Öztürk & Engin

2019).

Live coloration. Body pale gray to pale brown with 5 whitish saddles and dark mottled pattern (Fig. 62), the

first narrow light saddle, at the origin of first dorsal fin, is followed by 2 broader ones below second dorsal fin, and

2 on caudal peduncle. Body with 5 dark midlateral blotches, sometimes reduced to double black dots, extend

from below the above light saddles, the last blotch appearing as a horizontal black mark on caudal-fin base.

Cheeks pale gray, with only scattered black dots. A dark patch around upper pectoral-fin base divides in 2 oblique

branches over upper and lower opercle and preopercle. Dorsal fins with three horizontal pale brown to pinkish lines

(often with rows of black dots) alternating with three whitish to light blue lines.

Similar species. Pomatoschistus bathi, P. nanus, P. marmoratus, P. microps.

Habitat. Infralittoral species, on coarse sand containing biogenic calcareous fragments and shell gravel with

coralline deposits (Öztürk & Engin 2019), sand at 1–20 m depth (Patzner 2021).

Geographic distribution. Mediterranean. Known from southern France, the Adriatic Sea and the Sea of Marmara

(summarized by Patzner 2021).

FIGURE 62. Live coloration in its natural habitat of Pomatoschistus adriaticus. Photo by S. Guerrieri, Rijeka, Croatia, northern

Adriatic Sea.

KOVAČIĆ ET AL.

Pomatoschistus bathi Miller, 1982 (Fig. 63)—Bath’s Goby

Pomatoschistus bathi Miller, 1982: 6; type locality: Sea of Marmara, Turkey, Bandýrma, Erdek.

Size. Maximum size 3.2 cm total length (Miller 1986).

Morphology. D VI (V–VI) + I,7–9; A I,6–9; P 13–17 (Miller 1986). Small goby with subcylindrical body,

laterally compressed towards caudal fin, head slightly depressed and distinctly small compared to thickness of

body (Fig. 63). Snout moderately long. Eyes dorsolateral, interorbital space moderately narrow. Caudal peduncle

slender, clearly lower than body depth. The first dorsal fin about equal or shorter than the second dorsal fin, with more

or less rounded distal margin. Caudal fin rounded. Scales present on body, usually visible on close-up photographs

at least dorsally. Predorsal area naked.

Live coloration. Body from beige to light brown with pale saddles and mottled patterning dorsally and white

ventrally, with vertical, iridescent, pale gray streaks. Five small pale dorsal saddles ending ventrally above the

midline not linked with a midlateral irregularly shaped, horizontally elongate, black spot below (Fig. 63).

Each black lateral spot with a short ventral black extension, except the last, pear-shaped, at caudal-fin base. Short

dark dash from the center of the snout to the upper lip, not always visible. Indistinct gray to dark vertical bar

extending from ventral edge of eye to corner of mouth; cheek pale (Fig. 63). On side of head, between opercle

and upper pectoral-fin base, a horizontal, Y-shaped brown to reddish marking, branching forward (Fig.

63).

Similar species. Pomatoschistus nanus, P. quagga, P. adriaticus, Pseudaphya ferreri.

Habitat. Infralittoral species, on sand and gravel to 15 m depth (Engin et al. 2018a).

Geographic distribution. Mediterranean and Black Sea. In the Mediterranean Sea, there are records from the

northwestern Mediterranean, Adriatic Sea, Aegean Sea and Sea of Marmara (Miller 1986), also Sicily (Giacobbe et

al. 2017). In the Black Sea it was recorded from the eastern coast (Vasil’eva & Bogorodskii 2004).

FIGURE 63. Live coloration in its natural habitat of Pomatoschistus bathi. Photo by R. Pillon, Rab Island, Croatia, northern

Adriatic Sea.

Pomatoschistus knerii (Steindachner, 1861) (Fig. 64)—Kner’s Goby

Gobius knerii Steindachner, 1861: 287, Pl.1 (fig. 1); type locality: Adriatic Sea, Croatia, Hvar.

Size. Maximum size 4 cm total length (Miller 1986).

Morphology. D VI (VI–VII) + I,9–11; A I,8–10; P 17–20 (Miller 1986). Small goby with subcylindrical body,

laterally compressed towards caudal fin, head moderately small. Typically short and rounded snout. Eyes lateral,

interorbital space wide. Caudal peduncle slender, clearly lower than body depth. Dorsal fins of similar height,

the first dorsal fin with more or less rounded margin and no elongated spines. Caudal fin truncate to slightly

emarginate. Scales present on body, usually visible on photographs at least dorsally. Predorsal area naked (Miller

1986).

Live coloration. Background color beige to reddish orange, with alternating narrow dark and whitish

vertical bars, more numerous in males. Irregular dark markings along back. Sometimes 4 pale dorsal saddles, first

below origin of second dorsal fin, last on rear caudal peduncle; a large red or blackish triangular spot on base of

caudal fin (Fig. 64). May be 2 red or dark bars below eye, first reaching corner of mouth, second on cheek. First

dorsal fin with three oblique orange stripes, second sometimes with series of brown spots (Miller 1986; Kovačić

et al. 2017b; present work, Fig. 64).

Similar species. Pomatoschistus bathi, P. quagga.

Habitat. Infralittoral species, mostly encountered between 0–10 m depth (Kovačić 2003), but known down to 45

m (Kovačić et al. 2017b), often hovering 10–50 cm above the substrata, on sand near rock and Cymodocea meadows

or among boulders and scree (Kovačić 2003; Patzner 2021).

Geographic distribution. Mediterranean. Known from Giglio Island in the Tyrrhenian Sea and from the Adriatic

Sea (Miller 1986; Kovačić 2003).

FIGURE 64. Live coloration in aquariumof Pomatoschistus knerii. Photo by J. Vukić, Krk Island, Croatia, northern Adriatic

Sea.

KOVAČIĆ ET AL.

Pomatoschistus marmoratus (Risso, 1810) (Fig. 65)—Marbled Goby

Atherina marmoratus Risso, 1810: 339; type locality: northwestern Mediterranean Sea, France, Nice.

Size. Maximum size 6.5 cm total length (Miller 1986).

Morphology. D V (V–VII) + I,8–10; A I,8–10; P 17–21 (Miller 1986). Moderately small goby with subcylindrical

body, laterally compressed towards caudal fin, head slightly depressed and moderately large. Snout moderately

large. Eyes dorsolateral, interorbital space moderately narrow. Caudal peduncle long and slender, clearly lower

than body depth. Dorsal fins of similar height, the first dorsal fin may be slightly higher, with more or less rounded

margin and no elongate spines. Caudal fin rounded. Scales present on body, sometimes visible on photographs, at

least dorsally. Predorsal area naked (Miller 1986).

FIGURE 65. Live coloration in its natural habitat of Pomatoschistus marmoratus. A) Male and B) female, photos by R. Pillon,

Rab Island, Croatia, northern Adriatic Sea. C) Female, photo by J. Renoult, Thau lagoon, Southern France.

Live coloration. Background a pale sandy color, with a more or less marbled pattern of dark and white markings.

On pale fine sand, light colors prevail, with almost no recognizable patterns (Figs. 65a and 65b). On coarser

sediments, more contrasting patterns are usually displayed: 6 pale saddles on the back, 1 below first dorsal fin, 2

below second dorsal fin, 2 on caudal peduncle and one, narrower and sometimes faint, on nape above pectoral-fin

base; variable number of faint rounded dark spots along midline, the last longitudinal and more obvious at caudal-

fin base (Fig. 65c). A preorbital dark bar, especially obvious in females, extends from each eye, through upper lip,

to connect with that from the opposite side, forming a distinctive dark chin spot (Fig. 65b). Often a whitish to pale

blue dot on nape, just behind eyes, generally more or less circled with an interrupted dark line, often like between

brackets (such a marking may also appear in P. microps or Deltentosteus quadrimaculatus). In males, cheeks often

densely sprinkled with small black dots; first dorsal fin may be large, with brownish and pale stripes, and a blue-

black spot on its rear upper corner (Fig. 65a).

Similar species. Pomatoschistus adriaticus, P. microps, P. minutus.

Habitat. Infralittoral species, often common on shallow soft bottoms (mostly fine to coarse sand) from 0.3–20

m depth (Apostolou et al. 2011; Pillon et al. 2015; Patzner 2021), may occur to 60–70 m (Miller 1986).

Geographic distribution. Black Sea, Mediterranean and adjacent Atlantic Ocean. Widespread in the Mediterranean

from Gibraltar to Levant, also common in the Black Sea (Miller 1986; Boltachev & Karpova 2017). Along the south

Mediterranean coast between the Alboran Sea and Suez Canal in Egypt (Miller 1986) the species was confirmed

from northern and southern Tunisia (Mejri et al. 2009a). In the Atlantic Ocean, known from Gibraltar to south

Portuguese coast (Miller 1986).

Pomatoschistus microps (Krøyer, 1838) (Fig. 66)—Common Goby

Gobius microps Krøyer, 1838: 172, Pl. 102 (fig. 3); type locality: England, Somersetshire, Weston-super-Mare.

Size. Maximum size 6.4 cm total length (Miller 1986).

Morphology. D VI (V–VII) + I,8–11; A I,7–10; P 15–20 (Miller 1986). Moderately small goby with subcylindrical

body, laterally compressed towards caudal fin, head slightly depressed and moderately large. Snout moderately large

and pointed. Eyes dorsolateral, interorbital space moderately narrow. Caudal peduncle long and slender, clearly

lower than body depth. Dorsal fins of similar height, the first dorsal fin with more or less rounded margin and no

elongated spines. Caudal fin rounded. Scales present on body, sometimes visible on photographs at least dorsally.

