taxonID	type	description	language	source
492D87AAFFB3FFFC018CFB17639BFA7F.taxon	discussion	Discussion In this informal clade, we combine Bollmannia and the fossil otolith-based genus Brassoichthys n. gen. that share a specific otolith morphology characterized by an otolith shape that is longer than high, or as long as high, and that show a relatively flat inner face with a relatively large, somewhat deepened sulcus and a large subcaudal iugum.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB3FFFC018CFA576785F978.taxon	discussion	Discussion Fricke et al. (2023) list 11 valid extant Bollmannia species, four in the West Atlantic, seven in the East Pacific. Otoliths are available from all species except B. gomezi Acero, 1981, from the Pacific coast of southern Colombia. We distinguish four groups (otolith plexus) within the genus based on certain otolith patterns: Te Bollmannia boqueronensis otolith plexus containing the extant B. boqueronensis Evermann & Marsh, 1899 (Fig. 15 a), from the West Atlantic; and B. marginalis Ginsburg, 1939 (Fig. 15 k), and B. ocellata Gilbert, 1892 (Fig. 15 l – m), from the East Pacific. Te otoliths of this plexus are characterized by a relatively high predorsal angle and a compressed shape (OL: OH = 1.0 – 1.1).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB3FFFC018CFA576785F978.taxon	description	Te Bollmannia eigenmannorum otolith plexus containing the extant B. eigenmannorum (Garman, 1896) (Fig. 21 e) from the West Atlantic. Tese otoliths differ from all the others in the nearly rectangular outline and the rather short and narrow sulcus (OL: SuL> 1.9 vs <1.9 in other Bollmannia species). Tere is no comprehensive phylogenetic evaluation of Bollmannia species. Van Tassell et al. (2012), however, reviewed the Atlantic species of the genus. Tey found that B. eigenmannorum was the most distinct of the Atlantic species and commented that Ginsburg considered separating it into its own subgenus. In the review of Van Tassell et al. (2012), B. communis and B. litura were considered closely related (in agreement with comments they mentioned from Ginsburg). In fact, the otoliths of both species are so similar that they may not be distinguishable. Ginsburg (1939) considered B. marginalis from the Pacific to be most closely related to the Atlantic B. litura, while we have B. marginalis in the B. boqueronensis plexus. Ginsburg (1939) also compared B. chlamydes, B. macropoma, and B. umbrosa. Acero (1981) related B. macropoma, B. stigmatura, and B. gomezi. Te otolith plexus grouping is consistent with results from the review of Van Tassell et al. (2012) and Acero (1981) and also appears to be mostly consistent with comments by Ginsburg (1939) commented in Van Tassell et al. (2012).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB3FFFE0236F89663C1FADD.taxon	description	Figure 15 a – h Material (fossil specimens) 92 specimens: Zanclean (49 specimens): Atlantic Panama, Cayo Agua FM, 2 specimens PPP 1276; Venezuela, Cubagua FM, 2 specimens PPP 2554; 3 specimens PPP 2555; 1 specimen PPP 2556 (figured specimen NMB P 15474); 1 specimen PPP 2557; 21 specimens PPP 2568; 19 specimens PPP 2571 (figured specimen NMB P 15473). Piacenzian (3 specimens): Atlantic Panama, Escudo de Veraguas FM, 2 specimens PPP 2181; 1 specimen PPP 2182 (figured specimen NMB P 15472). Gelasian (40 specimens): Atlantic Panama, Escudo de Veraguas FM, 7 specimens PPP 2170 (figured specimens NMB P 15471); Bastimentos FM, Ground Creek Unit, 12 specimens PPP 1252; 1 specimen PPP 1304; Atlantic Costa Rica, Moin FM, 4 specimens PPP 1773; 1 specimen PPP 2038; 5 specimens PPP 3259. Calabrian: Atlantic Panama, Swan Cay FM, 4 specimens PPP 2221; Venezuela, Cumaná FM, 6 specimens PPP 3038. Description Otolith size up to 3 mm in length. OL: OH = 1.0 – 1.08; OH: OT = 2.9 – 3.3. Otolith shape nearly rectangular, predorsal angle high, broadly rounded, postdorsal angle higher than predorsal angle, postdorsal projection blunt, only slightly longer than postventral angle, not bent outward. Ventral rim straight, horizontal. Anterior and posterior rims nearly vertical; preventral angle blunt, nearly orthogonal; posterior rim with distinct indentation at its middle. All rims smooth, dorsal and anterior rims slightly undulating, weakly crenulated in small specimens (Fig. 15 e – h). Inner face flat in vertical direction and slightly bent in horizontal direction. Sulcus sole-shaped with low ostial lobe, deepened, inclined at 10 – 15 °. OL: SuL = 1.7 – 1.9. Subcaudal iugum large, broad. Dorsal depression wide, relatively deep and close to sulcus with distinct crista superior. Ventral furrow distinct, moderately close to ventral rim of otolith. Outer face distinctly convex, strongest inframedian, relatively smooth with few faint furrows on dorsal field.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB3FFFE0236F89663C1FADD.taxon	discussion	Discussion Bollmannia boqueronensis has been an inconspicuous and apparently widely distributed species in the tropical Atlantic since at least Early Pliocene. For comparison with coeval species, see below. Extant specimens of B. boqueronensis were collected over sandy and muddy bottoms and flat areas adjacent to coral reefs at 8.4 – 73 m throughout the Caribbean and the eastern Gulf of Mexico (Van Tassell et al., 2012).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB1FFFE018CFA3461CCFED8.taxon	description	Figure 15 i – j Material 13 specimens: Zanclean: Atlantic Panama, Cayo Agua FM, 7 specimens PPP 1276; upper Cayo Agua FM, 6 specimens PPP 1188 (figured specimens NMB P 15475). Discussion A few specimens from the Pliocene of the Atlantic side of Panama differ from the typical B. boqueronensis in the completely smooth outline and could possibly represent a different species. However, all specimens available are smaller than 1.5 mm in length, likely represent juveniles and thus do not qualify for a proper identification.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB1FFFE0236FE376669FAD1.taxon	description	Figure 15 l – p Material (fossil specimens) 9 specimens: Messinian (Bollmannia cf. ocellata): Pacific Panama, Chucunaque FM, 1 specimen PPP 1576, 1 specimen PPP 1578 (figured specimen NMB P 15477). Calabrian: Pacific Panama, Armueles FM, 6 specimens PPP 3235 (figured specimen NMB P 15478); 1 specimen PPP 3236 (figured specimen NMB P 15476). Discussion Otoliths of B. ocellata resemble those of B. boqueronensis described above but differ in a few subtle characters: a more strongly and regularly crenulated dorsal rim, the slightly projecting preventral angle and the caudal tip being slightly bent upward, a feature commonly observed in a number of Bollmannia species. Te otoliths of B. ocellata also resemble those of B. marginalis (Fig. 15 k), but differ in the more intense crenulation of the dorsal rim and the slightly upward bent caudal tip. Te two figured specimens from the Calabrian of Armuelles (Fig. 15 n – o) are well preserved but small (OL = 1.5 mm), probably from juveniles. Te single large specimen from the Messinian of Darien (Fig. 15 p) is rather poorly preserved and therefore only tentatively allocated. Bollmannia ocellata today is found from the Gulf of California to Ecuador at depths of 10 – 120 m on muddy bottoms (Froese & Pauly, 2023).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB1FFF80236FA2A60F2FED8.taxon	description	Figure 16 a – b 2010 Bollmannia nubila — Schwarzhans: pl. 101, Fig. 1. 2014 Gobiidae, genera and species indet. 1 — Aguilera et al.: Fig. 12 / 5 – 7. Material 9 specimens: holotype (refigured from Schwarzhans, 2010) from the Langhian of Dingden, northern Germany, North Sea Basin (SMF PO 83052). Aquitanian to Burdigalian: Brazil, Pirabas FM, 8 specimens (1 specimen refigured from Aguilera et al., 2014 as Gobiidae indet.) (MPEG- 1820 - V). Description Otolith size up to 2 mm in length. OL: OH = 1.05 – 1.1; OH: OT = 3.0 – 3.2. Otolith shape nearly rectangular, predorsal angle high, nearly as high as postdorsal angle, postdorsal projection blunt, not longer than postventral angle, not bent outward. Ventral rim straight, horizontal, smooth. Anterior and posterior rims nearly vertical; preventral angle blunt, nearly orthogonal; posterior rim with broad, shallow indentation at its middle. All rims smooth, dorsal rim occasionally crenulated. Inner face nearly flat. Sulcus sole-shaped but very narrow with low ostial lobe, slightly deepened, inclined at 10 – 15 °. OL: SuL = 1.65 – 1.85. Subcaudal iugum long, relatively narrow. Dorsal depression wide, indistinct. Ventral furrow distinct, close to ventral rim of otolith. Outer face distinctly convex, strongest inframedian, relatively smooth with few short furrows on dorsal field.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB1FFF80236FA2A60F2FED8.taxon	discussion	Discussion Bollmannia nubila is the earliest record of the genus and now known from two rather extreme geographic positions: the unique type from the North Sea Basin and the now-assigned referred specimens from Brazil. Bollmannia nubila is readily distinguished from its congeners by the very narrow sulcus, the relatively narrow subcaudal iugum, and the high predorsal angle. It resembles B. eigenmannorum. It is therefore possible that B. nubila is positioned relatively closely to the origin of the genus and thus combines characteristics of the B. boqueronensis plexus and the B. eigenmannorum plexus.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB7FFF8018CFE3061E5FCD8.taxon	description	Figure 16 c – g Holotype NMB P 15479 (Fig. 16 c), Tortonian (NN 10), Yaviza FM, Panama, Darien, Río Tuquesa, PPP 1609. Paratypes 6 specimens: Tortonian: Tuira FM, 3 specimens PPP 1137 (NMB P 15480 - 82); Yaviza FM, 1 specimen PPP 1566 (NMB P 15483). Messinian: Trinidad, Manzanilla FM, 2 specimens PPP 2666 (NMB P 15484 - 85). Referred material 5 specimens: Tortonian: Panama, 2 specimens same data as holotype; Yaviza FM, 1 specimen PPP 1566. Messinian: Trinidad, Manzanilla FM, 2 specimens PPP 2666. Etymology Superlative of ornatus (Latin) = ornamented, referring to the strongly crenulated dorsal rim. Diagnosis OL: OH = 1.0 – 1.1. Maximal size 2.35 mm (holotype). Predorsal rim high; postdorsal region distinctly higher. Dorsal rim strongly crenulated, with radial furrows extending on inner and outer faces. Sulcus narrow. Subcaudal iugum long, narrow. Description Otolith size up to 2.35 mm in length (holotype). OH: OT = 3.0 – 3.2. Otolith shape rectangular with elevated postdorsal region, predorsal angle high, rounded, distinctly lower than postdorsal angle, postdorsal projection blunt, slightly longer than postventral angle, not bent outward. Ventral rim straight, horizontal, smooth or slightly undulating. Anterior and posterior rims nearly vertical; preventral angle blunt; posterior rim with distinct, mostly sharp indentation at its middle. All rims slightly undulating to strongly crenulated, particularly dorsal rim. Inner face nearly flat in vertical direction, slightly bent in horizontal direction. Sulcus sole-shaped, narrow with low ostial lobe, slightly deepened, inclined at 10 – 15 °. OL: SuL = 1.8 – 1.9 (rarely 2.1, Fig. 16 f). Subcaudal iugum long, relatively narrow. Dorsal depression wide, relatively short and moderately indistinct, with some radial furrows ingressing from the marginal crenulation, close to central portion of sulcus. Ventral furrow distinct, relatively close to ventral rim of otolith. Outer face distinctly convex, strongest inframedian, with several strong radial furrows related to marginal crenulation.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB7FFF8018CFE3061E5FCD8.taxon	discussion	Discussion Bollmannia ornatissima is a relatively uncommon species in the Late Miocene of Pacific Panama (Darien) but has also been found in time-equivalent strata of Trinidad. Te specimens from Trinidad (Fig. 16 f – g) differ from the Panamanian ones in the relatively short sulcus (OL: SuL = 1.9 – 2.1 vs 1.8 – 1.9), but we do not consider this difference as warranting separation on species level, particularly since the few Trinidadian specimens are also relatively small (OL = 1.3 mm and 1.45 mm in length). Bollmannia ornatissima is best recognized by its intense marginal crenulation, particularly the dorsal rim, and the relatively narrow subcaudal iugum. It resembles B. nubila in the latter trait and the narrow sulcus but shows a much more pronounced postdorsal region. Bollmannia ornatissima differs from other species in the plexus such as B. marginalis and B. ocellata in the stronger crenulation of the dorsal rim, the narrower sulcus, and the narrower subcaudal iugum.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB7FFF90236FC306083FB98.taxon	description	Figure 16 h – o Holotype NMB P 681 (Fig. 16 h), Burdigalian (N 7, NN 4), Brasso FM, Trinidad, Tamanaquito River, KR 6666. Paratypes 7 specimens: Burdigalian (N 7): Trinidad, Brasso FM, 4 specimens same data as holotype (NMB P 15486 - 89). Burdigalian (N 8): Venezuela, Cantaure FM, 1 specimen PPP 2546 (NMB P 15492). Langhian: Trinidad, upper Brasso FM (N 10 - 12), 2 specimens KR 9244 (NMB P 15490 - 91). Referred material 390 specimens: Burdigalian: Trinidad, Brasso FM, 82 specimens same data as holotype; Venezuela, Cantaure FM, 16 specimens PPP 2543; 43 specimens PPP 2545; 235 specimens PPP 2546; 3 specimens PPP 2539. Langhian: Trinidad, upper Brasso FM, 11 specimens KR 9244. Etymology After Trinidad, the country of origin of the holotype. Diagnosis OL: OH = 1.0 – 1.05. Maximal size 2.2 mm. Predorsal rim high; postdorsal region somewhat higher. Dorsal rim smooth to slightly undulating or crenulated. Sulcus wide, relatively deep. Subcaudal iugum long, narrow. Description Otolith size up to 2.2 mm in length (holotype 2.1 mm). OH: OT = 3.0 – 3.2. Otolith shape nearly rectangular with moderately elevated postdorsal region, predorsal angle high, rounded, slightly lower than postdorsal angle, postdorsal projection blunt, not or slightly longer than postventral angle, not bent outward. Ventral rim straight, horizontal, smooth. Anterior and posterior rims nearly vertical; preventral angle blunt; posterior rim with distinct, broad indentation at its middle. All rims smooth, dorsal rim undulating to mildly crenulated. Inner face nearly flat in vertical direction, slightly bent in horizontal direction. Sulcus sole-shaped, broad but with low ostial lobe, distinctly deepened, inclined at 12 – 17 °. OL: SuL = 1.75 – 1.85. Subcaudal iugum long, relatively narrow. Dorsal depression wide, relatively long, distinct, moderately close to sulcus with broad crista superior. Ventral furrow distinct, moderately close to ventral rim of otolith. Outer face distinctly convex, strongest inframedian, more or less smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB7FFF90236FC306083FB98.taxon	discussion	Discussion Bollmannia trinidadensis is best recognized by its wide and rather deep sulcus in combination with a relatively narrow subcaudal iugum and a very weak marginal ornamentation. It is a common species in the late Burdigalian of Trinidad (Fig. 16 h – m, o) and the Cantaure Formation (Langhian) of Venezuela (Fig. 16 o).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB6FFF9018CFB30670BFCB8.taxon	description	Figure 17 a – d Material (fossil specimens) 11 specimens: Calabrian: Pacific Panama, Armuelles FM, 2 specimens PPP 3235 (figured specimens NMB P 15493 – 94), 9 specimens PPP 3236. Description Otolith size up to 4.2 mm in length. OL: OH = 1.15 – 1.4, increasing with size; OH: OT = 2.9 – 3.5, decreasing with size. Predorsal angle low, often depressed, postdorsal angle expanded, postdorsal projection blunt, only slightly longer than postventral angle, not or very slightly bent outward. Ventral rim straight, horizontal. Anterior rim inclined, with projecting preventral angle; posterior rim nearly vertical, with broad, wide indentation at its middle. All rims smooth, except dorsal rim undulating and increasingly intense crenulated in large specimens (Fig. 17 b). Inner face flat in vertical direction and slightly bent in horizontal direction. Sulcus sole-shaped with low ostial lobe, moderately deepened, inclined at 15 – 18 °. Caudal tip rounded, slightly bent upward. OL: SuL = 1.65 – 1.95. Subcaudal iugum long, relatively narrow. Dorsal depression wide, relatively deep and close to sulcus with distinct crista superior. Ventral furrow distinct, relatively distant from ventral rim of otolith. Outer face moderately to distinctly convex, decreasingly convex with size, strongest inframedian, relatively smooth with few furrows on dorsal field. Discussion Only relatively small specimens are found in the Calabrian of Pacific Panama, which are here attributed to B. chlamydes (Fig. 17 c – d). Tey differ from the large extant ones (Fig. 17 a – b) in the lower ratio OL: OH (1.15 – 1.2 vs 1.25 – 1.4) and in being thicker (OH: OT <3.1 vs> 3.2). We contribute these differences to allometric growth. Today, B. chlamydes is reported from Pacific Colombia to northern Peru over muddy bottoms at depths of 10 – 120 m (Froese & Pauly, 2023).