Gobiidae

Kovačić, Marcelo, Svensen, Rudolf, Renoult, Julien P. & Patzner, Robert A., 2025, Checklist, key and latitudinal distribution of the eastern Atlantic marine gobies along the European coast (Actinopterygii: Gobiidae and Oxudercidae), Zootaxa 5609 (1), pp. 1-28 : 10-14

Checklist of Gobiidae and Oxudercidae of the Atlantic coast of Europe

The checklist includes 30 native marine species of gobies (Actinopterii: Gobiidae and Oxudercidae ) confirmed from the Atlantic coast of Europe, alongside four alien species recorded in transitional waters or in the low salinity waters of the Baltic Sea ( Table 1 ). Of the 34 species, 22 are confirmed based on both morphological and genetic data, and with specimen(s) stored in a collection. Eight of these 22 species have their type locality in the area, with type material currently stored in collections ( Table 1 ). For one of these 22 species, Pomatoschistus marmoratus (Risso, 1810) , the morphological data were not published, but one of the authors (MK) conducted a morphological study of the material stored in a public collection ( Table 1 ). Four species are confirmed by a combination of published morphological or genetic data and independent specimen(s) stored in a collection, while another four are confirmed solely by published morphological or genetic data. Two species are confirmed based only on photographic evidence ( Table 1 ). Lastly, two common and long-known species from the area surprisingly lack published morphological evidence for their populations along the Atlantic coast of Europe; they are confirmed based on criterion of material stored in collections ( Table 1 ).

Key to the species of Gobiidae and Oxudercidae of the Atlantic coast of Europe

Alien species in the key are marked with an asterisk (*).

1a Head canals with additional pore π on snout ( Fig. 1 )..................................... Tridentiger bifasciatus * ( Fig. 6a )

1b Head canals without pore π on snout.......................................................................... 2

2a (1) First dorsal fin absent, or with 3 or fewer spines ( Fig. 7 )............................. Crystallogobius linearis ( Fig. 6b )

2b (1) First dorsal fin with 4 or more spines....................................................................... 3

3a (2) Anterior nostril long, projecting forward clearly in front of lips......................... Proterorhinus cf. nasalis ( Fig. 6c )

3b (2) Anterior nostril short, at best overlying posterior edge of upper lip when depressed.................................. 4

4a (3) First dorsal fin with 7 or 8 spines.......................................................................... 5

4b (3) First dorsal fin with 6 or fewer spines...................................................................... 6 5a (4) Scales absent on body.............................................................. Gobiosoma bosc * ( Fig. 6d )

5b (4) Scales present on body...................................................... Pomatoschistus flavescens ( Fig. 8a )

6a (4) Suborbital sensory papillae with row a below eye present ( Fig. 1b )............................................... 7

6b (4) Suborbital sensory papillae without row a below eye ( Figs. 1b and 9 ).............................................21

7a (6) Pelvic disc without anterior transverse membrane (frenum) ( Fig. 5 )............................................... 8

7b (6) Pelvic disc with anterior transverse membrane (frenum) ( Fig. 5 ).................................................. 9

8a (7) Anal fin I/7–9, pectoral fin 17–21............................................. Lebetus scorpioides ( Fig. 6e and 6f )

8b (7) Anal fin I/4–6, pectoral fin 14–17................................................ Lebetus guilleti ( Fig. 6g and 6h )

9a (7) Head canals and pores absent ( Figs. 1a and 10 ).............................................................. 10

9b (7) Head canals and pores present ( Figs. 1a and 11 )............................................................. 12

10a (9) Suborbital transverse rows of sensory papillae present ( Fig. 10 )................................ Aphia minuta ( Fig. 8b )

10b (9) Suborbital transverse rows of sensory papillae absent, suborbital rows only longitudinal ( Fig. 12 )..................... 11

11a (10) Pectoral fin 22............................................................... Lesueurigobius sanzi ( Fig. 8c )

