taxonID	type	description	language	source
03ED255D3B3EFFAB99BF681AFE345066.taxon	description	Figure 10 a – d 1980 Panturichthys subglaber (Schubert, 1906) — Nolf & Martinell, pl. 1 Figs. 3, 6 (4, 5?). 1989 Panturichthys subglaber (Schubert, 1906) — Nolf & Cappetta: pl. 1, Fig. 8. 1998 Panturichthys subglaber (Schubert, 1906) — Nolf, Mané & Lopez, pl. 1, Fig. 4. 2020 Panturichthys sp. — Agiadi & Albano: Figs. 4.3. 2022 Panturichthys subglaber (Schubert, 1906) — van Hinsbergh & Hoedemakers: pl. 1, Fig. 3. Material 4 specimens SMF PO 101.184, Zanclean, Dar bel Hamri. Discussion Heterenchelyid otoliths have a simplified otolith morphology with an oval shape, a convex inner face, a nearly flat outer face, and an essentially unstructured shallow sulcus that closely approaches the anterior rim or opens to it. Te specimens found in the Pliocene of the Mediterranean have traditionally been placed in P. subglaber (Schubert, 1906), which was originally described from the Middle Miocene of Austria. However, the Pliocene otoliths differ in being more slender (OL: OH = 1.45 – 1.6 vs. 1.3 – 1.4), in the lack of a rostrum (vs. present in most other cases), and in a shorter sulcus (OL: SuL = 1.5 – 1.75 vs. 1.4 – 1.5). Instead, the Pliocene otoliths match well with an otolith figured as Panturichthys sp. from the Holocene off Lebanon (Agiadi & Albano, 2020) and an otolith of the extant P. fowleri, which I had the opportunity to investigate in the collection of IRSNB courtesy of D. Nolf (Bruges). Otoliths of P. mauretanicus Pellegrin, 1930, an extant species living at the Northwest African shelf are more compressed (OL: OH = 1.35 vs. 1.45 – 1.6) (for figures see Nolf, 2013). Two further extant species occur along the tropical West African coast chiefly south of the equator are P. isognathus Poll, 1953 and P. longus (Ehrenbaum, 1915). Teir otoliths are not known. An otolith identified as Panturichthys sp. in Schwarzhans (2013 a) from dredges in the Gulf of Guinea may represent one of the two species and is distinguished by its very short, narrow and tapering sulcus. In the light of the occurrence of this morphotype in the Early Pliocene of the Mediterranean as well, all Pliocene records, including the finds in Morocco, may represent the extant P. fowleri except for two figured specimens in Nolf and Martinell (1980), which are more compressed but cannot be evaluated based on the documentation. Because of some remaining uncertainty related to the fact that no otoliths are known of the two extant tropical West African species, I describe the Pliocene otoliths as P. cf. fowleri.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B3EFFB599BF6DFAFEE75707.taxon	description	Figure 9 a – e 1981 Pterothrissus darbelhamriensis — Schwarzhans: Figs. 6 – 8.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B3EFFB599BF6DFAFEE75707.taxon	materials_examined	Material 80 specimens, Zanclean, 79 from Dar bel Hamri (refigured holotype SMF P 6234, and refigured paratypes SMF P 6235 - 6237) and one from Asilah.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B3EFFB599BF6DFAFEE75707.taxon	discussion	Discussion Pterothrissus darbelhamriensis is known from the Zanclean and Piacenzian of Atlantic Morocco and Portugal and has recently also been found in the Piacenzian of Estepona near Málaga, Andalusia, Spain by van Hinsbergh & Hoedemakers. It differs from the coeval Mediterranean species P. compactus Schwarzhans, 1981 in the thinner appearance and the rectangular outline. Otoliths of P. compactus from the Zanclean of Le Puget, southern France (Fig. 9 f – g, j) and from the Tortonian of Stazzano, Piedmont, Italy (Fig. 9 h, j) are figured for comparison.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B20FFAB99BF6EFBFF1D53A6.taxon	description	Figure 10 e 2022 Echelus myrus (Linnaeus, 1758) — van Hinsbergh & Hoedemakers: pl. 1, Fig. 5. Material 1 specimen SMF PO 101.216, Zanclean, Dar bel Hamri.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B20FFAA9A056A9BFF235006.taxon	description	Figure 10 l 2017 Gnathophis mystax (Delaroche, 1809) — Lin et al.: pl. 2, Fig. 2 J, K. 2022 Gnathophis mystax (Delaroche, 1809) — van Hinsbergh and Hoedemakers: pl. 3, Figs. 1, 2. Material 2 specimens, Zanclean, Dar bel Hamri (SMF PO 101.188). Discussion An extant species that has been recorded in the Mediterranean since Late Miocene. A second, extinct species (Gnathophis kanazawai Nolf & Martinell, 1980) was described from the Early Pliocene of Catalonia, Spain.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B20FFAB99BF6CBBFB7D5206.taxon	description	Figure 10 f – h 2022 Saurenchelys silex — van Hinsbergh and Hoedemakers: pl. 1, Figs. 7, 8. Material 2 specimens SMF PO 101.185 (Fig. 10 f) and 2 juvenile specimens SMF PO 101.186 (Fig. 10 g – h), Zanclean, Dar bel Hamri. Description Small otoliths up to 3 mm in length are slen- der (OL: OH = 2.0 – 2.15), specimens from 4 to 6 mm in length are more compressed (OL: OH = 1.65 – 1.8); OH: OT = 2.1 – 2.2. Dorsal and ventral rims shallow, regular. Anterior tip slightly pointed; posterior tip blunt with rounded angles at junction with dorsal and ventral rims. All rims smooth. Inner face flat with slightly inclined, centrally positioned, elongate oval and shallow sulcus filled with a single colliculum. OL: SuL = 2.5 – 2.6. Sulcus not extending anterior of colliculum. No dorsal depression and no ventral furrow. Outer face smooth, moderately convex, thickest behind the middle. Discussion Nettastomatid otoliths show a low degree of morphological diversity, which renders identification difficult. Otoliths of representatives of all extant genera in the family are figured in Schwarzhans (2019 b). Tose of Saurenchelys most closely resemble in otolith and sulcus shape. Saurenchelys cancrivora Peters 1864 is the only extant species in the genus occurring off West Africa (and in the Indo-West Pacific). Two extant otoliths figured (Fig. 10 i, j) show a distinct variability in the shape of the postdorsal angle, which is pronounced in one specimen (Fig. 10 i) and completely rounded in the other (Fig. 10 j). Te index OL: OH ranges from 1.62 to 1.75. Saurenchelys silex described by van Hinsbergh & Hoedemakers, 2022, from the Piacenzian of Estepona near Málaga, Spain shows similar proportions in specimens of 4 – 5.5 mm in length, which is also matched with large specimens from Dar bel Hamri (Fig. 10 e), but differs in the deepened ventral rim and the posterior thickening of the outer face. Two smaller specimens (3 mm in length) from Dar bel Hamri, however, are considerably more slender in shape and could potentially represent a second species. However, it is likely that this difference is due to allometric ontogenetic growth. Despite the lack of specimens of intermediate sizes the small specimens are, therefore, also placed in S. silex.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B21FFAA99BF6E9CFE245526.taxon	description	Figure 10 m 2019 b Japonoconger africanus (Poll, 1953) — Schwarzhans: pl. 6, Figs. 11, 12. 2022 Japonoconger africanus (Poll, 1953) — van Hinsbergh & Hoedemakers: pl. 3, Figs. 12 – 14 (see there for further references). Material 1 specimen, Zanclean, Dar bel Hamri (SMF PO 101.189).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B21FFAA99BF6E9CFE245526.taxon	discussion	Discussion Japonoconger africanus today is found at a depth of 250 – 650 m off tropical West Africa from Gabon to Angola. During the Pliocene, it was much more widely distributed and has regularly been reported from the Mediterranean and now also from Atlantic Morocco. In all these fossil locations, however, it is rare, except for the Piacenzian of Estepona near Málaga, Spain (van Hinsbergh & Hoedemakers, 2022).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B21FFAA99BF6BFCFE595606.taxon	description	Figure 10 n Material 2 specimens, Zanclean, Dar bel Hamri (SMF PO 101.273).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B21FFAA99BF6BFCFE595606.taxon	discussion	Discussion Paraxenomystax is commonly recorded as a junior synonym of Xenomystax Gilbert, 1891 (see Fricke et al. 2022). Te otoliths of the type species P. bidentatus show a centrally positioned colliculum with an anterior spur and thus differ from those of other Xenomystax species (see Lin et al., 2017 and Schwarzhans, 2019 b for extant otoliths). I, therefore, keep Paraxenomystax provisionally valid here. Te single extant species, Paraxenomystax bidentatus is only known from the tropical West Atlantic. Te fossil record from Morocco is only tentatively placed in the extant species because of the geographic difference and only two specimens being available from Morocco.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B21FFAA9A056E3BFB1754A6.taxon	description	Figure 11 d – e 1906 Otolithus (Ophidiidarum) pantanellii — Bassoli & Schubert (in Bassoli): pl. 1, Figs. 41 – 42. 1983 Hildebrandia pantanellii (Bassoli & Schubert, 1906) — Nolf & Steurbaut: pl. 1, Figs. 4 – 9 (see there for further references) 1989 Hildebrandia pantanellii (Bassoli & Schubert, 1906) — Nolf & Cappetta: pl. 2, Figs. 1 – 4. 2013 a Rhynchoconger pantanellii (Bassoli & Schubert, 1906) — Schwarzhans: pl. 1, Fig. 14. 2017 Rhynchoconger pantanellii (Bassoli & Schubert, 1906) — Lin et al.: Fig. 2 F. 2022 Rhynchoconger pantanellii (Bassoli & Schubert, 1906) –– van Hinsbergh & Hoedemakers: pl. 3, fig. 16. Material 117 specimens, Zanclean, 116 from Dar bel Hamri (figured specimens SMF PO 101.190) and one from Asilah.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B21FFAA9A056E3BFB1754A6.taxon	discussion	Discussion Rhynchoconger pantanellii is relatively common and widely recorded from the Tortonian to the Zanclean of the Mediterranean and the Zanclean of Atlantic Morocco. No extant species of Rhynchoconger have been recorded from the East Atlantic, but Smith (1990, in Quéro et al.) mentioned unidentified larval Rhynchoconger specimens from the Gulf of Guinea to Angola. Schwarzhans (2013 a) found Rhynchoconger otoliths from adult specimens in dredged sediments of Holocene age off Ivory Coast that indicate that an extant eastern Atlantic species of Rhynchoconger indeed exists and that it closely resembles the fossil R. pantanellii.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B21FFA89A05693BFF355786.taxon	description	Figure 11 a – c 1978 a Gnathophis sp. — Schwarzhans: Figs. 55, 139. Holotype SMF PO 101.191 (Fig. 11 b), Agrigento, below Rupe Athenae, Sicily, Italy, Zanclean. Paratype Two specimens: one specimen MGPT-PU 130452, Borelli, Piedmont, Italy, late Tortonian; one specimen SMF PO 101.336, Orciano near Pisa, Tuscany, Italy, Zanclean. Etymology In honor of Giorgio Carnevale, Torino, Italy, for his many contributions to the understanding of fossil bony fishes. Diagnosis OL: OH = 1.8 – 2.1; OH: OT = 2.5 – 3.1. Dorsal rim anteriorly depressed and generally low; ventral rim shallow, regularly curved. Ostium closely approaching anterior rim of otolith. Description Slender, large, elegant otoliths reaching sizes of 17 mm in length (holotype). OL: OH = 1.8 – 2.1, increasing with size; OH: OT = 2.5 – 3.1, decreasing with size. Dorsal rim shallow, anteriorly depressed and irregular, highest behind middle of otolith. Ventral rim shallow, smooth and very regularly curved. Anterior tip rounded, posterior tip slightly tapering. Inner face convex, with a long, nearly horizontal and relatively narrow sulcus. Ostium marked against cauda at ventral sulcus rim by slight indentation, its anterior tip reaching close to anterior rim of otolith. Ostial channel narrow, short, positioned perpendicular on rear part of ostium. Cauda narrow, straight. Single colliculum contiguous from ostium, including ostial channel and cauda. OL: SuL = 1.6 – 1.7; CaL: OsL = 1.3 – 1.4. Dorsal depression narrow, relatively shallow and with indistinct margins; no ventral furrow. Outer face smooth, flat or with moderate central umbo. Discussion Te presence of a second species of Rhynchoconger in the young Neogene of the Mediterranean was already recognized by Schwarzhans (1978 a), who refrained from describing a new species based on a single large specimen. With the finding of two additional specimens of smaller and intermediate size, it is now clear that the morphological characteristics are stable through ontogeny. Te species is much rarer than R. pantanellii and differs mainly in being distinctly more elongate (OL: OH = 1.8 – 2.1 vs. <1.6), the depressed predorsal rim and the ostium approaching the anterior rim of the otolith more closely than in R. pantanellii. Its stratigraphic range in the Mediterranean reaches from the late Tortonian to the Zanclean. It is described here because it is considered relevant for the faunal evaluation discussed later.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B23FFA89A056D3BFBCC55E6.taxon	description	Figure 11 f Material 2 specimens, SMF PO 101.192, Dar bel Hamri, Zanclean. Discussion Te two, large, and thick otoliths of 11.4 and 12.4 mm in length resemble closely the extant specimen figured by Veen and Hoedemakers (2005; pl. 2, Fig. 2). Te outline of the otolith and its proportions match well, but Veen & Hoedemakers did not figure the ventral face with the diagnostic sulcus-like mesial depression and the mesial inward curve (terminology of Ohe in Aguilera et al., 2013). In the case of the specimen from Morocco, this inward curve is relatively long and narrow and only slightly inclined, which would probably represent a distinctive feature. Terefore, this specimen is only tentatively placed in C. latiscutatus. Today, C. latiscutatus occurs south of the northwest African upwelling system from Senegal to Angola.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B23FFA899BF681BFA6F5066.taxon	materials_examined	Material Tree clupeid otolith fragments from Dar bel Hamri that cannot be identified to genus or species level.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B23FFAF9A056AFBFF145026.taxon	description	Figure 12 d Material 2 fragmented specimens, SMF PO 101.193, Dar bel Hamri, Zanclean. Discussion Te figured specimen lacks most of the dorsal portion but is the better preserved one of the two fragments. Te regularly curved and somewhat undulating ventral rim is similar to the appearance in extant otoliths of H. hygomi. Tis species has commonly been recorded from the Mediterranean Pliocene following the synonymization of the fossil otolith-based Hygophum agrigentense Schwarzhans, 1978, with the extant H. hygomi by Brzobohatý and Nolf (1996). It is likely that the specimens from Dar bel Hamri represent the same species.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B24FFAF99BF6BBBFEFB5786.taxon	description	Figure 12 e 1971 Ceratoscopelus maderensis (Lowe, 1839) — Weiler: pl. 1, Fig. 10. 1978 a Ceratoscopelus maderensis (Lowe, 1839) — Schwarzhans: Figs. 33 – 35, 37. 1986 Ceratoscopelus maderensis (Lowe, 1839) — Schwarzhans: Fig. 22. 1989 Ceratoscopelus maderensis (Lowe, 1839) — Nolf & Cappetta: pl. 6, Figs. 17 – 20. Material 1 specimen, SMF PO 101.195, Dar bel Hamri, Zanclean.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B24FFAF9A056CBBFC605586.taxon	description	Figure 12 g 2022 Diaphus adenomus Gilbert, 1905 — van Hinsbergh & Hoedemakers: pl. 11, Figs. 9 – 11 (see there for further references). Material 4 specimens SMF PO 101.274, Dar bel Hamri, Zanclean.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B24FFAF99BF6EFBFEA952A6.taxon	description	Figure 12 c. 1978 a Gymnoscopelus fitchi — Schwarzhans: Figs. 30, 31 1, 129. 2022 Myctophum fitchi (Schwarzhans, 1978) — Carnevale & Schwarzhans: Fig. 8 I – K (see there for further references). 2022 Myctophum fitchi (Schwarzhans, 1978) — van Hinsbergh & Hoedemakers: pl. 12, Figs. 19 – 25. Material 4 specimens, Zanclean, 2 specimens Dar bel Hamri (SMF PO 101.194), 1 specimen Jebel Zebbouj, 1 specimen Asilah.