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB6FFFB0236FC90607FFE38.taxon	description	Figure 17 e – j 1992 Gobiidae sp. 2 — Nolf & Stringer: pl. 17, Fig. 13. Material (fossil specimens) 48 specimens: Messinian: 1 specimen Dominican Republic, Gurabo FM, NMB 15864 (NMB P 407); Zanclean, Atlantic Panama: 35 specimens; 2 specimens Shark Hole FM, PPP 2217; 25 specimens Cayo Agua FM, 24 specimens PPP 1276 (NMB P 15496), 1 specimen PPP 2222; 8 specimens Cubagua FM, 5 specimens PPP 2570, 3 specimens PPP 2571 (NMB P 15498); 1 specimen Piacenzian, Atlantic Costa Rica, Rio Banano FM, PPP 3245 (NMB P 15495); 11 specimens Gelasian: 7 specimens Atlantic Panama, Bastimentos FM, Ground Creek Unit, 5 specimens PPP 1256, 2 specimens PPP 3204; 4 specimens, Atlantic Costa Rica, Moin FM, 1 specimen PPP 1776 (NMB P 15497), 2 specimens PPP 2032, 1 specimen PPP 2038. Discussion Otoliths of B. communis are similar to those of B. chlamydes (see above) from the East Pacific and those of the West Atlantic B. litura (Fig. 17 k – l). Tey differ from B. chlamydes in the less projecting preventral angle, the higher predorsal angle, the often slightly projecting postdorsal angle and in the generally less strongly developed ornamentation of the dorsal rim. Furthermore, otoliths of B. communis remain thicker at large sizes compared to B. chlamydes (OH: OT = 2.75 – 3.0 vs 3.2 – 3.5 at sizes> 2.8 mm in length). We were unable to identify a reliable morphological difference between the otoliths of B. communis and B. litura. Extant specimens of B. communis were collected on mud bottoms at depths of 18 – 77 m (Van Tassell et al., 2012). Tey were primarily taken from the Gulf of Mexico and off Puerto Rico and French Guiana (Van Tassell et al., 2012). Bollmannia litura was taken at depths of 12.8 – 71 m over mud and mixed-mud bottoms in the western Caribbean (Van Tassell et al., 2012).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB4FFFB018CFE1061F1FE58.taxon	description	Figure 17 m – q Holotype NMB P 15499 (Fig. 17 m), Zanclean, Cubagua FM above Cerro Negro Member, Venezuela, Isla Cubagua, PPP 2568. Paratypes 11 specimens: Zanclean, Cubagua FM above Cerro Negro Member, Venezuela, Isla Cubagua, 2 specimens PPP 2568 (NMB P 15500 - 01), 6 specimens PPP 2571 (NMB P 15502 - 04), 3 specimens PPP 2572 (NMB P 15508 - 10). Etymology After the Cubagua Formation, the type formation. Diagnosis OL: OH = 1.1 – 1.15. Maximal size 3.9 mm (holotype). Predorsal rim slanted; postdorsal region slightly elevated. Dorsal rim irregularly undulating or crenulated. Preventral angle slightly projecting. Sulcus wide; OL: SuL = 1.85 – 1.95. Subcaudal iugum long, moderately wide. Description Otolith size up to 4.0 mm in length (holotype 3.9 mm). OH: OT = 3.1 – 3.3. Predorsal angle low, often depressed, postdorsal angle expanded, postdorsal projection blunt, only slightly longer than postventral angle, not bent outward. Ventral rim straight, horizontal. Anterior rim inclined, with slightly projecting preventral angle; posterior rim nearly vertical, with broad, wide and rather deep indentation at its middle. All rims smooth, except dorsal rim irregularly undulating or crenulated. Inner face almost entirely flat. Sulcus sole-shaped with low ostial lobe, moderately deepened, inclined at 12 – 15 °. Caudal tip rounded, slightly bent upward. OL: SuL = 1.85 – 1.95. Subcaudal iugum long, moderately wide. Dorsal depression wide, indistinct and close to sulcus with distinct crista superior. Ventral furrow distinct, moderately distant from ventral rim of otolith. Outer face moderately to distinctly convex, strongest inframedian, relatively smooth with few furrows on dorsal field.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB4FFFB018CFE1061F1FE58.taxon	discussion	Discussion Bollmannia cubaguana is very similar to B. chlamydes differing only in few subtle characters, primarily in being less elongate in specimens larger than 3 mm in length (OL: OH 1.1 – 1.15 vs 1.25 – 1.4) and the less strongly inclined sulcus (12 – 15 ° vs 15 – 18 °). Presumably, B. cubaguana represents an extinct geminate species in the West Atlantic to the extant East Pacific B. chlamydes. Te largest specimen (holotype) of B. cubaguana shows a slightly middorsal concavity, similar to but much less strongly developed as in otoliths of the B. stigmatura plexus (see below).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB4FFC50236FDB062E1FEF8.taxon	description	Figure 18 a – n 1908 Otolithus (Gobius) gatunensis — Schubert: Fig. 7.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB4FFC50236FDB062E1FEF8.taxon	materials_examined	Material 188 specimens, Tortonian: 2 specimens Trinidad, Tamana FM, KR 25956 (NMB P 541 and P 511); 15 specimens Ecuador, Angostura FM, 3 specimens PPP 3290, 2 specimens PPP 3297 (NMB P 15511), 5 specimens PPP 3301 (NMB P 15512), 2 specimens PPP 3369, 1 specimen PPP 3372, 2 specimens PPP 3465; 54 specimens Pacific Panama, Darien, Tuira FM, 7 specimens PPP 1132 (figured specimen NMB P 15522), 14 specimens PPP 1137, 1 specimen PPP 1139 (NMB P 15520), 4 specimens, PPP 1593, 2 specimens 1142, 4 specimens PPP 1627, 21 specimens PPP 1163, 1 specimen PPP 1164; 57 specimens Pacific Panama, Darien, Yaviza FM, 5 specimens PPP 1607, 5 specimens PPP 1609, 3 specimens PPP 1528, 3 specimens PPP 1145, 12 specimens PPP 1620, 6 specimens PPP 1149 (figured specimen NMB P 15521), 9 specimens PPP 1171, 1 specimen PPP 1565, 13 specimens PPP 1566 (figured specimen NMB P 15519); 47 specimens Atlantic Panama, Gatun FM, 1 specimen PPP 1973, 5 specimens PPP 2165 (NMB P 15514), 1 specimens PPP 2166 (NMB P 15513), 28 specimens PPP 2167 (figured specimens NMB P 15515 - 16), 12 specimens PPP 2168 (figured specimen NMB P 15517). 13 specimens Messinian: Chucunaque FM, Pacific Panama, Darien, PPP 1612 (figured specimen NMB P 15518), Description Otolith size up to 3.1 mm in length. OL: OH = 1.02 – 1.12; OH: OT = 3.1 – 3.4. Predorsal angle relatively high, postdorsal angle expanded, postdorsal projection rounded, projecting beyond postventral angle, slightly bent outward. Ventral rim straight, horizontal. Anterior rim nearly vertical, with nearly orthogonal preventral angle; posterior rim slightly inclined, with broad and rather deep indentation at its middle. Ventral rim smooth, dorsal rim irregularly undulating or crenulated. Inner face flat in vertical direction and slightly bent in horizontal direction. Sulcus sole-shaped with moderate ostial lobe, moderately deepened, inclined at 10 – 15 °. Caudal tip rounded, slightly bent upward. OL: SuL = 1.65 – 1.85. Subcaudal iugum long, moderately wide. Dorsal depression wide, distinct and close to sulcus with distinct crista superior. Ventral furrow distinct, relatively distant from ventral rim of otolith. Outer face moderately to distinctly convex, strongest inframedian, with radial furrows on dorsal field.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFB4FFC50236FDB062E1FEF8.taxon	discussion	Discussion Bollmannia gatunensis is a common species during the Tortonian from Ecuador in the East Pacific to Trinidad in the West Atlantic. Bollmannia gatunensis is very similar to B. communis but tends to be thinner (OH: OT = 3.1 – 3.4 vs 2.75 – 3.0) and shows a more strongly ornamented dorsal rim and more distinct postdorsal projection. No doubt, both species are closely related.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF8AFFC60236FF506782FE78.taxon	description	Figure 19 a – e Holotype NMB P 15523 (Fig. 19 a), Messinian, Onzole FM, Ecuador, PPP 3314. Paratypes 5 specimens: 1 specimen Tortonian, Pacific Panama, Darien, Yaviza FM PPP 1620 (NMB P 15527); 2 specimens Tortonian, Ecuador, Angostura FM, 1 specimen PPP 3480 (NMB P 15526), 1 specimen PPP 3465 (NMB P 15528; 2 specimens same data as holotype (NMB P 15524 - 25). Referred specimens 18 specimens: 3 specimens Tortonian, Pacific Panama, Yaviza FM, 1 specimen PPP 1620, 2 specimens PPP 1566; 3 specimens Tortonian, Ecuador, Angostura FM, 1 specimen PPP 3439, 2 specimens PPP 3465; 3 specimens Messinian, Ecuador, Onzole FM PPP 3314; 9 specimens Messinian, Pacific Panama, Darien, Chucunaque FM, 6 specimens PPP 1576, 3 specimens PPP 1578. Etymology From propensus (Latin) = inclined, skewed, referring to the parallelogram-like shape of the otolith. Diagnosis OL: OH = 1.2 – 1.3. Maximal size 4.6 mm (holotype). Predorsal rim rounded or depressed; postdorsal region slightly elevated. Preventral projection and postdorsal projection strong giving otolith a parallelogram-like shape. Dorsal rim irregularly undulating or slightly crenulated. Sulcus large; OL: SuL = 1.65 – 1.85. Subcaudal iugum long, wide. Description Otolith size up to 4.6 mm in length (holotype). OH: OT = 3.1 – 3.4 (one specimen of Fig. 19 d OH: OT = 2.7). Predorsal angle low, slanted or depressed, postdorsal angle slightly expanded. Postdorsal projection projecting well beyond postventral angle, slightly bent outward; preventral angle distinctly projecting; both resulting in parallelogram-like otolith shape. Ventral rim straight, horizontal. Anterior rim inclined, posterior rim inclined, with variable, broad and shallow or no indentation at its middle. Ventral rim smooth, dorsal rim irregularly undulating or slightly crenulated. Inner face flat in vertical direction and distinctly bent in horizontal direction. Sulcus sole-shaped with moderate ostial lobe and narrow cauda, moderately deepened, inclined at 12 – 15 °. Caudal tip rounded, slightly bent upward. OL: SuL = 1.65 – 1.85. Subcaudal iugum long, wide. Dorsal depression wide, rather indistinct and close to sulcus with moderately developed crista superior. Ventral furrow distinct, relatively distant from ventral rim of otolith. Outer face nearly flat with moderate subcentral umbo (only one specimen with strongly convex outer face — Fig. 19 d), few radial furrows on dorsal field close to otolith margin.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF8AFFC60236FF506782FE78.taxon	discussion	Discussion Bollmannia propensa is easily recognized by its parallelogram-like shape, thin appearance and distinctly bent inner face in horizontal direction. It appears to be restricted to the late Miocene of the East Pacific. It resembles most the extant B. umbrosa (Fig. 19 f) but differs in the more slender shape (OL: OH = 1.2 – 1.3 vs 1.1 – 1.15) and the distinctly expanded postdorsal projection.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF89FFC70236FE506019FD58.taxon	description	Figure 19 g – i 1976 Gobiidae — Nolf: pl. 9, Fig. 6. Holotype NMB P 15529 (Fig. 19 g), Tortonian, Ojo de Agua FM, Venezuela, NMB 15730. Paratypes 4 specimens: 2 specimens Langhian, Trinidad, Brasso FM, 1 specimen KR 7546 (NMB P 540), 1 specimen KR 7573 (NMB P 692); 2 specimens same data as holotype (NMB P 15530 - 31). Referred specimens 5 specimens same data as holotype. Etymology After Venezuela, the country of origin of the holotype. Diagnosis OL: OH = 1.07 – 1.12. Maximal size 3.8 mm (holotype). Predorsal angle high, rounded; postdorsal region elevated, distinctly and irregularly ornamented. Preventral angle slightly projecting; postdorsal projection slightly extending beyond postventral angle. Sulcus wide; OL: SuL = 1.7 – 1.8. Subcaudal iugum long, wide. Outer face with narrow but strongly projecting umbo close to ventral rim of otolith. Description Otolith size up to 3.8 mm in length (holotype). OH: OT = 2.6 – 3.0. Predorsal angle low, rounded, postdorsal angle expanded. Postdorsal projection projecting slightly beyond postventral angle, not bent outward; preventral angle slightly projecting. Ventral rim straight, horizontal. Anterior and posterior rims slightly inclined, posterior rim with relatively deep indentation at its middle. Ventral rim smooth, dorsal rim broadly and irregularly crenulated. Inner face nearly flat. Sulcus sole-shaped with moderate ostial lobe, moderately deepened, inclined at 14 – 20 °. Caudal tip rounded, slightly bent upward. OL: SuL = 1.7 – 1.8. Subcaudal iugum moderately long and wide. Dorsal depression wide, rather indistinct and close to sulcus with moderately developed crista superior. Ventral furrow distinct, relatively distant from ventral rim of otolith. Outer face with narrow, strongly projecting, subcentral umbo close to ventral rim of otolith, few radial furrows on dorsal field close to otolith margin.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF89FFC70236FE506019FD58.taxon	discussion	Discussion Bollmannia venezuelana is relatively inconspicuous in appearance and differs from fossil and extant congeners only in subtle characteristics. It is distinctly more compressed than B. propensa with less projecting preventral and postdorsal projections. Bollmannia venezuelana most closely resembles the extant B. umbrosa from the East Pacific but differs in the higher predorsal angle (vs depressed) and the distinct subcentral umbo on the outer face, a trait that is not seen in a similar expression in any of the other known Bollmannia species. Bollmannia venezuelana differs from B. gatunensis in the less prominent postdorsal projection and the more strongly inclined sulcus (14 – 20 ° vs 10 – 15 °).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF88FFC7018CFCB060E3FA98.taxon	description	Figure 18 o Material 2 specimens, Dominican Republic, Tortonian, Cercado FM, 1 specimen NMB 15912 (NMB P 396), 1 specimen NMB 16837 (NMB P 410). Discussion Two moderately large specimens (the figured one is 2.65 mm in length) differ from the otherwise similar B. gatunensis in the robust appearance and completely smooth rims without crenulation. It is possible that these specimens still fall within the variability of B. gatunensis, but because they too are found only in the north Dominican Basin it is also possible that they represent yet another species. More specimens from that region must be awaited before a reliable identification can be made.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF88FFC1018CFA30636DFA38.taxon	description	Figure 20 a – m Holotype NMB P 15537 (Fig. 20 g), Tortonian, Yaviza FM, Pacific Panama, Darien, PPP 1609. Paratypes 12 specimens: 4 specimens late Burdigalian, Venezuela, Cantaure FM, 1 specimen PPP 2545 (NMB P 15533), 2 specimens PPP 2546 (NMB P 15534 - 35), 1 specimen PPP 2539 (NMB P 15536); 2 specimens Langhian, Trinidad, Brasso FM, Cb. 1852 (NMB P 676 and NMB P 15532); 3 specimens same data as holotype, 1 specimen PPP 1607 (NMB P 15538), 2 specimens PPP 1620 (NMB P 15539 - 40); 1 specimen Tortonian, Trinidad, Manzanilla FM, PPP 2666 (NMB P 15543); 2 specimens Messinian, Venezuela, Isla Margarita, La Tejita FM, PPP 3087 (NMB P 15541 - 42). Referred specimens 88 specimens: 24 specimens late Burdigalian, Venezuela, Cantaure FM, 6 specimens PPP 2545, 18 specimens PPP 2546; 2 specimens Langhian, Trinidad, Brasso FM, Cb. 1852; 12 specimens Langhian, Venezuela, Socorro FM, 4 specimens PPP 2642, 8 specimens PPP 2644; 2 specimens Langhian, Pacific Panama, Darien, Tapaliza FM, 1 specimen PPP 1152, 1 specimen PPP 1538; 4 specimens Tortonian, Pacific Panama, Darien, Tuira FM, 1 specimen PPP 1134, 1 specimen PPP 1139, 2 specimens PPP 1627; 36 specimens Tortonian, Pacific Panama, Darien, Yaviza FM, 8 specimens PPP 1607, 1 specimen PPP 1528, 2 specimens PPP 1145, 11 specimens PPP 1620, 3 specimens PPP 1149, 8 specimens PPP 1171, 1 specimen PPP 1565, 2 specimens PPP 1566; 5 specimens Tortonian, Trinidad, Manzanilla FM, PPP 2666; 3 specimens Messinian, Venezuela, Isla Margarita, La Tejita FM, PPP 3087. Etymology Named in honor of Carole Baldwin (USNM, Washington D. C.) in recognition of her many contributions to the understanding of gobies from America. Diagnosis OL: OH = 1.05 – 1.12. Maximal size 2.6 mm. Dorsal rim with distinctive middorsal indentation. Predorsal angle high, postdorsal region elevated, preventral angle orthogonal; postdorsal projection slightly extending beyond postventral angle. Sulcus wide, deep; OL: SuL = 1.6 – 1.8. Subcaudal iugum short, wide. Sulcus inclination 8 – 16 °. Description Otolith size up to 2.6 mm in length (holotype 2.4 mm). OH: OT = 3.0 – 3.3. Dorsal rim with distinct, (See figure on next page.) Fig. 20 a – m Bollmannia baldwinae n. sp.; g holotype, NMB P 15537, Yaviza FM, Darien, Panama, PPP 1609; b – m paratypes, a – b Brasso FM, Langhian, Trinidad, Cb. 1852, a NMB P 676, b NMB P 15532; c Cantaure FM, El Hatillo- 1, Venezuela, PPP 2545, NMB P 15533; d, f Cantaure FM, Casa Cantaure, Venezuela, PPP 2546, NMB P 15534 - 35; e Cantaure FM, San José de Cocodite, Venezuela, PPP 2539, NMB P 15536; h – j Yaviza FM, Darien, Panama, h PPP 1607, NMB P 15538, i – j PPP 1620, NMB P 15539 - 40; k – l Tejita FM, Isla Margarita, Venezuela, PPP 3087, NMB P 15541 - 42; m Manzanilla FM, Trinidad, PPP 2666, NMB P 15543. n Bollmannia sp. 2, Bastimentos FM, Ground Creek Unit, Bastimentos, PPP 1256, NMB P 15544. o Bollmannia macropoma Gilbert, 1892, extant, CAS 44137, Cost Rica, Golfo de Nicoya, SL 86 mm. p Bollmannia stigmatura Gilbert, 1892, extant, USNM 116364, Pacific Columbia, Puerto Utria. q Bollmannia sp. 3, Socorro FM, Paraguaná, Venezuela, PPP 2642, NMB P 15545 somewhat variable but mostly sharp and deep middorsal indentation in front of postdorsal expansion. Predorsal angle high, rounded, postdorsal region expanded, often with steep anterior border. Postdorsal projection projecting slightly beyond postventral angle, not bent outward; preventral angle orthogonal. Ventral rim straight to slightly curved, horizontal. Anterior rim nearly vertical, posterior rim slightly inclined, with relatively broad indentation at its middle. Ventral rim smooth, dorsal rim smooth or irregularly undulating. Inner face nearly flat. Sulcus sole-shaped with moderate ostial lobe, deep, inclined at 8 – 16 °. Caudal tip rounded. OL: SuL = 1.6 – 1.8. Subcaudal iugum short, wide to almost as wide as long in cases. Dorsal depression wide, deep and mostly distinct, open to dorsal rim of otolith and close to sulcus with well-developed crista superior. Ventral furrow distinct, relatively close to ventral rim of otolith. Outer face broadly convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF88FFC1018CFA30636DFA38.taxon	discussion	Discussion Bollmannia baldwinae is readily distinguished from its fossil congeners by the distinct middorsal indentation. Te middorsal indentation varies somewhat in expression and strength from broadly concave to nearly orthogonally incised and usually increases in strength with size. Bollmannia baldwinae is distinguished from the extant congeners of the plexus B. macropoma (Fig. 20 o) and B. stigmatura (Fig. 20 p) in being more compressed (OL: OH = 1.05 – 1.12 vs 1.15 – 1.2) and showing a broader, more pronounced postdorsal region. Te short and wide subcaudal iugum is also characteristic of B. baldwinae. Bollmannia baldwinae is a widely distributed species in the East Pacific and West Atlantic with a long duration range from the late Burdigalian to the Messinian over about 11 to 12 mya. Te species is regarded as the earliest record in the Bollmannia stigmatura otolith plexus because of its conspicuous middorsal indentation.