11b (10) Pectoral fin 18–19........................................................... Lesueurigobius friesii ( Fig. 8d )

12a (9) Head canals with numerous extra pores (more than 22 pores) ( Fig. 13 )........................................... 13

12b (9) Head canals with usual number (22 or fewer) of pores ( Figs. 1a and 14 )......................................... 14

13a (12) Second dorsal fin I/8–9; anal fin I/9....................................... Deltentosteus quadrimaculatus ( Fig. 8e )

13b (12) Second dorsal fin I/10–11; anal fin I/10......................................... Deltentosteus colonianus ( Fig. 8f )

14a (12) Suborbital transverse rows of sensory papillae absent ( Fig. 14 )................................................ 15

14b (12) Suborbital transverse rows of sensory papillae present ( Fig. 11 )............................................... 16

15a (14) Anal fin I/6–7; scales in transverse series 5–6 ( Fig. 2 )..................................... Buenia massutii ( Fig. 8g )

15b (14) Anal fin I/9; scales in transverse series 7 ( Fig. 2 )......................................... Buenia jeffreysii ( Fig. 8h )

16a (14) Longitudinal scale count 34–52 ( Fig. 2 ).................................................................. 17

16b (14) Longitudinal scale count 55–75 ( Fig. 2 ).................................................................. 19

17a (16) First dorsal fin base naked posteriorly to the last spine ( Fig. 3 )...................... Pomatoschistus microps ( Fig. 15a )

17b (16) First dorsal fin base scaled at least in part ( Fig. 3 ).......................................................... 18

18a (17) First dorsal fin base naked backwards to the last spine ( Fig. 3 ), rear edge of pelvic fin anterior membrane (frenum) crenate to villose ( Fig. 5 )........................................................ Pomatoschistus marmoratus ( Fig. 15b )

18b (17) First dorsal fin base scaled to spine III–IV ( Fig. 3 ), rear edge of pelvic fin anterior membrane (frenum) smooth ( Fig. 5 )............................................................................... Pomatoschistus pictus ( Fig. 15c )

19a (16) Suborbital sensory papillae row b ends anteriorly below posterior part of eye, pectoral fin 16–18 ( Fig. 1b ).................................................................................... Pomatoschistus norvegicus ( Fig. 15d )

19b (16) Suborbital sensory papillae row b ends anteriorly below anterior part of eye, pectoral fin 18–21 ( Fig. 1b ).............. 20

20a (19) Only most posterior suborbital transverse row c extending below row d ( Fig. 1b )....... Pomatoschistus minutus ( Fig. 15e )

20b (19) Three suborbital transverse rows, including most posterior row c, extending below row d ( Fig. 1b )............................................................................................. Pomatoschistus lozanoi ( Fig. 15f )

21a (6) All three head canals present ( Fig. 1a ).................................................................... 22

21b (6) At least one of the three head canals absent ( Fig. 1a )........................................................ 34

22a (21) Anterior dorsal row g of sensory papillae ends anterior to lateral end of row o ( Figs. 1b and 8 )............................................................................................... Thorogobius ephippiatus ( Fig. 15g )

22b (21) Anterior dorsal row g of sensory papillae ends behind or on lateral end of row o ( Fig. 1b ).......................... 23

23a (22) Seven suborbital transverse rows c of sensory papillae ( Fig. 1b )............................................... 24

23b (22) Six suborbital transverse row c of sensory papillae ( Fig. 1b ).................................................. 26

24a (23) Predorsal area scaled ( Fig. 3 ).............................................. Neogobius melanostomus * ( Fig. 15h )

24b (23) Predorsal area naked ( Fig. 3 )........................................................ 25 ( Zebrus sp. , Fig. 16a )

25a (24) Depth of the pelvic anterior membrane at midline more than 0.5 of the depth of the pelvic spinous ray ( Fig. 5 ), four to five lateral dark bands in front of the vertical of the second dorsal fin.................................... Zebrus pallaoroi