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B24FFAD9A056BDBFE025406.taxon	description	Figure 12 h – l 2000 Diaphus cavallonis — Brzobohatý & Nolf: pl. 5, Figs. 7 – 14 (see there for further references).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B24FFAD9A056BDBFE025406.taxon	materials_examined	Material 146 specimens, figured specimens SMF PO 101.197, Dar bel Hamri, Zanclean.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B24FFAD9A056BDBFE025406.taxon	discussion	Discussion Diaphus cavallonis has 8 to 12 delicate denticles along the ventral rim, according to van Hinsbergh and Hoedemakers (2022), based on more than 1000 specimens from Spain. Schwarzhans & Aguilera (2013) noted 10 to 12 denticles on much fewer specimens, which may reflect the predominant range. In the case of the Moroccan specimens, the denticles are eroded except for few specimens (e. g., Fig. 10 i, k, l), which show 8 – 11 denticles. Te Moroccan specimens show a considerable variability in respect to the predorsal rim which can be shallow (Fig. 12 l) or expanded (Fig. 12 h, i), the shape of the posterior rim which can be rounded (Fig. 12 i) or relatively blunt (Fig. 12 l) and the opening of the ostium. Te variability appears to be within the range observed in the types of Brzobohatý and Nolf (2000) and figures shown in Schwarzhans & Aguilera (2013) and van Hinsbergh and Hoedemakers (2022). Te specimens currently available to me do not warrant a further investigation of the nature of the species but indicate that a detailed review of its morphologic limit may be required. Diaphus cavallonis was apparently widely distributed in the North Atlantic and the Mediterranean during the Zanclean; the oldest records in the Mediterranean date from the Tortonian (Lin et al., 2015, 2017). Van Hinsbergh and Hoedemakers (2022) found that D. cavallonis is lacking in the Piacenzian of the Mediterranean and instead is replaced by D. postcavallonis van Hinsbergh & Hoedemakers, 2022. In contrast, Agathangelou et al. (2022) reported D. cavallonis from the Piacenzian of Cyprus. Diaphus cavallonis is the second largest Diaphus species in Dar bel Hamri, reaching 5 mm in length, surpassed only by D. adenomus (see above) with 6 mm in length.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B26FFAD99BF6A5BFAE65286.taxon	description	Figure 12 m – o 1986 Diaphus dirknolfi — Schwarzhans: pl. 3, Figs. 34, 35.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B26FFAD99BF6A5BFAE65286.taxon	materials_examined	Material 90 specimens, Zanclean: 84 specimens Dar bel Hamri (figured specimen SMF PO 101.198), 4 specimens Sidi Mohamed ech Chleuh (SMF PO 101.199), 2 specimens Asilah.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B26FFAD99BF6A5BFAE65286.taxon	discussion	Discussion Diaphus dirknolfi was originally established on specimens from Dar bel Hamri with incompletely preserved denticles along the ventral rim, which resulted in a too high count of 12 – 14 denticles. Te species was redefined in 2022 by Carnevale and Schwarzhans (2022) based on better preserved specimens from the type locality and the nearby location Sidi Mohammed ech Chleuh, which yielded uneroded specimens. Te recount of the denticles along the ventral rim resulted in 6 – 8. Van Hinsbergh and Hoedemakers (2022) reported 7 – 11 denticles. Tese records show that the number of denticles is quite variable in this species, but mostly seems to fall in the range of 7 – 9. Obviously, this characteristic cannot be used as a distinguishing trait between D. dirknolfi, the Early to Middle Miocene D. hataii (Ohe & Araki, 1973), and the extant D. dumerilii (Bleeker, 1856), but other characteristics such as the proportional length of rostrum to antirostrum and the proportions of ostium and cauda remain valid (see Carnevale & Schwarzhans, 2022, for details). Diaphus dirknolfi is known from the latest Messinian (Lago Mare interval) to the Piacenzian of the Mediterranean and the Zanclean of Morocco.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B26FFAC9A056CDBFD9D50E6.taxon	description	Figure 13 a – c 2013 c Diaphus draconis — Schwarzhans: pl. 4, Fig. 8 – 12. 2019 a Diaphus draconis Schwarzhans, 2013 — Schwarzhans: Fig. 57.1 – 3. 2022 Diaphus cf. draconis Schwarzhans, 2013 — Carnevale & Schwarzhans: Fig. 7 Q – T. Material 3 specimens Kef Nsour, Messinian (SMF PO 101.200), 5 specimens Sidi Mohamed ech Chleuh, Zanclean, possibly reworked from Messinian.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B26FFAC9A056CDBFD9D50E6.taxon	discussion	Discussion Diaphus draconis is an inconspicuous small species reaching about 2 mm in length in the Moroccan localities. It can be easily confused with small specimens of larger species (see extensive discussion in Carnevale & Schwarzhans, 2022). It may be recognized best by its 4 – 6 widely spaced strong denticles along the ventral rim. Diaphus draconis was originally described from the Serravallian / Tortonian of Gabon (Schwarzhans, 2013 c) and also recorded from the Tortonian of New Zealand (Schwarzhans, 2019 a), indicating a wide geographic distribution. Specimens from the latest Messinian Lago Mare phase of Italy were only tentatively allocated (Carnevale & Schwarzhans, 2022). In the Rharb Basin of Morocco, the best preserved specimens were obtained from the basal levels of Kef Nsour, which are supposed to be Messinian in age. Specimens from the basal level of Sidi Mohammed ech Chleuh are slightly leached, possibly indicating an origin from reworked Messinian strata below. Tere are no positive records from the Zanclean.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B27FFAC99BF6EFBFB2E5666.taxon	description	Figure 13 d – j 2022 Diaphus taaningi Norman, 1930 — van Hinsbergh & Hoedemakers: pl. 8, Figs. 1 – 5. Holotype SMF PO 101.201 (Fig. 13 f), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 7 specimens, 4 specimens SMF PO 101.202, Dar bel Hamri, Zanclean, 1 specimen SMF PO 101.203, Kef Nsour, Messinian, 2 specimens SMF PO. 101.204, Sidi Mohamed ech Cleuch, Zanclean. Further material 291 specimens, 23 Kef Nsour and Chaba Kaudiat el Mogen, Messinian, 213 specimens Dar bel Hamri, Zanclean, 38 specimens Sidi Mohamed ech Chleuh, Zanclean, 17 specimens Jebel Zebbouj, Zanclean. Etymology After Maghreb, the name of the northwestern region of Africa. Diagnosis OL: OH = 1.1 – 1.2; OH: OT = 3.25 – 3.35. Ventral rim regularly and deeply curved, with 6 – 9 denticles. OCL: CCL = 1.95 – 2.7. Inner face convex; outer face with distinct postcentral umbo. Description Relatively small otoliths reaching sizes of 2.5 mm in length (holotype 2.3 mm). Dorsal rim high, regularly curved or anteriorly expanded; posteriorly rounded or with mild, short depression. Ventral rim deep, regularly curved, with 6 – 9 denticles (mostly 7 – 8). Rostrum slightly longer than antirostrum, 8 – 15 % of OL, ratio rostrum to antirostrum about 2: 1. Posterior rim broadly rounded, blunt. Inner face distinctly convex in horizontal direction, less bent in vertical direction. Sulcus long, slightly supramedian, shallow, straight, OL: SuL = 1.25 – 1.35. Ostium slightly wider than cauda and twice to 2.5 times the length of the cauda (OCL: CCL = 1.95 – 2.7). Upper margin of ostium straight. Cauda short, with equally long caudal pseudocolliculum. Dorsal depression wide; ventral furrow mostly distinct, relatively close to ventral rim of otolith. Outer face relatively smooth, with distinct, broad postcentral umbo. Discussion Many small and somewhat inconspicuous otoliths from the Miocene and Pliocene from the Mediterranean and Paratethys have been placed in the extant Diaphus taaningi Norman, 1930, by Brzobohatý and Nolf (2000). Diaphus taaningi is a pseudoceanic species occurring in the tropical West and East Atlantic (Froese & Pauly, 2022). In dredge samples from the Gulf of Guinea and in a comparative morphological study of extant otoliths of the genera Diaphus, Idiolychnus, and Lobianchia, Schwarzhans (2013 a, 2013 b) figured several extant otoliths of D. taaningi from both areas of its distribution and found slight but consistent differences between the West and East Atlantic populations. Schwarzhans & Aguilera (2013) concluded on this basis that none of the fossil otoliths figured as D. taaningi until then actually pertains to this species and also figured presumably valid D. taaningi otoliths from the Gelasian of Atlantic Panama. More recently, van Hinsbergh and Hoedemakers (2022) figured, without description or explanation, Diaphus taaningi otoliths from the Zanclean and Piacenzian of Estepona near Málaga, Spain. In the late Tortonian / early Messinian and Zanclean of the locations in the Rharb Basin of Morocco studied here, the otoliths of Diaphus maghrebensis represent the most common myctophid species. Tey clearly represent the same species as the otoliths figured by van Hinsbergh & Hoedemakers as D. taaningi from Estepona near Málaga, but reach larger sizes (2.5 mm in length vs. <2 mm). Terefore, I assume that these otoliths from Morocco represent true adult forms of the species in question. In many aspects, they resemble extant D. taaningi otoliths (Fig. 13 k – m), which reach nearly 3 mm in length, but they also differ in some consistent aspects as follows (D. maghrebensis first, D. taaningi second): OH: OT = 3.25 – 3.35 vs. 4.3 – 4.5, OCL: CCL = 1.95 – 2.7 vs. 1.1 – 1.4, outer face with distinct postcentral umbo vs. flat or with shallow umbo. Te relatively significant difference of the ratio of the ostium to the cauda length in the two species may even indicate that they are not closely related. In fact, I do not know of any the extant species that may be related. Diaphus maghrebensis is known from the Messinian and Zanclean of Morocco and from the Zanclean and Piacenzian of the Mediterranean. Earlier records of so-called Diaphus taaningi otoliths from the Tortonian of the Mediterranean require specific review; referenced records from the Badenian of the Paratethys have recently been re-identified as Diaphus cassidiformis (Frost, 1933) (see Schwarzhans & Radwańska, 2022).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B29FFA299BF69FBFF1D5606.taxon	materials_examined	Material A single, rather eroded specimen from Dar bel Hamri.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B29FFA29A056C9BFC8E57C6.taxon	description	Figure 14 j – l Material 9 fragmentary specimens, 1 specimen from Kef Nsour, SMF PO 101.208, Messinian; 8 specimens Dar bel Hamri, figured specimens SMF PO 101.209, Zanclean. Discussion None of the specimens available are complete, the best preserved ones are from two-thirds of a specimen. Te depressed predorsal projection and the rounded anterior tip are considered typical for M. polli in otoliths of Merluccius species from the East Atlantic. Te most complete specimen of Fig. 14 j is 18 mm in length and reconstructed for the broken part would have been about 24 mm in length. Te largest, not figured fragmented specimen may have been 33 mm in length when complete. For comparison, extant otoliths are figured of M. merluccius (Linnaeus, 1758) (Fig. 14 g), M. senegalensis Cadenat, 1950 (Fig. 14 h), and M. polli (Fig. 14 i). Today, M. polli is known from the west African shelf off Mauritania to Angola from 29 ° to 19 ° S in deep water from 50 to 550 m.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B29FFA29A056E1BFB5350A6.taxon	description	Figure 14 e Material 10 specimens (figured specimens SMF PO 101.207), Dar bel Hamri, Zanclean.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B29FFA29A056D7BFC605246.taxon	description	Figure 14 f Material 1 specimen SMF PO 101.275, Dar bel Hamri, Zanclean.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B29FFA09A05681BFD9355E6.taxon	description	Figure 14 m – p 2010 Paratrisopterus glaber — Schwarzhans: pl. 30, Figs. 12 – 15.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B29FFA09A05681BFD9355E6.taxon	materials_examined	Material 72 specimens, 2 specimens, Kef Nsour, Messinian, 64 specimens, Dar bel Hamri (figured specimens SMF PO 101.210), Zanclean, 2 specimens Sidid Mohamed ech Chleuh, Zanclean, 3 specimens, Jebel Zebbouj, Zanclean. 1 specimen, Asilah, Zanclean.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B29FFA09A05681BFD9355E6.taxon	discussion	Discussion Paratrisopterus glaber differs from the Middle Miocene species P. brinki (Posthumus, 1923) in its more compressed shape and relatively smooth outer face. It is, however, not as compressed as the rare Middle Miocene P. globosus (Posthumus, 1923). Paratrisopterus glaber was described from the Late Miocene of the North Sea Basin but was apparently more widely distributed in the European seas. It has recently been recorded from the Zanclean and Piacenzian of Spain, where it occurs in parallel with P. labiatus (Schubert, 1905) in the Zanclean. Te occurrence in the Moroccan Early Pliocene fills a geographic gap in the distribution. Otoliths have been described in situ by Landini and Sorbini (1999) from a fish interpreted as Gadiculus labiatus from the Early Pleistocene of Italy. According to the otoliths figured in this article, the species represents P. glaber.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B2BFFA099BF6ABBFEFB5606.taxon	description	Figure 15 a 2006 Carapus sp. — Nolf & Girone: pl. 1, Fig. 10. 2017 Carapus acus (Brünnich, 1768) — Lin et al.: Fig. 9 D – E. 2019 a Carapus acus (Brünnich, 1768) — Agiadi et al.: Fig. 3 G. Material A single somewhat eroded specimen of 2.8 mm in length (SMF PO 101.211) from Dar bel Hamri, Zanclean.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B2BFFA09A056E1BFBD554C6.taxon	description	Figure 15 b – e 1978 a Carapus acus (Brünnich, 1768) — Schwarzhans Fig. 112.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B2BFFA09A056E1BFBD554C6.taxon	materials_examined	Material 24 specimens, Zanclean, 23 specimens Dar bel Hamri (figured specimens SMF PO 101.212), 1 specimen Sidi Mohamed ech Chleuh (SMF PO 101.213). Discussion Tere has been some confusion about the identity of carapid otoliths of the genus Echiodon in the Pliocene of the Mediterranean. A large series of extant otoliths of E. dentatus has been figured in Lombarte et al. (2006), and one of them is refigured here (Fig. 15 e). Tese otoliths are characterized by being relatively thin, having a relatively narrow sulcus terminating at some distance from the anterior rim of the otolith, and a flat ventral otolith rim. Some specimens found in Morocco and recorded in the literature from the Mediterranean (see synonymy listing) match the morphology of the extant specimens very well. However, there is also a coeval second species known from the Zanclean of the Mediterranean and Morocco that represents a different morphotype and has been described as Carapus praeimberbis by Weiler (1971) but is sometimes confused with E. dentatus (see below). Echiodon dentatus differs from E. praeimberbis by the narrower and shorter sulcus (although in large specimens of E. dentatus, the sulcus increases in size; see Tuset et al., 2008), the unexpanded (vs. expanded) ventral otolith rim, and the outer face projecting over the dorsal rim of the otolith (vs. unexpanded).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B2BFFA79A056AA8FE025466.taxon	description	Figure 15 f – i 1971 Carapus praeimberbis — Weiler: pl. 2, Fig. 35. 1978 a Echiodon drumondii Tompson, 1837 — Schwarzhans: Fig. 113.