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF8EFFC1018CFA10619AFE98.taxon	description	Figure 20 n Material 1 specimen Gelasian, Atlantic Panama, Bastimentos FM, Ground Creek Unit, PPP 1256 (NMB P 15544). Discussion A single, rather poorly preserved otolith from the early Pleistocene of Bastimentos Island, Atlantic Panama, of 2.8 mm in length shows a mild middorsal indentation. Tis could either represent a somewhat unusual variation of a known extant Atlantic species such as B. communis or indicate that species of the Bollmannia stigmatura otolith plexus which nowadays is restricted to the East Pacific also existed in the West Atlantic until the Gelasian.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF8EFFC10236FEF06172FD58.taxon	description	Figure 20 q Material 3 specimens: 1 specimen Langhian, Venezuela, Socorro FM, PPP 2642 (NMB P 15545); 2 specimens Messinian, Pacific Panama, Darien, Chucunaque FM, 1 specimen PPP 1576, 1 specimen PPP 1578. Discussion Tree poorly preserved, slender otoliths with a broad middorsal concavity could possible represent another species of the Bollmannia stigmatura otolith plexus.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF8EFFC30236FCF0635FFCB8.taxon	description	Figure 21 a – d Holotype NMB P 15529 (Fig. 21 a), Tortonian, Angostura FM, Ecuador, PPP 3465. Paratypes 3 specimens: 2 specimens Tortonian, Atlantic Panama, Gatun FM, PPP 2167 (NMB P 15548 - 49); 1 specimen Messinian, Onzole FM, Ecuador, NMB 19084 (NMB P 15547). Etymology After the type formation, the Angostura FM of Tortonian age in Ecuador. Diagnosis OL: OH = 1.05 – 1.15. Maximal size 2.25 mm (holotype). Otolith shape nearly rectangular with rounded dorsal rim. Predorsal angle high, postdorsal region not significantly elevated, preventral angle not projecting, postdorsal projection not extending beyond postventral angle. Sulcus narrow, small, with very low ostial lobe; OL: SuL = 1.9 – 2.2. Subcaudal iugum long, relatively narrow. Sulcus inclination angle 7 – 12 °. Description Otolith size up to 2.25 mm in length (holotype). OH: OT = 2.7 – 2.9. Dorsal rim relatively low, regularly curved, with high, rounded predorsal angle and rounded postdorsal angle. Postdorsal projection blunt, not projecting beyond postventral angle, not bent outward; preventral angle orthogonal or rounded. Ventral rim straight, horizontal. Anterior and posterior rims nearly vertical, posterior rim with relatively weak indentation at its middle. All rims smooth, dorsal rim slightly undulating. Inner face nearly flat. Sulcus sole-shaped, narrow, small, with low ostial lobe, moderately deep, inclined at 7 – 12 °. Caudal tip rounded. OL: SuL = 1.9 – 2.2. Subcaudal iugum long, narrow, often relatively weak. Dorsal depression wide, deep and distinct, open to dorsal rim of otolith and moderately close to sulcus with well-developed crista superior. Ventral furrow distinct, gently curved and clipping preventral and postventral tips of otolith. Outer face broadly convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF8EFFC30236FCF0635FFCB8.taxon	discussion	Discussion Bollmannia angosturae is similar to the extant B. eigenmannorum (Fig. 21 e) sharing the small and narrow and only slightly inclined sulcus and the weak preventral and postdorsal projections. It differs from the extant West Atlantic species in being more elongate (OL: OH = 1.05 – 1.15 vs 1.0 – 1.05) and the relatively weak subcaudal iugum.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF8CFFC3018CFAD06759FA98.taxon	description	Figure 21 j – m Holotype NMB P 15551 (Fig. 21 j), late Burdigalian, Cantaure FM, Venezuela, PPP 2545. Paratypes 3 specimens: 2 specimens same data as holotype (NMB P 15552 - 53); 1 specimen Tortonian, Tuira FM, Pacific Panama, Darien, PPP 1593 (NMB P 15554). Referred specimens 1 specimen late Burdigalian, Cantaure FM, Venezuela, PPP 2546. Etymology After the Paraguaná Peninsula, Venezuela, where the type location is situated. Diagnosis OL: OH = 1.02 – 1.15. Maximal size 1.55 mm (holotype). Otolith shape nearly rectangular with rounded predorsal, preventral, postdorsal and postventral angles. Otolith thin (OH: OT = 3.3 – 3.5). Sulcus long, with wide ostium and narrow cauda; OL: SuL = 1.55 – 1.65. Subcaudal iugum long, relatively narrow. Sulcus inclination angle 15 – 22 °. Description Otolith size up to 1.55 mm in length (holotype). Dorsal rim relatively low, nearly straight, slightly ascending towards posterior. Postdorsal projection blunt, not projecting beyond postventral angle, not bent outward; preventral angle orthogonal. Ventral rim straight, horizontal. Anterior and posterior rims nearly vertical, posterior rim with weak indentation at its middle. All rims smooth. Inner face nearly flat. Sulcus sole-shaped, deep, inclined at 15 – 22 °; ostium wide, with moderate ostial lobe, cauda narrow, straight. OL: SuL = 1.55 – 1.65. Subcaudal iugum long, narrow, often relatively weak. Dorsal depression indistinct, with weak crista superior. Ventral furrow distinct, gently curved and clipping preventral and postventral tips of otolith. Outer face nearly flat, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF8CFFC3018CFAD06759FA98.taxon	discussion	Discussion Bollmannia? paraguanaensis is only tentatively placed in the genus Bollmannia, primarily because of the long subcaudal iugum, which, however, is often rather poorly developed / visible. It differs from typical Bollmannia otoliths in the thin appearance caused by the nearly flat outer face and the relatively long sulcus. Te single late Miocene specimen from Darien is more compressed than the ones from the early Miocene Cantaure Formation (OL: OH = 1.02 vs 1.07 – 1.15) indicating that potentially two separate species could be involved.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF8CFFC3018CFC906010FAF8.taxon	description	Figure 21 f – i Material 6 specimens Tortonian, Atlantic Costa Rica, Uscari FM, PPP 3270 (NMB P 15550). Discussion Six small specimens (<1.4 mm in length) from the Uscari Formation of Costa Rica resemble the single known otolith of the extant B. eigenmannorum (Fig. 21 e). However, the extant specimen is affected by formalin, and does not reveal all pertinent characteristics for comparison with the fossil otoliths. Te allocation is therefore provisional. Extant specimens of B. eigenmannorum have been taken over muddy bottoms at depths of 36.5 – 170 m from the Florida Keys, throughout the Gulf of Mexico to Venezuela (Van Tassell et al., 2012).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF8CFFCC0236FAF06011FD98.taxon	materials_examined	Type species Brassoichthys tornabenei n. sp. Etymology Named after the type formation, the Brasso Formation of Trinidad. Diagnosis A fossil otolith-based genus of the Gobiosomatini, Microgobius Group, Bollmannia clade with the following combination of characters. Otoliths nearly quadratic in outline with rounded dorsal rim and slightly expanded postdorsal projection; OL: OH = 0.98 – 1.03. Inner face flat; outer face mildly convex, smooth. Sulcus relatively large and wide but with low ostial lobe; OL: SuL = 1.7 – 1.85; sulcus inclination angle 7 – 10 °. Subcaudal iugum moderately large. Dorsal depression deep; ventral furrow distinct, gently curving.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF8CFFCC0236FAF06011FD98.taxon	discussion	Discussion Brassoichthys is an easily recognizable otolith pattern that occupies a morphological position between the Bollmannia type and the Microgobius type pattern. Te nearly equal OL: OH ratio of around 1.0 matches this intermediate position, with Bollmannia otoliths being more slender and those of Microgobius more compressed and high-bodied. Te sulcus shape and expression of the subcaudal iugum matches those of both clades; the low sulcus inclination angle is to the lower end of the range found in the putatively related taxa.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF83FFCC018CFD106361FCDF.taxon	description	Figure 21 n – q Holotype NMB P 15555 (Fig. 21 j), Langhian, Brasso FM, Trinidad, Cb. 1852.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF83FFCE0236FF506005FE9E.taxon	discussion	Discussion Te genus Akko was reviewed by Van Tassell and Baldwin (2004) and was deemed to contain three species: Akko dionaea Birdsong & Robbins, 1995, from the Atlantic off northern Brazil; A brevis (Günther, 1896) in the Pacific from El Salvador to Peru; and A. rossi Van Tassell & Baldwin, 2004, based on a single specimen from off El Salvador. Akko is characterized by a number of autapomorphic characters and its relationships have long been enigmatic. Van Tassell and Baldwin (2004) placed Akko in the Microgobius group of the Gobiosomatini and compared it to Bollmannia and Parrella. Tis placement is apparently supported by molecular phylogenetic analysis shown by Tornabene and Van Tassell (2014) and Tornabene et al. (2016). In the latter article and Tornabene et al. (2022), Akko is shown in a clade further containing Parrella and Microgobius. Based on otoliths we consider Akko and Parrella to constitute a separate clade, closely related, however, to a clade combining all Microgobius species. Fishes of the genus Akko have a secretive lifestyle in burrows or covered in mud in the substrate at depth down to about 20 m (Froese & Pauly, 2023). Te reduced eye size may be indicative of living in muddy environments or being hidden in the substrate (Birdsong & Robins, 1995) Teir large, caninoid teeth suggest that they could be ambush hunters. Otoliths are available from two of the three extant species of Akko, namely A. dionaea (Fig. 22 b – c) and A. brevis (Fig. 22 a), and of two of the five species of Parrrella, namely P. ginsburgi Wade, 1946 (Fig. 22 h) and P. lucretiae (Eigenmann & Eigenmann, 1888) (Fig. 22 n – o) both from the East Pacific. In addition, we place two new fossil otolith-based genera in this clade, namely Cubaguanichthys n. gen. and Proparrella n. gen. Te otoliths of these genera are characterized by being slightly longer than or equally long as high, showing a recessed preventral angle (except in Cubaguanichthys, where it is projecting), lack or weak indentation of the posterior rim which is nearly vertical, a sole-shaped sulcus with low ostial lobe, and a distinct subcaudal iugum (sulcus reduced in size in (See figure on next page.) Fig. 22 a Akko brevis (GÜnther, 1864), extant, USNM 368666, 13 ° 41 ’ N 90 ° 00 ’ W, 13 m. b – c Akko dionaea Birdsong & Robins, 1995, extant, paratypes, USNM 329525, Brazil, 02 ° 32 ’ N 49 ° 57 ’ W, SL 80 and 83 mm. d – e Akko canoa n. sp.; d holotype, NMB P 15556, Canoa FM, Ecuador, PPP 3363; e paratype, Canoa FM, Ecuador, NMB 19146, NMB P 15557. f Akko lobata n. sp., holotype, NMB P 15558, Yaviza FM, Darien, Panama, PPP 1566. g Cubaguanichthys lanceolatus n. gen., n. sp., holotype, NMB P 15559, Cubagua FM, Venezuela, PPP 2568. h – i Parrella ginsburgi Wade, 1946; h extant, CAS ex SU 46827, Costa Rica, Gulf of Nicoya; i Armuelles FM, Pacific Panama, PPP 3236, NMB P 15560. j Parrella sp., Swan Cay FM, Swan Cay, Atlantic Panama, PPP 2221, NMB P 15561. k – m Parrella lucida n. sp.; k holotype, NMB P 690, Manzanilla FM, Trinidad, K 9833; l – m paratypes, l Shark Hole FM, Valiente Peninsula, Panama, PPP 2217, NMB P 15562, m Cubagua FM, Venezuela, PPP 2653, NMB P 15563. n – o Parrella lucretiae (Eigenmann & Eigenmann, 1888), extant, USNM 322734, 08 ° 06 ’ N 80 ° 31 ’ W	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF83FFCE0236FF506005FE9E.taxon	description	Akko). Otoliths of the proposed Antilligobius and Microgobius clades are higher than long.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF81FFCE018CFE36618AFE9E.taxon	description	Figure 22 d – e Holotype NMB P 15556 (Fig. 22 d), Gelasian, Canoa FM, Ecuador, PPP 3363. Paratypes 1 specimen, Gelasian, Canoa FM, Ecuador, NMB 19146 (NMB P 15557). Etymology After the type formation, the Canoa FM of Gelasian age in Ecuador. Diagnosis OL: OH = 1.04 – 1.08. Maximal size 2.6 mm (holotype). Otolith shape nearly rectangular with postdorsal rim slightly elevated. Predorsal angle high, preventral angle orthogonal; posterior rim nearly vertical, without indentation and no marked postdorsal projection. Sulcus narrow, moderately small, with very low ostial lobe; OL: SuL = 1.8 – 1.9. Subcaudal iugum short, wide. Sulcus inclination angle 16 – 22 °. Description Otolith size up to 2.6 mm in length (holotype). OH: OT = 2.7. Dorsal rim relatively low, smooth, nearly straight, slightly ascending towards posterior. Predorsal angle high, rounded, no distinct postdorsal projection; preventral and postventral angles orthogonal. Ventral rim straight, horizontal. Anterior and posterior rims nearly vertical, posterior rim without marked indentation at its middle. All rims smooth. Inner face flat in vertical direction, slightly bent in horizontal direction. Sulcus sole-shaped, moderately deep, narrow, inclined at 16 – 22 °; low ostial lobe, cauda with rounded tip slightly bent upward. OL: SuL = 1.8 – 1.9. Subcaudal iugum short, wide. Dorsal depression indistinct, with indiscernible crista superior. Ventral furrow distinct, half-moon shaped and regularly curved at moderate distance from ventral rim of otolith. Outer face broadly convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF81FFCE018CFE36618AFE9E.taxon	discussion	Discussion Akko canoa resembles the otoliths of the two known species in shape and sulcus organization but differs in being thicker (OH: OT = 2.7 vs 3.0 – 3.6), the larger sulcus (OL: SuL = 1.8 – 1.9 vs 2.0 – 2.2) and the less strongly bent inner face which in the two extant species studied shows a distinct outward bent posterior region of the otolith. Akko canoa shows that a further species of the genus existed in the East Pacific other than the extant species.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF81FFCE0236FEF666B5F9FE.taxon	description	Figure 22 f Holotype NMB P 15558 (Fig. 22 f), Tortonian, Yaviza FM, Pacific Panama, Darien, PPP 1566. Etymology From lobatus (Latin) = lobed, referring to the continuously curved anterior and dorsal rims. Diagnosis OL: OH = 1.03. Size 1.75 mm (holotype). Anterior and dorsal rim continuously curved. Posterior rim nearly vertical, with weak and broad indentation. Sulcus narrow, moderately large, with moderate ostial lobe; OL: SuL = 1.6. Subcaudal iugum short, wide. Sulcus inclination angle 18 °. Description Otolith size up to 2.6 mm in length (holotype). OH: OT = 2.8. High-bodied otolith; anterior and dorsal rim continuously curved, postdorsal rim slightly elevated and with small indentation in front of postdorsal tip. Preventral angle rounded orthogonal; posterior rim nearly vertical, with weak and broad indentation and weak, blunt postdorsal projection. Ventral rim straight, horizontal. All rims smooth, dorsal rim slightly undulating. Inner face nearly flat. Sulcus sole-shaped, moderately deep, long, inclined at 18 °; moderate ostial lobe, cauda with rounded tip. OL: SuL = 1.6. Subcaudal iugum short, wide. Dorsal depression distinct, with poorly defined crista superior. Ventral furrow distinct, regularly curved close to ventral rim of otolith. Outer face broadly convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF81FFCE0236FEF666B5F9FE.taxon	discussion	Discussion Akko lobata represents what one might envisage as a plesiomorphic otolith morphology for the genus where the sulcus is still relatively large, the ventral furrow close to the ventral rim of the otolith and the inner face is nearly flat. Tis character status is also observed in otoliths of Bollmannia and Microgobius.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF81FFCF0236F9D66399FBDE.taxon	materials_examined	Type species Cubaguanichthys lanceolatus n. sp. Etymology Named after the type formation, the Cubagua Formation of Venezuela. Diagnosis A fossil otolith-based genus of the Gobiosomatini, Microgobius Group, Akko clade with the following combination of characters. Ventral rim straight, horizontal and posterior rim straight, vertical forming orthogonal angle; dorsal rim regularly curved, anteriorly depressed; preventral projection pointed, distinct; OL: OH = 1.2. Outer face convex, smooth. Sulcus long, narrow, with very low ostial lobe; OL: SuL = 1.8; sulcus inclination angle 16 °. Subcaudal iugum short, wide, oval shaped. Ventral furrow distinct, close to ventral rim of otolith.