25b (24) Depth of the pelvic anterior membrane at midline less than 0.5 of the depth of the pelvic spinous ray ( Fig. 5 ), three or less lateral dark bands in front of the vertical of the second dorsal fin...................................... Zebrus zebrus

26a (23) Anterior oculoscapular head canal with short side branch to pore α below eye ( Fig. 1a )....... Gobius paganellus ( Fig. 16b )

26b (23) Anterior oculoscapular head canal with pore α at rear of orbit ( Fig. 1a )......................................... 27

27a (26) Oculoscapular row xa of sensory papillae extending forward to pore β or anteriorly to it ( Fig. 1 )................................................................................................... Gobius cruentatus ( Fig. 16c )

27b (26) Oculoscapular row xa of sensory papillae ending forward posteriorly to pore β ( Fig. 1 )............................ 28

28a (27) Longitudinal scale count ≥50 ( Fig. 2 ).................................................................... 29

28b (27) Longitudinal scale count <50 ( Fig. 2 )................................................................... 30

29a (28) Head and body variegated, finely mottled greenish on beige, with fairly invariable pattern of large and irregular dark midlateral blotches. The mottled pattern and midlateral blotches visible as dark grey or brown in preserved material................................................................................................ Gobius cobitis ( Fig. 16d )

29b (28) Head and body uniformly pale gray, greenish gray or light brown, covered with longitudinal lines of brown dots. The brown dots also visible in preserved specimens............................................. Gobius incognitus ( Fig. 16e )

30a (28) Predorsal area naked ( Fig. 3 )........................................................ Gobius roulei ( Fig. 16f )

30b (28) Predorsal area scaled ( Fig. 3 ).......................................................................... 31

31a (30) Suborbital row d continuous ( Fig. 1b ).................................................. Gobius niger ( Fig. 16g )

31b (30) Suborbital row d divided ( Fig. 1b )...................................................................... 32

32a (31) Length of pelvic disc emargination more than 1/8 the disc length ( Fig. 4 ).............. Gobius xanthocephalus ( Fig. 16h )

32b (31) Pelvic disc complete or length of emargination less than or equal 1/8 the disc length ( Fig. 4 )........................ 33

33a (32) Pectoral fin 16–17................................................................. Gobius couchi ( Fig. 17a )

33b (32) Pectoral fin 19–21.............................................................. Gobius gasteveni ( Fig. 17b )

34a (21) All scales ctenoid; no pectoral fin bar; intense dark spot on lower anterior of operculum..... Chromogobius britoi ( Fig. 17c )

34b (21) Scales mostly cycloid, ctenoid only posteriorly; pectoral-fin bar present; spot on lower anterior corner of operculum absent......................................................................... Chromogobius zebratus ( Fig. 17d )

Figures

FIGURE 1. Terminology and position of head canal, pores, and papillae rows of lateral-line system. A) Head canals with pores. Head anterior oculoscapular canal pores π, σ, λ, κ, ω, α, β, ρ; posterior oculoscapular canal pores θ, τ; preopercular canal pores γ, δ, ε. B) Rows of sensory papillae. Preorbital: r, sa, sb, sc, ja, jb, jc, jd, suborbital: a, b, c (if c rows transverse: c1, c2, c3, c4, c5, c6...), d; preoperculo-mandibular: e, i, f; oculoscapular: xa, xb, ta, tp, z, q, y, ka, kb, kc, la, lb; opercular: ot, os, oi; anterior dorsal: n, o, g, m, h; interorbital: p. Modified from Kovačić (2020) by M. Kovačić.

FIGURE 2. Count of scales in lateral and transverse series, marked as a pale stripe. Modified from Kovačić (2020) by M. Kovačić.

FIGURE 3. Location of regions where the presence or absence of scales serves as a character in the key. The example of (a) partially scaled cheek and completely scaled: (b) opercle, (c) predorsal area, (d) the base of the first dorsal fin between spine I and spine VI and (e) base of the second dorsal fin from the first spine to the posteriormost ray. From Kovačić (2020).