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B2BFFA79A056AA8FE025466.taxon	materials_examined	Material 79 specimens, Dar bel Hamri, Zanclean (figured specimens SMF PO 101.214). Discussion Otoliths of E. praeimberbis differ from those of E. dentatus in being more thickset, showing a distinctly wider and larger sulcus that reaches close to the anterior and dorsal rims of the otolith and a deepened, curved ventral otolith rim. Except for the deepened ventral rim, these otoliths resemble those of the extant E. drummondii from the northern North Sea and Norwegian Sea. Schwarzhans (2013 a) recorded otoliths of Echiodon sp. from Holocene dredge samples off West Africa between Guinea and Nigeria, which are refigured here for comparison (Fig. 15 j – k). Tese otoliths resemble E. praeimberbis even more closely in the expanded ventral rim and differ only in the pointed anterior tip and the reduced, rounded posterior tip. No Echiodon species is recorded from tropical West Africa today. Tus, the Holocene dredge samples indicate that an undescribed Echiodon species may exist in West Africa (Schwarzhans, 2013 a), and E. praeimberbis would be the fossil species most closely related to it. Nolf (2013) considered all records of E. praeimberbis to represent the extant E. dentatus, but this concept is probably a result of the presence of two different species occurring coevally and mostly in small numbers. However, van Hinsbergh and Hoedemakers (2022) listed 328 specimens from Estepona near Málaga, Spain, mostly from the Piacenzian. Te two specimens figured pertain to E. praeimberbis, but it is quite possible that the assemblage also contained E. dentatus.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B2CFFA799BF693BFAD554C6.taxon	description	Figure 16 a 1998 Brotula aff. multibarbata Temminck & Schlegel, 1846 — Nolf, Mañe & Lopez: pl. 5, Fig. 6. 2006 Brotula cf. multibarbata Temminck & Schlegel, 1846 — Nolf & Girone: pl. 1, Fig. 14, pl. 4, Fig. 18. 2022 Brotula cf. multibarbata Temminck & Schlegel, 1846 — van Hinsbergh & Hoedemakers: pl. 18, Figs. 8, 9. Material 5 specimens SMF PO 101.215, Dar bel Hamri, Zanclean. Discussion Incomplete, eroded, or small otoliths of Brotula have occasionally been reported from the Pliocene of the Mediterranean (see synonymy listing). Te large, figured specimen from Morocco is well preserved but lacks the entire rear part of the otolith. Within the extant species of Brotula, two slightly different otolith morphotypes are recognized. One morphotype is characterized by elongate otoliths and a long ostium (CaL: OsL = 1.4 – 1.6), comprising the extant species B. barbata (Bloch & Schneider, 1801) and B. clarkae Hubbs, 1944 (see Nolf, 1980, for figures). Te other morphotype has relatively more compressed otoliths with a shorter ostium (CaL: OsL ≅ 2.0), comprising B. multibarbata (Temminck & Schlegel, 1846) (see Nolf, 1980, for figures), B. flaviviridis Greenfield, 2005, B. ordwayi Hildebrand & Barton, 1949, and B. townsendi Fowler, 1900. Brotula barbata is the only one of these species living today in the Atlantic. Te fossil otoliths from the Pliocene of the Mediterranean and Morocco, however, represent the second morphotype and have usually been tentatively associated with B. multibarbata. Today, B. multibarbata is known from the Indo-West Pacific, with its closest occurrence to the fossil specimens in southeastern Africa. I follow previous records in recording the Moroccan Pliocene otoliths as B. aff. multibrabata. Te only known large and complete otolith specimen figured from B. aff. multibrabata is by Nolf et al. (1998) from the Pliocene of southern Spain, and it confirms both the short ostium and the moderately elongate shape. Without a review of all relevant fossil specimens from Europe, however, it is not clear whether it could represent an extinct species of Brotula.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B2CFFA79A05691BFC6B5606.taxon	description	Figure 16 b – g Holotype SMF PO 101.217 (Fig. 16 b), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 5 specimens SMF PO 101.218, same data as holotype.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B2EFFA59A056D3BFB0453A6.taxon	description	Figure 16 j Material A single, slightly eroded otolith from the Zanclean of Dar bel Hamri (SMF PO 101.276) that is too poorly preserved for further identification.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B2EFF9B9A056CFBFF1454A6.taxon	description	Figure 17 a – d Holotype SMF PO 101.219 (Fig. 17 b), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 5 specimens SMF PO 101.277, same data as holotype. Further material 5, mostly fragmentary or eroded specimen, Dar bel Hamri, Zanclean. Etymology Named in honor of Fritz von der Hocht (Kerpen) who alerted me to this unique location and collected many otoliths of the studied assemblage including the type specimens of this species. Diagnosis Large otoliths to 20 mm in length. OL: OH increasing from 1.05 at 6 mm in length to 1.35 at 20 mm in length. Dorsal rim strongly lobate; ventral rim flattened at midsection. OsL: CaL = 1.10 – 1.17. Cauda slightly upward directed, with tapering, symmetrical tip. Description Large otoliths reaching at least 20 mm in length (holotype). OL: OH distinctly allometric, increasing from 1.05 at 6 mm in length to 1.35 in holotype; OH: OT 3.0 – 3.3. Rostrum and posterior tip moderately strong, superior, with rounded tips, nearly symmetrical. Excisura and antirostrum very small. Dorsal rim anteriorly and posteriorly shortened, relatively shallow, deeply and coarsely lobate, usually highest at predorsal angle. Ventral rim moderately deep, with nearly straight, inclined pre- and postventral rims and flattened midsection, relatively smooth. Inner face distinctly convex, with supramedian positioned, large and wide sulcus. OsL: CaL = 1.13 – 1.17 in large and medium sized specimens (Fig. 17 a – c), not measurable in smallest specimen (Fig. 17 d) because of incomplete rostrum but probably 1.10 or smaller. Ostium slightly upward oriented, its ventral margin deeply lobed, slightly narrowing toward anterior. OsL: OsH = 1.45 – 1.75. Cauda slightly upward oriented, nearly symmetrically tapering. Dorsal depression marked by distinct crista superior. Ventral furrow indistinct, very close to ventral rim of otolith if discernable. Outer face flat in large specimens (Fig. 17 a 2, b 2) to distinctly convex in small specimens (Fig. 17 d 2), smooth. Discussion Te genus Centroberxy is not currently known from the Atlantic; the nearest extant occurrence is C. spinosus (Gilchrist, 1903) from eastern South Africa. Te current center of distribution of the genus is seas around Australia and New Zealand (four of seven extant species), but this distribution must be regarded as a secondary geographic relict distribution (Schwarzhans & Jagt, 2021). So far, C. manens Nolf & Brzobohatý, 2004, from the Early Miocene of Italy and the North Sea Basin represented the last record in the Atlantic, but it is from a different phylogenetic lineage than the specimens from the Pliocene of Morocco. Most extant species of Centroberyx grow to large sizes (50 to 65 cm TL; see Froese & Pauly, 2022) and their otoliths grow to sizes of more than 22 mm in length. For comparison, two extant otoliths are figured of C. affinis (Günther, 1859) reflecting a large specimen near 45 mm TL (Fig. 17 e) and a smaller one of unrecorded fish length (Fig. 17 f). Smale et al. (1995) figured two otoliths of C. spinosus one 10.6 mm in length from a specimen of 15.2 cm SL, which is near the maximum size of the species (20 cm TL), and the other 7.1 mm in length from a specimen of 12 cm TL. Centroberyx spinosus is a relatively small species in the genus. Centroberyx vonderhochti resembles C. spinosus but differs in having attained at least twice the size of the extant species, the flattened midventral rim, the very intensely lobate dorsal rim, and the ostium being slightly longer than the cauda (OsL; CaL = 1.1 – 1.17 vs. 1.0 – 1.05). Most likely, C. vonderhochti and C. spinosus represent vicariant species.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B10FF9B9A056CDBFC6B5446.taxon	description	Figure 17 i – l Holotype SMF PO 101.220 (Fig. 17 i), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 4 specimens SMF PO 101.221, same data as holotype.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B10FF9B9A056E9BFB6D5206.taxon	description	Figure 17 g Material 1 specimen SMF PO 101.278, Dar bel Hamri, Zanclean. Discussion Te only available specimen is relatively well preserved except for the eroded rostrum, and resembles well the extant specimens figured of S. hastatum by Lombarte et al. (2006). Today, S. hastatum is the only species of the genus known in the eastern Atlantic from Portugal to Angola (Froese & Pauly, 2022).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B12FF989A056ADBFD395266.taxon	description	Figure 18 f – j? 1997 Deltentosteus sp. — Nolf & Marques da Silva: pl. 3, Figs. 7 – 9. 2010 Lesueurigobius sp. 2 — Girone, Nolf & Cavallo: Fig. 10 c 1 – 4. 2020 Lesueurigobius stazzanensis — Schwarzhans, Agiadi & Carnevale: Fig. 6 AG – AP.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B12FF989A056ADBFD395266.taxon	materials_examined	Material 46 specimens Dar bel Hamri, Zanclean (figured specimens SMF PO 101.224).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B12FF989A056ADBFD395266.taxon	discussion	Discussion Lesueurigobius stazzanensis is known from the Tortonian to Piacenzian of the Mediterranean, the Zanclean of Atlantic Morocco, and possibly the Piacenzian of Portugal. It differs from the otoliths of other extant and fossil Lesueurigobius otoliths except L. heterofasciatus Maul, 1971, in being more elongate (OL: OH = 1.05 – 1.17 vs. <1.0). Te drawing of an extant specimen of L. heterofasciatus provided by D. Nolf (Bruges) shows similar proportions (OL: OH = 1.22) but with a pronounced preventral projection (vs. blunt anterior rim in L. stazzanensis) and a regularly curved dorsal rim (vs. pronounced predorsal and postdorsal angles in L. stazzanensis).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B13FF989A056D7BFB995486.taxon	description	Figure 19 a – c 2022 Buenia pulvinus — van Hinsbergh & Hoedemakers: pl. 20, Figs. 6 – 13. Material 9 specimens, Dar bel Hamri, Zanclean (figured specimens SMF PO 101.226). Discussion Te genus Buenia shows a remarkable diversity in otolith-based taxa in the Pliocene of the Mediterranean (Schwarzhans et al., 2020; van Hinsbergh & Hoedemakers, 2022). Two of the extant species (B. affinis Iljin, 1930, and B. massutii Kovačić, Ordines & Schliewen, 2017) have also been recorded as fossils from the Pliocene, in addition to the two extinct species B. pisiformis Schwarzhans et al., 2020, and B. pulvinus van Hinsbergh & Hoedemakers, 2022. Van Hinsbergh & Hoedemakers compared B. pulvinus with the extant B. affinis and mentioned the smaller size of the sulcus as the main distinguishing characteristic. Tis characteristic is shared by the Moroccan otoliths, which do not, however, exhibit any postdorsal projection as described by van Hinsbergh & Hoedemakers. I attribute the latter difference to the smaller size of the Moroccan otoliths compared to those from Estepona near Málaga.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B13FF9899BF6CBBFB7F50E6.taxon	description	Figure 18 k – m 2013 a Nematogobius maindroni (Sauvage, 1880) — Schwarzhans: pl. 13, Figs. 1, 2. Material 2 specimens (SMF PO 101.225), Dar bel Hamri, Zanclean. Discussion Extant otoliths of Vanneaugobius are not well known, and those of other West African gobies such as Didogobius, Ebomegobius, and Wheelerigobius are still unknown. A poorly preserved specimen of V. canariensis Van Tassell, Miller & Brito, 1987, extracted from USNM 298,746 and a small specimen of V. dollfusi Brownell, 1978, figured in Lombarte et al. (2006) suggest that these otoliths could belong to an unknown species of the genus Vanneaugobius. Te specimens from the Zanclean of Morocco are characterized by a relatively flat and smooth inner face bearing a small, slightly inclined sulcus with low ostial lobe and no subcaudal iugum. Te same otolith morphotype has also been dredged from Holocene sediments of Guinea and Ivory Coast and has been erroneously identified as Nematogobius maindroni (Fig. 18 m is refigured from Schwarzhans, 2013 a) based on small specimens (the only ones then available). New otoliths of both nominal species extracted from larger specimens exhibited a distinctly different morphology reflecting a remarkable ontogenetic allometry possibly accentuated also by different habitats and water chemistry, as the small otoliths were obtained from freshwater specimens. Te true identity of the Pliocene Moroccan and Holocene dredged Gulf of Guinea specimens remains somewhat obscure, since it is not possible to discern from the few extant otoliths known of Vanneaugobius whether they have or lack a subcaudal iugum.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B13FF9E9A05691BFE7F5762.taxon	description	Figure 19 f – j Holotype SMF PO 101.227 (Fig. 19 f), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 4 specimens SMF PO 101.228, same data as holotype. Further material 21 specimens, Zanclean, 20 specimens same data as holotype, 1 specimen Asilah. Etymology From planus (Latin) = flat, referring to the relatively flat inner face. Diagnosis OL: OH = 1.02 – 1.1. Dorsal rim regularly curved, smooth. Sulcus shape very asymmetrical with a wide ostium and a narrow, tapering and pointed cauda. Ostium inclined at 22 – 25 °. Description Small otoliths with rounded shape without prominent angles or projections but resembling a well-rounded rectangle. Size up to 2 mm in length (holotype 1.8 mm). OH: OT = 2.7 – 3.0. Ventral rim shallow, nearly straight; dorsal rim slightly elevated with well-rounded pre- and postdorsal angles at about same level. Anterior and posterior rims nearly vertical. All rims smooth. Inner face relatively flat, only slightly convex, and rather smooth. Sulcus positioned slightly eccentrically towards anterior and very asymmetrical with a wide ostium and a narrow cauda. Ostium dorsally and ventrally widened, steeply inclined at 22 – 25 °, with angular or rounded anterior tip. Ostial-caudal joint marked by angle on ventral sulcus margin. Cauda narrow and short, with tapering, pointed tip, inclined at 7 – 12 °. OL: SuL = 1.85 – 2.1; OsL: CaL = 1.85 – 2.2; OsH: CaH = 2.5 – 2.8. Subcaudal iugum very feeble and small. Dorsal depression small, cup like, high above sulcus and dorsally open; ventral furrow weak, close to ventral rim of otolith, fading upwards anteriorly and posteriorly. Outer face evenly but relatively faintly convex, smooth.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B13FF9E9A05691BFE7F5762.taxon	discussion	Discussion Deltentosteus planus was apparently a relatively small species with otoliths not exceeding 2 mm in length. Te otoliths are nevertheless easily recognized by the nearly flat inner face, the smooth, rounded rectangular outline, and the specific asymmetric sulcus. Of the two extant species, it most resembles D. collonianus (Risso, 1820) (Fig. 18 e) in the relatively flat inner face and the low OL: OH ratio known from the Mediterranean and adjacent part of the Atlantic. Deltentosteus planus differs from D. collonianus in the smooth, rounded rectangular outline (vs. bulged dorsal rim and distinct crenulation and undulations), the distinct shape of the sulcus, and the rather steep inclination of the ostium (22 – 25 ° vs. 15 °). Furthermore, otoliths of D. planus do not reach the size of its extant congener.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B15FF9E9A056CDBFB5B5486.taxon	description	Figure 20 a – c 1955 Eucitharus balearicus — Bauza-Rullan: pl. 8, Fig. 16.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B15FF9E9A056CDBFB5B5486.taxon	materials_examined	Material 8 specimens, Dar bel Hamri, Zanclean (figured specimens SMF PO 101.230).