Predorsal area, nape and most of the area under the first dorsal fin naked.

Live coloration. Background color light gray to beige, with many small, irregular, white or more or less dark

dots and spots. Depending on the environment, light colors may prevail, with almost no recognizable patterns (Fig.

66b). Or there can be more contrasting patterns: usually 5, sometimes ill-defined whitish dorsal saddles, 1 below

first dorsal-fin, 2 below second dorsal fin, 2 on caudal peduncle (Fig. 66a), and in some instances, a narrower

and often fainter sixth saddle on nape above pectoral-fin base; sometimes a midlateral series of dark blotches,

often ill defined; in some instances, a deep (i.e., not superficial) oblique blackish blotch between abdominal cavity

and base of first dorsal fin. Typically 2 suborbital dark bars, one from anterior edge of eye to upper lip, a second

one, less obvious and more irregular, from rear edge of eye to just behind corner of mouth (Fig. 66a). Sometimes

a whitish dot on nape, just behind eyes, more or less edged in dark pigment (quite similar to P. marmoratus and

Deltentosteus quadrimaculatus). In males, first dorsal fin with alternate red-brown and pale longitudinal stripes, and

an elongate black or bluish spot in its rear part, but not close to the edge (Fig. 66a).

Similar species. Pomatoschistus adriaticus, P. marmoratus, P. minutus.

Habitat. Infralittoral species, on soft bottoms from a few centimeters to 10 m depth, mainly in brackish, low

salinity water (Renoult et al. 2016; Patzner 2021).

Geographic distribution. Mediterranean and Atlantic Ocean. Restricted in the Mediterranean to the northwestern

coast of the western Mediterranean (Miller 1986). The easternmost and southernmost Mediterranean records are

from Sicily (Tiralongo et al. 2019). In the Atlantic Ocean, known from the Canary Islands to Norway (Miller

1986).

KOVAČIĆ ET AL.

FIGURE 66. Live coloration in its natural habitat of Pomatoschistus microps, A) male, B) female. Photos by R. Groenveld,

Netherlands.

Pomatoschistus minutus (Pallas, 1770) (Fig. 67)—Sand Goby

Gobius minutus Pallas, 1770: 4; type locality: Belgian Sea.

Size. Maximum size 9.5 cm total length (Miller 1986).

Morphology. D VI (VI–VII) + I,10–12; A I,9–12; P 18–21 (Miller 1986). Medium-sized goby with subcylindrical

body, laterally compressed towards caudal fin, head slightly depressed and moderately large. Snout moderately

large and pointed. Eyes dorsolateral, interorbital space quite narrow. Caudal peduncle long and slender, clearly

lower than body depth. Dorsal fins of similar height, the first dorsal fin with more or less rounded margin and no

elongated spines. Caudal fin rounded. Scales present on body and on predorsal area at least posteriorly, usually

visible on photographs at least dorsally.

FIGURE 67. Live coloration in its natural habitat of Pomatoschistus minutus. A) Espiguette sand bank, Gulf of Lion, South

France, Mediterranean, photo by T. Menut. B) Stavanger, Norway, Atlantic Ocean, photo by R. Svensen.

Live coloration. Background color pale with irregular reticulation and light brown, white or gray spots. Usually

light colored with no recognizable pattern (Fig. 67a), but sometimes with more contrasting markings: 7, often ill-

defined whitish dorsal saddles, the first, narrower and often fainter than the others, on nape above pectoral-fin

base, 2 below first dorsal-fin, 2 below second dorsal fin, 2 on caudal peduncle (Fig. 67); variable number of faint

brown spots and vertical lines along midline, with one horizontal darker horizontal T-shaped blotch on caudal-fin

base. One suborbital dark bar, often obvious, between eye and side of upper lip. No pale dot on nape behind eyes.

First dorsal fin poorly pigmented, often with a blue-black spot on the distal rear corner, close to the edge,

including in females (Fig. 67b).

KOVAČIĆ ET AL.

Similar species. Pomatoschistus marmoratus, P. microps, P. norvegicus.

Habitat. Infralittoral and circalittoral species, on soft bottoms from less than 1–70 m depth (Miller 1986).

Geographic distribution. Atlantic, Mediterranean and Black Sea. Recorded in the Mediterranean along the north

coast from the Alboran Sea to Aegean Sea (Miller 1986; Özen et al. 2007), also common in the Black Sea (Miller

1986). In the Atlantic Ocean known from Gibraltar to Norway (Miller 1986).

Pomatoschistus nanus Engin & Seyhan, 2017 (Fig. 68)—Pygmaean Goby

Pomatoschistus nanus Engin & Seyhan, 2017: 3; type locality: Turkey, Antalya, Kas, Camel Reef.

Size. Maximum size 2.0 cm total length (Engin & Seyhan 2017).

Morphology. D VI + I,8; A I,9–11; P 15–16 (Engin & Seyhan 2017). Very small goby with subcylindrical body,

laterally compressed towards caudal fin, head slightly depressed and moderately small. Snout pointed. Eyes large.

Caudal peduncle slender, lower than body depth. Dorsal fins of similar height, the first dorsal fin with more or less

rounded margin and no elongated spines. Caudal fin slightly emarginate. Scales present, but not usually visible on

photographs.

Live coloration. Body mostly translucent (Fig. 68); most visible chromatophores are deep inside the body,

arranged along the vertebral column in a white line interspersed with dark markings and then, three longer dark

lines and finally an elongate spot on caudal-fin base. Superimposed on this deep pattern are 4 very thin dorsal

white saddles and a fifth broader saddle on caudal peduncle; a midlateral dark spot, that may slightly extend

ventrally, below each saddle. Body whitish ventrally, with vertical, iridescent, pale gray streaks. Cheeks pale.

Usually no horizontal Y-shaped dark marking on opercle. On nape, 3 whitish to white markings just behind eyes

with 2 dark longitudinal streaks in between, and underlined with dark on sides (Engin & Seyhan 2017; present

work).

Similar species. Pomatoschistus bathi, Pseudaphya ferreri.

Habitat. Infralittoral and circalittoral species, on coarse sand close to rocky reefs at depths of 29–41 m (Engin

& Seyhan 2017).

Geographic distribution. Eastern Mediterranean. Known only from the northern coast of the Levantine Sea

(Engin & Seyhan 2017).

FIGURE 68. Live coloration in its natural habitat of Pomatoschistus nanus. Photo by S. Engin, Kaş, Antalya, Turkey.

Pomatoschistus norvegicus (Collett, 1902) (Fig. 69)—Norway Goby

Gobius minutus norvegicus Collett, 1902: 54; type locality: Norway, Breviksfjord.

Size. Maximum size 6.5 cm total length (Miller 1986).

Morphology. D VI + I,8–10; A I,8–11; P 16–18 (Miller 1986). Moderately small goby with subcylindrical body,

laterally compressed towards caudal fin, head slightly depressed and moderately large. Snout moderately large and

pointed. Eyes dorsolateral, interorbital space moderately narrow. Caudal peduncle long and slender, clearly lower

than body depth. Dorsal fins of similar height, the first dorsal fin with more or less rounded margin and no elongated

spines. Caudal fin rounded (Miller 1986). Scales present on body, possibly visible on photographs, at least dorsally.

Predorsal area scaled at least posteriorly.

Live coloration. Body pale fawn, somewhat translucent, covered with faint darker reticulation and

scattered ferruginous to orange dots (Fig. 69). Often a lateral series of dark blotches, largest one below first dorsal

fin, and darker one on caudal-fin base. Usually a preorbital dark bar extending from the eye through upper lip, often

to the chin. Adult male with about 10–12 narrow dark bars across side of body, pigmented breast, and a black spot

bordered in blue dorsally and ventrally in rear part of the first dorsal fin (Miller 1986; Louisy 2015).

Similar species. Pomatoschistus marmoratus, P. microps, P. minutus.

Habitat. Infralittoral and circalittoral to bathyal species, from 18–325 m depth, soft bottom and mussel grounds

(Miller 1986).

Geographic distribution. Atlantic Ocean and Mediterranean. In the Mediterranean, along the north coast from

Alboran Sea to Aegean Sea (Miller 1986). Also recorded in the Adriatic Sea (Stefanni 2000). In the Atlantic Ocean

known from Normandy, France and Irish coast to Norway (Miller 1986).

FIGURE 69. Live coloration in its natural habitat of Pomatoschistus norvegicus from Atlantic Ocean. Photo by R. Svensen,

Ryfylke, Norway.

KOVAČIĆ ET AL.

Pomatoschistus quagga (Heckel, 1837) (Fig. 70)—Quagga Goby

Gobius quagga Heckel, 1837: 150, Pl. 9 (figs. 5–6); type locality: Mediterranean Sea, Italy, Sicily, Palermo.

Size. Maximum size 4.6 cm total length (Miller 1986).

Morphology. D VI (VI–VII) + I,9; A I,8–9; P 15 (Miller 1986). Small goby with subcylindrical body, laterally

compressed towards caudal fin. Small head with short pointed snout. Eyes lateral. Caudal peduncle long and

slender, clearly lower than body depth. Dorsal fins of similar height, the first dorsal fin with more or less rounded

margin and no elongated spines. Caudal fin truncate to slightly emarginate. Scales more or less visible on body

from reticulate pattern of pigmentation along scale edges.