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF81FFCF0236F9D66399FBDE.taxon	discussion	Discussion Cubaguanichthys has a highly distinctive otolith pattern that resembles otoliths of Akko in the straight, vertical posterior rim without indentation, the sulcus with a very low ostial lobe, and the wide, short subcaudal iugum. It differs in the more slender shape, the projecting preventral angle, and the long and narrow sulcus. Te allocation of Cubaguanichthys to the Microgobius group and Akko clade of the Gobiosomatini therefore remains tentative. It also seems possible that Cubaguanichthys represents an exotic faunal element in the tropical West Atlantic of the Late Pliocene that had emigrated from another region, but we know of no gobiid otolith morphology from outside of the American taxa that would show any significant similarity. Species A single species, C. lanceolatus, from the Early Pliocene, Zanclean, Cubagua FM, above Cerro Negro Member, of Venezuela.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF80FFCF018CFB36669BFEDE.taxon	description	Figure 22 g Holotype NMB P 15559 (Fig. 22 g), Zanclean, Cubagua FM, above Cerro Negro member, Venezuela, PPP 2568. Etymology From lanceolatus (Latin) = lanceolate, referring to the pointed preventral angle. Diagnosis See diagnosis of genus (monospecific genus). Description Otolith size 1.7 mm in length; OL: OH = 1.2; OH: OT = 2.5. Ventral rim straight, horizontal and posterior rim straight, vertical forming orthogonal angle. Dorsal rim regularly curved, anteriorly depressed, highest posteriorly at moderately developed postdorsal angle; no postdorsal projection. Preventral projection pointed, distinct; posterior rim without indentation. All rims smooth, slight concavity on dorsal rim behind obtuse postdorsal angle. Inner face slightly bent. Sulcus relatively long, very narrow, shallow, with very low ostial lobe; OL: SuL = 1.8; sulcus inclination angle 16 °. Subcaudal iugum short, wide, nearly oval in shape. Dorsal depression indistinct. Ventral furrow distinct, close to ventral rim of otolith. Outer face distinctly convex, more than inner face, smooth. Genus Parrella Ginsburg, 1938	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF80FFCF0236FE3666B6FC1E.taxon	description	Figure 22 h – i Material (fossil specimen) 1 specimen, Calabrian, Pacific Panama, Armuelles FM, PPP 323 (NMB P 15560). Discussion Te single fossil specimen from the late Early Pleistocene perfectly matches the otoliths of the extant specimens of this species (Fig. 22 h) that occurs in the same region. Typical is the recessed, rounded preventral angle, the high and projecting predorsal angle, the small concavity of the dorsal rim behind its highest point at the postdorsal angle, the shallow, even sulcus and the short but wide subcaudal iugum. Today, P. ginsburgi is found from the Gulf of California to Panama in nearshore, soft bottom habitats at depths at 1 – 10 m.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF80FFC80236FC76631CFC5E.taxon	description	Figure 22 k – m Holotype NMB P 690 (Fig. 22 k), Zanclean, Manzanilla FM, Telemaque Member, Trinidad, K 9833. Paratypes 3 specimens Zanclean: 2 specimens Shark Hole FM, Atlantic Panama, Valiente Peninsula, PPP 2217 (NMB P 15562); 1 specimen, Cubagua FM above Cerro Negro Member, Venezuela, PPP 2653 (NMB P 15563). Etymology From lucidus (Latin) = clear, referring to the clear allocation of the holotype to the genus Parrella. Diagnosis (holotype) OL: OH = 1.15. Maximal size 2.05 mm (holotype). Otolith shape dorsally expanded with predorsal angle and postdorsal projection projecting strongest. Sulcus narrow, short, with low ostial lobe; OL: SuL = 2.1. Subcaudal iugum moderately long and wide. Sulcus inclination angle 20 °. Description (holotype) Otolith size up to 2.05 mm in length (holotype). OH: OT = 2.9. Predorsal angle high, rather sharp and distinctly projecting; postdorsal projection also sharp and projecting resulting in longest axis of otolith shifted way above sulcus. Dorsal rim regularly curved, highest shortly behind its middle, undulating. Preventral and postventral angles rounded, recessed. Anterior rim straight, downward inclined; posterior rim straight, downward inclined, with slight concavity below pointed postdorsal projection. Ventral rim straight, horizontal. Inner face slightly convex. Sulcus sole-shaped, moderately deep, narrow, inclined at 20 °; low ostial lobe, cauda with rounded tip. OL: SuL = 2.1. Subcaudal iugum moderately long, wide. Dorsal depression broad, indistinct. Ventral furrow distinct, regularly curved, relatively close to ventral rim of otolith. Outer face broadly convex, slightly more than inner face, smooth. Remarks Te paratypes associated with this species are distinctly smaller and show less clear characteristics than the holotype, which we consider an ontogenetic effect. However, with more specimens being recovered it could also appear that they represent juveniles of another species.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF80FFC80236FC76631CFC5E.taxon	discussion	Discussion Parrella lucida differs from the known extant congeners in the otolith shape with the projecting predorsal and postdorsal regions and the relatively narrow sulcus. Today, a single species occurs in the West Atlantic, P. macropteryx Ginsburg, 1939, of which otoliths are not known.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF87FFC8018CFBB661AFFD5E.taxon	materials_examined	Type species Proparrella darienensis n. sp. Etymology Combination of pro (Latin) for before and the genus name Parrella, indicating its presumed plesiomorphic position as compared to Parrella. Diagnosis A fossil otolith-based genus of the Gobiosomatini, Microgobius Group, Akko clade with the following combination of characters. Small otoliths to about 2 mm in length and nearly quadratic otolith shape with slightly elevated postdorsal region; OL: OH = 1.0 – 1.1. Postdorsal rim with conspicuous narrow concavity behind postdorsal angle and before blunt, short postdorsal projection. Sulcus long, narrow, with low ostial lobe and long, narrow, straight, sometimes tapering cauda; OL: SuL very variable, ranging from 1.5 to 2.2; sulcus inclination angle 12 – 18 °. Subcaudal iugum moderately long, relatively narrow. Dorsal depression shifted dorsally with straight lower margin and broad crista superior. Ventral furrow distinct, close to ventral rim of otolith.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF87FFC8018CFBB661AFFD5E.taxon	discussion	Discussion Proparrella is best recognized by the characteristic shape of the postdorsal region with the narrow concavity in front of a short, blunt postdorsal projection and the, at first perhaps, inconspicuous shape of the cauda. Proparrella resembles otoliths of Akko as well as Parrella and is therefore considered to represent a fossil genus related to these taxa. It differs from Akko in the shape of the cauda, the broad crista superior, and the relatively long sulcus (except a few tentatively assigned specimens; see below), which it shares with the most plesiomorphic species in that genus, Akko lobata. Proparrella differs from Parrella in being more compressed and in the shape of the dorsal rim and the broad crista superior. It appears plausible that Proparrella represented an extinct branch as a sister taxon to Akko. Species At least two species: the widespread Proparrella darienensis n. sp., known from the Tortonian to Zanclean and distributed from Ecuador to Trinidad; and P. pusilla n. sp., known from the Tortonian and Messinian of Panama.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF87FFCA0236FCB66398FA18.taxon	description	Figure 23 a – m Holotype NMB P 15565 (Fig. 23 a), Tortonian, Tuira FM, Pacific Panama, Darien, PPP 1154. Paratypes 12 specimens: 9 specimens Tortonian: 3 specimens Angostura FM, Ecuador, 1 specimen PPP 3297 (NMB P 15572), 2 specimens NMB 19301 (NMB P 15576 - 77); 1 specimen Tuira FM, Pacific Panama, Darien, PPP 1163 (NMB P 15569); 2 specimens Yaviza FM, Pacific Panama, Darien, PPP 1566 (NMB P 15567 - 68); 3 specimens Gatun FM, Atlantic Panama, PPP 2168 (NMB P 15573 - 75). 3 specimens Zanclean: 1 specimen Cubagua FM, above Cerro Negro Member, Venezuela, PPP 2568 (NMB P 15566); 2 specimens Cayo Agua FM, Atlantic Panama, PPP 1276 (NMB P 15570 - 71). Referred specimens 198 specimens: 183 specimens Tortonian: 11 specimens Angostura FM, Ecuador, 4 specimens PPP 3297, 6 specimens PPP 3301, 1 specimen PPP 3465; 15 specimens Tuira FM, Pacific Panama, Darien, 6 specimens PPP 1139, 1 specimen PPP 1593, 1 specimen PPP 1627, 4 specimens PPP 1163, 3 specimens PPP 1164; 64 specimens Yaviza FM, Pacific Panama, Darien, 6 specimens PPP 1607, 1 specimen PPP 1609, 2 specimens PPP 1528, 1 specimen PPP 1145, 3 specimens PPP 1620, 36 specimens PPP 1171, 4 specimens PPP 1565, 11 specimens PPP 1566; 93 specimens Gatun FM, Atlantic Panama, PPP 2168. 6 specimens Messinian: 1 specimen Chucunaque FM, Pacific Panama, Darien, PPP 1578; 3 specimens Manzanilla FM, Montserrat Member, Trinidad, PPP 2666; 2 specimens, La Tejita FM, Venezuela, Isla Margarita, PPP 3087. 9 specimens Zanclean, Cubagua FM, above Cerro Negro Member, Venezuela, 3 specimens PPP 2568, 6 specimens PPP 2571. Etymology After Darien, Province in Panama, from which the holotype originates and which has yielded so many fossil gobiid otoliths. Diagnosis OL: OH = 1.04 – 1.1. Maximal size 2.0 mm. Otolith shape nearly quadratic with postdorsal region slightly elevated. Postdorsal rim with conspicuous narrow concavity behind postdorsal angle and before blunt, short postdorsal projection. Sulcus long, narrow, with low ostial lobe and long, narrow, straight, sometimes tapering cauda; OL: SuL = 1.5 – 1.75; sulcus inclination angle 12 – 18 °. Subcaudal iugum moderately long, relatively narrow. Dorsal depression shifted dorsally, with straight lower margin and broad crista superior. Ventral furrow distinct, close to ventral rim of otolith. Description Otolith size up to 2.0 mm in length (holotype 1.9 mm). OH: OT = 2.4 – 2.9. Dorsal rim anteriorly straight, ascending from high predorsal angle to slightly elevated postdorsal angle, followed by narrow, distinct postdorsal concavity just before narrow, blunt, short postdorsal projection. Preventral and postventral angles rounded, nearly orthogonal. Ventral rim straight to slightly bent, horizontal. Anterior and posterior rims nearly vertical or slightly inclined downward, posterior rim without or with shallow and broad indentation at its middle. All rims smooth. Inner face slightly convex. Sulcus sole-shaped, relatively deep, inclined at 12 – 18 °; ostium relatively wide but with moderate ostial lobe; cauda long, narrow, straight, sometimes with tapering tip. OL: SuL = 1.5 – 1.75. Subcaudal iugum moderately long, relatively narrow. Dorsal depression long, shifted dorsally, with straight lower margin and broad crista superior. Ventral furrow distinct, regularly curved and relatively close to ventral rim of otolith. Outer face broadly convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF87FFCA0236FCB66398FA18.taxon	discussion	Discussion Proparrella darienensis is a common, widespread and relatively small species of the Late Miocene and Early Pliocene that is best recognized by the specific shape of the dorsal rim, the overall quadratic shape and the shape of the cauda.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF85FFCA018CFA7066F5FEF8.taxon	description	Figure 23 n – p Material 4 specimens Zanclean, Cubagua FM, above Cerro Negro Member, Venezuela, PPP 2568 (figured specimens NMB P 15583 - 85). Discussion A few specimens from the Zanclean of the Cubagua FM in Venezuela differ from the typical P. darienensis specimens in the relatively short sulcus (OL: SuL = 2.0 – 2.2 vs 1.5 – 1.75) which may indicate the presence of a further species of Proparrella. However, the few specimens are only 1.1 to 1.4 mm in length and therefore do not warrant a clear definition.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF85FFCB0236FED0608EFE18.taxon	description	Figure 23 q – r Holotype NMB P 15578 (Fig. 23 r), Tortonian, Gatun FM, Atlantic Panama, PPP 2168. Paratypes 4 specimens: 3 specimens same data as holotype (NMB P 15579 - 81); 1 specimen, Messinian, Chucunaque FM, Pacific Panama, Darien, PPP 1576 (NMB P 15582). Referred specimens 5 specimens Messinian, Chucunaque FM, Pacific Panama, Darien, 2 specimens PPP 1576, 3 specimens PPP 1578. Etymology From pusillus (Latin) = teeny, small, referring to the small size of the specimens of this species. Diagnosis OL: OH = 1.03 – 1.08. Maximal size 1.4 mm. Otolith shape nearly quadratic with similarly high and projecting predorsal angle and postdorsal projection. Dorsal rim relatively low with weak concavity behind postdorsal angle. Sulcus long, narrow, with low ostial lobe and long, narrow, straight cauda; OL: SuL = 1.75 – 2.0; sulcus inclination angle 12 – 14 °. Subcaudal iugum moderately long, narrow. Dorsal depression indistinct. Ventral furrow distinct, close to ventral rim of otolith. Description Otolith size up to 1.4 mm in length (holotype). OH: OT = 2.7 – 2.9. Dorsal rim anteriorly straight, posteriorly slightly concave, with broadly rounded, nearly centrally positioned postdorsal angle. Predorsal angle projecting, high, at about the level and strength of postdorsal projection resulting in nearly symmetrical otolith outline. Preventral and postventral angles rounded, slightly recessed. Ventral rim straight, horizontal. Anterior and posterior rims nearly straight, slightly inclined downward. All rims smooth. Inner face nearly flat. Sulcus sole-shaped, relatively deep, inclined at 12 – 14 °; ostium moderately wide with moderate ostial lobe; cauda long, narrow, straight. OL: SuL = 1.75 – 2.0. Subcaudal iugum moderately long, narrow, sometimes rather indistinct. Dorsal depression shallow, indistinct. Ventral furrow distinct, regularly curved and relatively close to ventral rim of otolith. Outer face broadly convex, smooth. Discussion Despite of its small size, the otoliths of P. pusilla are relatively constant in expression and are therefore believed to represent a small species rather than juvenile specimens. Te highly distinctive otolith outline and the relatively short sulcus distinguish it from its parallel occurring congener, P. darienensis.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF84FFCB018CFAD066A3FB38.taxon	description	Figure 24 a – i Material (fossil specimens) 137 specimens: 1 specimen late Zanclean, upper Cayo Agua FM, Atlantic Panama, PPP 1188 (NMB P 15592); 9 specimens Piacenzian, Escudo de Veraguas FM, Atlantic Panama: 6 specimens PPP 1241, 3 specimens PPP 2169. 97 specimens Gelasian: 36 specimens Escudo de Veraguas FM, Atlantic Panama, 34 specimens PPP 2170 (figured specimens NMB P 15587 - 89), 1 specimen PPP 2173, 1 specimen PPP 2297; 47 specimens Bastimentos FM, Ground Creek Unit, Atlantic Panama, 18 specimens PPP 1252, 12 specimens PPP 1256 (figured specimen NMB P 15586), 27 specimens PPP 3204 (figured specimen NMB P 15591); 4 specimens Moin FM, Atlantic Costa Rica, PPP 2032. 30 specimens Calabrian, Swan Cay FM, Atlantic Panama, 24 specimens PPP 2221 (figured specimen NMB P 15590), 6 specimens PPP 2531. Description Otolith size up to 1.6 mm in length; OL: OH = 0.92 – 1.02; OH: OT = 3.2 – 3.7. Otolith shape high-bodied with predorsal angle positioned high and at same level as postdorsal tip; preventral and postventral angles well developed and longest horizontal section of otolith between them. Dorsal rim distinctly curved, sometimes irregularly undulating or highest at broad postdorsal angle. Ventral rim straight, horizontal. Anterior and posterior rims steep, dorsally inclined, with weak and broad central concavities. All rims smooth. Inner face flat with region above ventral furrow and below dorsal depression slightly bulged. Sulcus small, narrow, relatively deep, with low ostial lobe; sulcus inclination angle 6 – 12 °; OL: SuL = 1.9 – 2.1. Subcaudal iugum moderately long, wide. Dorsal depression large, wide, somewhat variably expressed. Ventral furrow distinct, regularly curved, relatively close to ventral rim of otolith. Outer face variably convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF84FFCB018CFAD066A3FB38.taxon	discussion	Discussion Antilligobius nikkiae is a common species in many of the deeper water sediments in the western Caribbean from Late Pliocene through Pleistocene. Today, Antilligobius nikkiae occurs hovering above the substrate in small schools at depth of about 100 to 200 m in the Caribbean (Tornabene et al., 2016).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF84FFCB018CFE706097FAB8.taxon	discussion	Discussion Te genera Antilligobius and Palatogobius represent a clade of mesophotic gobies in the tropical West Atlantic generally occurring at depths of about 100 m to 300 m (Tornabene & Baldwin, 2017; Tornabene et al., 2016). Tey are shown as the sister group to genera here combined in the Akko and Microgobius clades of Tornabene et al. (2016) or the Bollmannia clade (Tornabene et al., 2022). Antilligobius is monospecific with A. nikkiae Van Tassell & Colin, 2012 (Fig. 24 a – b), and Palatogobius contains three species, two of which are small, and otoliths are known from the largest, P. grandoculus Greenfield, 2002 (Fig. 25 a). Te otoliths of the two genera share certain features that, despite their generally small size, mostly distinguish them well from juveniles of other gobiid taxa of the region. Tese features are the high-bodied shape with an OL: OH ratio of about 1.0 or less, a low sulcus inclination angle of 12 ° or less (mostly <10 °), and the stretch from the preventral to the postventral angle being longer than from the high-positioned predorsal angle to the postdorsal tip (no distinct postdorsal projection developed). Tis morphological pattern is reasonably similar to otoliths of the Akko and Microgobius clades and thus supports the phylogenetic analysis of Tornabene et al. (2016).