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B15FF9E9A056CDBFB5B5486.taxon	discussion	Discussion Citharus balearicus differs from otoliths of the extant C. linguatula (Linnaeus, 1758) in the lower expression of the postdorsal angle.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B15FF9E99BF697EFBCC5246.taxon	description	Figure 19 d, k – m 2020 Deltentosteus quadrimaculatus (Valenciennes, 1837) — Schwarzhans, Agiadi & Carnevale: Fig. 12 AE – AJ (see there for further references). 2022 Deltentosteus quadrimaculatus (Valenciennes, 1837) — van Hinsbergh & Hoedemakers: pl. 20, Figs. 23 – 26. Material 43 specimens, Dar bel Hamri, Zanclean (figured specimens SMF PO 101.229). Discussion Otoliths of D. quadrimaculatus have frequently been retrieved from Pliocene rocks from Mediterranean realms. Tey are easily distinguishable from D. planus by the more elongate shape in adults (OL: OH = 1.15), while small specimens at a size of about 1.05 are similar in proportions (see extant specimen in Fig. 19 d and small fossil specimen in 19 k). Te sulcus is larger in D. quadrimaculatus than in D. planus, with a weaker asymmetry of the ostium to cauda, and the inner face is distinctly convex.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B15FF9C9A05695BFD915386.taxon	description	Figure 20 d – g	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B15FF9C9A05695BFD915386.taxon	materials_examined	Material 31 specimens, Zanclean, 29 specimens Dar bel Hamri (figured specimens SMF PO 101.231), 1 specimen Sidi Mohamed ech Chleuh, 1 specimen Asilah.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B15FF9C9A05695BFD915386.taxon	discussion	Discussion Arnoglossus kokeni is recognized by its nearly perfectly square shape with a ratio OL: OH just slightly above 1.0 (1.1 – 1.2). Arnoglossus kokeni is known in the Mediterranean from the Tortonian to the Piacentian and from Atlantic Morocco from the Zanclean.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B17FF9C99BF6DDBFD635406.taxon	description	Figure 20 h – k 1999 Arnoglossus quadratus — Schwarzhans: Fig. 365 – 373.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B17FF9C99BF6DDBFD635406.taxon	materials_examined	Material 18 specimens, Dar bel Hamri, Zanclean, photographed holotype (Fig. 20 j) SMF P 9317 and paratypes SMF P 9318.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B17FF9C99BF6DDBFD635406.taxon	discussion	Discussion Arnoglossus quadratus differs from the coeval A. kokeni in the more elongate, rectangular shape with a ratio OL: OH ranging from 1.25 to 1.3. It has been found also with few specimens in the Pliocene of Estepona near Málaga (van Hinsbergh & Hoedemakers, 2022).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B17FF9C9A056D7BFC385566.taxon	description	Figure 21 e – f Material 1 specimen SMF PO 101.232, Dar bel Hamri, Zanclean. Discussion Otoliths of D. hexophthalma are relatively thin, oval in shape with a ratio OL: OH of 1,25 – 1.4 (corrected from Schwarzhans, 1999) and a relatively thin sulcus. A recent specimen is figured for comparison (Fig. 21 e). Te single fossil specimen from Morocco (Fig. 21 f) resembles the extant specimens in all morphological aspects but is distinctly thicker (OH: OT = 2.7 vs. 3.5 – 4.5), and is, therefore, only tentatively placed in the same species.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B17FF9C99BF6A9CFAB250E6.taxon	description	Figure 20 l – o 1999 Laeops rharbensis — Schwarzhans: Fig. 444 – 447. Material 7 specimens, Dar bel Hamri, Zanclean, photographed paratypes SMF P 9320.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B17FF9C99BF6A9CFAB250E6.taxon	discussion	Discussion Laeops rharbensis resembles Arnoglossus kokeni in proportions and general appearance, but differs in being more compressed (OL: OH = 1.0 – 1.1 vs. 1.1 – 1.2), in the more rounded and expanded ventral rim, and in the higher positioned anterior rostrum-like tip of the otolith. Te generic allocation to the genus Laeops, which today only lives in the Indo-Pacific, is poorly constrained by means of otolith morphological characters. If verified, it would represent the second taxon in the Zanclean of northwestern Morocco with clear Indo-Pacific affinities, the other being Brotula multibarbata (see above).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B17FF939A056BBBFDA65798.taxon	description	Figure 21 i – j 1978 Microchirus variegatus (Donovan, 1808) — Nolf: pl. 7, Fig. 18. 1989 Microchirus variegatus (Donovan, 1808) — Nolf & Cappetta: pl. 18, Fig. 18. 2019 b Microchirus variegatus (Donovan, 1808) — Agiadi et al.: Fig. 4 l. 2022 Microchirus variegatus (Donovan, 1808) — van Hinsbergh & Hoedemakers: pl. 27, Figs. 13 – 15. Material 3 specimens SMF PO 101.280, Dar bel Hamri, Zanclean. Discussion Otoliths of M. variegatus are recognized by their rounded trinagular outline with a deep ventral rim and a shallow dorsal rim and the distinctly convex inner face. Te species has been regularly recorded from Pliocene and Pleistocene rocks of the North Sea Basin and the Mediterranean. Miocene records may represent a different species (see Schwarzhans, 1999).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B18FF9299BF6874FEA650A6.taxon	description	Figure 21 a – d 1999 Quenselia cornuta — Schwarzhans: Fig. 751 – 758. Material 35 specimens, Dar bel Hamri, Zanclean, photographed holotype (Fig. 21 b) SMF P 9325 and paratypes SMF P 9326.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B18FF9299BF6874FEA650A6.taxon	discussion	Discussion Quenselia cornuta is readily recognized by its horn-like predorsal lobe positioned near the anterior end of the otoliths and the fact that the cauda is longer than the ostium. Both are unusual characteristics and the proportions of the sulcus could in fact indicate that Q. cornuta could belong to an extinct lineage / genus. Quenselia cornuta has only been recorded from the Pliocene of Atlantic Morocco.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B19FF929A056D61FB95552F.taxon	description	Figure 21 l – o 1999 Cynoglossus obliqueventralis — Schwarzhans: Fig. 905 – 906.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B19FF929A056D61FB95552F.taxon	materials_examined	Material 7 specimens, Dar bel Hamri, Zanclean, photographed holotype (Fig. 21 j) SMF P 9327 and paratype SMF P 9328, and 5 newly collected specimens (figured specimens SMF PO 101.281).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B19FF9299BF6B5BFD7A5766.taxon	description	Figure 21 h Material 1 specimen SMF PO 101.235, Dar bel Hamri, Zanclean. Discussion Otoliths of S. kleinii are recognized by their compressed shaped (OL: OH = 1.1 – 1.15) in combination with somewhat undulating otolith rims and a pointed postdorsal projection. Synapturichthys kleinii is the only species of the genus occurring today in the Mediterranean and along the East Atlantic coast to South Africa into the western Indian Ocean to off Durban.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B19FF9299BF69BBFA9850E6.taxon	description	Figure 21 k Material 1 specimen SMF PO 101.236, Dar bel Hamri, Zanclean. Discussion Otoliths of V. chirophthalma are recognized by their compressed, high-bodied shape (OL: OH = 1.0 – 1.15) combined with a flat inner face, a convex outer face and a small, morphologically much reduced sulcus. Today, V. chirophthalma occurs only along the tropical coast of West Africa from Guinea-Bissau to Angloa.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B1BFF9099BF6E9BFBF850E6.taxon	description	Figure 22 e – g Holotype SMF PO 101.238 (Fig. 22 e), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 8 specimens SMF PO 101.282, same data as holotype. Etymology From Rharb, the geographical region in northwest Morocco where Dar bel Hamri is located. Diagnosis OL: OH = 1.95 – 2.1. Ventral rim shallow, smooth; dorsal rim shallow, coarsely crenulated. Rostrum relatively short, 20 – 25 % OL. Ostium short, CaL: OsL = 2.9 – 3.3. Curvature of caudal tip 40 – 50 °, terminating very close to postventral rim, caudal tip rounded and slightly widened. Description Relatively large and slender otoliths up to 9 mm in length (holotype 8.25 mm). OH: OT = 3.0 – 3.4. Ventral rim very shallow, regularly curved, smooth; dorsal rim shallow as well but intensely and irregularly crenulated, without discernable pre- or mediodorsal angles and variably developed postdorsal angle. Rostrum relatively short (completely preserved only in specimens of Fig. 22 f, g); antirostrum and excisura weak. Posterior rim blunt (Fig. 22 e) or oblique (Fig. 22 f, g). Inner face slightly convex with slightly supramedian positioned, deep, relatively wide and long sulcus. Ostium short, narrow, only slightly widened ventrally. Cauda long, straight, its tip bend downward with the downward oriented section nearly straight; caudal tip broadly rounded, slightly widened, terminating very close to postventral otolith rim. Dorsal depression narrow, ventrally marked by crista superior, dorsal margin indistinct; ventral field smooth, with feeble ventral furrow close to ventral rim of otolith. Outer face flat to slightly concave, relatively smooth. Discussion Tese otoliths are placed in the genus Caranx because of their resemblence with otoliths of the extant C. hippos (Linnaeus, 1766) and C. rhonchus Geoffroy Saint-Hilaire, 1817 as figured in Lombarte et al. (2006). It differs from extant and coeval Trachurus otoliths in the low ventral and dorsal rims. Caranx rharbensis is distinguished from said extant Caranx species in the short rostrum and rounded and widened caudal tip, and from parallel occurring Trachurus species (see below) in the slender shape (OL: OH = 1.95 – 2.1 vs. 1.55 – 1.9), the shallow dorsal and ventral rims and the rounded and widened caudal tip.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B1BFF979A056D3BFD4E5619.taxon	description	Figure 22 b – d Holotype SMF PO 101.283 (Fig. 22 b), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 11 specimens SMF PO 101.284, same data as holotype. Etymology From insectus (Latin) = notched, referring to the characteristic postdorsal notch. Diagnosis OL: OH = 1.55 – 1.7. Ventral rim with obtuse, rounded midventral angle; dorsal rim with broad lobe behind its middle followed by distinct notch. Rostrum relatively short, 15 % OL. Ostium short, CaL: OsL = 2.9 – 3.6. Curvature of caudal tip 37 – 45 °, terminating close to postventral rim. Description Moderately large and thin otoliths up to 6.9 mm in length (holotype). OH: OT = 4.0 – 4.5. Ventral rim moderately deep, anterior and posterior regions nearly straight, inclined, with rounded midventral angle, smooth or finely crenulated; dorsal rim anteriorly depressed, rising to broad lobe slightly behind middle follwed by deep and distinct notch and slightly expanded postdorsal section, irregularly undulating. Rostrum relatively short (completely preserved only in specimens of Fig. 22 b, d); antirostrum and excisura very weak. Posterior rim angular. Inner face slightly convex with distinctly supramedian positioned, deep, relatively narrow and long sulcus. Ostium short, narrow, only slightly widened ventrally. Cauda long, straight, its tip bend downward; caudal tip not widened, terminating close to postventral otolith rim. Dorsal depression narrow, ventrally marked by crista superior, dorsal margin indistinct; ventral field with feeble ventral furrow distant from ventral rim of otolith, smooth above ventral furrow, slightly plicated below. Outer face flat to slightly concave, relatively smooth. Discussion Tis species is recognized as a member of the genus Trachurus by the otolith shape with the midventral angle and the shape and proportions of the sulcus. It differs from the four extant congeners occurring in the eastern Atlantic in the short rostrum and the peculiar shape of the dorsal rim with the broad lobe behind the middle and the postdorsal notch. Otoliths of the parallel occurrig T. mediterraneus are further distinguished by the regularly rounded and intensely crenulated dorsal rim.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B1CFF969A0569B4FE545086.taxon	description	Figure 22 h – k Material 32 specimens, figured specimens SMF PO 101.239, Dar bel Hamri, Zanclean. Discussion Otoliths of T. mediterraneus differ from otoliths of T. trachurus in being slightly more compressed, showing a much stronger and finer crenulation of the dorsal and ventral rims and a more rounded postdorsal angle. Both species have been commonly recorded from Pliocene and Pleistocene sediments in the Mediterranean (e. g., Agiadi et al., 2019 a).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B1DFF9699BF695BFB1657A6.taxon	description	Figure 23 a – c Holotype SMF PO 101.285 (Fig. 23 a), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 3 specimens SMF PO 101.286, same data as holotype. Etymology Named after the Oued Beth along which the outcrops occur from which the otoliths were obtained. Diagnosis OL: OH = 1.5 – 1.6. Dorsal rim irregularly undulating, highest anteriorly; ventral rim regularly curved. Cauda distinctly inclined at angle of 50 – 65 °. Description Relatively large and elongate otoliths with overall oval shape reaching 9.5 mm in length (holotype). OH: OT = 3.0. Dorsal rim gently curved and irregularly undulating, highest anteriorly, with rounded postdorsal region. Ventral rim relatively shallow, gently and regularly curved. Rostrum short, blunt to broadly rounded; no or very weak excisura or antirostrum. Posterior tip broadly rounded, slightly inferior. Inner face distinctly convex, with slightly supramedian positioned, moderately deepened sulcus. Ostium short, its ventral rim box-shaped, the dorsal rim short and upward directed. Cauda long, narrow, reaching close to posterior tip of otolith, distinctly curved toward its tip. CaL: OsL = 1.75 – 1.85; OsH: CaH = 1.8 – 2.0. Dorsal field narrow, with indistinct, narrow depression; ventral field smooth without ventral furrow. Outer face concave, with small central umbo, smooth. Discussion Tese otoliths readily differ from the more common haemulid otoliths at Dar bel Hamri (Pomadasys incisus and P. zemmourensis n. sp.) in the more slender shape, the shallower and more gently curved ventral rim and the more strongly bent caudal tip. Parapristipoma bethensis is characterized through its relatively elongate shape and the gently curved ventral rim as a memebr of the genus Parapristipoma. Tree of four extant species of Parapristipoma are known from the east Atlantic. Otoliths are known from all of them (see Lombarte et al., 2006 and Nolf et al., 2009 for figures). Otoliths of Parapristipoma bethensis are less slender than those of P. humile (Bowdich, 1825) and the inner and outer faces are less strongly curved than in P. humile and P. octolineatum (Valenciennes, 1833) and the ostium is relatively shorter and wider. Parapristipoma bethensis resembles most P. macrops (Pellegrin, 1912) but differs in the rounded posterior rim (vs. tapering) and the broadly undulating anterior dorsal rim (vs. finely crenulated; see Lombarte et a., 2006 for figures of otoliths of P. macrops).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B1DFF9699BF6D9BFD8D5486.taxon	description	Figure 22 l – o 1906 Ot. (Sparidarum) mutinensis. Bassoli: pl. 2, Fig. 36.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B1DFF9699BF6D9BFD8D5486.taxon	materials_examined	Material 101 specimens, Zanclean, 99 specimens, Dar bel Hamri (figured specimens SMF PO 101.240), 2 specimens Jebel Zebbouj.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B1DFF9699BF6D9BFD8D5486.taxon	discussion	Discussion Verilus mutinensis is a long ranging and rather common species in the Mediterranean from the Tortonian (Bassoli, 1906) well into the Pleistocene (Agiadi et al., 2018). Today, the genus Verilus is not present anymore in the East Atlantic or Mediterranean.