Live coloration. Background color pale beige to light gray, sometimes partly translucent, with variable dark or

white patterns. When discernible, vertical black streaks (4–5 in males, 3 in females), restricted to the ventral

two-thirds of body, typical for this species. Often 5 pale to golden dorsal saddles, the first, small and often

faint, below first dorsal fin, the fifth just above the dark midlateral triangular blotch at caudal-fin base (Fig.

70). In translucent individuals, 4 deep elongated dark blotches run along the vertebral column (in addition to the

caudal spot). Cheeks usually pale without dark markings. A dark, reddish or pinkish oblique stripe on gill cover,

sometimes bifurcating on its lower part, somewhat reminiscent of the Y-mark of P. bathi.

Similar species. Pomatoschistus bathi, P. knerii, Pseudaphya ferreri.

Habitat. Infralittoral species observed between 2–35 m depth, often in schools of ten to several hundred

specimens, swimming up to 1 m above ground or resting on the substrate. They can be found on bedrock with

macroalgae, more or less covered with sand, on coarse to muddy sand, or near Posidonia seagrass beds (Kovačić

2003, Louisy, unpublished data).

Geographic distribution. A Mediterranean species, known along the north coast from Alboran Sea to Aegean

Sea (summarized by Patzner 2021).

FIGURE 70. Live coloration in its natural habitat of Pomatoschistus quagga. Photo by R. Pillon, Rab Island, Croatia, northern

Adriatic Sea.

Pomatoschistus tortonesei Miller, 1968 (Fig. 71)—Tortonese’s Goby

Pomatoschistus tortonesei Miller, 1968: 221; type locality: Mediterranean Sea, Italy, Sicily, Marsala.

Size. Maximum size 3.7 cm total length (Miller 1986).

Morphology. D VI (V–VI) + I,6–8; A I,6–8; P 17–20 (Miller 1986). Small goby, with relatively robust

subcylindrical body, laterally compressed towards caudal fin. Head slightly depressed and moderately large. Snout

short. Eyes dorsolateral, interorbital space moderately narrow. Caudal peduncle slender, distinctly lower than body

depth, more slender in females. First dorsal fin about equal to or lower than second dorsal fin, with more or less

rounded margin. Caudal fin rounded. Scales visible on dead specimens from reticulate pattern of pigmentation along

scale edges, visibility on live specimens unknown (Miller 1986).

Freshly dead coloration. Body fawn with darker ferruginous reticulation. Usually 2 suborbital bars (with a

pale area in between), one from anterior edge of eye to upper lip, a second one from rear edge of eye to corner of

mouth; occasionally a dark bar at the junction of preopercle and opercle. Females with 3 conspicuous thin dark

bars on posterior body (behind anus); bright yellow branchiostegal membrane (under head) and sometimes a

yellow abdomen; a dark mark on chin (Fig. 71). Males with numerous, poorly-defined, vertical dark markings

across sides; dark pelvic fins, breast and branchiostegal membrane (Miller 1986).

Similar species. P. marmoratus, P. microps, P. quagga.

Habitat. Infralittoral species, in lagoons, from brackish to slightly hypersaline, in shallows on sand near seagrass

(Miller 1986).

Geographic distribution. Mediterranean. Recorded from the central and southern Mediterranean from Sicily,

Libya and Tunisia (Miller 1986; Mejri et al. 2009b).

FIGURE 71. Freshly dead coloration of Pomatoschistus tortonesei female. Photo by R. Mejri, exact locality unknown,

Tunisia.

Pseudaphya ferreri (de Buen & Fage, 1908) (Fig. 30)—Ferrer’s Goby

Aphya ferreri de Buen & Fage, 1908: 105, Fig. 1–5; type locality: western Mediterranean Sea, Spain, vicinity of Mahon.

Size. Maximum known size 3.5 cm total length (Miller 1986)

Morphology. D V + I,7–10; A I,9–10; P 15–16 (Miller 1973). Pedomorphic habitus. Body laterally compressed,

slender compared to other gobies. Large lateral eyes, wide interorbital space. Mouth superior, oblique, tip of lower

jaw reaching the horizontal level of pupil. Caudal peduncle slender, distinctly lower than body depth. The first

dorsal fin about equal to or lower than second dorsal fin, with more or less rounded margin. Dorsal fins widely

separate with a broad interdorsal space. Pelvic disc complete with anterior membrane. Caudal fin truncate to slightly

emarginate. Scales present, but usually not visible on photographs.

Live coloration. Body semitranslucent with 4 elongate black markings along vertebral column and a

large, conspicuous, triangular black spot at the base of caudal fin (Fig. 30). A series of small black dots along

lower margin of posterior body; white superficial speckles and markings in variable amounts.

Similar species. Pomatoschistus quagga, P. bathi.

Habitat. Pelagic and neritic species, recorded over coarse sand, muddy sand or partially sand covered bedrock

from 2.5–20 m depth (Miller 1986; Kovačić 2003; Le Bris et al. 2016).

Geographic distribution. Mediterranean. Recorded from the northwestern Mediterranean, the Adriatic Sea,

northeastern Aegean Sea and northern Sinai, Egypt (Miller 1986; Kovačić & Golani 2007b; Engin et al. 2018a).

KOVAČIĆ ET AL.

Silhouettea aegyptia (Chabanaud, 1933) (Fig. 72)—Egyptian Goby

Gobius lesueuri aegyptius Chabanaud, 1933: 11; type locality: Egypt, Lake Timsah.

Size. Attains about 5 cm total length.

Morphology. D VI + I,9–11; A I,11–12; P 13–15. Body elongate, its depth 6.0–7.0 in standard length. Head with

short snout. Eyes dorsolateral, about upper one-third elevated above dorsal profile of head. Caudal peduncle

slender, lower than body depth. First dorsal fin higher than second fin, the first spine longest, slightly prolonged

in males, giving the fin a triangular shape. Pelvic fins united to form a disc, reaching origin of anal fin. Caudal fin

rounded, slightly shorter than head. Scales on body ctenoid, no scales on head, nape, and prepectoral and prepelvic

areas. Scales visible on dead specimens from reticulate pattern of pigmentation along scale edges, visibility on live

specimens unknown.

Freshly dead coloration. Finely mottled light brown dorsally, grading to white ventrally on head and abdomen,

with a midlateral series of 9–10 gray to blackish blotches about eye size or smaller, the first 2 beneath pectoral

fin and the last on caudal-fin base, the penultimate being the smallest; most of these blotches are bisected by a

vertical, slightly curved, brown-orange line (Fig. 72). A series of similar curved lines below the midlateral row;

scattered shorter brownish yellow dashes dorsally. First dorsal fin with three oblique dark crossbands; rays of

dorsal three-fourths of caudal fin with aligned small dark dots.

Similar species. Pomatoschistus marmoratus, P. microps.

Habitat. Inhabits silty sand or mud bottom, generally at depths of less than 1 m. Able to quickly bury itself into

the sediment to level of eyes; tolerates a wide range of salinities (Miller & Fouda 1986).

Geographic distribution. Originally restricted to the Red Sea. In the Mediterranean, it has been recorded from

Ashdod and Bardawil Lagoon (Miller & Fouda 1986; Golani 2021) and Syria (Ali 2018).

FIGURE 72. Freshly dead coloration of Silhouettea aegyptia. Photo by S.V. Bogorodsky, Al Lith, Saudi Arabia, Red Sea.

Speleogobius llorisi Kovačić, Ordines & Schliewen, 2016 (Fig. 13)—Lloris’ Grotto Goby

Speleogobius llorisi Kovačić, Ordines & Schliewen, 2016: 302, Figs. 2–5; type locality: western Mediterranean Sea, Spain,

Balearic Islands, Mallorca.

Size. Maximum size about 3 cm total length.

Morphology. D VI + I,6–7; A I,5–7; P 15. Body slender, laterally compressed. Head long, slightly depressed.

Snout pointed, equal to or longer than eye diameter. Second dorsal and anal fins with short base and low ray

counts (see fin meristics). Caudal peduncle slender, lower than body depth. Dorsal fins of similar height, first dorsal

fin with more or less rounded margin. Caudal fin truncate (Kovačić et al. 2016; Engin et al. 2017; Kovačić &

Glavičić 2019). Scales present on body and on predorsal area, poorly visible on photographs.

Live coloration. The general hue may vary from beige to brown to reddish brown depending on the environment

and the fish’s mood. Predorsal area reddish brown to whitish brown, underside of head, cheek and opercle mostly

red with poorly visible white parts (Engin et al. 2017), occasionally also yellowish (Kovačić & Glavičić 2019).

Three dark bars on dorsal half of body, alternating with 4 pale areas, these 3 bars connected to 4 dark blotches

on lower side of body, one more midlateral blotch on the posterior ventral part of caudal peduncle (Duhau-Spielmann

et al. 2021) (Fig. 13). Caudal peduncle uniformly pale dorsally, dark ventrally. Dark bars and blotches may be

quite diffuse in males, resulting in most of the body being rather uniformly dark, except caudal peduncle dorsally.

Dorsal fins with a wide beige to whitish outer margin.

Similar species. Speleogobius trigloides.

Habitat. Circalittoral species, known from 32–69 m depth on rhodolith beds or detritic coarse sand and gravel

(Kovačić et al. 2016; Engin et al. 2017; Kovačić & Glavičić 2019; Duhau-Spielmann et al. 2021; J. Renoult,

unpublished observation).

Geographic distribution. Northern Mediterranean, presently known from the Balearic Islands (Kovačić et al.