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF84FFD50236FB106031FAB8.taxon	description	Figure 24 j – s Holotype NMB P 15593 (Fig. 24 j), Piacenzian, Escudo de Veraguas FM, Atlantic Panama, PPP 1241. Paratypes 9 specimens: 2 specimens Tortonian: 1 specimen Angostura FM, Ecuador, 1 specimen PPP 3301 (NMB P 15602); 1 specimen Tuira FM, Pacific Panama, Darien, PPP 1132 (NMB P 15594). 1 specimen Messinian, Chucunaque FM, Pacific Panama, Darien, PPP 1578 (NMB P 15599). 1 specimen Zanclean, Cubagua FM, Cerro Negro Member, Venezuela, PPP 3055 (NMB P 15601). 1 specimen, Gelasian, upper Escudo de Veraguas FM, Atlantic Panama, PPP 2170 (NMB P 15600). 4 specimens Gelasian, Bastimentos FM, Ground Creek Unit, Atlantic Panama, PPP 3202 (NMB P 15595 - 98). Referred specimens 141 specimens: 34 specimens Tortonian: 10 specimens Tuira FM, Pacific Panama, Darien, 3 specimens PPP 1132, 6 specimens PPP 1134, 1 specimen PPP 1164; 8 specimens Yaviza FM, Pacific Panama, Darien, PPP 1566; 16 specimens Gatun FM, Atlantic Panama, PPP 2167. 5 specimens Messinian, Chucunaque FM, Pacific Panama, Darien, PPP 1578. 6 specimens Zanclean: 3 specimens Cubagua FM, Venezuela, 1 specimen PPP 2556, 2 specimens PPP 3055; 3 specimens Springvale FM, Trinidad, KR 1216. 8 specimens Piacenzian, Escudo de Veraguas FM, Atlantic Panama, PPP 1241. 88 specimens Gelasian: 15 specimens, Gelasian, upper Escudo de Veraguas FM, Atlantic Panama, PPP 2170; 52 specimens Bastimentos FM, Ground Creek Unit, Atlantic Panama, 40 specimens PPP 3202, 3 specimens PPP 1256, 9 specimens PPP 3204; 21 specimens Moin FM, Atlantic Costa Rica, 4 specimens PPP 2032, 4 specimens PPP 2038, 7 specimens PPP 3259, 2 specimens PPP 3260, 4 specimens PPP 3262. Etymology Named in honor of Laurel Collins (Florida International University) in recognition of her important contribution to the understanding of the geology and paleontology of the Isthmus of Panama. Diagnosis OL: OH = 1.05 – 1.1; OH: OT = 2.65 – 2.8. Maximal size 1.3 mm. Otolith shape nearly quadratic with predorsal angle positioned high and at same level as postdorsal tip. Sulcus short, narrow, with very low ostial lobe; OL: SuL = 1.85 – 2.05; sulcus inclination angle 6 – 12 °. Subcaudal iugum moderately long, relatively narrow. Dorsal depression indistinct, small. Ventral furrow distinct, regularly curving, close to ventral rim of otolith. Description Otolith size up to 1.3 mm in length (holotype 1.25 mm); Otolith shape high-bodied with predorsal angle positioned high and at same level as postdorsal tip; preventral and postventral angles well developed. Dorsal rim variably curved or relatively flat, with variable postdorsal tip, smooth. Ventral rim straight, horizontal. Anterior and posterior rims steep, without or with weak and broad central concavities. All rims smooth. Inner face flat with region above ventral furrow and below dorsal depression slightly bulged. Sulcus small, narrow, relatively deep, with very low ostial lobe; sulcus inclination angle 6 – 12 °; OL: SuL = 1.85 – 2.05. Subcaudal iugum moderately long, relatively narrow. Dorsal depression indistinct, mostly small. Ventral furrow distinct, regularly curved, relatively close to ventral rim of otolith. Outer face variably convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF84FFD50236FB106031FAB8.taxon	discussion	Discussion Antilligobius collinsae differs in only few subtle characters from the extant A. nikkiae, i. e., the slightly less compressed appearance (OL: OH 1.05 – 1.1 vs 0.92 – 1.02) and in being thicker (OH: OT = 2.65 – 2.8 vs 3.2 – 3.7). Antilligobius collinsae appears to represent a plesiomorphic counterpart to the extant species and occurs earlier in the stratigraphy in the Tortonian (Late Miocene) but shows a considerable stratigraphic overlap with A. nikkiae over the Piacenzian and Gelasian.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9AFFD5018CFAD061E1FA98.taxon	description	Figure 25 a – g Material (fossil specimens) 28 specimens: 19 specimens Zanclean, Cubagua FM, Venezuela: 4 specimens PPP 2554, 9 specimens PPP 3055 (figured specimen NMB P 15608), 6 specimens PPP 3057 (figured specimens NMB P 15603 - 04). 9 specimens Gelasian, Bastimentos FM, Ground Creek Unit, Atlantic Panama: 7 specimens PPP 1252 (figured specimen NMB P 15607), 2 specimens PPP 3204 (NMB P 15605 - 06). Description Otolith size up to 1.3 mm in length; OL: OH = 0.92 – 1.0; OH: OT = 2.8 – 3.2. Otolith shape high-bodied with relatively regularly curved dorsal rim without prominent predorsal or postdorsal angles, highest at its middle. Ventral rim straight, horizontal. Preventral and postventral angles distinct, longest section of otolith between them. Anterior and posterior rims steep, dorsally inclined. All rims smooth. Inner face flat with region above ventral furrow and below dorsal depression slightly bulged. Sulcus small, narrow, relatively deep, with very low ostial lobe; sulcus inclination angle 2 – 7 °; OL: SuL = 2.1 – 2.6. Subcaudal iugum moderately long, relatively narrow. Dorsal depression large, wide, somewhat variably expressed. Ventral furrow distinct, regularly curved, moderately close to ventral rim of otolith. Outer face convex, smooth. Discussion Te otoliths of Palatogobius grandoculus are quite similar to those of Antilligobius nikkiae, differing in the more regularly curved dorsal rim without marked angles, the even smaller sulcus (OL: SuL = 2.1 – 2.6 vs 1.9 – 2.1) and the less inclined sulcus (2 – 7 ° vs 6 – 12 °). Te species occurs regularly but never very common since the Early Pliocene in deeper water sediments of Venezuela and Atlantic Panama. Today, Palatogobius grandoculus occurs hovering above the substrate in small schools at depth of about 220 to 280 m in the Caribbean (Tornabene et al., 2016).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9AFFD70236FAF0602FFA38.taxon	description	Figure 25 h Holotype NMB P 15609 (Fig. 25 h), Zanclean, Cayo Agua FM, Atlantic Panama, PPP 2240. (See figure on next page.) Fig. 25 a – g Palatogobius grandoculus Greenfield, 2002; a extant, USNM 438754, 15 ° 34 ’ N 61 ° 28 ’ W, 194 m; b – c Cubagua FM, Venezuela, PPP 3057, NMB P 15603 - 04; d, e Bastimentos FM, Ground Creek Unit, Bocas del Toro, Panama, PPP 3204, NMB P 15605 - 06; f Cubagua FM, Venezuela, PPP 3055, NMB P 15608; g Bastimentos FM, Ground Creek Unit, Bastimentos Island, Panama, PPP 1252, NMB P 15607. h Palatogobius magnus n. sp., holotype (h 1 SEM picture), Cayo Agua FM, Atlantic Panama, PPP 2240, NMB P 15609. i – k Palatogobius pacificus n. sp., Armuelles FM, Pacific Panama; k holotype, NMB P 15610, PPP 3240, i – j paratypes, PPP 3237, NMB P 15611 - 12. l – q Palatogobius vantasselli n. sp., Manzanilla FM, Trinidad, PPP 2666; l holotype, NMB P 15613; m – q paratypes, NMB P 15614 - 18. r – u Palatogobius sp. 1, Cubagua FM, Venezuela, PPP 2571, NMB P 15619 - 22. v Palatogobius sp. 2, Tuira FM, Darien, Panama, PPP 1137, NMB P 15623 Etymology From magnus (Latin) = large, referring to the large size of the otolith. Diagnosis Otolith size 1.7 mm in length. OL: OH = 0.92; OH: OT = 3.0. Otolith shape high-bodied with relatively regularly curved dorsal rim, highest at its middle. Inner face completely flat. Sulcus deep, short, relatively wide, with very low ostial lobe; OL: SuL = 2.0; sulcus inclination angle 11 °. Subcaudal iugum moderately long, relatively narrow. Dorsal depression wide, with indistinct margins. Ventral furrow distinct, regularly curving, relatively distant from ventral rim of otolith. Description Otolith size 1.7 mm in length. Otolith shape high-bodied with relatively regularly curved dorsal rim; dorsal rim highest at its middle, with rounded postdorsal tip and high, very gently rounded predorsal angle. Anterior and posterior rims near vertical; posterior rim with very wide and shallow concavity. Ventral rim straight, horizontal. Preventral angle well-developed, postventral angle broadly rounded. All rims smooth. Inner face completely flat. Sulcus small, wide, deep, with very low ostial lobe; sulcus inclination angle 11 °; OL: SuL = 2.0. Subcaudal iugum moderately long, well-developed and relatively wide. Dorsal depression large, wide, with indistinct margins, reaching close to sulcus. Ventral furrow distinct, regularly curved, relatively distant from ventral rim of otolith. Outer face evenly convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9AFFD70236FAF0602FFA38.taxon	discussion	Discussion Palatogobius magnus is remarkable for its size, which immediately puts it apart from the extant P. grandoculus and the other two species, of which otoliths are not known, but which are distinctly smaller even than P. grandoculus. Te otolith and sulcus shape, however, characterize the single, very well preserved holotype unequivocally in the genus Palatogobius.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF98FFD7018CFA10660DFAF8.taxon	description	Figure 25 i – k Holotype NMB P 15610 (Fig. 25 k), Calabrian, Armuelles FM, Pacific Panama, PPP 3240. Paratypes 2 specimens, Calabrian, Armuelles FM, Pacific Panama, PPP 3237 (NMB P 15611 - 12). Referred specimen 1 specimen same data as holotype. Etymology Referring to the occurrence in the Pacific Ocean. Diagnosis Otolith size up to 2.35 mm in length (holotype). OL: OH = 0.95 – 1.05; OH: OT = 2.6 – 3.0. Otolith shape high-bodied with regularly curved dorsal rim, highest at its middle. Inner face completely flat. Sulcus deep, short, narrow, with very low ostial lobe; OL: SuL = 1.85 – 2.05; sulcus inclination angle 5 – 10 °. Subcaudal iugum short, very wide. Dorsal depression wide, with indistinct margins. Ventral furrow distinct, regularly curving, relatively close to ventral rim of otolith. Description Otolith size up to 2.35 mm in length (holotype). Otolith shape high-bodied with regularly curved dorsal rim without prominent angles; dorsal rim highest at its middle or slightly behind. Anterior rim slightly inclined, posterior rim near vertical, with wide, indistinct concavity. Ventral rim straight or slightly bent, horizontal. Preventral angle well-developed, postventral angle rounded. All rims smooth. Inner face completely flat. Sulcus small, narrow, deep, with very low ostial lobe; sulcus inclination angle 5 – 10 °; OL: SuL = 1.85 – 2.05. Subcaudal iugum moderately long, well-developed and very wide. Dorsal depression large, wide, with indistinct margins, not reaching close to sulcus. Ventral furrow distinct, regularly curved, relatively close to ventral rim of otolith. Outer face evenly convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF98FFD7018CFA10660DFAF8.taxon	discussion	Discussion Palatogobius pacificus represents the only species of the genus in the East Pacific while all extant species are strictly West Atlantic. It is also the largest species in the genus, larger even than P. magnus from the Early Pliocene of the Atlantic side of Panama. Palatogobius pacificus differs from P. magnus in the more regularly curved dorsal rim, and the narrower sulcus. Both species likely represented geminate species on either side of the Isthmus of Panama.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF98FFD00236FAD0636EF9B8.taxon	description	Figure 25 l – q Holotype NMB P 15613 (Fig. 25 l), Messinian, Manzanilla FM, Montserrat Member, Trinidad, PPP 2666. Paratypes 5 specimens same data as holotype (NMB P 15614 - 18). Referred specimen 51 specimens: 1 specimen Tortonian: Manzanilla FM, San José Member, Trinidad, PPP 2669. 50 specimens Messinian: 45 specimens same data as holotype; 5 specimens Manzanilla FM, Montserrat Member, Trinidad, PPP 2677. Etymology Named in honor of James Van Tassell (New York) in recognition of his many contributions to the understanding of gobies from America. Diagnosis Otolith size up to 1.32 mm in length. OL: OH = 0.95 – 1.05; OH: OT = 2.8 – 3.2. Otolith shape high-bodied with regularly curved dorsal rim, highest near its middle and symmetrically developed predorsal angle and short postdorsal projection. Inner face completely flat. Sulcus deep, short, narrow, with low ostial lobe; OL: SuL = 1.9 – 2.0; sulcus inclination angle 10 – 15 °. Subcaudal iugum long, narrow. Dorsal depression small, with distinct margins, not reaching close to sulcus. Ventral furrow distinct, close to ventral rim of otolith. Description Otolith size up to 1.32 mm in length (holotype 1.28 mm). Otolith shape high-bodied with regularly curved dorsal rim highest near its middle. Predorsal angle and postdorsal projection similarly developed and nearly symmetrical. Anterior rim slightly inclined, posterior rim near vertical, both with wide, shallow central concavity. Ventral rim straight, horizontal. Preventral angle well-developed, sometimes projecting, postventral angle less pronounced. All rims smooth. Inner face completely flat. Sulcus small, moderately narrow, deep, with low ostial lobe; sulcus inclination angle 10 – 15 °; OL: SuL = 1.9 – 2.0. Subcaudal iugum long, narrow. Dorsal depression relatively small, cup-shaped, not reaching close to sulcus. Ventral furrow distinct, regularly curved, relatively close to ventral rim of otolith. Outer face evenly convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF98FFD00236FAD0636EF9B8.taxon	discussion	Discussion Palatogobius vantasselli differs from its later congeners in the more pronounced predorsal angle and postdorsal projection, the mild concavities at the anterior and posterior margins, the steeper sulcus inclination angle (10 – 15 ° vs 2 – 11 °) and the relatively narrow subcaudal iugum. Tese features are all considered plesiomorphic for the genus and would thus indicate that P. vantasselli might be positioned near the base of the Antilligobius clade.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9FFFD00236FCF0671DFBD8.taxon	discussion	Discussion In this informal clade we combine Microgobius and the fossil otolith-based genus Magnogobius n. gen. that share a specific otolith morphology characterized by an otolith shape which is higher than long, the inner face being more or less strongly bent in the horizontal direction, and a large sulcus, often with a distinct ostial lobe and with a large subcaudal iugum.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9FFFD0018CF9906637FE38.taxon	description	Figure 25 r – u Material 65 specimens Zanclean, Cubagua FM, Venezuela, 5 specimens PPP 2556, 3 specimens PPP 2563, 4 specimens PPP 3057, 31 specimens PPP 2568, 22 specimens PPP 2571 (figured specimens NMB P 15619 - 22). Discussion Te Cubagua FM of Venezuela has yielded a number of small gobiid otoliths not exceeding 0.9 mm in length that appear to represent a species of Palatogobius. In the light the Early Pliocene age and the fact that otoliths of two very small extant Palatogobius living in the Caribbean are not known, these otoliths are here referred to in open nomenclature.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9FFFD00236FE106715FC98.taxon	description	Figure 25 v Material 1 specimen Tortonian, Tuira FM, Pacific Panama, Darien, PPP 1137 (NMB P 15623). Discussion A single small (1 mm long), high-bodied otolith (OL: OH = 0.94) from the Tortonian of Darien may possibly represent another species of Palatogobius similar to P. vantasselli but because of its small size and singular occurrence we refrain from description.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9FFFD10236FB3063F4FB38.taxon	materials_examined	Type species Magnogobius grandis n. sp. Etymology Combination of magnus (Latin) for large and the genus name Gobius, indicating the large size of the otoliths that infers also a large size of the fishes from which they originated, and also as antagonism to the name of the putatively related genus Microgobius. Diagnosis A fossil otolith-based genus of the Gobiosomatini, Microgobius Group, Microgobius clade with the following combination of characters. Large otoliths to about 3.3 mm in length and high-bodied with an elevated postdorsal region and a broad predorsal angle and postdorsal projection; OL: OH = 0.83 – 0.97. Sulcus large, with a wide ostium and a pronounced ostial lobe; cauda long, narrow, straight, slightly deepened; OL: SuL = 1.3 – 1.5; sulcus inclination angle 12 – 20 °. Subcaudal iugum very long, distinct, narrow. Dorsal depression small, only above central part of sulcus. Ventral furrow distinct, anteriorly not extending beyond ostium.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9FFFD10236FB3063F4FB38.taxon	discussion	Discussion Magnogobius is recognized by its large size in combination with a high-bodied shape, the large sulcus with a distinct ostial lobe, the narrow, long, and distinct subcaudal iugum, and the small dorsal depression. Te otolith size and the expression of the ostial lobe and the subcaudal iugum are the main features distinguishing Magnogobius from the otoliths of the putatively related Microgobius. Otoliths are known from all other large-growing extant American gobioid genera, and none of them resemble Magnogobius in any significant way. It cannot, however, be entirely ruled out that Magnogobius is of an exotic origin in the Miocene and Pliocene of the West Atlantic and that the genus is related to one outside of America. For instance, otoliths of the West African Porogobius schlegeli (Günther, 1861) somewhat resemble Magnogobius in the distinct and long subcaudal iugum and the distinct ostial lobe, but they have a massive postdorsal projection and a concave outer face not matched in Magnogobius otoliths. We therefore interpret Magnogobius as representing an extinct genus of the Gobiosomatini, Microgobius group, and possibly related to the genus Microgobius. Species Two species: Magnogobius costaricensis n. sp. from the Tortonian of Panama, Darien to the Piacenzian of Atlantic Costa Rica and Panama, and Magnogobius grandis n. sp. from the Tortonian of Panama, Darien and the Tortonian to Gelasian of the Dominican Republic, Atlantic Costa Rica and Panama.