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B1EFF9599BF6FDBFE185506.taxon	description	Figure 23 d-h 2010 Pomadasys incisus (Bowdich, 1825) — Schwarzhans: pl. 78, Figs. 1 – 6 (see there for further references). 2022 Pomadasys incisus (Bowdich, 1825) — van Hinsbergh & Hoedemakers: pl. 31, Fig. 1. Material 518 specimens, Dar bel Hamri, Zanclean (figured specimens SMF PO 101.241).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B1EFF9599BF6FDBFE185506.taxon	discussion	Discussion Pomadasys incisus is one of the most common species in Dar bel Hamri. It is also a long-ranging species that has been recorded since the late Early Miocene (Schwarzhans, 2010, and references therein). Te otoliths of P. incisus are also remarkable for a pronounced late ontogenetic morphological change whereby large otoliths (14 mm in length, Fig. 23 d) become increasingly more elongate than smaller ones (8.7 – 11.8 mm in length, Fig. 23 e – h) (see Lombarte et al., 2006, for figures of extant otoliths). Tis effect is also documented in the ontogenetic sequence depicted in Lombarte et al. (2006). Today, P. incisus is distributed from the Strait of Gibraltar to Angola and is also known from the western Mediterranean (Froese & Pauly, 2022).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B1EFF9599BF6B5BFF0654C6.taxon	description	Figure 23 j – l Holotype SMF PO 101.242 (Fig. 23 j), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 6 specimens SMF PO 101.243, same data as holotype.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B1EFF8B9A0569BBFB805266.taxon	description	Figure 24 i – n 1989 Cepola rubescens Linnaeus, 1766 — Nolf & Cappetta: pl. 16, Fig. 4.? 2000 Cepola rubescens Linnaeus, 1766 — Nolf & Girone: pl. 4, Fig. 22. Holotype SMF PO 101.245 (Fig. 24 i), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 5 specimens SMF PO 101.246, same data as holotype. Further material 16 specimens, Zanclean, 5 specimens same data as holotype, 2 specimens Sidi Mohamed ech Chleuh, 1 specimen Asilah. Etymology Named in honor of Antoni Lombarte (Barcelona) in recognition of his contribution to the knowledge of extant otoliths. Diagnosis OL: OH = 2.03 – 2.1. Dorsal and ventral rims shallow. Cauda very narrow. Ostial colliculum terminating at some distance from anterior opening. Collum narrow. Description Slender, thin otoliths up to 4.2 mm in length (holotype). OH: OT = 2.8 – 3.0. Dorsal and ventral rims shallow. Dorsal rim nearly straight in central section, with weak or indiscernible postdorsal angle and broadly rounded predorsal angle. Ventral rim regularly curved, sometimes flattened at its center. Anterior tip pointed in large specimens, less in smaller ones; posterior tip rounded in small specimens, becoming more pointed in large ones but less pointed than anterior tip. All rims smooth or slightly undulating. Inner face distinctly convex, with slightly supramedian positioned narrow sulcus. OL: SuL = 1.4 – 1.5. Sulcus anteriorly open, but ostial colliculum not reaching anterior rim of otolith but terminating at some distance from it. Cauda very small and narrow, slightly shifted upwards. OsL: CaL = 2.1 – 2.5; OsH: CaH = 1.6 – 2.0. Cauda somewhat deepened with caudal colliculum less well defined than ostial colliculum. Dorsal depression indistinct; ventral furrow moderately developed, distant from ventral rim of otolith, anteriorly departing from ventral rim and leading to anterior tip of ostial colliculum. Outer face flat to slightly concave, smooth. Discussion Te genus Cepola currently contains five recognized valid recent species. Otoliths are known from four of those (except C. australis Ogilby, 1899) and are figured here for comparison: Cepola macrophthalma (Linnaeus, 1758), known from the northeastern Atlantic and the Mediterranean (Fig. 24 a); C. pauciradiata Cadenat, 1950, known from West Africa from Mauritania to Angola (Fig. 24 e – f); C. schlegelii Bleeker, 1854, known from Indonesia and the West Pacific (Fig. 24 g); and C. haastii (Hector, 1881) from New Zealand (Fig. 24 h). Te differences between the otoliths of these species are subtle and concern otolith proportions, course of the dorsal rim, and details of the sulcus. One important characteristic is the position of the ostial colliculum, which usually terminates at some distance from the anterior rim of the otolith in all species except C. macrophthalma, where it reaches the anterior rim of the otolith or approaches very closely. Cepola lombartei resembles the extant West African C. pauciradiata in otolith shape, but is more slender (OL: OH = 2.03 – 2.1 vs. 1.85 – 1.9) and has a narrower cauda. It also lacks the well-developed postdorsal angle of C. pauciradiata. Te Indo-West Pacific species (C. schlegelii and C. haastii) show a distinctly shorter ostial colliculum. It appears that C. lombartei has also been found in the Pliocene and possibly Pleistocene of the Mediterranean. Tese interpretations are based on published drawings (see synonymy listing) showing likewise slender otoliths with the ostial colliculum detached from the anterior rim of the otolith and are to be regarded tentative at present until revision.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B00FF8A9A056C7BFEBB5066.taxon	description	Figure 24 a – d 1980 Cepola macrophthalma (Linnaeus, 1758) — Nolf & Martinell: pl. 4, Figs. 25, 26. 1994 Cepola rubescens Linnaeus, 1766 — Nolf & Cavallo: pl. 7, Fig. 7. 1998 Cepola rubescens Linnaeus, 1766 — Nolf, Mané & Lopez: pl. 7, Fig. 12. 2019 a Cepola macrophthalma (Linnaeus, 1758) — Agiadi et al.: Fig. 5 G. 2022 Cepola macrophthalma (Linnaeus, 1758) — van Hinsbergh & Hoedemakers: pl. 30, Figs. 3 – 5. Material 27 specimens, Zanclean, 25 specimens Dar bel Hamri (figured specimens SMF PO 101.247), 1 specimen Sidi Mohamed ech Chleuh, 1 specimen Asilah. Discussion For differentiation of C. macrophthalma otoliths from C. lombartei, see above. Cepola macrophthalma otoliths have been referred to since Early Miocene from the North Sea Basin and the Mediterranean but are in much need of revision. Most or all of the Miocene specimens probably represent different species for which at least three nominal species names are available: C. praerubescens Bassoli, 1906 (Tortonian of Italy), C. voeslauensis Schubert, 1907 (Badenian of Austria), and C. multicrenata Radwańska, 1984 (Badenian of Poland). For further discussion, see Schwarzhans (2014). Terefore, I have accepted only Pliocene and younger otoliths as valid references here.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B01FF8A99BF6B5BFEFB5466.taxon	description	Figure 24 p Material 1 specimen, SMF PO 101.249, Dar bel Hamri, Zanclean.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B01FF8A99BF693BFB725006.taxon	description	Figure 25 a 2022 Trachinus sp. — van Hinsbergh & Hoedemakers: pl. 28, Figs. 7, 8. Material 3 specimens, Dar bel Hamri, Zanclean (figured specimen SMF PO 101.250).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B01FF8A99BF693BFB725006.taxon	discussion	Discussion Trachinus armatus today lives along the shores of West Africa from Senegal to Angola (Schwarzhans & Kovalchuk, 2022) but has also been tentatively recorded from the Middle Miocene of the Aquitaine Basin (Steurbaut, 1984). See Schwarzhans (2019 c) for figures of extant trachinid otoliths.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B01FF8A9A056E5BFC6B53C6.taxon	description	Figure 25 b – d Holotype SMF PO 101.251 (Fig. 25 b), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 4 specimens SMF PO 101.252, same data as holotype.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B02FF889A0569ADFF065066.taxon	description	Figure 25 e – h Holotype SMF PO 101.253 (Fig. 25 g), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 7 specimens SMF PO 101.254, same data as holotype.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B03FF889A056EDCFBCA5466.taxon	description	Figure 25 i 2010 Uranoscopus ciabatta — Girone, Nolf & Cavallo: Fig. 12 b 1 – 12 b 2. 2022 Uranoscopus sp. — van Hinsbergh & Hoedemakers: pl. 28, Fig. 11. Material 2 specimens SMF PO 101.287, Dar bel Hamri, Zanclean.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B03FF889A056EDCFBCA5466.taxon	discussion	Discussion Uranoscopus ciabatta was established based on two compact and thick otoliths from the pre-evaporitic Messinian of Piedmont, Italy, which fall out of the variation breadth observed in the extant U. scaber (see Girone et al., 2010). Van Hinsbergh and Hoedemakers (2022) described a unique very compressed otolith from the Piacenzian of Estepona as Uranoscopus sp. because of uncertainties in respect to the degree of variability known from the extant U. scaber. Now, with two more specimens of such compressed shape from the Zanclean of Morocco, these together with the Messinian and Piacenzian specimens are regarded as representing a single species, i. e., U. ciabatta.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B03FF8E9A056A7CFEE35786.taxon	description	Figure 25 p – q Holotype SMF PO 101.258 (Fig. 25 p), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratypes 3 specimens SMF PO 101.288, same data as holotype. Etymology Named in honor of Kristiaan Hoedemakers (Mortsel, Belgium) for his contribution to the knowledge of fossil otoliths. Diagnosis OL: OH = 1.8 – 1.9; OH: OT = 2.45. Dorsal rim with broad middorsal expansion. Sulcus deepened, anteriorly closed, with clearly distinct ostium and cauda. Anterior and posterior tips symmetrical, moderately pointed. OL: SuL = 1.75 – 1.9; OsL: CaL = 1.8 – 2.2. Broad dorsal field with many radial furrows. Description Robust, large otoliths up to 8.5 mm in length (holotype). Dorsal rim high, with broad mediodorsal expansion and with flat predorsal and slightly curved postdorsal regions, almost triangular in shape, somewhat irregular. Ventral rim shallower than dorsal rim, regularly curved. Anterior tip inferior, strongly projecting, rounded. Posterior tip at level with anterior tip, symmetrical in expression but slightly less projecting. Inner face mildly convex, with short, deepened, anteriorly closed sulcus. Ostium about twice as long as cauda and slightly wider, somewhat downward inclined towards collum. Cauda terminating far from posterior tip of otolith, slightly upward shifted against ostium. Dorsal depression very wide, dorsally open, with many radial furrows. Ventral field with three transverse furrows below central part of ostium. Ventral furrow running on ventral rim of otolith except somewhat turning inwards posteriorly. Outer face flat to slightly convex, somewhat irregularly shaped. Discussion Uransocopus hoedemakersi represents a different morphotype in the highly diverse genus Uranoscopus from the other species described here. Te otoliths of the three tropical West African species — U. albesca Regan, 1915, U. cadenati Poll, 1959, and U. polli Cadenat, 1951 — are known (Schwarzhans, 2019 c) and also represent different morphotypes. Otoliths of U. albesca and U. cadenati are somewhat similar in otolith shape and in the reduced sulcus morphology, but show much further separation of the sulcus from the anterior rim and a small, completely unstructured sulcus. Te closest morphotype is found in U. archionema Regan, 1921 (Fig. 25 o), a species distributed in the southeastern Indian Ocean along East Africa from Kenya to South Africa, Madagascar, and Mauritius and Reunion (Froese & Pauly, 2022). Otolith shape, proportions, and robustness and the wide dorsal depression with radial furrows are all shared characteristics. Uranoscopus hoedemakersi differs from U. archionema in the shorter sulcus (OL: SuL = 1.75 – 1.9 vs. 1.4) and the clearly structured sulcus with welldefined ostium and cauda (vs. contiguous ostium and cauda). Both species are thought to represent a vicariant species pair.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B05FF8E99BF69DCFACE53C6.taxon	description	Figure 25 j – k 2022 Uranoscopus scaber Linnaeus, 1758 — van Hinsbergh & Hoedemakers: pl. 28, Fig. 10 (see there for further references). Material 20 specimens, Dar bel Hamri, Zanclean (figured specimens SMF PO 101.255).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B05FF8E99BF69DCFACE53C6.taxon	discussion	Discussion Uranoscopus scaber is known for its large range in variability (see figures and discussion in Girone et al., 2010 and figures in Lombarte et al., 2006). Most Uranoscopus otoliths found in Dar bel Hamri can be convincingly placed within the morphological limits depicted by Lombarte et al. (2006) for U. scaber.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B05FF8D9A056D9CFDB85086.taxon	description	Figure 25 l – n Holotype SMF PO 101.256 (Fig. 25 m), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 4 specimens SMF PO 101.257, same data as holotype. Etymology Named in honor of Victor van Hinsbergh (Leiden, the Netherlands) for his contribution to the knowledge of fossil otoliths. Diagnosis OL: OH = 2.15 – 2.35; OH: OT = 2.2 – 2.8. Rostrum massive, moderately long, 13 – 17 % OL. OsL: CaL = 1.1 – 1.4. Outer face flat to concave. Description Elongate, thin and rather large otoliths reaching 7.3 mm in length (holotype 6.05 mm). Dorsal rim relatively shallow, irregular, slightly undulating, highest at its middle, without prominent angles; ventral rim shallow, gently curved. Rostrum well developed, relatively long and massive, with rounded tip. Excisura wide, broadly concave; no or very feeble antirostrum. Posterior rim blunt, with central or inferior rounded tip. Inner face distinctly convex, with moderately long, slightly bent, shallow sulcus. Ostium slightly longer than cauda and slightly wider, its anterior opening indistinct. OL: SuL = 1.5 – 1.7. Distinction of ostium and cauda and of colliculi feeble. Dorsal depression wide, dorsally open, with indistinct margins. Ventral furrow rarely visible, then very close to ventral margin of otolith. Outer face flat to concave, smooth.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B05FF8D9A056D9CFDB85086.taxon	discussion	Discussion Uranoscopus vanhinsberghi is closely related to U. scaber and differs in being more slender and thinner and showing a more massive rostrum. Te species thus falls outside the range of variations shown in Lombarte et al. (2006) for U. scaber. Uranoscopus vanhinsberghi may represent a sympatric vicariant species to the extant U. scaber, known in parallel from Dar bel Hamri and today from the Mediterranean and in the adjacent Atlantic from the British Isles to Mauritania.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B06FF8D99BF6A9BFA705086.taxon	description	Figure 26 a – b 1997 Boops boops (Linnaeus, 1758) — Nolf & Marques da Silva: pl. 1, Fig. 13. 2022 Boops boops (Linnaeus, 1758) — van Hinsbergh & Hoedemakers: pl. 33, Figs. 1 – 4. Material 3 specimens, Zanclean, 2 specimens SMF PO 101.259, Dar bel Hamri, 1 specimen SMF PO 101. 260, Asilah. Discussion Otoliths of Boops boops are recognized by the highly characteristic dorsal rim with the sharply bordered, flat, box-shaped postdorsal expansion (see Nolf et al., 2009 for extant otoliths). Today, the species occurs in the Mediterranean and the East Atlantic from Norway to Angola. It has also been recorded as fossil from the Piacenzian of Portugal (Nolf & Marques da Silva, 1997).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B06FF8D9A056D1BFC975446.taxon	description	Figure 26 d – f 1997 Diplodus aff. bellottii (Steindachner, 1882) — Nolf & Marques da Silva: pl. 2, Figs. 13, 14. Material 20 specimens (figured specimens SMF PO 101.261), Dar bel Hamri, Zanclean. Discussion Otoliths of Diplodus bellottii differ from those of its congeners by the relatively straight and distinctly inclined cauda (see Lombarte et al., 2006, for figures of extant otoliths). Nolf and Marques da Silva (1997) reported this species tentatively because of incomplete preservation from the Piacenzian of Portugal, which, following the new specimens from Morocco, can now be allocated with more certainty. Today, Diplodus bellottii occurs from southern Spain (Malaga, Cadiz) to the Cape Verde islands (Froese & Pauly, 2022). Its occurrence in the Alboran Sea is considered to represent a recent immigration through the Strait of Gibraltar (Golani et al., 2021).