2016) to the Aegean Sea (Engin et al. 2017), including France (Duhau-Spielmann et al. 2021; J. Renoult unpublished

observation), Malta (Kovačić & Schembri 2019) and the Adriatic Sea (Kovačić & Glavičić 2019).

Speleogobius trigloides Zander & Jelinek, 1976 (Fig. 12)—Grotto Goby

Speleogobius trigloides Zander & Jelinek, 1976: 275, Figs. 4–5; type locality: Adriatic Sea, Croatia, Grotto of Banjole, near

Rovinj.

Size. Maximum size about 2.8 cm total length.

Morphology. D VI + I,6–7; A I,5–7; P 14–15. Body slender, laterally compressed. Head slightly depressed

behind eyes. Snout pointed, shorter than eye diameter. Caudal peduncle slender, lower than body depth. Second

dorsal and anal fins with short base and low ray counts (see fin meristics). Dorsal fins of similar height, the first

dorsal fin with more or less rounded margin and no elongated spines, except in breeding males that have both dorsal

fins higher (Zander & Jelinek 1976; Miller 1986; Kovačić et al. 2016; Engin et al. 2017). Caudal fin truncate.

Scales present on body and on predorsal area, mostly not visible on photographs.

Live coloration. Body primarily red (from rosy to yellow-orange to vermilion) with series of white blotches

along dorsal and ventral sides (Fig. 12), 5 irregular whitish dorsal saddles (first below first dorsal-fin origin, second

(wider) at interdorsal space, third below rear second dorsal fin, fourth at about middle of caudal peduncle and fifth

just before caudal-fin base); a series of 5–6 barely-defined and somewhat coalescing white to yellowish blotches

along ventral profile (first on belly below first dorsal fin, last on caudal peduncle). In males, body becoming yellow

posteriorly, or at least the caudal fin; white blotches often reduced, sometimes absent, especially the lower ones;

dorsal fins yellowish with at least one oblique pink to light purple stripe visible on each (Fig. 12a). In females,

background body color generally a uniform pinkish red (Fig. 12b). Dorsal fins mostly transparent, usually with an

oblique whitish stripe on each. While female colors and patterns appear to be quite constant, males may completely

lose their white or pale blotches (Zander & Jelinek 1976; Fesser 1980; Miller 1986; Louisy 2015; Kovačić et al.

2016; Engin et al. 2017).

Similar species. Speleogobius llorisi.

Habitat. Infralittoral to circalittoral species, known from 8–48 m depth, most often on coralligenous beds and

concretion, possibly on scree made of boulders, sometimes on deep rocks, on detritic coarse sand or in caves; may

swim a few centimeters from the substrate (Zander & Jelinek 1976; Miller 1986; Le Bris et al. 2015; Louisy 2015;

Engin et al. 2017; Bérenger et al. 2019 and pers. com.; Duhau et al. 2020a).

Geographic distribution. Mediterranean. Presently known from southern France (Duhau et al. 2015; Le Bris et

al. 2015; Bérenger et al. 2019 and pers. com.; Iglésias et al. 2020), to the Aegean Sea (Engin et al. 2017), including

the northern and southern Tyrrhenian Sea (Duhau et al., 2020a, 2020b) and the Adriatic Sea (Zander & Jelinek 1976;

Miller 1986; Kovačić 1997).

KOVAČIĆ ET AL.

Thorogobius ephippiatus (Lowe, 1839) (Fig. 5)—Leopard-spotted Goby

Gobius ephippiatus Lowe, 1839: 84, type locality: eastern Atlantic, Madeira.

Size. Known adult size to 11 cm total length.

Morphology. D VI + I,11; A I,10; P 17–18. Body relatively short, laterally compressed. Proportionately large

head with a moderately steep snout profile. Anterior nostril tubular, with no dermal process from its rim. Caudal

peduncle deep, but lower than body depth. Dorsal fins of similar height, the first dorsal fin with more or less rounded

margin. The uppermost pectoral-fin rays are within the membrane, there are no free rays (Miller 1969; Schultz

1975). Scales present on body, usually poorly visible on photographs. Predorsal area naked.

Live coloration. Ground coloration grayish with a blue-green sheen on the back, covered with large and

round dark spots (Fig. 5). Head, including the predorsal area, covered with brown to dark orange, smaller round

spots, usually lighter and more reddish than the body blotches. Five large, dark brown, brown-purple to black

midlateral blotches, circular in shape in the Mediterranean and 6 orange-brown blotches longer than deep in the

Atlantic form (entering the Mediterranean just east of the Strait of Gibraltar). Above midline, 7-15 dark blotches

smaller than midlateral blotches. First dorsal fin blue-gray with 2 brown transverse bands (sometimes faint or

limited to brown spots). Dorsal, caudal and anal fins with a whitish to light blue margin (Schultz 1975; Kovačić &

Svensen 2018).

Similar species. Thorogobius macrolepis (or occasionally orange-spotted T. ephippiatus).

Habitat. Infralittoral to circalittoral species, known from 2–156 m depth, on muddy sand, gravel or detritic

substrata near crevices, beneath overhangs, in deep gullies or caves (Miller 1984; Stern et al. 2018; Renoult et al.

2022).

Geographic distribution. Northern Mediterranean, presently known from almost every rocky shore between

the Strait of Gibraltar in Spain and the Strait of Dardanelles in Turkey, including the Ligurian, Thyrrhean, Adriatic

(Schultz 1975, Kovačić et al. 2012a, Trkov et al. 2019) and Aegean Seas (Gerovasileiou et al. 2015), as well as

the Levant basin eastwards to Cyprus and Israel (Stern et al. 2018). Probably present in the Black Sea; relatively

widespread in the northern East Atlantic.

Thorogobius macrolepis (Kolombatović, 1891) (Fig. 6)—Large-scaled Goby

Gobius macrolepis Kolombatović, 1891: 4; type locality: Adriatic Sea, Croatia, Dalmatia.

Size. Known adult size about 6 cm total length.

Morphology. D V–VI + I,10–12; A I,10; P 17–20. Body relatively short, laterally compressed. Head moderately

long, with a steep snout profile. Anterior nostril tubular, with no dermal process from its rim. Caudal peduncle deep,

but lower than body depth. Dorsal fins of similar height, the first dorsal fin with more or less rounded margin. The

uppermost pectoral-fin rays are within the membrane with no free rays (Miller 1969; Ahnelt & Kovačić 1997).

Scales present on body, usually poorly visible on photographs. Predorsal area naked.

Live coloration. Ground coloration white to light gray with a light blue sheen on the back and pearly white

markings ventrally. Orange to yellow-orange spots on body (Fig. 6). Five brick red to orange midlateral

blotches, 2 to 3 times longer than deeper, each blotch sometimes appearing as 2 contiguous spots. Interspaces

between blotches often pearly white. Many short line segments, horizontal or oblique, each of 3-5 light orange

dots, above the lateral midline. Head and predorsal area covered with many orange spots larger than the dots

of the back but smaller than the midlateral blotches. No V-shaped mark on snout (Fig. 6). One orange, oblique

preorbital bar extending onto the upper lip; otherwise the lips are white. Dorsal fins blue-gray with three orange

transverse bands and a broader whitish to light blue marginal band. Caudal fin also blue gray with 3-4 sinuous

orange bars (Ahnelt & Kovačić 1997).

Similar species. Gobius kolombatovici, Thorogobius ephippiatus.

Habitat. Infralittoral and circalittoral species, known from 6–60 m depth, on detritic coarse sand and gravel,

close to rocky shelters, typically at the foot of rocky walls (Glavičić & Kovačić 2016; Renoult et al. 2022).

Geographic distribution. Northern Mediterranean, presently known from continental Spain (off Barcelona,

Colección de referencia de otolitos, Instituto de Ciencias del Mar-CSIC via GBIF) and the Balearic Island of Ibiza

(Ahnelt & Kovačić 1997), between Marseilles in France and Tuscany in Italy including Monaco (Francour 2007),

Lipari Island (Italy; Muriel Duhau unpublished observation), the Adriatic (Kolombatović 1891; Ahnelt & Kovačić

1997; Guidetti et al. 2006; Trkov et al. 2019), the Turkish Aegean coast (Francour et al. 2007; Bilecenoglu & Yokes

2016), and further east in that country in the Levant basin (Bilecenoglu & Yokes 2016).

Tridentiger trigonocephalus (Gill, 1859) (Fig. 9)—Chameleon Goby

Triaenophorus trigonocephalus Gill, 1859: 18; type locality: Hong Kong.

Size. Reaches 11 cm total length.

Morphology. D VI + I,11–13; A I,10–12; P 19–20, single uppermost ray free. Body moderately elongate,

nearly round anteriorly and laterally compressed posteriorly, depth 5.3–5.8 in standard length. Head large, somewhat

depressed, with relatively short blunt snout. Caudal peduncle deep, but lower than body depth. First to fifth spines

of first dorsal fin subequal in length, none elongate. Pelvic fins joined to form a disc. Caudal fin rounded, shorter

than head (Boltachev & Karpova 2017). Scales present on body, usually poorly visible on photographs. No scales

on head and prepectoral area; predorsal area naked.

Live coloration. Body brown to beige, usually with 2 continuous or interrupted dark brown stripes (Fig.

9), first beginning from just above orbit, continuing along back and ending at upper caudal-fin base, second stripe

extending from snout through eye along mid-side of body ending at caudal-fin midbase. Side of head with small

pale spots. Another color form with stripes interrupted by pale interspaces, short brown bars on back and the

black spot on upper caudal peduncle visible. A third color form is blackish with indistinct stripes.