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9EFFD3018CFB1063FEFE38.taxon	description	Figure 26 a – g Holotype NMB P 15624 (Fig. 26 a), Piacenzian, Río Banano FM, Atlantic Costa Rica, PPP 1732. Paratypes 7 specimens: 5 specimens Piacenzian, Río Banano FM, Atlantic Costa Rica; 1 specimen PPP 676 (NMB P 15629); 4 specimens same data as holotype (NMB P 15625 - 28). 1 specimen late Zanclean, upper Cayo Agua FM, Atlantic Panama, PPP 1188 (NMB P 15630). 1 specimen Tortonian, Yaviza FM, Pacific Panama, Darien, PPP 1609 (NMB P 15631). Referred specimen 6 specimens: 1 specimen Tortonian, Yaviza FM, Pacific Panama, Darien, PPP 1609. 1 specimen Zanclean, upper Cayo Agua FM, Atlantic Panama, PPP 1188. 5 specimens Piacenzian: 4 specimens Río Banano FM, Atlantic Costa Rica, PPP 1732. Etymology Referring to Costa Rica where the holotype was found. Diagnosis Otolith size up to 2.25 mm in length. OL: OH = 0.9 – 0.97. Otolith shape high-bodied; dorsal rim irregularly curved, with pronounced postdorsal angle and slight postdorsal projection; predorsal angle slightly projecting. Inner face slightly convex. Sulcus long, moderately wide, with distinct ostial lobe and deepened cauda; OL: SuL = 1.35 – 1.5; sulcus inclination angle 12 – 17 °. Subcaudal iugum long, narrow. Dorsal depression small, oval, only above central part of sulcus. Description Otolith size up to 2.25 mm in length (holotype). OH: OT = 3.0 – 3.3. Otolith shape high-bodied with irregularly curved dorsal rim highest at its postdorsal angle. Predorsal angle rounded, often slightly projecting; postdorsal projection blunt or rounded, slightly projecting. Anterior and posterior rims vertical to slightly inclined towards ventral; posterior rim with or without shallow central concavity. Ventral rim straight to slightly bent, horizontal. Preventral angle recessed, postventral angle rounded. All rims smooth except dorsal rim often irregularly undulating. Inner face slightly convex, particularly in horizontal direction. Sulcus long, moderately wide, with distinct ostial lobe; cauda straight, with rounded tip, slightly upward bent and slightly deepened; sulcus inclination angle 12 – 17 °; OL: SuL = 1.35 – 1.5. Subcaudal iugum long, narrow, distinct. Dorsal depression small, oval, only above central part of sulcus. Ventral furrow distinct, moderately close to ventral rim of otolith, anteriorly terminating at tip of ostium. Outer face evenly convex, relatively smooth. (See figure on next page.) Fig. 26 a – g Magnogobius costaricensis n. gen., n. sp.; a holotype, NMB P 15624, Río Banano FM, Limón, Costa Rica, PPP 1732; b – g paratypes; b Río Banano FM, Limón, Costa Rica, PPP 676, NMB P 15629; c, f, g Río Banano FM, Limón, Costa Rica, PPP 1732, NMB P 15625 - 28; d Cayo Agua FM, Bocas del Toro, Panama, PPP 1188, NMB P 15630; e Yaviza FM, Darien, Panama, PPP 1609, NMB P 15631. h – o Magnogobius grandis n. gen., n. sp.; h holotype, NMB P 386, Cercado FM, Dominican Republic, NMB 16915; i – o paratypes; i, l Cercado FM, Dominican Republic, NMB 16918, i NMB P 386, l NMB P 15632; j Cayo Agua FM, Bocas del Toro, Panama, PPP 2222, NMB P 15633; k Cercado FM, Dominican Republic, NMB 16916, NMB P 403; m Río Banano FM, Limón, Costa Rica, PPP 1732, NMB P 15636; n Cayo Agua FM, Bocas del Toro, Panama, PPP 1188, NMB P 15637; o Tuira FM, Darien, Panama, PPP 1163, NMB P 15638	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9EFFD3018CFB1063FEFE38.taxon	discussion	Discussion Magnogobius costaricensis does not grow to the large sizes observed in M. grandis n. sp. and further differs in the narrower sulcus, the less strongly developed ostial lobe and the oval dorsal depression. Both species occur parallel in space and time.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9CFFD3018CFE10669BFB38.taxon	description	Figure 26 h – o 1992 Gobiidae sp. 1 — Nolf & Stringer: pl. 17, Fig. 12. Holotype NMB P 386 (Fig. 26 h), Tortonian, Cercado FM, Dominican Republic, NMB 16915. Paratypes 11 specimens: 8 specimens Tortonian: 7 specimens Cercado FM, Dominican Republic; 2 specimens NMB 16918 (NMB P 15632); 2 specimens, NMB 16916 (NMB P 403); 1 specimen NMB 16923 (NMB P 405); 2 specimens NMB 15903 (NMB P 389); 1 specimen Tuira FM, Pacific Panama, Darien, PPP 1163 (NMB P 15638). 2 specimen Zanclean, Cayo Agua FM, Atlantic Panama, 1 specimen PPP 2222 (NMB P 15633), 1 specimen PPP 1188 (NMB P 15637). 1 specimen Piacenzian, Río Banano FM, Atlantic Costa Rica, PPP 1732 (NMB P 15636). Referred specimen 22 specimens: 6 specimens Tortonian: 3 specimens Tuira FM, Pacific Panama, Darien, 1 specimen PPP 1627, 1 specimen PPP 1154, 1 specimen PPP 1164; 2 specimens Cercado FM, Dominican Republic, NMB 16923. 12 specimens Zanclean, Atlantic Panama: 1 specimen Shark Hole FM, PPP 2219; 11 specimens Cayo Agua FM, 6 specimens PPP 1276, 5 specimens, PPP 1188. 3 specimens Piacenzian, Río Banano FM, Atlantic Costa Rica, 1 specimen PPP 676, 2 specimens PPP 1732. 1 specimen Gelasian, Bastimentos FM, Ground Creek Unit, Atlantic Panama, PPP 1252. Etymology From grandis (Latin) large, referring to the large size of the otoliths. Diagnosis Otolith size up to 3.3 mm in length. OL: OH = 0.83 – 0.92. Otolith shape high-bodied; dorsal rim regularly curved, crenulated, with rounded postdorsal angle and slight postdorsal projection; predorsal angle slightly projecting. Inner face slightly convex. Sulcus long, wide, with strong ostial lobe and deepened cauda; OL: SuL = 1.35 – 1.5; sulcus inclination angle 15 – 20 °. Subcaudal iugum long, narrow, distinct. Dorsal depression small, cup-shaped, only above central part of sulcus and close to sulcus. Description Otolith size up to 3.3 mm in length (holotype 2.9 mm). OH: OT = 3.4 – 3.7. Otolith shape high-bodied with regularly curved dorsal rim highest at its postdorsal angle and usually crenulated. Predorsal angle rounded, often slightly projecting; postdorsal projection blunt or rounded, slightly projecting. Anterior and posterior rims vertical to slightly inclined towards ventral; posterior rim usually straight. Ventral rim straight to slightly bent, horizontal. Preventral angle recessed, postventral angle rounded. All rims crenulated to various extents. Inner face slightly convex, particularly in horizontal direction. Sulcus long, wide, with strong, often sharp ostial lobe; cauda straight, with rounded tip, slightly upward bent and slightly deepened; sulcus inclination angle 15 – 20 °; OL: SuL = 1.35 – 1.5. Subcaudal iugum long, narrow, distinct. Dorsal depression small, cup-shaped, only above central part of sulcus; separated from sulcus only by narrow crista superior. Ventral furrow distinct, moderately close to ventral rim of otolith, anteriorly terminating below tip of ostium. Outer face convex, intensely ornamented.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9CFFD3018CFE10669BFB38.taxon	discussion	Discussion Magnogobius grandis differs from M. costaricensis in the larger size the otoliths can attain, the larger sulcus with the more strongly developed ostial lobe, the more evenly curved and crenulated dorsal rim, the strongly ornamented outer face and the more steeply inclined sulcus (15 – 20 ° vs 12 – 17 °). Both species occur parallel in a number of locations.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9CFFDC0236FB106167FDBE.taxon	discussion	Discussion Fricke et al. (2023) list 15 valid extant Microgobius species, six in the West Atlantic and nine in the East Pacific. Otoliths are available from all Atlantic species and from the majority of Pacific species except for Microgobius cyclolepis Gilbert, 1890, M. emblematicus Jordan & Gilbert, 1882, M. miraflorensis Gilbert & Starks, 1904, and M. urraca Tornabene, van Tassell & Robertson, 2012. We distinguish four groups (otolith plexus) within the genus based on certain otolith patterns: Te Microgobius gulosus otolith plexus containing the extant M. gulosus (Girard, 1858) (Fig. 27 a) and M. microlepis Longley & Hildebrand, 1940 (Fig. 27 i), from the West Atlantic; and M. tabogensis Meek & Hildebrand, 1928 (Fig. 27 c – d), from the East Pacific. Te otoliths of this plexus are characterized by the lowest height (OL: OH = 0.86 – 0.97) and an almost rectangular outline.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9CFFDC0236FB106167FDBE.taxon	description	Te Microgobius carri otolith plexus containing the extant M. carri Fowler, 1945 (Fig. 28 a), M. meeki Evermann & Marsh, 1899 (Fig. 28 i), M. signatus Poey, 1876 (Fig. 28 d), and M. thalassinus Jordan & Gilbert, 1883 (Fig. 29 h – i), from the West Atlantic; and M. brevispinis Ginsburg, 1939 (drawing of otolith provided by D. Nolf), and M. crocatus Birdsong, 1968 (Fig. 28 j – l), from the East Pacific. Te otoliths of this plexus are distinctly more high-bodied than those of the Microgobius gulosus plexus (OL: OH = 0.77 – 0.88 vs 0.86 – 0.97) and are usually associated with a highly curved dorsal rim.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF9CFFDC0236FB106167FDBE.taxon	description	Te composition of the Microgobius otoliths in these four otolith plexuses is only meant to be for comparison purposes and does not imply in any way possible phylogenetic relationships within the genus. In his review of the genus, Birdsong (1981) commented on an unpublished study by Ginsburg that was to define seven subgenera. Birdsong himself, however, stated that he has “ come to many tentative conclusions, but only one seems worth mentioning: I don’t know how they are related to each other. ” In a study about the speciation in Gobiosomatini that also included molecular phylogenetic data, Rüber et al. (2003) presented a cladogram containing six Microgobius species plus one Microgobius sp. Teir phylogenetic association of the six species of Microgobius partly coincides with the otolith plexus grouping and partly does not.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF93FFDC0236FDD666D2FB3E.taxon	description	Figure 27 a – b Material 2 specimens late Zanclean, upper Cayo Agua FM, Atlantic Panama, PPP 1188 (NMB P 15639).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF93FFDC0236FDD666D2FB3E.taxon	discussion	Discussion Microgobius gulosus has the most compressed otoliths known so far from extant species in the genus, with an OL: OH ratio of 0.95 – 0.97, caused primarily by the rather low dorsal rim, and a relatively small sulcus (OL: SuL = 1.65 – 1.85). Two well-preserved specimens from the Late Pliocene of Atlantic Panama are reported south of the southernmost extant occurrence, which according to Froese and Pauly (2023) is the Yucatan Peninsula of Mexico. Microgobius gulosus is known to commonly live in burrows (Birdsong, 1981).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF93FFDE0236FB166307F9F8.taxon	description	Figure 27 e – h 1998 Gobiidae indet. — Nolf & Aguilera: pl. 15, Fig. 9 – 11. Holotype NMB P 15641 (Fig. 27 f), Burdigalian, Cantaure FM, Venezuela, PPP 2539. Paratypes 6 specimens: 2 specimens Burdigalian, same data as holotype (NMB P 15642 - 43). 4 specimens Tortonian: 1 specimen Gatun FM, Atlantic Panama, PPP 2165 (See figure on next page.) Fig. 27 a – b Microgobius gulosus (Girard, 1858); a extant, CAS ex SU 5507, Galveston Bay; b Cayo Agua FM, Bocas del Toro, Panama, PPP 1188, NMB P 15639. c – d Microgobius tabogensis Meek & Hildebrand, 1928, extant, USNM 367369, El Salvador, 13 ° 10 ’ N 88 ° 28 ’ W, SL 35 and 36 mm. e – h Microgobius cantaurensis n. sp.; f holotype, NMB P 15641, Cantaure FM, Paraguaná, Venezuela, PPP 2539; e, g – h paratypes; e Gatun FM, Colon, Panama, PPP 2165, NMB P 15647; g Cantaure FM, Paraguaná, Venezuela, PPP 2539, NMB P 15642; h Tuira FM, Darien, Panama, PPP 1606, NMB P 15644. i – l Microgobius microlepis Longley & Hildebrand, 1940; i extant, FSBC 34988, 27 ° 844 N 80 ° 44 W, SL 31 mm; j – l Cumaná FM, Venezuela, PPP 3038, NMB P 15648 - 50. m – s Microgobius verecundus n. sp.; n holotype, NMB P 15651, Tuira FM, Darien, Panama, PPP 1163; m, o – s paratypes; m, p, q Angostura FM, Ecuador, PPP 3301, NMB P 15652 - 54; o, r, s Tuira FM, Darien, Panama, PPP 1163, NMB P 15655 - 57 (NMB P 15647); 3 specimens Tuira FM, Pacific Panama, Darien, PPP 1606 (NMB P 15644 - 46). Referred specimen 17 specimens Tortonian, Gatun FM, Atlantic Panama, 3 specimens PPP 2165, 2 specimens PPP 2167, 12 specimens PPP 2168. Etymology Referring to the Cantaure FM of Venezuela, the type formation. Diagnosis Otolith size up to 1.7 mm in length. OL: OH = 0.95 – 0.97. Otolith shape nearly rectangular; dorsal rim irregularly curved, low, with pronounced postdorsal projection. Inner face slightly convex. Sulcus long, moderately wide, with distinct ostial lobe and deepened cauda; OL: SuL = 1.4 – 1.55; sulcus inclination angle 12 – 15 °. Subcaudal iugum long, narrow. Ventral furrow very close and parallel to ventral rim of otolith. Description Otolith size up to 1.7 mm in length (holotype). OH: OT = 3.4 – 3.5. Otolith shape nearly rectangular with irregularly curved and undulating dorsal rim highest at its postdorsal angle. Predorsal angle rounded; postdorsal projection rounded, slightly projecting. Anterior rim near vertical, posterior rim slightly inclined towards ventral; posterior rim with weak central concavity. Ventral rim straight to slightly bent, horizontal. Preventral angle orthogonal, postventral angle orthogonal, rounded. All rims smooth except dorsal rim. Inner face slightly convex, particularly in horizontal direction. Sulcus long, moderately wide, with distinct ostial lobe; cauda straight, with rounded tip, slightly deepened; sulcus inclination angle 12 – 15 °; OL: SuL = 1.4 – 1.55. Subcaudal iugum long, narrow. Dorsal depression small, usually indistinct. Ventral furrow distinct, very close and parallel to ventral rim of otolith. Outer face evenly convex or with slight ventral umbo, relatively smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF93FFDE0236FB166307F9F8.taxon	discussion	Discussion Microgobius cantaurensis closely resembles the extant and Late Pliocene M. gulosus particularly in otolith shape and proportions. It differs from M. gulosus in the much larger sulcus (OL: SuL = 1.4 – 1.55 vs 1.65 – 1.85) and the more strongly developed postdorsal projection.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF91FFDE018CF9D0617DFDD8.taxon	description	Figure 27 i – l Material 31 specimens Calabrian, Cumaná FM, Venezuela, PPP 3038 (figured specimens PPP 15648 - 50). Discussion Otoliths of M. microlepis are recognized by their nearly rectangular shape characterized by near vertical anterior and posterior rims and a relatively shallow and gently curved dorsal rim (OL: OH = 0.85 – 0.92). It is further characterized by a relatively short sulcus with tapering cauda and triangular ostial lobe, while the ventral sulcus margin is nearly straight. Tese relatively characteristic otoliths have only been found in the Pleistocene, Calabrian of Venezuela, which is south of the present distribution of the species in the Gulf of Mexico and the western Caribbean along the coasts Yucatan and Belize.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF91FFDF0236FD3063E4FB7D.taxon	description	Figure 27 m – s Holotype NMB P 15651 (Fig. 27 n), Tortonian, Tuira FM, Pacific Panama, Darien, PPP 1163. Paratypes 7 specimens Tortonian: 4 specimens same data as holotype (NMB P 15655 - 58); 3 specimens Angostura FM, Ecuador, PPP 3301 (NMB P 15652 - 54). Referred specimen 112 specimens: 111 specimens Tortonian: 19 specimens Angostura FM, Ecuador, 7 specimens PPP 3297, 12 specimens PPP 3301; 33 specimens Tuira FM, Pacific Panama, Darien, 2 specimens PPP 1132, 11 specimens PPP 1134, 7 specimens PPP 1137, 2 specimens PPP 1606, 5 specimens PPP 1997, 3 specimens PPP 1142, 1 specimen PPP 1154, 1 specimen PPP 1163, 1 specimen PPP 1164; 31 specimens Yaviza FM, Pacific Panama, Darien, 3 specimens PPP 1607, 9 specimens PPP 1609, 3 specimens PPP 1528, 4 specimens PPP 1171, 5 specimens PPP 1570, 2 specimens PPP 1565, 5 specimens PPP 1566; 13 specimens Gatun FM, Atlantic Panama, PPP 2168; 13 specimens Manzanilla FM, San José Member, Trinidad, 1 specimen PPP 2669, 1 specimen PPP 2670, 10 specimens PPP 2671, 1 specimen PPP 2676. 3 specimens Messinian, Chucunaque FM, Pacific Panama, Darien, PPP 1576, 2 specimens PPP 1612. Etymology From verecundus (Latin) = modest, referring to the unspectacular morphology of the otoliths of this species. Diagnosis Otolith size up to 1.8 mm in length. OL: OH = 0.86 – 0.9. Otolith shape nearly rectangular; dorsal rim irregularly curved, with weak postdorsal projection. Sulcus long, relatively narrow, with relatively low ostial lobe and deepened cauda; OL: SuL = 1.55 – 1.75; sulcus inclination angle 15 – 20 °. Subcaudal iugum moderately long, narrow. Ventral furrow very close to ventral rim of otolith, relatively regularly curved. Description Otolith size up to 1.8 mm in length (holotype 1.75 mm). OH: OT = 3.5 – 3.8. Otolith shape nearly rectangular with irregularly curved, slanting dorsal rim highest at its postdorsal angle. Predorsal angle rounded; postdorsal projection rounded, slightly projecting. Anterior rim near vertical, posterior rim slightly inclined towards ventral; posterior rim with weak central concavity. Ventral rim straight, horizontal. Preventral and postventral angles rounded. All rims irregularly undulating or, rarely, crenulated (Fig. 27 m). Inner face flat in vertical direction and slightly bent in horizontal direction. Sulcus long, relatively narrow, with relatively low ostial lobe; cauda straight, with rounded or tapering tip, slightly deepened; sulcus inclination angle 15 – 20 °; OL: SuL = 1.55 – 1.75. Subcaudal iugum moderately long, not reaching posterior tip of cauda, narrow. Dorsal depression small to moderate in size, with indistinct margins. Ventral furrow distinct, very close to ventral rim of otolith and relatively regularly curved. Outer face with slight ventral umbo, moderately ornamented.