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B06FF8D9A056A5BFB9B57C6.taxon	description	Figure 26 c 2022 Oblada melanura (Linnaeus, 1758) — van Hinsbergh & Hoedemakers: pl. 33, Figs. 12, 13. Material 7 specimens, Dar bel Hamri, Zanclean (figured specimen SMF PO 101.262).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B06FF8D99BF6D1BFD655406.taxon	discussion	Remarks Te Sparidae represent the family with the largest diversity in Dar bel Hamri with 15 species. Te distinction of the otoliths of the many species often depends on rather subtle traits such as proportions of the otolith or the sulcus, curvature of inner and outer face, details of the otolith outline particularly of the dorsal rim and curvature of the caudal tip. Many small or poorly preserved specimens in the collection from Dar bel Hamri cannot be identified to species level and therefore are omitted from the description. Fortunately, many large and well preserved sparid otoliths exist as well from Dar bel Hamri and allow recognition of taxa. Te allocation of the otoliths to genera of the Sparidae is also a delicate task and often depends on direct comparison with extant species since no useful traits have been identified for a definition of genera by means of otoliths. Some genera contain rather different otolith morphological patterns, for instance Pagellus or Dentex. In the case of Dentex, the highly diverse otolith patterns strongly support the separation of the formal genus in independant clades as depicted in Chiba et al. (2009) and Santini et al. (2014). However, a character analysis of sparid otoliths has not yet been done, but see also comment to Dentex.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B06FF8C9A0569DBFD6A53C6.taxon	description	Figure 26 j 1980 Pagellus cf. acarne (Risso, 1827) — Nolf & Martinell: pl. 4, Fig. 13. 2022 Pagellus acarne (Risso, 1827) — van Hinsbergh & Hoedemakers: pl. 34, Figs. 3 – 8. Material 7 specimens, Dar bel Hamri, Zanclean (figured specimen SMF PO 101.289). Discussion Otoliths of Pagellus acarne resemble those of Diplodus bellottii in shape and proportions but differ in the relatively longer cauda (CaL: OsL = 1.4 – 1.5 vs. 1.1 – 1.3) and the fine crenulation of all otolith rims.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B07FF8C99BF6D9BFE8A5426.taxon	description	Figure 26 k 1997 Pagellus aff. bellottii Steindachner, 1882 — Nolf & Marques da Silva: pl. 2, Figs. 3, 4. 2022 Pagellus cf. bellottii Steindachner, 1882 — van Hinsbergh & Hoedemakers: pl. 36, Fig. 7. Material 13 specimens, Dar bel Hamri, Zanclean (figured specimen SMF PO 101.263). Discussion Otoliths of Pagellus bellottii are characterized by a relatively high body and pronounced mid- und postdorsal angles (see Lombarte et al., 2006 for figures of extant otoliths). Tey resemble P. erythrinus but are more compressed. Today, Pagellus bellottii occurs from the Strait of Gibraltar to Angola (Froese & Pauly, 2022). Its occurrence in the Mediterranean is considered to represent a recent immigration through the Strait of Gibraltar (Golani et al., 2021).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B07FF8C99BF6ABBFBDD5366.taxon	description	Figure 26 g – i 1979 Pagellus aff. bogaraveo (Brünnich, 1768) — Lanckneus & Nolf: pl. 3, Fig. 2. 2022 Pagellus bogaraveo (Brünnich, 1768) — van Hinsbergh & Hoedemakers: pl. 34, Fg. 9 (? 11 – 13, non 10). Material 26 specimens, Dar bel Hamri, Zanclean (figured specimens SMF PO 101.264). Discussion Large specimens of P. bogaraveo are characterized by a slender shape, rough crenulation of the dorsal rim, and a distinctive twist of the otolith along the long axis (Fig. 26 g 2). Te rostrum is sometimes bent downward at its tip (Fig. 26 h). However, otoliths of P. bogaraveo show a pronounced ontogenetic allometry, and smaller specimens are sometimes difficult to distinguish from other Pagellus species, notably P. erythrinus. A large ontogenetic series of extant otoliths is depicted by Lombarte et al. (2006). Today, P. bogaraveo is distributed from Norway to Mauritania and in the western Mediterranean (Froese & Pauly, 2022).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B07FF8C9A056D7BFAEB57C6.taxon	description	Figure 26 l 1979 Pagellus aff. erythrinus (Linnaeus, 1758) — Lanckneus & Nolf: pl. 3, Fig. 4. 1980 Pagellus aff. erythrinus (Linnaeus, 1758) — Nolf & Martinell: pl. 4, Figs. 14, 15. 1988 Pagellus aff. erythrinus (Linnaeus, 1758) — Nolf & Cappetta: pl. 15, Fig. 10. 1994 Pagellus erythrinus (Linnaeus, 1758) — Nolf & Cavallo: pl. 7, Fig. 5. 1997 Pagellus erythrinus (Linnaeus, 1758) — Nolf & Marques da Silva: pl. 2, Figs. 6, 7. 2022 Pagellus erythrinus (Linnaeus, 1758) — van Hinsbergh & Hoedemakers: pl. 36, Figs. 4 – 6.? 2022 Pagellus bogaraveo (Brünnich, 1768) — van Hinsbergh & Hoedemakers: pl. 34, Fig. 10 non 9, 11 – 13). Material 41 specimens, Dar bel Hamri, Zanclean (figured specimen SMF PO 101.265). Discussion Otoliths of P. erythrinus are amongst the most common sparids recorded from the European Pliocene. Tey differ from otoliths of P. bellottii primarily in being slightly more elongate (OL: OH = 1.4 vs. 1.3). Today, the species is distributed from Norway to Guinea Bissau and in the Mediterranean (Froese & Pauly, 2022).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B07FF829A0569DCFEA65326.taxon	description	Figure 26 m – o Material 39 specimens, Dar bel Hamri, Zanclean (figured specimen SMF PO 101.290). Discussion Otoliths of Pagrus pagrus are very similar to those of Pagellus erythrinus differing mainly in being slightly more elongate (OL: OH = 1.45 – 1.5 vs 1.4), which is partly counterbalanced by a more pronounced middorsal projection (Fig. 26 m). Pagrus pagrus, Pagellus erythrinus and Pagellus bellottii are resolved in one clade in the molecular phylogenetic studies in Chiba et al. (2009) and Santini et al. (2014). Otolith morphology confirms such interrelationships.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B09FF8299BF6DFBFE1C54A6.taxon	description	Figure 27 d 2022 Spicara alta (Osório, 1917) — van Hinsbergh & Hoedemakers: pl. 36, Figs. 8, 9. Material 24 specimens, Dar bel Hamri, Zanclean (figured specimen SMF PO 101.291). Discussion Fossil otoliths of S. alta were first described by van Hinsbergh and Hoedemakers (2022) from the Piacenzian of Estepona, Spain. It is relatively common in the Zanclean of Dar bel Hamri, and as commented by van Hinsbergh & Hoedemakers can be easily confused with small specimens of species of Opsodentex. Tey differ from O. angolensis in the lesser curvature of the inner face and the lesser curvature of the caudal tip. In the molecular phylogenetic analysis of Chiba et al. (2009), Spicara alta resolves in a clade also containing species of Dentex which are here placed in Opsodentex. Such a relationship of Spicara alta would be consistent with its otolith morphology. Spicara alta today lives along the tropical west African coast from Senegal to southern Angola (Froese & Pauly, 2022).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B09FF8299BF693BFC7350E6.taxon	description	Figure 27 a 2022 Spicara smaris (Linnaeus, 1758) — van Hinsbergh & Hoedemakers: pl. 35, Figs. 3 – 6. Material 2 specimens SMF PO 101.292, Dar bel Hamri, Zanclean. Discussion Two relatively small (4.5 mm in length) and slender otoliths (OL: OH = 1.75) are interpreted to represent S. smaris. Tey are further characterized by a low curvature of the inner face and a nearly straight caudal tip. For figures of extant otoliths of Spicara otoliths see Nolf et al. (2009). Spicara smaris occurs today in the Mediterranean and adjacent region of the northeast Atlantic.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B09FF819A056D3BFE655346.taxon	description	Figure 27 b – c Holotype SMF PO 101.293 (Fig. 27 c), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 4 specimens SMF PO 101.294, same data as holotype. Etymology From “ Mauretania tingitana ” (Latin) = the Roman name for the northwestern Moroccan region, deduced from Tingis (Latin), the ancient name of Tanger. Diagnosis OL: OH = 1.75 to approximately 2.0. Ventral rim very shallow, regularly curved; dorsal rim with low predorsal and distinct postdorsal angle; posterior rim slanted, concave. Rostrum relatively short, 15 % OL. CaL: OsL = 1.6. Curvature of caudal tip 45 – 55 °, terminating moderately close to postventral rim. Description Moderately large, elongate and thin otoliths up to at least 8 mm in length judging from the incomplete specimen of Fig. 27 b (holotype 5.7 mm). OH: OT = 3.4 – 3.7. Ventral rim shallow, regularly curved, smooth; dorsal rim with low predorsal angle, more pronounced postdorsal angle, slightly to irregularly undulating. Rostrum relatively short (completely preserved only in specimens of Fig. 27 c); no antirostrum or excisura. Posterior rim slanted, concave below postdorsal angle, with rounded or projecting inferior tip. Inner face slightly convex with distinctly supramedian positioned, moderately deep, long sulcus. Ostium short, wide. Cauda long, anteriorly straight, its tip bend downward at 45 to 55 °; caudal tip tapering or rounded, terminating moderately close to postventral otolith rim. Dorsal depression narrow, short, ventrally marked by crista superior, dorsal margin indistinct; ventral field with feeble ventral furrow close to ventral rim of otolith. Outer face distinctly concave, relatively smooth. Discussion Te genus Spondyliosoma contains two species today, S. cantharus (Linnaeus, 1758) in the Mediterranean and northeast Atlantic from the British Isles to Namibia (Froese & Pauly, 2022; for extant otoliths see Lombarte et al., 2006 and Nolf et al., 2009) and S. emarginatum (Valenciennes, 1830) in the southwestern Indian Ocean along the shores of South Africa and Madagascar (Froese & Pauly, 2022; for extant otoliths see Smale et al. 1995). Spondyliosoma tingitana resembles the otoliths of both extant species, but differ in the relatively strongly developed postdorsal angle, concave posterior rim and in being less strongly bent.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B0AFF8199BF691CFB8150A6.taxon	description	Figure 27 e – f Material 12 specimens, Dar bel Hamri, Zanclean (figured specimens SMF PO 101.266). Discussion Otoliths of D. canariensis are similar to those of D. dentex and differ in only a few subtle features: OL: OH ratio (1.5 – 1.6 vs. 1.65 – 1.7; D. canariensis first), being more strongly bent along the long axis of the otolith, a blunter posterior tip, and a more strongly bent caudal tip. However, many of these characteristics appear to be considerably variable and thus may only be useful for distinction in combination (see Lombarte et al., 2006, for extant otoliths of both species). Dentex canariensis occurs in the tropical East Atlantic from Cabo Bojador to Angola (Froese & Pauly, 2022).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B0AFF8199BF6D1BFE855746.taxon	discussion	Remarks Te genus Dentex has been subdivided into several subgenera, most of which are regarded as valid genera in the recent literature (Fricke et al., 2022). Molecular studies (Chiba et al., 2009 and Santini et al., 2014) have consistently shown the genus Dentex to be polyphyletic. Te species currently allocated to Dentex (Froese & Pauly, 2022) contain two distinct otolith morphologies, one with more elongate otoliths and the other with more compressed otoliths (Nolf, 1979). As far as their otoliths are known, the distinction of the two otolith morphotypes reflects the the clustering of the molecular phylogeny. Te type species, Dentex dentex, belongs to the elongate otolith morphotype. Te compressed otolith morphotype is found in the species D. angolensis, D. congoensis, D. macrophthalmus and D. maroccanus (see Nolf, 1979). Dentex macrophthalmus is the type species of Opsodentex, established as subgenus of Dentex by Fowler (1925). In the light of the congruence of molecular and otolith morphology data I, therefore, propose to use Opsodentex as a valid genus for the above mentioned four nominal Dentex species, the fossil otolith-based Dentex gregarius (Koken, 1891) and Opsodentex mordax n. sp. described in the following. Fossil otolith-based evidence shows that these two morphotypes belong to lineages that have been separate since at least the Late Oligocene (Schwarzhans, 1994, 2010). It should be noted, however, that distinction of Opsodentex, Polysteganus and Evynnis by means of otolths is complex and requires further investigation.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B0AFF819A056D3CFC5452E6.taxon	description	Figure 27 h – i Material 19 specimens, Dar bel Hamri, Zanclean (figured specimens SMF PO 101.267). Discussion For distinction from otoliths of D. canariensis, see above. Dentex dentex is distributed in the Mediterranean and northeastern Atlantic from the British Isles to Mauritania (Froese & Pauly, 2022); today, it has only a small area of overlap with the vicariant D. canariensis (see above).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B0AFF819A056B3BFA705666.taxon	description	Figure 27 g Material 14 specimens, Dar bel Hamri, Zanclean (figured specimen SMF PO 101.268). Discussion Te distinction of the four extant species in Opsodentex (O. angolensis, O. congoensis, O. macrophthalmus, O. maroccanus) is difficult and may not be possible in many cases. Veen and Hoedemakers (2005) commented that these species could not be distinguished and combined their otoliths in a “ group de Dentex maroccanus. ” Van Hinsbergh and Hoedemakers (2022), however, distinguish otoliths of O. macrophthalmus and O. maroccanus in the Pliocene of Estepona near Málaga without giving a rationale. It appears that they concluded that O. macrophthalmus has more stretched otoliths than O. maroccanus, a view that I follow. Opsodentex angolensis is similar to the otoliths of O. macrophthalmus but differs in a more strongly bent caudal tip (up to 60 °), although this is not evident in all cases studied by me or figured in the literature (see Veen & Hoedemakers, 2005, and Lombarte et al., 2006). Today, O. angolensis is distributed from Mauritania to Angola (Froese & Pauly, 2022).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B0AFF879A05687BFF0650A6.taxon	description	Figure 27 j – m Holotype SMF PO 101.269 (Fig. 27 j), Dar bel Hamri, coquina at river level of Oued Beth, Zanclean. Paratype 6 specimens SMF PO 101.295, same data as holotype.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B0CFF879A056D1BFBB35786.taxon	description	Figure 28 a – d 1993 Afroscion trewavasae — Schwarzhans: Fig. 246 – 250.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B0CFF879A056D1BFBB35786.taxon	materials_examined	Material 1564 specimens, Dar bel Hamri, Zanclean, figured specimens are photographs of holotype, SMF P 8226 (Fig. 28 b) and paratypes, SMF P 8227 (Fig. 28 a, c – d).	