Similar species. Gobius vittatus.

Habitat. Inhabits sand and stone bottoms in marine and brackish waters at depths of 1–5 m.

Geographic distribution. Native distribution range is confined to Japan, Korea and China. Introduced to eastern

Pacific, Australia and Black Sea in the Sevastopol Bay (Boltachev & Karpova 2017). In the Mediterranean there is

a single record from Ashdod, Israel (Goren et al. 2009).

Trypauchen vagina (Bloch & Schneider, 1801) (Fig. 1)—Pink Worm Goby

Gobius vagina Bloch & Schneider, 1801: 73; type locality: India, Tranquebar [Tharangambadi].

Size. Reaches 23 cm total length.

Morphology. D VI, 40–50; A39–47; P 15–20. Body very elongate and laterally compressed. A pouch-like

cavity just above operculum. Head laterally compressed with reduced eyes. Dorsal and anal fins low, long

and confluent with caudal fin. Pelvic fins fused medially, but connecting membrane between innermost rays

emarginated posteriorly (Murdy 2006). Scales on body cycloid, no scales on head. Scales small, visible on body and

predorsal area as pattern of pale dots.

Fresh coloration. Head and body entirely reddish or pinkish red (Fig. 1a).

Similar species. None of Mediterranean Gobiidae.

Habitat. Inhabits muddy or silty sand bottom in bays and brackish estuaries at depths of 10–90 m (Murdy 2006;

Ergüden et al. 2018).

Geographic distribution. Recorded from the Arabian Gulf east to Vietnam, Taiwan and the Philippines, unknown

from the Red Sea. In the Mediterranean it has been collected in Israel (Salameh et al. 2010) and Turkey (Ergüden

et al. 2018).

Vanderhorstia mertensi Klausewitz, 1974 (Fig. 4)—Mertens’ Shrimpgoby

Vanderhorstia mertensi Klausewitz, 1974: 210; type locality: Red Sea, Gulf of Aqaba, Egypt, Marsa Murach.

Size. Largest specimen 9.8 cm total length.

Morphology. D VI + I,16; A I,17–18; P 18. Body slender, its depth 6.2–7.3 in standard length. Head subcylindrical,

with short snout and large eyes elevated above dorsal profile. Caudal peduncle deep, but lower than body depth, and

KOVAČIĆ ET AL.

short compared to the elongate body shape. First dorsal fin moderately high, the fourth spine the longest, the

third to fifth spines filamentous. Pelvic fins united, not reaching anus. Caudal fin lanceolate, longer than head

length (Goren 1979). Scales visible on body only from reticulate pigmentation along scale edges, no scales on head

and predorsal area.

Live coloration. Body pale to grayish blue-green dorsally, most scales with a blackish edge and a yellow spot

varying from semicircular to covering most of scale, grading to white on abdomen; yellow-spotted scales grouping

anteriorly into vertically elongate yellow blotches; 5 large dark brown spots in a longitudinal row on side

of body, some of them diffuse (Fig. 4), the first below first dorsal fin and vertically elongate. A small black spot

behind upper end of gill opening, and another of the same size dorsally on opercle. Head and nape with numerous

dark-edged yellow spots. First dorsal fin with 2 rows of large yellow-orange spots. Second dorsal and caudal

fins with yellow spots basally on membranes.

Similar species. Cryptocentrus caeruleopunctatus, C. steinhardti.

Habitat. Inhabits silty sand bottom in bays and lagoons at depths of 2–52 m (deepest record from the

Mediterranean). Lives in symbiotic association with a snapping shrimp (Alpheus sp.) (Louisy 2015; Louisy et al.

2016; Tiralongo & Pillon 2019).

Geographic distribution. Known from the Red Sea to the Gulf of Oman. In the Mediterranean it has been

recorded in Greece (Tiralongo & Pillon 2019), Israel (Goren et al. 2013) and Turkey (Bilecenoglu et al. 2008).

Vanneaugobius dollfusi Brownell, 1978 (Fig. 19)—Dollfus’ Goby

Vanneaugobius dollfusi Brownell, 1978: 124, Figs. 1, 2, 4; type locality: Morocco, off Agadir.

Size. Maximum known size about 3.9 cm total length (Kovačić 2008b).

Morphology. D VI–VII + I,9–10; A I,8–9; P 16–17. A small species with a moderately elongated body. Caudal

peduncle deep, almost as deep as body. Dorsal fins of similar height, the first dorsal fin with more or less rounded

margin, can be pointed in males with the first spine longest (see Francour et al. 2010*). Caudal fin rounded, 2 large

spinous scales on upper and lower fin base (barely visible, even in close-up photographs). Pelvic fins are separated,

also hardly visible from lateral view. Scales present on body, usually poorly visible on photographs. Predorsal area

naked (Kovačić 2008b).

Live coloration. Head and body grayish with pale orange, reddish, rarely pinkish blotches and speckles, or

indistinct bars of same colors on a paler background (Fig. 19), sometimes faded depending on the environment and

the mood of the fish. Head mostly orange with irregular pale markings or speckles; the tubular anterior nostrils

are white. Usually 8–9 pale areas along back (from nape to caudal peduncle), sometimes becoming small white

saddles. A series of typical short pearly white bars arranged in 3-5 pairs on lower body (depending on how much

visible they are), usually interspersed with midlateral orange elongated blotches. First dorsal fin translucent with

orange-red dots along spines. Sometimes an elongate dark spot on anterior basal part of first dorsal fin, but not

always visible in the field; this dark spot on the first dorsal fin is present in both sexes, but seems more commonly

displayed by females (Van Tassell et al. 1988; Kovačić 2008b; Louisy 2015).

Similar species. Odondebuenia balearica, Corcyrogobius liechtensteini.

Habitat. Observed or collected underwater by SCUBA diving between 27–60 m depth, often found lying on

sediment at the foot of rocky or coralligenous walls or slopes, quickly hiding in a nearby cavity or shelter (Kovačić

2008b; Francour et al. 2010*; Le Bris et al. 2015; Glavičić & Kovačić 2016; Glavičić et al. 2016; Dubas et al.

2018). Most of the collected specimens were caught deeper, mostly with beam trawls between 54–160 m depth,

on different habitats: coralligenous or rocky grounds, maërl, deep algal beds, shell debris, sand, mud (Pallaoro &

Kovačić 2000; Ahnelt & Dorda 2004; Ordines et al. 2019a).

Geographic distribution. Eastern Atlantic and northern Mediterranean species. Types collected in Agadir,

Atlantic coast of Morocco (Brownell 1978). Known in the Mediterranean from the Balearic Islands (Ordines et al.

2019) to the Aegean Sea, Peloponnese (Ahnelt & Dorda 2004); records include France, Gulf of Lion (Le Bris et al.

2015), French Riviera (Duhau et al. 2020c), Corsica (Francour et al. 2010*), Italy, Strait of Sicily (Consoli et al.

2019), Malta (Kovačić & Schembri 2019) and the Adriatic Sea (Pallaoro & Kovačić 2000; Ahnelt & Dorda 2004;

Kovačić et al. 2012b).

* Although misidentified in the cited report, three good photographs allow for a positive identification of V.

dollfusi (which makes them the first French record of this species). See also Louisy (2015).

Zebrus pallaoroi Kovačić, Šanda & Vukić, 2021 (Fig. 29)—Pallaoro’s Goby

Zebrus pallaoroi Kovačić, Šanda & Vukić, 2021: 9, Figs. 1A, 2A–C, 3; type locality: Adriatic Sea, Montenegro, Boka Kotorska,

Kostanjica.

Note: This species was described recently and we lack information on its aspect and coloration in life; the present

account is thus provisional and may not allow for a fully reliable visual identification and differentiation from Z.

zebrus.

Size. Maximum known size 4.1 cm total length.

Morphology. D VI + I,10–11; A I,8–9; P 17. Small goby with a body more slender than in Zebrus zebrus, a large

and somewhat flattened head, and a deep caudal peduncle of almost same depth as body. Eyes relatively small, snout

slightly longer than eye diameter. Head wide, with large prominent cheeks (Fig. 29c): head width at opercle level

about 80–90 % of head length (compared to 50–75 % in Z. zebrus). Anterior nostril a tube with tentacle (Fig.

29c); posterior nostril tube only slightly shorter (more than 80 % of the anterior). Dorsal fins of similar height, the

first dorsal fin with more or less rounded margin. The uppermost pectoral-fin rays partly free from membrane

(Kovačić et al. 2021; present work). Caudal fin rounded. Body scaled, nape and predorsal area naked. Scales more

or less visible on body from reticulate pigmentation along scale edges.

Coloration. In dead or preserved specimens, body with 10−11 pale bars along side (Figs. 29a and 29b), first

in front of the first dorsal fin, last at the end of the second dorsal fin. Very little is known for sure about this species’

in vivo coloration. Overall live coloration dark to pale brownish or greenish with thin, alternating, pale and dark

bars (*). Usually, there are at least 4 pale bars under the first dorsal fin (Figs. 29a and 29b), but this pattern may

disappear, the sides of body becoming uniformly colored or mottled dark. When present, the crescent band on

head (broad pale transverse band across anterior nape extending on the sides reaching pectoral-fin bases) does not

extend forward to eyes. First dorsal fin usually with a whitish longitudinal band, topped with a contrasting orange-

yellow to pinkish distal half. Second dorsal fin with dark spots seldom coalescing in lines.

(*) The pattern of lateral bars visible on dead or stressed animals do not exactly match the number and location

of lateral markings usually displayed underwater (these 2 pattern types may superimpose to various extents).