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF91FFDF0236FD3063E4FB7D.taxon	discussion	Discussion Microgobius verecundus is an inconspicuous species that is more compressed than M. cantaurensis (OL: OH = 0.86 – 0.9 vs 0.95 – 0.97) and is distinguished from M. microlepis by the larger size of the sulcus (OL: SuL = 1.55 – 1.75 vs 1.8 – 1.9) and its different shape. It is a common and widespread species during the Tortonian from Ecuador to Trinidad.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF90FFDF018CFA966063F8DE.taxon	description	Figure 28 a – c Material 6 specimens Gelasian, Moin FM, Atlantic Costa Rica, PPP 1773 (figured specimens NMB P 15659 - 60). Discussion Te otoliths of M. carri are moderately compressed (OL: OH = 0.85 – 0.87) and characterized by an inclined predorsal rim and pronounced, high postdorsal angle. Anterior and posterior rims are almost vertical. Another typical feature is the long sulcus (OL: SuL = 1.35 – 1.45), with a wide ostium and rounded ostial lobe, and a sulcus inclination angle of 15 – 20 °. Today, M. carri occurs in moderately deep water (6 to 21 m) in the vicinity of coral reefs (Birdsong, 1981). Te only identified fossil occurrence in the Early Pleistocene of Costa Rica is consistent with the extant distribution pattern.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF90FFD90236FF5663FDFE98.taxon	description	Figure 28 e – h Holotype NMB P 15661 (Fig. 28 e), Tortonian, Tuira FM, Pacific Panama, Darien, PPP 1163. Paratypes 6 specimens Tortonian: 5 specimens Pacific Panama, Darien: 4 specimens Tuira FM, 2 specimens PPP 1593 (NMB P 15662 - 63), 1 specimen PPP 1154 (NMB P 15664), 1 specimen PPP 1137 (NMB P 15666); 1 specimen Yaviza FM, PPP 1565 (NMB P 15667); 1 specimen Manzanilla FM, San José Member, Trinidad, PPP 2674 (NMB P 15665). Referred specimen 10 specimens Tortonian: 7 specimens Tuira FM, Pacific Panama, Darien, 1 specimen PPP 1137, 6 specimens PPP 1163; 3 specimens Gatun FM, Atlantic Panama, PPP 2165. Etymology Named after the Chocó Indians living in the Chocó Province in Darien, Panama. Diagnosis Otolith size up to 1.4 mm in length. OL: OH = 0.75 – 0.8. Otolith shape high-bodied; dorsal rim high, with weak postdorsal projection. Sulcus moderately long and wide, with relatively broad ostial lobe; OL: SuL = 1.45 – 1.65; sulcus inclination angle 16 – 18 °. Subcaudal iugum long, narrow. Ventral furrow moderately close to ventral rim of otolith, regularly curved. Description Otolith size up to 1.4 mm in length (holotype 1.35 mm). OH: OT = 3.5 – 4.3. Otolith shape high-bodied, with high dorsal rim and prominent middorsal angle. Predorsal angle rounded; postdorsal region rounded, not projecting. Anterior and posterior rims near vertical; posterior rim with weak central concavity. Ventral rim slightly curved, horizontal. Preventral and postventral angles rounded. All rims smooth, dorsal rims sometimes undulating or crenulated (Fig. 28 f). Inner face flat in vertical direction and slightly bent in horizontal direction. Sulcus moderately long and wide, with broad ostial lobe; cauda straight, with rounded tip, not particularly deepened; sulcus inclination angle 16 – 18 °; OL: SuL = 1.45 – 1.65. Subcaudal iugum long, narrow, may expand over posterior tip of cauda in juvenile specimens (Fig. 28 h). Dorsal depression moderate in size, with indistinct margins. Ventral furrow distinct, moderately close to ventral rim of otolith and regularly curved. Outer face with slight ventral umbo, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF90FFD90236FF5663FDFE98.taxon	discussion	Discussion Microgobius chocorum resembles the extant M. carri and M. crocatus. It differs from M. carri in the more high-bodied shape (OL: OH = 0.75 – 0.8 vs 0.85 – 0.87) and from M. crocatus in the distinct middorsal angle (vs regularly curved) and in being more compressed (OL: OH = 0.75 – 0.8 vs 0.83 – 0.88). Otoliths of the extant M. meeki (Fig. 28 i) and M. signatus (Fig. 28 d) are similarly compressed, as far as can be judged from the available, somewhat formalin-eroded specimens. Otoliths of M. meeki differ from M. chocorum in the high postdorsal angle and the steeply inclined sulcus (25 ° vs 16 – 18 °) and those of M. signatus in the small sulcus (OL: SuL = 1.8 vs 1.45 – 1.65) and the low sulcus inclination angle (about 10 ° vs 16 – 18 °).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF96FFD9018CFEF063E3FBD8.taxon	description	Figure 28 j – o Material 20 specimens Tortonian, Pacific Panama, Darien: 17 specimens Tuira FM, PPP 1137 (figured specimens NMB P 15668 - 70); 3 specimens Yaviza FM, 1 specimen PPP 1609, 2 specimens PPP 1528. Discussion Te otoliths of M. crocatus are relatively unremarkable and are best recognized by the regularly curved, high, and evenly crenulated or undulating dorsal rim that distinguishes them from otoliths of M. carri and M. chocorum. Microgobius crocatus occurs today in mangrove environments in the tropical East Pacific (Birsdong, 1981). Its fossil occurrence is rare, concentrated in a single location in the Tortonian of Darien, Río Tuquesa (PPP 1137), which has been described as “ clayey siltstone and bioclastic and lithic sandstone, rich in bivalves, especially Pecten ” (Coates et al., 2004). Pecten clams are not specific or even common in mangrove environments, and the cause for the singular abundance of M. crocatus in that location is therefore unknown. Moreover, M. crocatus represents the earliest extant goby species identified by otoliths in tropical America.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF96FFD9018CFB306641FA18.taxon	description	Figure 29 a – g Holotype NMB P 15671 (Fig. 29 b), Calabrian, Cumaná FM, Venezuela, PPP 3038. Paratypes 7 specimens same data as holotype (NMB P 15672 - 78). Referred specimen 43 specimens same data as holotype. Etymology Named after the type formation, the Cumaná FM. Diagnosis Otolith size up to 1.9 mm in length. OL: OH = 0.79 – 0.93. Otolith shape high-bodied; dorsal rim high, with angular, high predorsal angle, obtuse middorsal angle and short but angular to pointed postdorsal projection. Sulcus relatively short and narrow, with low ostial lobe and tapering caudal tip; OL: SuL = 1.7 – 1.8; sulcus inclination angle 13 – 17 °. Subcaudal iugum long, narrow. Ventral furrow close to ventral rim of otolith, regularly curved. Description Otolith size up to 1.9 mm in length (holotype 1.4 mm). OL: OH = 0.79 – 0.93, mostly 0.79 – 0.87, but largest specimen 0.93 (Fig. 29 a); OH: OT = 3.4 – 4.0. Otolith shape high-bodied. Dorsal rim with high, angular to slightly rounded predorsal angle, obtuse middorsal angle and short but angular or pointed postdorsal projection. Anterior rim vertical, with broad, shallow concavity at level of ostium; posterior rim vertical to slightly inclined and broadly concave. Ventral rim flat to slightly curved, horizontal. Preventral angle orthogonal, postventral angle rounded. All rims smooth, except dorsal rim intensely lobed in largest specimen (Fig. 29 a). Inner face flat in vertical direction and slightly bent in horizontal direction. Sulcus moderately long and wide, with broad ostial lobe; cauda straight, with rounded tip, not particularly deepened; sulcus inclination angle 13 – 17 °; OL: SuL = 1.7 – 1.8. Subcaudal iugum long, narrow, may expand over posterior tip of cauda in juvenile specimens (Fig. 28 h). Dorsal depression moderate in size, with indistinct margins. Ventral furrow distinct, moderately close to ventral rim of otolith and regularly curved. Outer face with slight ventral umbo, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF96FFD9018CFB306641FA18.taxon	discussion	Discussion Microgobius cumana is recognized by the high-bodied otolith with its characteristic shape of the dorsal rim, the relatively short sulcus (OL: SuL = 1.7 – 1.8), the low ostial lobe and tapering caudal tip. In the latter two characters it resembles otoliths of M. microlepis with which it occurs parallel at the same location, but differs in the specific shape of the dorsal rim and the more compressed appearance (OL: OH = 0.79 – 0.87, rarely 0.93 vs 0.85 – 0.92) and the more steeply inclined sulcus (13 – 17 ° vs 9 – 11 °). Microgobius cumana also resembles the otoliths of extant M. thalassinus (Fig. 29 h – i) from the West Atlantic and M. brevispinis (according to drawings provided by D. Nolf) from the East Pacific, but differs in the specific shape of the dorsal rim and the broadly concave posterior rim (vs vertical).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF96FFDB0236FA7060F9F8B8.taxon	description	Figure 29 j – q Holotype NMB P 15679 (Fig. 29 l), Tortonian, Angostura FM, Ecuador, NMB 19301. Paratypes 7 specimens Tortonian, Angostura FM, Ecuador: 1 specimen PPP 3369 (NMB P 15680), 2 specimens PPP 3444 (NMB P 15681 - 82), 4 specimens PPP 3301 (NMB P 15683 - 86). Referred specimen 95 specimens Tortonian: 94 specimens Angostura FM, Ecuador: 3 specimens NMB 19301, 28 specimens PPP 3297, 51 specimens PPP 3301, 10 specimens PPP 3369, 3 specimens PPP 3372, 1 specimen PPP 3444; 1 specimen Tuira FM, Pacific Panama, Darien, PPP 1132. Etymology Named after Ecuador, the country from which the type specimens originate. Diagnosis Otolith size up to 2.0 mm in length. OL: OH = 0.78 – 0.9. Otolith shape high-bodied; dorsal rim high, regularly crenulated, with rounded, obtuse middorsal angle and short rounded postdorsal projection. Sulcus moderately large and wide, with broad ostial lobe and rounded caudal tip; OL: SuL = 1.5 – 1.7; sulcus inclination angle 15 – 20 °. Subcaudal iugum long, moderately wide. Ventral furrow distant from ventral rim of otolith, regularly curved. Description Otolith size up to 2.0 mm in length (holotype 1.75 mm). OH: OT = 3.9 – 4.2. Otolith shape high-bodied. Dorsal rim high, regularly crenulated, with broadly rounded predorsal angle, obtuse to rounded middorsal angle and short broadly rounded postdorsal projection. Anterior rim near vertical or slightly inclined towards dorsal; posterior rim vertical to slightly inclined and with indistinct central concavity. Ventral rim flat to slightly curved, horizontal. Preventral angle orthogonal, postventral angle rounded. All rims slightly undulating and dorsal rim regularly crenulated. Inner face flat in vertical direction and slightly bent in horizontal direction. Sulcus moderately long and wide, with broad ostial lobe; cauda straight, with rounded tip, slightly deepened; sulcus inclination angle 15 – 20 °; OL: SuL = 1.5 – 1.7. Subcaudal iugum long, moderately wide, may expand over posterior tip of cauda in juvenile specimens (Fig. 29 q). Dorsal depression moderate in size, with indistinct margins. Ventral furrow distinct, distant ventral rim of otolith and regularly curved. Outer face with slight ventral umbo, slightly ornamented.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF96FFDB0236FA7060F9F8B8.taxon	discussion	Discussion Microgobius ecuadorensis is characterized by its broadly crenulated dorsal rim, which distinguishes it from most of its congeners except for M. crocatus and M. verecundus. It differs from M. crocatus in the higher and less regularly curved dorsal rim and the shorter sulcus (OL: SuL = 1.5 – 1.7 vs 1.4 – 1.55). Microgobius ecuadorensis differs from the parallel occurring M. verecundus in being mostly more compressed (OL: OH = 0.78 – 0.9 vs 0.86 – 0.9) and thinner (OH: OT = 3.9 – 4.2 vs 3.5 – 3.8) and the usually more regularly developed crenulation of the dorsal rim.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF94FFDB0236FF50673FF9F8.taxon	description	Figure 28 p 2014 Gobiidae, Genera and species indet. 3 — Aguilera et al.: Fig. 12 / 10. Holotype MPEG- 1947 - V (Fig. 28 p), Aquitanian to Burdigalian, Pirabas FM, Brazil. Etymology Named after the type formation, the Pirabas FM. Diagnosis Otolith size 1.45 mm in length. OL: OH = 0.88. Otolith shape nearly rectangular; dorsal rim shallow, nearly flat and horizontal. Sulcus moderately long and narrow, with relatively low ostial lobe; OL: SuL = 1.45; sulcus inclination angle 16 °. Subcaudal iugum long, narrow. Ventral furrow close to ventral rim of otolith. Description Otolith size 1.45 mm in length (holotype). OH: OT = 4.4. Otolith shape nearly rectangular, with low, nearly flat and horizontal dorsal rim and high, rounded predorsal angle and orthogonal postdorsal projection. Anterior and posterior rims near vertical; posterior rim with very weak central concavity. Ventral rim slightly relatively straight, horizontal. Preventral and postventral angles rounded orthogonal. All rims slightly undulating. Inner face almost flat in vertical direction and slightly bent in horizontal direction. Sulcus moderately long and narrow, with relatively low and broad ostial lobe; cauda straight, with rounded tip, not particularly deepened; sulcus inclination angle 16 °; OL: SuL = 1.45. Subcaudal iugum long, narrow. Dorsal depression not discernable. Ventral furrow distinct, parallel and close to ventral rim of otolith. Outer face relatively flat and smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF94FFDB0236FF50673FF9F8.taxon	discussion	Discussion Microgobius pirabasensis is recognized by its nearly flat, horizontal dorsal rim resulting in an almost perfect compressed rectangular otolith shape. Other characteristical features like the low ostial lobe and the ventral furrow running close and parallel to the ventral rim of the otolith are considered plesiomorphic.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF94FFA40236F9D061CBFC18.taxon	description	Figure 30 a – g 2014 Gobiidae, Genera and species indet. 2 — Aguilera et al.: Fig. 12 / 8 – 12 / 9. Holotype NMB P 15687 (Fig. 30 b), Burdigalian, Cantaure FM, Venezuela, PPP 2539. Paratypes 12 specimens: 8 specimens Burdigalian, Cantaure FM, Venezuela: 5 specimens same data as holotype (NMB P 15688 - 92), 3 specimens PPP 2545 (NMB P 15693 - 95). 3 specimens Burdigalian, Balboa FM, Dominican Republic, NMB 17290 (NMB P 400). 1 specimen Aquitanian to Burdigalian, Pirabas FM, Brazil (MPEG- 1946 - V). Referred specimen 238 specimens: 233 specimens Burdigalian, Cantaure FM, Venezuela: 87 specimens same data as holotype, 4 specimens PPP 2543, 135 specimens PPP 2545, 10 specimens PPP 2546. 1 specimen Burdigalian, Brasso FM, Trinidad KR 6666. 1 specimen Aquitanian to Burdigalian, Pirabas FM, Brazil. Etymology Named in honor of Ross Robertson (Panama City, Republic of Panama) for his contribution to the knowledge of the fishes of the tropical East Pacific. Diagnosis Otolith size up to 1.8 mm in length. OL: OH = 0.83 – 0.88. Otolith shape high-bodied; dorsal rim high, relatively smooth, with rounded, obtuse postdorsal angle and short rounded postdorsal projection. Anterior rim ventrally expanded below ostium. Sulcus moderately large and wide, with broad ostial lobe and rounded caudal tip; OL: SuL = 1.4 – 1.5; sulcus inclination angle 15 – 22 °. Subcaudal iugum moderately long, narrow. Ventral furrow close to ventral rim of otolith, regularly curved. Description Otolith size up to 1.8 mm in length (holotype). OH: OT = 3.6 – 3.9. Otolith shape high-bodied. Dorsal rim high, regularly curved, with broadly rounded predorsal angle and broadly rounded middorsal angle, and short broadly rounded postdorsal projection. Anterior rim slightly inclined towards dorsal, with bulge or rounded projection ventral of ostial tip in specimens larger than 1.5 mm in length (Fig. 30 a, b, g); posterior rim vertical to slightly inclined and with indistinct central concavity. Ventral rim flat to slightly curved, horizontal. Preventral angle recessed below bulge of anterior rim, postventral angle rounded. All rims smooth to slightly undulating. Inner face nearly flat in vertical direction and slightly bent in horizontal direction. Sulcus moderately large and wide, with broad ostial lobe; cauda straight, with rounded tip, slightly deepened; sulcus inclination angle 15 – 22 °; OL: SuL = 1.4 – 1.5. Subcaudal iugum moderately long, narrow. Dorsal depression small, with indistinct margins. Ventral furrow distinct, close ventral rim of otolith and regularly curved. Outer face nearly flat or with slight ventral umbo, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFF94FFA40236F9D061CBFC18.taxon	discussion	Discussion Microgobius robertsoni is characterized by the rather smooth otolith rims and, in larger otoliths longer than 1.5 mm, the bulge at the anterior rim below the ostium. Tese are also the characters distinguishing M. robertsoni from M. ecuadorensis. Microgobius robertsoni differs from the parallel occurring M. cantaurensis in the more compressed shape (OL: OH = 0.83 – 0.88 vs 0.95 – 0.97) and steeper inclined sulcus (15 – 22 ° vs 12 – 15 °), and from M. verecundus in the larger sulcus (OL: SuL 1.4 – 1.5 vs 1.55 – 1.75).