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B0CFF879A056D1BFBB35786.taxon	discussion	Discussion Afroscion trewavasae is the most common species at Dar bel Hamri. I know of no other fossil otolith location where a sciaenid species represents the most common species. Furthermore, the majority of specimens are large, in the size category of 8 to 14 mm in length, while specimens 5 mm in length (Fig. 28 d) or smaller are rare. At Vale de Freixo (Portugal, Piacenzian) Nolf and Marques da Silva (1997) recorded A. trewavasae as one of the most common species in their assemblage, trailing only a goby (Deltentosteus sp.) and an ophidiid (Ophidion rochei), but they described a more continuous ontogenetic sequence starting with specimens slightly over 1 mm in length. Afroscion contains a single extant species, A. thorpei (Smith, 1977), distributed in southeastern Africa from Mozambique to Algoa Bay, where juveniles are found on sand and mud and adults predominantly on reefs (Sasaki in Heemstra et al. 2022). Even though obviously not reef-associated, it appears likely that the fossil northeastern Atlantic A. trewavasae lived in different environments during its ontogeny. Afroscion is often placed as a junior synonym of Argyrosomus de La Pylaie, 1835, since Sasaki and Kailola (1988), but in the light of its distinct lineage and disjunctive distribution pattern, I consider it valid.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B0CFF869A05681BFD765286.taxon	description	Figure 28 e Material 1 specimen, SMF PO 101.296, Dar bel Hamri, Zanclean. Discussion In a recent analysis of Song et al. (2017) the hitherto widely distributed East Atlantic / Indo-West Pacific species A. aequidens (Cuvier, 1830) has been subdivided into four geographically separated species. Specimens from West Africa are placed in A. macrolepis, while A. aeqidens is restricted to South Africa. Terefore, specimens figured from South Africa in Schwarzhans (1993) and Smale et al. (1995) would represent A. aequidens and specimens figured in Lombarte et al. (2006) from Mauritania may represent A. macrolepis. However, Song et al. only studied A. macrolepis specimens from Angola and Namibia and, therefore, the identity of further northerly specimens remains tentative. Te unique specimen from Dar bel Hamri is consistent with the morphology of the extant specimens from Mauritania figured by Lombarte et al. and should represent the same species. Song et al. figured an otolith of A. microlepis Song et al., 2017, but a comprehensive analysis of the otoliths of the various extant species of the genus is still outstanding.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B0DFF8699BF6B1BFEF054A6.taxon	description	Figure 28 g Material 1 specimen, SMF PO 101.270, Dar bel Hamri, Zanclean. Discussion Te single, relatively large otolith of 8.6 mm in length is very similar in all morphological aspects to the extant P. typus known from Mauritania to Angola (see Schwarzhans, 1993 for figures of extant specimens). Te identification, however, is only tentative because of some abrasion of the anterior otolith rim and in the area of the caudal tip.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B0DFF859A056B1BFE565086.taxon	description	Figure 28 i – l 1993 Pteroscion maroccanus — Schwarzhans: Fig. 289 – 292. 2003 Pteroscion peli (Bleeker, 1863) — Mendiola & Martínez: Fig. 9: 4 – 6, pl. 13, Fig. 289 – 291. 2003 Pteroscion guardamarensis — Mendiola & Martínez: Fig. 9: 7 – 9, pl. 12, Figs. 1 – 8, pl. 15, Figs. 1 – 8, pl. 16, Figs. 1 – 2. 2003 Pteroscion sp. 1 — Mendiola & Martínez: pl. 12, Figs. 9 – 16, pl. 16, Figs. 3, 4. Material 46 specimens, Dar bel Hamri, Zanclean, figured specimens are photographs of holotype of P. maroccanus, SMF P 8599 (Fig. 28 h) and paratypes, SMF P 8600 (Fig. 28 i – k). Discussion I agree with Mendiola and Martínez (2003) and in the light of the many extant and fossil species they figured that P. maroccanus is synonymous with the extant P. peli. However, I also agree with the view of the reviewer of their manuscript, D. Nolf (as mentioned in their acknowledgements), that the species they described from the Pliocene of Spain as P. guardamarensis represent a further synonym of P. peli. Te species was found to be by far the most common sciaenid in the lower Pliocene rocks of Guardamar in southern Spain (Mendiola & Martínez, 2003). Today, Pteroscion peli is distributed from Senegal to Namibia.	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
03ED255D3B0EFFF599BF6D1BFB415405.taxon	description	Figure 28 f 1993 Umbrina aff. canariensis Valenciennes, 1843 — Schwarzhans: Fig. 88 – 90. 2003 Umbrina canariensis Valenciennes, 1843 — Mendiola & Martínez: pl. 2, Figs. 3 – 7, pl. 3, Figs. 1, 2. 2003 Umbrina aff. canariensis Valenciennes, 1843 — Mendiola & Martínez: pl. 4, Figs. 1 – 4. Material 43 specimens, Dar bel Hamri, Zanclean (figured specimen SMF PO 101.272). Discussion Several large and well preserved specimens have now been obtained from Dar bel Hamri confirming the identitiy of these otoliths, which previously have been only tentatively allocated as U. aff. canariensis (see Schwarzhans, 1993). Diversification level and environmental assessment Ninety-six otolith-based species have been identified based on 4375 specimens collected from four locations along the Oued Beth and one location near Asilah (Table 1), whereby 4250 otoliths were obtained from a single location, the coquina at the base of section 1, approximately 1.5 km south of Dar bel Hamri, with 95 species. Terefore, only location 1 near Dar bel Hamri qualifies for a quantitative assessment of the bony fish fauna. However, only subsample 1 with 1202 specimens can be quantitatively evaluated because subsample 2 was screened off at 2 mm mesh diameter and thus small species are not adequately represented in it. Percentages in the following therefore reflect only subsample 1 (Table 1). Te two most common species are Afroscion trewavasae (19.55 %) and Diaphus maghrebensis (16.39 %), together accounting for 35.94 % of the entire assemblage (Table 1). Te diversification index is 31 of the most common species to reach the threshold of 90 % of all identified species. Tis is an exceptionally high diversification index and in fact the highest ever recorded in any otolith assemblage. Tis high diversity is probably a result of several interrelating effects, such as environment, sedimentary facies, water depth, rich and diverse food supply at various trophic levels, and the biogeographic situation (see below). Te coquina of Dar bel Hamri, from where the otoliths were obtained, has been deposited in a subaqueous environment, probably at moderate depth on the middle to lower shelf. Coquina accumulations observed in Late Miocene / Early Pliocene sediments of Portugal, the Huelva Basin in Spain, and the Rharb Basin in Morocco have been interpreted by Gonzales Delgado et al. (1994) as caused by winnowing from storm activities in shallow marine environments below the wave base. However, the composition of the otolith assemblage points to a deeper environment in a middle to lower shelf position that would be below the storm wave base. Te majority of the 43 extant fishes occurring as fossils in Dar bel Hamri occur at water depth of 50 to 100 m (55.8 %), another 18.6 % in water shallower than 50 m and the remainder in water deeper than 100 m (Table 2). About 58 % of the 43 extant fishes occurring as fossils in Dar bel Hamri have a demersal life habitat (Table 2) making them relatively reliable depth indicators. Te winnowing responsible for the formation of the coquina and the accumulation of fossils in the sediment at Dar bel Hamri instead could have been caused by subaqueous currents, possibly as a result of tidal reflux. Such explanation is supported by the report of a Late Miocene / Early Pliocene channel observed on seismic sections due east of Dar bel Hamri (Capella et al., 2017). Te current and winnowing at the base of the early Zanclean may also have been responsible for some reworking and erosion of directly underlying sediments of (late) Messinian age in the area. Furthermore, the current activity could have led to an increase in the availability of food in the trophic chain. In combination with the water activity this in turn could have led to an unusually high percentage of large, adult fishes as expressed in the high percentage of relatively large otoliths in the coquina. Te most abundant species, Afroscion trewavasae, is a sciaenid of a group that today feeds primarily on small, mostly nektonic fishes and large invertebrates. Te abundance of such a fish, particularly with adult specimens, is unusual and supports the hypothesis of an increased food supply in the depositional environment. Te abundance of large otoliths also increases the potential for their identification, which is somewhat counterbalanced by a nearly ubiquitous mild to moderate mechanical erosion, which one would expect in such a facies. Te other, stratigraphically comparable locations at Sidi Mohamed ech Chleuh, Kef Nsour and Jebel Zebbouj have yielded much fewer otoliths and are dominated by myctophids. Te myctophid otoliths in these locations were mostly well preserved including the frail denticles along the ventral rim of Diaphus otoliths, and thus indicate no or very little erosion in a more calm and possibly also deeper environment. Asilah has yielded even fewer otoliths, which are often somewhat leached hampering identification. Tis small assemblage contains only species also known from Dar bel Hamri. Lyellian percentage Fourty-three species identified by means of otoliths in the Early Pliocene of Dar bel Hamri are also known from today. Tis compares to 39 fossil species and 14 that cannot be identified to species level. Disregarding the latter which could either represent extant or extinct species, the Lyellian percentage thus amounts to 52.4 %. A calculation on the same basis for the otolith assemblage recently described from the Pliocene of Estepona near Málaga, southern Spain, by van Hinsbergh and Hoedemakers (2022) arrives at a Lyellian ratio of about 67 %. An evaluation of the Lyellian percentage for goby otoliths in the Mediterranean arrived at 70 % (Schwarzhans et al., 2020). Overall, the Lyellian percentage of otolith associations in the Zanclean of the Mediterranean is in the order of 72 % and for the Piacenzian 78 %. However, articulated skeletons exhibit a lower Lyellian percentage in the Early Pliocene of the Mediterranean than otoliths (Landini & Sorbini, 1992, 2005), which may be regarded as an expression of the generally more conservative approach in otolith research. Overall, the Lyellian percentages of Early Pliocene otolith associations vary between about 50 % (tropical West Atlantic Sciaenidae, Morocco) and about 75 % (New Zealand) (see Schwarzhans, 2019 a). Tus, the Moroccan otolith-based fauna described here is remarkable for a comparably low Lyellian percentage, which will be discussed further in the chapter below about a potential “ Maghrebian bioprovince ” during the Pliocene. Te ratio of extant versus extinct species is unevenly distributed among the families represented. Considering families with at least three identifiable species, one can note that extinct taxa prevail in the Myctophidae (5 extinct species in a total of 8 species), Gobiidae (4 in 5), Bothidae (3 extinct species), Trachinidae (with 2 extinct species in 3) and Uranoscopidae (3 extinct species in 4). I interpret the low yield of extant species in the Early Pliocene of these families as an indication of a dynamic speciation in the recent geologic past. Te dynamic recent speciation is matched with observations made about Pliocene myctophid otoliths from tropical America (Schwarzhans & Aguilera, 2013). In respect to gobies, the extant – extinct ratio is somewhat mixed. Schwarzhans et al. (2020) calculated a Lyellian percentage of about 70 % for the Gobiidae in the Early Pliocene of the Mediterranean. A calculation from the species list provided by van Hinsbergh and Hoedemakers (2022) for the Pliocene of Estepona near Málaga, southern Spain, arrived at about 62 %. Tus, the yield of extinct gobiid species in Dar bel Hamri is distinctly higher than in Mediterranean locations, even though all except one of the species (Deltentosteus planus) found in Morocco are also known from the Mediterranean. I find no ready explanation for this discrepancy. Conversely, the yield of extinct species is low in the Congridae (1 extinct species in 4), Carapidae (1 extinct species in 3), Soleidae, and Sciaenidae (1 extinct species in 5) and particularly in the Sparidae (2 extinct species in 15). Te high percentage of extant species in the Early Pliocene record of these families could indicate that much of the speciation that led to the extant fauna occurred earlier. However, the low yield of extinct taxa in the family with the most species in Dar bel Hamri, the Sparidae, could have a different cause. Otoliths of the Myctophidae (Brzobohatý & Nolf, 1996, 2000; Schwarzhans, 2013 b; Schwarzhans & Aguilera, 2013) and of the Gobiidae (Schwarzhans et al., 2020; Bratishko et al., 2023 ms, and literature cited in both studies) have been studied extensively and have in part been calibrated by finds of otoliths in situ (Bedini et al., 1986; Reichenbacher & Bannikov, 2021, 2022; Schwarzhans et al., 2017). As a result, the character analysis of otoliths of these two groups is more advanced than in Sparidae and aids the recognition of species. Comparable studies of otoliths are lacking in the case of the Sparidae, and it is, therefore, possible that the species identification is not as accurate as in the Myctophidae and Gobiidae and could in fact be too conservative. Comparison with Mediterranean assemblages from the Pliocene: implications for fish remigration from the Northeast Atlantic into the Mediterranean after the Messinian Crisis Te terminal Miocene in the Mediterranean is well known for the late Messinian Salinity Crisis (MSC) that was caused by the closure of the Mediterranean Sea to the northeastern Atlantic in the West during that time. Both seas became fully reconnected again beginning with the Pliocene. Since the first article on the MSC (“ When the Mediterranean dried up ” by Hsü, 1972), a large body of research has been published on the subject, and what precisely happened — whether marine life was extinguished in the Mediterranean during the event, the water budget, base level drop, and so on — is still very much under discussion (e. g., Roveri et al., 2014, 2016; Ben- Mosche et al., 2020; Gvirtzman et al., 2022; and literature cited in these articles). Recently, Carnevale and Schwarzhans (2022) showed that stenohaline marine fishes lived in the Mediterranean through the MSC, at least periodically, as evidenced by articulated fish skeletons and otoliths, which is in contrast to a study by Andreetto et al. (2021), who postulated that all marine fossils found in sediments of the MSC interval resulted from reworking of pre-MSC rocks. Te reworking hypothesis cannot be maintained for the explanation of the presence of articulated skeletons of marine fishes in MSC rocks. Another subject of continued dispute is the role of re-flooding of the Mediterranean with the onset of the Early Pliocene (Zanclean) (Bache et al., 2012; Garcia-Castellanos et al., 2009; Micallef et al., 2018; and literature cited in these articles). During the Pliocene, the Rharb Basin was a funnel-shaped embayment that represented the relict of the last connection of the Atlantic with the Mediterranean (Fig. 29) (Achalhi et al., 2016; Capella et al., 2017; de Weger et al., 2020 a, 2020 b, 2020 c; Flecker et al., 2015; Flinch, 1993; Martin et al., 2014; Pérez-Asensio, 2021). Te Rharb Basin is ideally situated to investigate the composition of the fauna during the late stage of the Atlantic – Mediterranean connection and the fauna during the re-flooding when the Rharb Basin represented the reservoir from which a proportion of the remigration must have been recruited. Little is known so far about the Late Miocene fish fauna of the Rharb Basin from the few otoliths collected from outcrops at Kef Nsour and Chaba Kaudiat el Mogen, which are all known in the Early Pliocene as