Similar species. Zebrus zebrus, Millerigobius macrocephalus, Chromogobius zebratus.

Habitat. Shallow infralittoral species, known from 0–1 m depth between gravel and small boulders (Kovačić et

al. 2021, 2022). Also found at 3 m depth (J. Renoult, unpublished data).

Geographic distribution. Mediterranean. Recorded in the southern, central and northern part along the eastern

Adriatic Sea, the northern part of the Ionian Sea, and the northern and western parts of the Aegean Sea (Kovačić et

al. 2021, 2022). Also known from Palavas-les-Flots, southern France (J. Renoult, unpublished data).

Zebrus zebrus (Risso, 1827) (Fig. 28)—Zebra Goby

Gobius zebrus Risso, 1827: 282; type locality: northwestern Mediterranean Sea, France, Gross Island, Banyuls sur Mer.

Note: This species closely resembles the recently described Z. pallaoroi, which live coloration is almost unknown,

and some photographs attributed to Z. zebrus may actually depict Z. pallaoroi; the present account is thus provisional

and may not allow for a fully reliable visual differentiation of the two Zebrus species.

Size. Maximum known size about 5.5 cm total length (6.1 cm in the Atlantic, Nieto & Alberto 1992).

Morphology. D V–VI + I,11; A I,7–10; P 16–18. Small goby with a stocky body, a relatively large head and

a deep caudal peduncle. Eyes quite large, snout slightly shorter than eye diameter. Head narrower than in Z.

pallaoroi: head width at opercle level about 50–75 % of head length (vs. 80–90 % in Z. pallaoroi). Anterior nostril

a tube with tentacle; posterior nostril tube clearly shorter (less than 40 % of the anterior). Caudal peduncle deep,

almost same depth as body. Dorsal fins of similar height, the first dorsal fin with more or less rounded margin.

Uppermost pectoral-fin ray may be almost free from membrane (Miller 1977; Louisy 2015, Kovačić et al.

KOVAČIĆ ET AL.

2021; present work). Caudal fin rounded. Body scaled, nape and predorsal area naked. Scales more or less visible

on body from reticulate pigmentation along scale edges.

Live coloration. Head and body with a beige, brownish or greenish background. Body with 6–9 pale bars

along side (Fig. 28a), mostly visible on dorsal half (Miller 1977, Kovačić et al. 2021) (*). In the wild, unstressed

animals often display pale dorsal saddles while the rest of body is more or less uniformly colored or mottled with

dark pigment (Figs. 28b); upper lateral dark markings reach to the base of dorsal fins. When present, the

crescent band on head (broad pale transverse band across anterior nape extending on the sides until pectoral fin

bases) does not extend forward to eyes (Fig. 28b); it is usually quite well defined in shape and contours. First

dorsal fin usually with 2 dark horizontal lines (the lower one often disrupted) with a whitish band in between, outer

margin of the fin often with a reddish or yellowish tint. Second dorsal fin with brownish to reddish spots that may

coalesce into slightly oblique lines.

(*) The pattern of lateral bars visible on dead or stressed animals do not exactly match the number and location

of lateral markings usually displayed underwater (these two pattern types may superimpose to various extents).

Similar species. Zebrus pallaoroi, Millerigobius macrocephalus, Chromogobius zebratus.

Habitat. Infralittoral species most often encountered between 0.1–3 m depth (Miller 1977) but also recorded

down to 13 m (Engin et al. 2018a) and 36 m (Kesici & Dalyan 2019). Occurs in a variety of inshore habitats, under

stones or in rock clefts and cavities, in Cymodocea or Posidonia seagrass meadows (sheltering below shells), often

in coastal lagoons (Miller 1977, Patzner 1999a, Kovačić et al. 2012a, Trkov et al. 2019). May be found in intertidal

pools (Ahnelt 1990, Nieto & Alberto 1992, Trkov et al. 2019), and juveniles sometimes under sea urchins (Patzner

1999b).

Geographic distribution. Recorded along the Atlantic coastline of southwest Spain and south Portugal (Nieto

& Alberto 1992; Kovačić et al. 2021) and along northern Mediterranean coasts (Miller 1986), from Alicante to the

eastern Aegean Sea (Bogorodsky et al. 2010), the Levantine Sea from Rhodes (Miller 1977) to Syria (Saad 2005),

and from the southern Mediterranean in Tunisia and Libya (Mejri et al. 2007). It has also been recorded from

the Bosphorus Strait (Kesici & Dalyan 2019) and the Black Sea (Kovačić & Engin 2009; Boltachev & Karpova

2017).

Discussion

Positive identification from photographs has been rarely used in studies aiming at clarifying the geographic and

ecological distribution of gobies. A few exceptions for the Mediterranean and European waters include the records

of G. steinitzi by Kovačić et al. (2011), T. ephippiatus by Kovačić & Svensen (2018) and the study by Ragkousis et

al. (2021) for several cryptobenthic gobies. For these findings, based exclusively on photographs of specimens that

were not collected, the diagnosis necessarily relied on a combination of characters all visible on the photographs

that are necessary but sufficient to identify individuals (Kovačić & Svensen 2018). In the examples cited above, the

diagnosis allowed identifying individuals down to species level while considering all confamiliar species occurring

in the area as alternatives (Kovačić & Svensen 2018). Designing an effective diagnosis for species identification

based on color patterns of live individuals requires sorting out the various characters listed in the species descriptions,

redescriptions and published reports, which can be a complex task. The present key, even with only 41 out of

66 species of marine gobies from the Mediterranean continental shelf that can be positively identified down to

species level from photographs, should foster the use of underwater photographs to expand knowledge on the

geographic and ecological distributions of gobiid species in the Mediterranean. It is also the first attempt of its kind

for gobies in general, the most species-rich fish family worldwide present mostly in depths reachable by underwater

photographers.

Some of the species positively identified to the species level by the key are still considered rare, or at least

elusive, having a limited known geographic distribution or being known from only very few records. These include

some recently described species like Hazeus ingressus, Lebetus patzneri, Speleogobius llorisi or Zebrus pallaoroi

(Kovačić et al. 2016; Engin et al. 2018; Schliewen et al. 2019; Kovačić et al. 2021). The key provided by this study

could help unambiguously identifying new records for these species that could ultimately change our perception of

their status.

In published results of underwater visual censuses, gobiid species have been mostly addressed by the generic

or family name, or alternatively only a few most common and largest gobiid species have been recognized (Relini

et al. 1998; Guidetti & Bussotti 2002; Azzuro et al. 2013). Consequently, the real diversity of gobies and their

influence on the total species richness in the littoral Mediterranean fish assemblages has thus far probably remained

underestimated. The application of the present guide, with the key and the short accounts, should increase the

amount of gobiid species recognized in the underwater visual censuses and improve knowledge of this species-

richest fish family of the region.

Acknowledgements

We are very grateful to Helen Larson, who has improved and corrected this work by her gobiological knowledge and

with her editing skills. We are grateful to our colleagues, ichthyologists and underwater photographers, who kindly

provided photographs for the present work: C. Balisky, L. Bérenger, M. Colombo, D. Darom, M. Duhau, A. Falleni,

R. Groeneveld, S. Guerrieri, B. Guichard, J. Herler, S. Iglésias, Z. Jaafar, A. Kassar, G. Kunz, S. Le Bris, R. Mejri,

T. Menut, F. Ordines, A. Pagano, J. Randall, X. Rufray, Sascha Schulz/iNaturalist Australasian Fishes Project, C.

Telford, L.S. Tocino, R. Tournier-Broer, J. Vukić. Part of the information used in this work comes from observations

made by amateur volunteers and collected through participatory science programs such as the Fish Watch Forum

(www.fish-watch.org) and iNaturalist (www.inaturalist.org); we gratefully acknowledge their contributions. Several

photographs and observations from France could be made thanks to the financial support of UMS PatriNat/OFB/

MNHN and to a grant of the Gulf of Lion natural marine park.

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New records of rarely reported species in the Mediterranean Sea (March 2025)

Article

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Mar 2025

This collective study presents information on fifteen (15) species recorded across five countries (Greece, Italy, Slovenia, Spain, and Türkiye) and four (4) ecoregions, ranging from the Alboran Sea to the Aegean Sea. The recorded species belong to three phyla: Mollusca (3 species), Arthropoda (3 species), and Chordata (9 species). Among mollusks, Martadoris mediterranea is reported for the first time in the Aegean Sea at two different locations, while Opisthoteuthis calypso and Octopoteuthis sicula are newly recorded in the western Mediterranean. Regarding arthropods, Ocypode cursor is reported from the Tyrrhenian coast of Sicily, Pachygrapsus transversus is recorded for the first time in the Tyrrhenian Sea, and Cancer pagurus in the Sea of Marmara. Among chordates, Lobotes surinamensis and Zu cristatus are newly recorded in the Alboran Sea, while Callionymus fasciatus and Symphodus doderleini are reported from Slovenian coastal waters. Tripterygion delaisi and Gymnothorax unicolor are documented for the first time in the waters of the Sea of Marmara and Crete, respectively. Additionally, new records of rarely reported fish species include Arctozenus risso, Chromogobius zebratus, and Epinephelus aeneus in Turkish, Spanish, and Italian waters, respectively.