FIGURE 4. Shape of pelvic fins. A) Pelvin fins almost completely separated, B) -D) pelvic fins fused into a pelvic disc. Pelvic disc B) rounded, C) truncate and D) emarginate. Pelvic disc emargination (e) is calculated as e=(b-a)/b, (a) the shortest branch of the fifth branched ray measured from the base of this ray, (b) the longest branch of the fourth branched ray measured from the base of this ray. From Kovačić (2020).

FIGURE 5. Pelvic disc anterior membrane (frenum). The membrane can be A) absent, or B) and C) present, B) without lateral lobes, or C) with lateral lobes. Absence or presence of anterior membrane in B) and lateral lobe in C) marked with a grey arrow. Pelvic disc anterior membrane height in midline (a) compared to the length of the spinous ray (b) in red. From Kovačić (2020).

FIGURE 6. A) Tridentiger bifasciatus (H.-B. Jeon). B) Crystallogobius linearis (R. Svensen). C) Proterorhinus cf. nasalis from the northern coast of Estonia, in the Gulf of Finland (K. Kurina). D) Gobiosoma bosc (C. Crippen). E) Lebetus scorpioides male (R. Svensen). F) Lebetus scorpioides female (R. Svensen). G) Lebetus guilleti male (J. Herler). H) Lebetus guilleti female (J. Herler).

FIGURE 7. Example of the first dorsal fin with two spines in Crystallogobius linearis. Material stained. From Kovačić (2020).

FIGURE 8. A) Pomatoschistus flavescens (R. Svensen). B) Aphia minuta (R. Svensen). C) Lesueurigobius sanzi (A. Kassar). D) Lesueurigobius friesii (R. Svensen). E) Deltentosteus quadrimaculatus (R. Pillon). F) Deltentosteus colonianus (S. le Bris). G) Buenia massutii (P. Girard). H) Buenia jeffreysii (R. Svensen).

FIGURE 9. Example of suborbital sensory papillae without row a below eye in Thorogobius ephippiatus. Photo by M. Kovačić.

FIGURE 10. Example of head canals and pores absent and suborbital transverse rows of sensory papillae present in Aphia minuta. Material stained. Photo and modification by M. Kovačić.

FIGURE 11. Example of head canals and pores, and suborbital transverse rows of sensory papillae both present in Pomatoschistus sp. Material stained. Photo and modification by M. Kovačić.

FIGURE 12. Example of suborbital transverse rows of sensory papillae absent in Lesueurigobius friesii. Photo and modification by M. Kovačić.

FIGURE 13. Example of the head canals with numerous extra pores in Deltentosteus quadrimaculatus. Photo and modification by M. Kovačić.

FIGURE 14. Example of suborbital transverse rows of sensory papillae absent in Buenia jeffreysii. Photo and modification by M. Kovačić.

FIGURE 15.A) Pomatoschistusmicrops (J. Renoult). B) Pomatoschistusmarmoratus (R. Pillon).C) Pomatoschistus pictus (R. Svensen). D) Pomatoschistus norvegicus (R. Svensen). E) Pomatoschistus minutus (R. Svensen). F) Pomatoschistus lozanoi (S. Iglesias). G) Thorogobius ephippiatus (R. Svensen). H) Neogobius melanostomus (S. Bogorodsky).

FIGURE 16. A) Zebrus sp. from Playa de Hondarraitz, Hendaye, France (A. Herrero). B) Gobius paganellus (R. Svensen). C) Gobius cruentatus (R.A. Patzner). D) Gobius cobitis (R.A. Patzner). E) Gobius incognitus (R. Svensen). F) Gobius roulei (R.A. Patzner). G) Gobius niger (R.A. Patzner). H) Gobius xanthocephalus (R.A. Patzner).

FIGURE 17. A) Gobius couchi (S. Guerrieri). B) Gobius gasteveni (J. Renoult). C) Chromogobius britoi (D. Rabeling). D) Chromogobius zebratus (R.A. Patzner).