well, except for Diaphus draconis. Te Early Pliocene fish fauna from Dar bel Hamri and the other studied locations is significant and allows for a correlation with time-equivalent assemblages from the Mediterranean. Indeed, there have been many studies about Pliocene otolith associations in the Mediterranean realm (Fig. 30): from Italy for example Dieni (1968), Weiler (1971), Schwarzhans (1978 a), Anfossi and Mosna (1979), Nolf and Cappetta (1988), Girone (2006), Nolf and Girone (2006); from Greece Agiadi et al., (2013, 2017, 2019 a, 2019 b); from southern France Schwarzhans (1986), Nolf and Cappetta (1988); from southern Spain Nolf and Martinell (1980), Nolf et al. (1988), Mendiola and Martínez (2003), van Hinsbergh and Hoedemakers (2022). Tese many articles contrast with only two otolith associations described from the adjacent Atlantic: one from the Piacenzian of central Portugal described by Nolf and da Silva (1997) and the other this study from Morocco. Many of the Zanclean otolith assemblages described from the Mediterranean realms originate from deep marine settings with predominant meso- to bathypelagic and bathydemersal fishes, while the Portuguese and Moroccan faunas are characterized by middle to lower shelf fishes associated with a few upper slope elements. Te assemblage described from Estepona in southern Spain by van Hinsbergh and Hoedemakers (2022) is the most important for comparison, because it is rich (209 species, 107 thereof in the Zanclean), includes faunal elements of the lower shelf similar to Morocco, and is located just about 50 km to the east of the Strait of Gibraltar. Te nearest location studied in Morocco is Asilah, about 50 km to the west of the Strait of Gibraltar and about 130 km from Estepona. Dar bel Hamri is 190 km south of the Strait of Gibraltar and about 260 km from Estepona. However, Dar bel Hamri is much closer to Estepona than the other nearest Mediterranean locations (450 km to Guardamar, described by Mendiola & Martínez, 2003, and 850 km to Papiol near Barcelona, described by Nolf et al., 1998). Fifty of the 82 identified species in the Early Pliocene of the Rharb Basin have also been recorded from time-equivalent strata of the Mediterranean (Table 1; 61 %), thereof 39 species in the Zanclean and / or Piacenzian of Estepona (Fig. 29). Tis ratio is consistent with the similarity coefficient calculated by Ben Moussa (1994) for Pliocene bivalves of Morocco in comparison with Mediterranean localities. Nevertheless, the fish fauna shows a lower correlation than one might expect over such a small distance and under the consideration of the recruitment of the Mediterranean fauna from the adjacent Northeast Atlantic during the Early Pliocene re-flooding event. It appears that not all species of the Northeast Atlantic were actually able to migrate into the Mediterranean, and furthermore there could be environmental differences at play in the locations that could mask faunal exchange. However, it is worth elucidating those 32 species that have not been found in the Mediterranean. Tirteen of those represent extant species, of which today 5 occur in the Atlantic off Morocco, 7 primarily south of the Mauritanian upwelling zone in the tropical East Atlantic and 1 in the tropical West Atlantic. Another 19 species, including all of the new species, could be considered potentially endemic to the area (a “ Maghrebian bioprovince ”, see below). Pterothrissus darbelhamriensis of those potentially endemic species may in fact represent an allopatric vicariant species of the Mediterranean P. compactus at the time. Pterothrissus darbelhamriensis was already considered by Schwarzhans (1981) as vicariant to P. compactus from the Mediterranean (known from the Tortonian [Fig. 9 h, j] until Zanclean [Fig. 9 f – g, i]). It has now also been identified from the Piacenzian of Estepona (van Hinsbergh & Hoedemakers, 2022), but this is not considered contradictory to the interpretation, since it could represent a late immigrant from the Northeast Atlantic into the adjacent region of the Mediterranean. Te recognition of a vicariant species in the Northeast Atlantic and the Mediterranean in the Zanclean (and ideally since the Late Miocene) supports the hypothesis of in situ survival of marine fish taxa in the Mediterranean, but it is so far restricted to a single conclusive case. Te more important outcome of the correlation of Northeast Atlantic and Mediterranean fishes of the Early Pliocene times is that a substantial number of Atlantic fish taxa apparently did not migrate into the Mediterranean during the re-flooding. Tis non-migratory effect could be caused by a warmer climate in the Rharb Basin than at the Strait of Gibraltar, as can be inferred from the presence of some southern elements then present in Morocco and putative endemic species. Tey are primarily the species with tropical East Atlantic affinities that have not been found in the Mediterranean. Comparison with extant fish faunas and its implications Te otolith association of the Early Pliocene of Dar bel Hamri and nearby locations contains 43 species that still exist today. Te majority of them are distributed over the subtropical to temperature zones (74.4 %), 30.2 % thereof exclusively in the subtropical zone (Table 2). Te distribution of only 20.9 % of the species range into the temperate zone, and 25.6 % are exclusively tropical (Table 2). Tirty-two species (74.4 %) of the extant fishes occurring as fossils in Dar bel Hamri live today in the same general area (i. e., in the Northeast Atlantic off the coasts of Morocco) (Table 3); and 9 species (20.9 %) live today in the tropical East Atlantic, chiefly south of Mauretania. Clearly, the high content of persistent species indicates a high degree of continuity in the faunal composition of species occurring today and during the Early Pliocene in the region, but tropical West African species, which do not live off Morocco anymore or occur rarely as stray specimens, also contributed a significant component. Teir presence indicates that the water temperatures in the sea off northwestern Morocco were significantly warmer in the Early Pliocene than they are today. Similar conclusions were drawn by Avila et al. (2016) on the basis of molluscs found in Late Miocene and Early Pliocene sediments on the Azores and many previous studies of molluscs cited therein. If the sea off northwestern Africa was warmer in the Early Pliocene than it is today, this must have had consequences for the paleo-currents in the region. Today’s faunal provinces along the northwestern coast of Africa are driven by the cool Canary Current, the permanent coastal upwelling between Cape Yubi and Cape Blanc, and further offshore by the position of the Intertropical Convergence Zone (Matsuzaki et al., 2011; Michel et al., 2011). Te change from a tropical sea off Morocco to the current subtropical situation dominated by the cool Canary Current and separation from the tropical West African sea by the coastal upwelling system probably occurred during the Late Pliocene. Studies of the mollusc faunas from the Mediterranean, the Azores and the Canary Islands have revealed the disappearance of tropical taxa between 4.2 and 3.0 Ma (i. e., latest during the mid-Piacenzian cooling event) (Avila et al., 2016, and literature cited therein). Even though indications for upwelling along the Northwestern African coast appear to have been present since the Early to Middle Miocene (Diester-Haas & Schrader, 1979), one has to assume that during the Late Miocene and Early Pliocene its intensity was insufficient to keep (some) tropical West African fishes from living in the Rharb Basin. However, some degree of separation still must have been effective, since most Early Pliocene fishes exhibit a clear subtropical Northeast Atlanto-Mediterranean relationship. In respect to the total of 74 biogeographically interpretable species (excluding fossil mesopelagic and bathydemersal fishes of uncertain biogeographic affinities), 42 species corresponding to 56.8 % are identical or related to extant species in the same area (Table 3). Twenty-three species (31.1 %) are identical or related to tropical East Atlantic species (Table 3). Nineteen species (25.6 % of above 74 species) are putative endemics during the Early Pliocene of Dar bel Hamri; they are mostly related to extant Mediterranean, tropical East Atlantic and South African species (Table 3). Of about 600 marine bony fish species recorded from Morocco, the Canary Islands, Madeira and the Azores today, only about 30 (5 %) represent northward extensions of species with primarily tropical East Atlantic distribution patterns (calculated from Froese & Pauly, 2022). Te relationships of the 31 extinct otolith-based fish species at Dar bel Hamri (of the 74 biogeographically interpretable species) on a stand-alone basis are naturally less certain. Te largest group among them is 19 species (25.6 %) that have not been recorded from Mediterranean localities and hence are considered of potentially endemic nature (Table 3). When investigating the probable provenances of the more exotic species (Table 3; Fig. 31), some more remote relationships are notable. Paratrisopterus glaber shows the closest relationship with northern Atlantic / North Sea Basin taxa. Tree species (4.1 %) exhibit affinities to the tropical West Atlantic (Paraxenomystax cf. bidentatus, Myripristis ouarredi and Verilus mutinensis), three (4.1 %) to South Africa (Centroberyx vonderhochti, Uranoscopus hoedemakersi and Afroscion trewavasae), and two (2.6 %), namely Brotula aff. multibarbata and Laeops rharbensis, to the Indo-West Pacific (Table 3). A somewhat special case is that of the genus Rhynchoconger which has had a long history in Europe since Eocene times. Its latest representatives are R. carnevalei in the Early Pliocene of the Mediterranean and R. pantanellii in the Pliocene of the Mediterranean and Northeast Atlantic. Te nearest occurrence of the genus today is in the central West Atlantic, but otoliths found in Holocene dredge samples in the Gulf of Guinea (Schwarzhans, 2013 a) indicate that the disappearance of the genus from the East Atlantic is either very recent or false. Te occurrence of another species with a West Atlantic relationship, Verilus mutinensis, is of a different nature, as it probably represents a vicariant East Atlantic / Mediterranean species ranging from the Late Miocene Tortonian (Bassoli, 1906) to the middle Pleistocene Calabrian (Agiadi et al., 2018) before the lineage became extinct in the East Atlantic. As for the Indo-West Pacific affinity, Brotula aff. multibarbata could possibly represent a fossil species (subject to further material becoming available) and in any case represents a clade not present anymore in the Atlantic (Froese & Pauly, 2022). Te South African link is particularly interesting and perhaps unexpected, as it also contains the most common otolith-based species (Afroscion trewavasae) of the entire assemblage. Te extant A. thorpei is geographically restricted to a rather small area in southeast Africa (Sasaki in Heemstra et al. 2022). Te occurrence of A. trewavasae in Morocco and Portugal clearly represented a vicariant species, and the distribution of Afroscion in southeast Africa must thus be considered a relict occurrence. Tis coincides with South Africa representing a classical region for (secondary) endemism (http: // lntre asures. com / rsa. html). Te same explanation may hold for Uranoscopus hoedemakersi, which is thought to represent a vicariant species to the extant U. archionema off southeastern Africa. Centroberyx is a slightly different case as it is today known from seven species with a disjunctive distribution pattern, richest in Australia and New Zealand with an outlier species each in Japan, Taiwan, and South Africa (Froese & Pauly, 2022). Centroberyx is a genus with a long history reaching back into Cretaceous times and is well known from European basins until the Eocene, after which it becomes rather sparse (e. g., Schwarzhans & Jagt, 2021). Its current distribution is clearly a relict of a formerly much wider pattern. In Europe, the last record so far was C. manens Nolf & Brzobohatý, 2004, from the Middle Miocene. Te large specimens of Centroberyx vonderhochti in the Pliocene of the Rharb Basin represent a different lineage from C. manens, that is probably related to the extant C. spinosus (Gilchrist, 1903) from South Africa (see Schwarzhans & Jagt, 2021). Was there a “ Maghrebian bioprovince ” during the Early Pliocene? Today’s fish fauna in the seas around Morocco and the Macaronesian archipelago (Canary Islands, Madeira, Azores) includes about 600 species, of which about 30 species (5 %) can be considered endemic and another 5 % as primarily tropical West African fishes that also occur rarely to the north of the Mauritanian upwelling system (calculated from Froese & Pauly, 2022). Tis bioprovince has been named the subtropical Mediterranean – Moroccan Province by Avila et al. (2016). Most of the 560 or so non-endemic indigenous fishes have distribution ranges northward to the Bay of Biscay or the British Isles, into the Mediterranean, and / or southward to various extents. Te situation was quite different in the Early Pliocene (and Late Miocene), when the Moroccan coast was under warmer climatic conditions and the Transition Zone was located further north. Based on analysis of the molluscan fauna, Avila et al. (2016) postulated a Pliocene Mediterranean – West African Province that stretched from the Azores to the southern tip of Portugal and incorporated the entire Mediterranean and the Gulf of Guinea. Te fish fauna from the Rharb Basin, however, indicates that some degree of differentiation existed in this large area at least toward the Mediterranean, and is also inferred toward tropical West Africa (Fig. 32). Te Early Pliocene otolith-based fish fauna of the Mediterranean is exceptionally well known and includes more than 200 identified taxa. Te recent monograph of the otolith assemblage from Estepona in southwest Spain not far from the Strait of Gibraltar is particularly important for correlation. Only half (52 %) of the otolith taxa identified in the Rharb Basin were also identified from the Pliocene in the Mediterranean. Nineteen species in the Rharb Basin are potential endemics, which is much higher than endemic species in the region today (25.6 % vs. 5 %), even taking into account uncertainties in the fossil data coverage. Tere is no comparable Early Pliocene otolith assemblage known from the Gulf of Guinea realms, the nearest being from the Middle Miocene of Gabon (Schwarzhans, 2013 c). However, it can be stated that the tropical West African influence in the Early Pliocene fish fauna of the Rharb Basin is significant at about 31.1 % (species also occurring today and those related to extant West African species). Tis compares to about 5 % of species off Morocco today shared with tropical West Africa. Tus, the assessment of the otolith-based fish fauna in the Rharb Basin does not justify combining it with the Early Pliocene Mediterranean nor with tropical West African fauna, the latter deduced from the extant faunal composition. I therefore postulate the presence of an Atlantic Moroccan bioprovince during the Early Pliocene, and potentially Late Miocene, which I propose to name the “ Maghrebian bioprovince. ”	en	Schwarzhans, Werner (2023): Geology and stratigraphy of the Neogene section along the Oued Beth between Dar bel Hamri and El Kansera (Rharb Basin, northwestern Morocco) and its otolith-based fish fauna: a faunal inventory for the Early. Swiss Journal of Palaeontology (4) 142 (1): 1-85, DOI: 10.1186/s13358-023-00268-4, URL: https://doi.org/10.1186/s13358-023-00268-4