The northernmost finding of Gobius gasteveni Miller, 1974 (Teleostei: Gobiidae) and the first record for the Netherlands Ichthyological note -First records

Article

Feb 2025
CYBIUM

La présence la plus septentrionale de Gobius gaste-veni Miller, 1974 et le premier signalement pour les Pays-Bas Un premier signalement de Gobius gasteveni a été rapporté pour les Pays-Bas, ce qui étend de manière significative la distribution géographique connue de cette espèce. L'espèce a été obser-vée sur l'épave du SS Trevier, située à environ 30 miles à l'ouest d'iJmuiden dans la mer du nord, à une profondeur d'environ 28 m. Gobius gasteveni a été identifié à partir de photographies, en utili-sant une identification rigoureuse de l'espèce basée sur les caractè-res morphologiques et de coloration visibles sur les photographies.

Testing of the Taxonomic Diagnosis of Zebrus pallaoroi Kovačić, Šanda & Vukić, 2021 (Actinopteri: Gobiiformes: Gobiidae), on a Large Sample from the Western Mediterranean

Article

Full-text available

Dec 2024

A large sample of 65 individuals of the recently described goby Zebrus pallaoroi was collected in France. The species identity of these individuals was confirmed based on morphology. In addition, the species identity was validated through genetic analysis for one of the two specimens collected from the new depth records for the species. The diagnostic characters of Z. pallaoroi, originally based on a limited number of type specimens, were tested on this larger sample and critically analyzed. The diagnostic characters of Z. pallaoroi were revised, and recommendations were made for improving the study of diagnostic characters in gobiid species description, particularly when based on small sample sizes. The record of Z. pallaoroi in the western Mediterranean significantly expands its known geographic range, increases the maximum recorded depth of the species, and reveals syntopic co-occurrence with its phylogenetically closest relatives.

Convergent evolution in shape in European lineages of gobies

Article

Full-text available

Nov 2024

During their radiation, certain groups of animals evolved significant phenotypic disparity (morphological diversity), enabling them to thrive in diverse environments. Adaptations to the same type of environment can lead to convergent evolution in function and morphology. However, well-documented examples in repeated adaptations of teleost fishes to different habitats, which are not primarily related to trophic specialization, are still scarce. Gobies are a remarkable fish group, exhibiting a great species diversity, morphological variability, and extraordinary ability to colonize very different environments. A variety of lifestyles and body forms evolved also in European lineages of gobies. We conducted two-dimensional geometric morphometric and phylomorphospace analyses in European lineages of gobies and evaluated the extent of convergent evolution in shape associated with adaptation to various habitats. Our analyses revealed the change in shape along the nektonic-cryptobenthic axis, from very slender head and body to stout body and wide head. We showed convergent evolution related to mode of locomotion in the given habitat in four ecological groups: nektonic, hyperbenthic, cryptobenthic and freshwater gobies. Gobies, therefore, emerge as a highly diversified lineage with unique lifestyle variations, offering invaluable insights into filling of ecomorphological space and mechanisms of adaptation to various aquatic environments with distinct locomotion requirements.

New records of the recently described Pomatoschistus nanus Engin & Seyhan, 2017 (Teleostei: Gobiidae)

Article

Full-text available

May 2024
CYBIUM

Nouvelles données pour l'espèce récemment décrite Pomatoschistus nanus Engin & Seyhan, 2017 (Teleostei : Gobii-dae). de nouvelles données sur le gobie Pomatoschistus nanus engin & Seyhan, 2017 récemment décrit de Méditerranée sont présentées. L'espèce est identifiée grâce à la génétique, la morphologie et la coloration. Les données de Malte et de Chypre étendent significa-tivement l'aire de distribution connue de l'espèce. la donnée de Malte étend également la limite bathymétrique supérieure ainsi que la taille maximale connue pour l'espèce, actuellement considérée comme la plus petite de Méditerranée. la variabilité morphologi-que de P. nanus est également discutée.

Primera cita de un ejemplar de góbido del género Zebrus (de Buen,1930) en el mar Cantábrico

Article

Oct 2024

Alfredo Herrero

En las últimas décadas, se viene registrando en el Atlántico noreste un cada vez mayor número de citas de especies ictiológicas de aguas cálidas más meridionales. Diversos trabajos relacionan este fenómeno con el aumento de la temperatura de las costas europeas como consecuencia del calentamiento global. En este contexto, se documenta para el mar Cantábrico la primera cita de un ejemplar de góbido del género Zebrus de distribución principalmente mediterránea y con tan solo unas pocas citas en el océano Atlántico.

Checklist, key and latitudinal distribution of the eastern Atlantic marine gobies along the European coast (Actinopterygii: Gobiidae and Oxudercidae)

Article

Mar 2025
ZOOTAXA

Marcelo Kovačić

The checklist of 34 marine gobiid species (Actinopterii: Gobiidae and Oxudercidae) with confirmed records is compiled for the Atlantic coast of Europe, including four alien gobiid species found in transitional waters or in the low salinity waters of the Baltic Sea. An identification key for the gobiid species known from the area is also provided. The latitudinal distribution of native goby species in this area is presented and discussed. Finally, the second northernmost record of a Gobiidae fish is reported, corresponding to Pomatoschistus minutus (Pallas, 1770) from Tromsø, Norway.

Gobies on the Turkish coast of the Black Sea, and first record of the yellow-headed goby, Gobius xanthocephalus Heymer & Zander, 1992

Article

Aug 2024

First record of leopard-spotted goby Thorogobius ephippiatus (Gobiiformes, Gobiidae) above the Arctic circle

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Jul 2024

Fishes in Marine Caves

Article

Full-text available

Jun 2024

Fishes in marine caves have attracted limited attention by the scientific community in comparison to subterranean fishes which have lost eyes and pigmentation. They constitute a largely unexplored component of marine fish diversity, except for the relatively well-studied marine caves of the Mediterranean Sea. These habitats are characterized by steep environmental gradients of decreasing light and decreasing water exchange. The fishes recorded so far in marine caves are not exclusive residents of this habitat and they are also present at least in the other mesolithial habitats. In the Mediterranean marine caves, 132 fishes have been recorded to date, representing about 17% of the total Mediterranean fish species richness. Most of these species are reported from the anterior cave zones where some light still exists, while a smaller number of species are known from the totally dark zones. Among them, 27.3% are accidental visitors, 53.8% are the regular mesolithial visitors and switchers between mesolithion and open water, 5.3% are permanent residents of the mesolithion, but also occur in other habitats, and 13.6% are exclusive permanent residents of mesolithion. Some mesolithial exclusive permanent residents recorded in marine caves share similar morphology, probably as adaptations to these habitats.

New records of recently described Zebrus pallaoroi (Actinopterygii: Gobiiformes: Gobiidae) with notes on its morphology, ecology, and molecular identification

Article

Full-text available

Mar 2022

The family Gobiidae represents the species-richest family of fishes in the Mediterranean Sea. However, many Mediterranean gobiid species are poorly known with just one or few species records in total. Among those species, the recently described Zebrus pallaoroi Kovačić, Šanda et Vukić, 2021 has been known only from the localities in the species description: the southern Adriatic, northern Ionian, and northern and western Aegean seas. The additional specimens of this species were collected in central and northern Adriatic Sea. The identification was confirmed by morphological and genetic methods. The morphological and preserved coloration characters were compared with the data from the limited type material and the alternative preserved coloration diagnostic character was suggested. The first data on alive and freshly dead coloration are reported. The phylogenetic analysis was performed on sequenced COI , cytochrome b, and rhodopsin genes. The analyzed molecular markers placed Z. pallaoroi in a distinct clade within Zebrus – Millerigobius group. The preference of this species for the very shallow habitat is confirmed and discussed.

Cryptocentrus steinhardti (Actinopterygii; Gobiidae): a new species of shrimp-goby, and a new invasive to the Mediterranean Sea

Article

Full-text available

Sep 2021

A new species of shrimp-goby was collected at depths of 60–80 m off the southern Israeli Mediterranean coast. A unique ‘DNA barcoding’ signature (mtDNA COI and Cytb ) revealed that it differs from any other previously bar-coded goby species clustered phylogenetically with the shrimp-gobies group, in which Cryptocentrus is the most speciose genus. A morphological study supported the assignment of the fish to Cryptocentrus and differentiated the new species from its congeners. The species is described here as Cryptocentrus steinhardti n. sp. However, the present phylogenetic analysis demonstrates a paraphyly of Cryptocentrus and emphasizes the need for revision of the genus based on integrating morphological and genetic characteristics. This finding constitutes the third record of an invasive shrimp goby in the Mediterranean Sea. An intriguing ecological issue arises regarding the possible formation of a fish-shrimp symbiosis in a newly invaded territory. Describing an alien tropical species in the Mediterranean prior to its discovery in native distribution is an unusual event, although not the first such case. Several similar examples are provided in the present article.

A checklist of the gobioid fishes of the world 2021

Article

Full-text available

Jul 2021

Paolo Parenti

Rarely Reported Cryptobenthic Fish in Marine Caves of the Eastern Mediterranean Sea

Article

Full-text available

May 2021

Data on the distribution and ecology of cryptobenthic fish of marine caves in the Mediterranean Sea are extremely scarce but necessary for scientists and marine managers alike in order to understand these fish’s ecological role and assess their conservation status. Broadscale surveys by implementing underwater visual census and photographic sampling in marine caves of the northeastern Mediterranean Sea, within different expeditions during the last 5 years, brought to light new records of eight rarely reported cryptobenthic fish species. To a smaller extent, complementary citizen science data from diving professionals of Crete were used to fill distribution gaps. A total of 36 new records (66 individuals) from 18 marine caves and caverns of the Aegean and northeastern Levantine Seas were assembled, belonging to the gobies Corcyrogobius liechtensteini, Didogobius splechtnai, Gammogobius steinitzi, and Thorogobius ephippiatus, the blenny Microlipophrys nigriceps, the tripterygiid Tripterygion melanurum, the speleophilic bythitid Grammonus ater, and the gobiesocid Lepadogaster cf. lepadogaster. The above species have been rarely reported from the Eastern Mediterranean Sea, with D. splechtnai and G. steinitzi being recorded for the first and second time from Greek waters, respectively, while L. cf. lepadogaster constitutes the second record of a clingfish species in a marine cave of the Aegean Sea. Interesting behavioral and ecological habits were also noted for some species, based on in situ observations and photographic evidence. Our study contributes to filling gaps in the knowledge of cave fish diversity and demonstrates that cryptobenthic mobile species in understudied cryptic habitats are more common than previously thought in the Mediterranean Sea.

Zebrus pallaoroi sp. nov.: a new species of goby (Actinopterygii: Gobiidae) from the Mediterranean Sea with a dna-based phylogenetic analysis of the Gobius-lineage

Article

Full-text available

Mar 2021
CONTRIB ZOOL

The gobies (Gobiidae) are the most diverse fish family in the Mediterranean Sea. Nevertheless, knowledge on their diversity, taxonomy, and phylogenetic relationships is still inadequate. The phylogenetic analyses reveal two genetically highly distinct clades among specimens identified as Zebrus zebrus . A new species, Zebrus pallaoroi sp. nov., is described based on an integrative approach. The neotype of Zebrus zebrus is designated. Genetic data confirm a pronounced level of divergence between Z. pallaoroi and Z. zebrus , with the mean genetic distance on cytochrome b being 18.1% and 1.07% on rhodopsin. Phylogenetic relationships within the Gobius -lineage were estimated on both markers. Morphologically, Z. pallaoroi is distinguished from the only congener Z. zebrus by having a snout longer than its eye, posterior nostril about 4/5–9/10 of the anterior nostril, eye diameter 4.3−4.7 in head length, ventrolateral head ridges transversally connected on the anterior side by a short transversal ridge, anterior membrane midline depth about 2/3 of the spinous ray, head canal pore α diameter about half of the distance between pore ρ and ρ 1 , suborbital sensory papillae row 5i going downwards to or near the level of row d , the distance between row 5i and row d absent or much smaller than the length of row 5i , and the body with ten to eleven vertical dark brown bands. Zebrus pallaoroi was recorded from the southern Adriatic, northern Ionian, and northern and western Aegean Seas, and is a cryptobenthic fish from very shallow waters.

New distributional records of Gobius bucchichi (Pisces, Gobiidae) from the Mediterranean Sea and in situ comparisons with Gobius incognitus

Article

Full-text available

Dec 2020

We report for the first time the presence of Gobius bucchichi from the coastal waters of three Mediter-ranean countries, namely Italy, Slovenia and Greece, thus increasing the knowledge on the distribution of this species in the Mediterranean Sea. We provide high quality in situ photos of the species, highlighting the main differences between similar species as a useful tool for in situ identification, which can be used for monitoring purposes. Habitat preferences, abundance and behaviour of G. bucchichi and similar species are also described. Comparisons with similar species, ecological notes, and the current distribution of G. bucchichi in the Mediterranean Sea are discussed. NUOVE SEGNALAZIONI DI GOBIUS BUCCHICHI (PISCES, GOBIIDAE) NEL MAR MEDITERRANEO E CONFRONTI IN SITU CON GOBIUS INCOGNITUS SINTESI Riportiamo per la prima volta la presenza di Gobius bucchichi nelle acque costiere di tre nazioni medi-terranee, ovvero Italia, Slovenia e Grecia, apportando nuove conoscenze sulla distribuzione di questa specie nel Mediterraneo. Forniamo anche foto della specie in ambiente di elevata qualità, evidenziando le principali differenze con le specie simili, fornendo quindi uno strumento utile per l'identificazione in situ della specie che può essere usato a fini di monitoraggio. Le preferenze di habitat, l'abbondanza e il comportamento di G. bucchichi e delle specie simili sono anch'essi descritti. Confronti con le specie simili, note ecologiche e l'attuale distribuzione di G. bucchichi nel Mediterraneo vengono discussi. Parole chiave: chiavi di identificazione, gobidi mediterranei, nuove segnalazioni, Adriatico, Egeo, fotografia subacquea ANNALES · Ser. hist. nat. · 30 · 2020 · 2

French ichthyological records for 2018

Article

Full-text available

Dec 2020
CYBIUM

Uncommon records of fishes may evidence local or global changes in fish composition resulting from environmental change or anthropogenic activities. Significant records of uncommon marine fishes, including migrant, non-native, cryptic, rare and threatened species, collected in french waters or by french vessels in European waters, observed by scuba divers or beachgoers, are reported for the year 2018. They include first, new, rare and unusual records for the following 52 species: Hexanchus griseus, Squatina squatina, Gymnura altavela, Acipenser sturio, A. gueldenstaedtii, Dalophis imberbis, Nemichthys curvirostris, Eurypharynx pelecanoides, Maulisia mauli, M. microlepis, Sagamichthys schnakenbecki, Melanostomias bartonbeani, Astronesthes niger*, Leptostomias gladiator*, Chlorophthalmus agassizi, Magnisudis atlantica, Evermannella balbo, egalecus glesne, Luvarus imperialis, Raniceps raninus, Fistularia cf. petimba*, Trigla lyra, Lepidotrigla dieuzeidei, Micrenophrys lilljeborgii, Howella atlantica*, Pomatomus saltatrix, Serranus cabrilla, Caranx crysos, Seriola rivoliana, Trachinotus ovatus, Lobotes surinamensis, Sarpa salpa, Chelon ramada, Pseudoscopelus altipinnis*, Trachinus draco, Parablennius pilicornis, P. ruber, Buenia affinis, B. jeffreysii, Chromogobius zebratus, Didogobius splechtnai, Gammogobius steinitzi, Gobius couchi, G. kolombatovici, Lebetus sp., Speleogobius trigloides, Thorogobius macrolepis, Vanneaugobius dollfusi, Siganus rivulatus*, Lepidocybium flavobrunneum, Hyperoglyphe sp. et Pegusa cf. nasuta, among which six, marked with an asterisk (*), represent additions to the Checklist of the marine fishes from metropolitan France and one, N. curvirostris, is a first record for the Mediterranean.

An updated Checklist of the Mediterranean fishes of Israel, with illustrations of recently recorded species and delineation of Lessepsian migrants

Article

Apr 2021

Daniel Golani

This checklist of the Mediterranean fishes of Israel enumerates 469 species which is an addition of 62 species since the previous checklist of 2005. This new checklist includes 58 Condrichthys and 411 Osteicthys species. Most newly-recorded species are of Red Sea origin (Lessepsian migrants)—38 species, 25 species are from previously poorly investigated habitats, mainly deep water, while two species reached the Mediterranean most likely by ballast water and two are aquaculture escapees. The dramatic increase in the number of Lessepsian migrants (an average of 2.5 species per year) is most likely due to the increased water influx between the Red Sea and the Mediterranean, following the recent opening of the new parallel, 72 km, “new canal” and the enlargement of other parts of the Suez Canal.

Gobius xoriguer, a new offshore Mediterranean goby (Gobiidae), and phylogenetic relationships within the genus Gobius

Article

Feb 2021
ICHTHYOL RES

Gobius xoriguer sp. nov., a new offshore species of goby (Teleostei: Gobiidae: Gobiinae) is described based on three specimens collected in 2010, 2012 and 2018 in the western Mediterranean, off Menorca (Spain), in the Gulf of Lion and off Corsica (France), at 51–104 m depth on coralline algae sea bed. It is easily distinguishable from its Atlantic-Mediterranean congeners by the combination of the following characters: large eyes, ~27-28% of head length; anterior nostril with a small triangular process; 14 soft rays on D2, 13 soft rays on A; enlarged first dorsal fin rays (adult males), with third D1 spine the longest, 23-27 % SL; uppermost P fin rays not free from membrane; long V-shape pelvic fins with vestigial frenum; 50-51 scales on LL; head oculoscapular canal with pores σ, λ, κ, ω, α, β, ρ, ρ1, ρ2, and preopercular canal with pores γ, δ, ε present; row x1 ending anteriorly behind pore β; a groove section between pore ρ and ρ1; suborbital row d discontinuous with large gap below suborbital rows 3 and 4; rows o separated; seven enlarged orange blotches on body side; white dotes on cheek and opercle on an orange background.With a known maximum size of 64 mm TL, it is among the smaller species of Gobius. Bayesian inference and Maximum likelihood phylogenetic tree topologies based on mitochondrial DNA COI sequences (barcoding region), including most Atlantic-Mediterranean Gobius species, support Gobius gasteveni Miller 1974 as the closest relative to Gobius xoriguer sp. nov. These sister species exhibit a high genetic divergence of 9.5% (uncorrected p-distance).

Checklist of gobies (Teleostei: Gobiidae) of the Mediterranean Sea and a key for species identification

Article

Nov 2020

Marcelo Kovačić

A checklist of 73 gobiid species (Teleostei: Gobiidae) recorded to date from the Mediterranean Sea is established following the evidence approach for checklists. The Mediterranean gobiofauna currently has 62 known native species and 11 alien species. An identification key to gobiid species known from the area is provided. The principles of character selection for the key are discussed.

Identification of Mediterranean marine gobies (Actinopterygii: Gobiidae) of the continental shelf from photographs of in situ individuals

Abstract

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