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFEBFFA60236FC706011FABF.taxon	description	Figure 30 h – l Holotype NMB P 15696 (Fig. 30 h), Burdigalian, Brasso FM, Trinidad, KR- 6666. Paratypes 5 specimens same data as holotype (NMB P 15697 - 701). Referred specimen 45 specimens same data as holotype. Etymology Named in memory of Karl Rohr, who has conducted important field reconnaissance work in Trinidad including the collecting of the otoliths of Microgobius rohri. Diagnosis Otolith size up to 1.6 mm in length. OL: OH = 0.83 – 0.9. Otolith shape high-bodied; dorsal rim high, relatively smooth, with rounded and broadly (See figure on next page.) Fig. 30 a – g Microgobius robertsoni n. sp.; b holotype, NMB P 15687, Cantaure FM, Paraguaná, Venezuela, PPP 2539; a, c – g paratypes; a, g same data as holotype, NMB P 15688 - 89; c – f Cantaure FM, Paraguaná, Venezuela, PPP 2545, NMB P 15693 - 95; e MPEG- 1946 - V (refigured from Aguilera et al., 2014, Fig. 12 / 9), Pirabas FM, Brazil. h – l Microgobius rohri n. sp., Brasso FM, Trinidad, KR- 6666; h holotype, NMB P 15696; i – l paratypes, NMB P 15697 - 15700. m Microgobius curtus Ginsburg, 1939, extant, USNM 293571, 08 ° 26 ’ N 78 ° 06 ’ W. n Microgobius erectus Ginsburg, 1938, extant, USNM 322733, Pacific Panama, 07 ° 25 ’ N 80 ° 11 ’ W, SL 55 mm. o – t Microgobius pezoldi n. sp.; p holotype, NMB P 15702, Chucunaque FM, Darien, Panama, PPP 1578; o, q – t paratypes; o Angostura FM, Ecuador, NMB 19301, NMB P 15703; q Tuira FM, Darien, Panama, PPP 1132, NMB P 15704; r Manzanilla FM, Trinidad, PPP 2674, NMB P 15705; s – t Manzanilla FM, Trinidad, PPP 2671, NMB P 15706 - 07 expanded postdorsal angle and short blunt postdorsal projection. Preventral and postventral angles projecting. Sulcus small, wide, with rounded ostial lobe and rounded caudal tip; OL: SuL = 1.7 – 1.9; sulcus inclination angle 8 – 12 °. Subcaudal iugum short, narrow. Ventral furrow distant from ventral rim of otolith, regularly curved. Outer face nearly flat. Description Otolith size up to 1.6 mm in length (holotype 1.5 mm). OH: OT = 3.9 – 4.4. Otolith shape high-bodied. Dorsal rim high, with rounded predorsal angle and rounded, broadly expanded postdorsal angle and short blunt postdorsal projection. Anterior and posterior rims slightly inclined towards dorsal, and with mild central concavities. Ventral rim flat to slightly curved, horizontal. Preventral and postventral angles protruding resulting distance between them clearly marking longest axis. All rims smooth or slightly and irregularly undulating. Inner face nearly flat in vertical direction and slightly bent in horizontal direction. Sulcus small and wide, with rounded ostial lobe; cauda straight, with rounded tip, slightly deepened; sulcus inclination angle 8 – 12 °; OL: SuL = 1.7 – 1.9. Subcaudal iugum short, not reaching caudal tip, narrow. Dorsal depression broad, with distinct ventral margin and broad crista superior. Ventral furrow distinct, relatively distant from ventral rim of otolith and regularly curved. Outer face nearly flat, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFEBFFA60236FC706011FABF.taxon	discussion	Discussion Microgobius rohri is easily distinguished from its congeners by the protruding preventral and postventral angles, the small sulcus and its low inclination angle. It likely represents an extinct lineage and could alternatively be placed in a separate genus.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE9FFA6018CFA90637DF8F8.taxon	description	Figure 31 x Material 2 specimens Calabrian, Armuelles FM, Pacific Panama, PPP 3236 (NMB P 15738). Discussion Two large and rather well-preserved otoliths from the upper Lower Pleistocene Armuelles FM of Pacific Panama resemble the Early Miocene Microgobius robertsoni in the presence of a bulge on the anterior rim just below the ostial tip. Tey differ, however, in the short subcaudal iugum terminating far from the posterior tip of the cauda and the short sulcus (OL: OH = 1.75 vs 1.4 – 1.5). Tis find indicates that this specific otolith morphology may be present in extant Microgobius species in the tropical East Pacific.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE9FFA70236FE9063AFFB98.taxon	description	Figure 30 o – t Holotype NMB P 15702 (Fig. 30 p), Messinian, Chucunaque FM, Pacific Panama, Darien, PPP 1578. Paratypes 7 specimens Tortonian: 1 specimen Angostura FM, Ecuador NMB 19301 (NMB P 15703); 3 specimens Tuira FM, Pacific Panama, Darien, 1 specimen PPP 1132 (NMB P 15704), 2 specimens PPP 1163 (NMB P 15708 – 09); 3 specimens Manzanilla FM, San José Member, Trinidad, 1 specimens PPP 2674 (NMB P 15705), 2 specimens PPP 2671 (NMB P 15706 – 07). Referred specimen 19 specimens: 16 specimens Tortonian: 3 specimens Angostura FM, Ecuador, 2 specimens NMB 19301, 1 specimen NMB 19081; 7 specimens Tuira FM, Pacific Panama, Darien, 1 specimen PPP 1137, 3 specimens PPP 1139, 1 specimen PPP 1593, 2 specimens PPP 1164; 1 specimens Yaviza FM, Pacific Panama, Darien, PPP 1171; 5 specimens Manzanilla FM, San José Member, 1 specimen PPP 2669, 1 specimen PPP 2670, 2 specimens PPP 2671, 1 specimen PPP 2676. 2 specimens Messinian, Chucunaque FM, Pacific Panama, Darien, PPP 1576. 1 specimen Piacenzian, Río Banano FM, Atlantic Costa Rica, PPP 1732. Etymology Named in honor of Frank Pezold (Corpus Christi, Texas), in recognition of his distribution to the knowledge of extant gobies from tropical America. Diagnosis Otolith size up to 1.75 mm in length. OL: OH = 0.85 – 0.93. Otolith shape high-bodied; dorsal rim shallow, broadly crenulated; predorsal angle protruding and on same level as slightly protruding postdorsal projection. Preventral and postventral angles rounded orthogonal. Sulcus long, narrow, with low ostial lobe and rounded caudal tip; OL: SuL = 1.45 – 1.65; sulcus inclination angle 11 – 15 °. Subcaudal iugum long, moderately narrow. Ventral furrow parallel and close to ventral rim of otolith. Description Otolith size up to 1.75 mm in length (holotype 1.2 mm). OH: OT = 3.0 – 3.2. Otolith shape high-bodied. Dorsal rim shallow, nearly flat, without prominent postdorsal angle, smooth or broadly crenulated; predorsal angle high, distinct, slightly protruding at same level as slightly protruding postdorsal projection. Longest horizontal axis of otolith shifted dorsally to predorsal angle – postdorsal projection level. Anterior and posterior rims slightly inclined towards ventral, straight, without marked concavities. Ventral rim flat to slightly curved, horizontal. Preventral and postventral angles recessed, rounded. All rims smooth except dorsal rim. Inner face nearly flat in vertical direction and slightly bent in horizontal direction. Sulcus long, narrow, with low ostial lobe; cauda straight, with rounded tip, slightly deepened; sulcus inclination angle 11 – 15 °; OL: SuL = 1.45 – 1.65. Subcaudal iugum long, moderately narrow. Dorsal depression indistinct. Ventral furrow distinct, relatively close and parallel to ventral rim of otolith. Outer face distinctly convex, with inframedian umbo, relatively smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE9FFA70236FE9063AFFB98.taxon	discussion	Discussion Microgobius pezoldi is a typical member of the M. erectus plexus, which is characterized by the slightly protruding predorsal angle and postdorsal projection, resulting in a dorsal shift of the longest axis of the otolith. It resembles M. curtus (Fig. 30 m) but tends to be less compressed (OL: OH = 0.85 – 0.93 vs 0.84). It is also similar to the extant M. erectus but has a more pronounced ostial lobe. Te shape of the postdorsal projection distinguishes it from both extant congeners. Microgobius pezoldi is a widely distributed species from the Tortonian of Ecuador to Trinidad and the Piacenzian of Costa Rica but is nowhere common.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE8FFA7018CFB30665EF898.taxon	description	Figure 31 a – e Holotype NMB P 15710 (Fig. 31 a), Zanclean, Cubagua FM, Venezuela, PPP 2549. Paratypes 5 specimens Zanclean, Cubagua FM, Venezuela, 1 specimen PPP 2568 (NMB P 15711), 2 specimens PPP 3057 (NMB P 15712 - 13), 1 specimen PPP 3066 (NMB P 15714), 1 specimen PPP 2549 (NMB P 15715). Referred specimen 71 specimens Zanclean: 54 specimens Cubagua FM, Venezuela, 5 specimens PPP 2549, 1 specimen PPP 2556, 6 specimens PPP 2557, 8 specimens PPP 3055, 7 specimens PPP 3057, 16 specimens PPP 2568, 4 specimens PPP 2570, 7 specimens PPP 2571; 7 specimens Cayo Agua FM, Atlantic Panama, PPP 1188; 9 specimens Springvale FM, Trinidad, KR- 1216; 1 specimen La Vela FM, Venezuela, PPP 2532. Etymology Named after the resemblance to otoliths of the northeastern Atlantic and Mediterranean genus Aphia. Diagnosis Otolith size up to 1.25 mm in length. OL: OH = 0.8 – 0.85. Otolith shape high-bodied; dorsal rim elevated; predorsal angle and postdorsal projection rounded, not protruding. Preventral angle orthogonal, postventral angle rounded. Sulcus short, narrow, with broad, rounded ostial lobe and cauda posteriorly reduced by ingressing subcaudal iugum; OL: SuL = 1.7 – 2.0; sulcus inclination angle 13 – 18 °. Subcaudal iugum long, wide, turning around caudal tip. Ventral furrow moderately close to ventral rim of otolith, regularly curved. Description Otolith size up to 1.25 mm in length (holotype 1.0 mm). OH: OT = 3.2 – 3.5. Otolith shape high-bodied. Dorsal rim high, often with obtuse, rounded middorsal angle; predorsal angle high, broadly rounded, not protruding, postdorsal projection rounded or blunt, not protruding. Anterior and posterior rims near vertical and relatively straight. Ventral rim flat to slightly curved, horizontal. Preventral angle orthogonal, postventral angle rounded. All rims smooth. Inner face completely flat. Sulcus small, short, narrow, with broad, rounded to triangular ostial lobe; cauda straight, short, with rounded or tapering tip; sulcus inclination angle 13 – 18 °; OL: SuL = 1.7 – 2.0. Subcaudal iugum long, wide, turning upward posteriorly and then ingressing into posterior tip of cauda to various extends. Dorsal depression indistinct. Ventral furrow distinct, moderately close to ventral rim of otolith and regularly curved. Outer face evenly convex, smooth. Discussion Te otoliths of Microgobius aphioides are smaller than most Microgobius otoliths and could be confused with otoliths of juveniles of other species. Te characteristic ingression of the subcaudal iugum into the rear part of the cauda is rarely observed in otoliths of juveniles of other species, but the shape of the sulcus with its low, rounded to triangular ostial lobe and the relatively wide subcaudal iugum distinguish M. aphioides from such otoliths of juveniles. Te smooth otolith rims and nearly vertical anterior and posterior rims distinguish M. aphioides from other species in the Microgobius camur plexus. Te ingression of the subcaudal iugum into the rear part of the cauda is stronger in M. aphioides than in M. glaber, resulting in a shorter sulcus (OL: SuL = 1.7 – 2.0 vs 1.5 – 1.6). Microgobius aphioides differs from M. camur in the relatively narrow ostium and the smooth otolith rims. Most similar morphologically are the otoliths of the small species M. praeglaber, which, however, is distinguished by its eccentrically forward-positioned sulcus and the curved ventral rim.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFEFFFA0018CFF506650FB38.taxon	description	Figure 31 f – k Holotype NMB P 15716 (Fig. 31 f), Gelasian, Escudo de Veraguas FM, Atlantic Panama, PPP 2171. Paratypes 5 specimens: 1 specimen Zanclean, Cayo Agua FM, Atlantic Panama, PPP 1275 (NMB P 15720). 1 specimen Piacenzian, Escudo de Veraguas FM, Atlantic Panama, PPP 1241 (NMB P 15718). 2 specimens Gelasian: 1 specimen Escudo de Veraguas FM, Atlantic Panama, PPP 2170 (NMB P 15717); 1 specimen Bastimentos FM, Ground Creek Unit, Atlantic Panama, PPP 1252 (NMB P 15719). 1 specimen Calabrian, Swan Cay FM, Atlantic Panama, PPP 2221 (NMB P 15721). Referred specimen 8 specimens: 2 specimens Piacenzian: 1 specimen Río Banano FM, Atlantic Costa Rica, PPP 3243; 1 specimen Escudo de Veraguas FM, Atlantic Panama, PPP 1241. 1 specimen Gelasian, Bastimentos FM, Ground Creek Unit, Atlantic Panama, PPP 1252. 5 specimens Calabrian: 1 specimen Bastimentos FM, Ground Creek Unit, Atlantic Panama, PPP 2249; 4 specimens Swan Cay FM, Atlantic Panama, PPP 2221. Etymology From camur (Latin) = hooked, curved, referring to the hooked shape of the subcaudal iugum. Diagnosis Otolith size up to 1.65 mm in length. OL: OH = 0.76 – 0.83. Otolith shape high-bodied; dorsal rim elevated, regularly crenulated; predorsal angle and postdorsal projection not protruding. Preventral and postventral angles rounded. Sulcus short, wide, with high, triangular ostial lobe and cauda posteriorly shortened by ingressing subcaudal iugum; OL: SuL = 1.65 – 2.05; sulcus inclination angle 15 – 20 °. Subcaudal iugum long, wide, hook-like extended into rear part of cauda. Ventral furrow close and parallel to ventral rim of otolith, regularly curved. Description Otolith size up to 1.65 mm in length (holotype). OH: OT = 3.3 – 4.0, decreasing with size. Otolith shape high-bodied. Dorsal rim high, rounded or with rounded middorsal angle; predorsal angle indistinct, postdorsal projection blunt, not protruding. Anterior rim nearly vertical, posterior rim slightly inclined toward ventral and straight. Ventral rim flat to slightly curved, horizontal. Preventral and postventral angles rounded. All rims more or less strongly crenulated or undulating. Inner face completely flat. Sulcus short, wide, particularly broad, triangular ostial lobe widened; cauda short, with rounded or tapering tip; sulcus inclination angle 15 – 20 °; OL: SuL very variable, depending on degree of ingression of subcaudal iugum, 1.65 – 2.05. Subcaudal iugum long, wide, usually ingressing into posterior portion of cauda to various extends: ranging from very strong and considerably shortening cauda (Fig. 31 f, g, j) to moderately strong ingressing (Fig. 31 i, k), however, almost not ingressing in rare cases (Fig. 31 h). Dorsal depression indistinct, wide. Ventral furrow mostly distinct, close and parallel to ventral rim of otolith and regularly curved. Outer face evenly convex, variably ornamented.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFEFFFA0018CFF506650FB38.taxon	discussion	Discussion Microgobius camur is the largest species in this otolith plexus and readily recognized by the combination of the intense crenulation or undulation of its rims, particularly the dorsal rim, the high triangular ostial lobe resulting in a wide sulcus and the large subcaudal iugum, which often very strongly ingresses into the rear part of the cauda thereby considerably shortening the length of the sulcus.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFEFFFA20236FB1063AAF8B8.taxon	description	Figure 31 l – p Holotype NMB P 15722 (Fig. 31 l), Messinian, Manzanilla FM, Montserrat Member, Trinidad, PPP 2666. Paratypes 9 specimens: 6 specimens Messinian, same data as holotype (NMB P 15723 - 28). 3 specimens Tortonian, Cercado FM, Dominican Republic, NMB 17003 (NMB P 392). (See figure on next page.) Fig. 31 a – e Microgobius aphioides n. sp., Cubagua FM, Venezuela; a holotype, NMB P 15710, PPP 2549; b – e paratypes; b PPP 2568, NMB P 15711; c PPP 3057, NMB P 15712; d PPP 3066, NMB P 15714; e PPP 2549, NMB P 15715. f – k Microgobius camur n. sp.; f holotype, NMB P 15716, Escudo de Veraguas FM, Bocas del Toro, Panama, PPP 2171; g – k paratypes; g Escudo de Veraguas FM, Bocas del Toro, Panama, PPP 2170, NMB P 15717; h Escudo de Veraguas FM, Bocas del Toro, Panama, PPP 1241, NMB P 15718; i Bastimentos FM, Ground Creek Unit, Bastimentos Island, Panama, PPP 1252, NMB P 15719; j Cayo Agua FM, Atlantic Panama, PPP 1276, NMB P 15720; k Swan Cay FM, Atlantic Panama, PPP 2221, NMB P 15721. l – p Microgobius glaber n. sp.; l holotype, NMB P 15722, Manzanilla FM, Trinidad, PPP 2666; m – p paratypes; m – o Cercado FM, Dominican Republic, NMB 17003, NMB P 392; p Manzanilla FM, Trinidad, PPP 2666, NMB P 15723. q – v Microgobius praeglaber n. sp.; q holotype, NMB P 15729, upper Brasso FM, Trinidad, KR- 9244; r – v paratypes; r – t upper Brasso FM, Trinidad, KR- 9244, NMB P 15730 - 32; u – v Gatun FM, Colon, Panama, PPP 2167, NMB P 15734 - 35. W Microgobius sp. 2, Armuelles FM, Pacific Panama, PPP 3236, NMB P 15737. X Microgobius sp. 1, Armuelles FM, Pacific Panama, PPP 3235, NMB P 15738 Referred specimen 50 specimens: 31 specimens Tortonian: 2 specimens Tuira FM, Pacific Panama, Darien, PPP 1134; 7 specimens Yaviza FM, Pacific Panama, Darien, PPP 1528; 10 specimens Gatun FM, Atlantic Panama, 2 specimens PPP 2165, 8 specimens PPP 2168; 5 specimens Manzanilla FM, San José Member, Trinidad, 1 specimen PPP 2671, 1 specimen PPP 2674, 3 specimens PPP 2676; 7 specimens Cercado FM, Dominican Republic, NMB 17003. 19 specimens Messinian, same data as holotype. Etymology From glaber (Latin) = smooth, referring to the smooth outline and surface of these otoliths. Diagnosis Otolith size up to 1.15 mm in length. OL: OH = 0.83 – 0.9. Otolith shape high-bodied; dorsal rim elevated, with rounded postdorsal angle; predorsal angle and postdorsal projection not protruding. Preventral and postventral angles rounded. Sulcus long, narrow, with moderately wide, angular ostial lobe and very narrow cauda; OL: SuL = 1.5 – 1.6; sulcus inclination angle 13 – 23 °. Subcaudal iugum long, narrow, slightly bent around rear part of cauda and ingressing cauda from ventral. Ventral furrow close and parallel to ventral rim of otolith, regularly curved. Inner and outer faces smoothened. Description Otolith size up to 1.15 mm in length (holotype 1.05 mm). OH: OT = 2.9 – 3.4. Otolith shape high-bodied. Dorsal rim high, rounded or with rounded postdorsal angle; predorsal angle broadly rounded, postdorsal projection blunt, not protruding. Anterior and posterior rims nearly vertical, nearly straight. Ventral rim slightly curved, horizontal. Preventral and postventral angles rounded. All rims smooth. Inner face flat in vertical direction and slightly bent in horizontal direction. Sulcus long, narrow, with moderately wide, angular ostial lobe and very narrow cauda, with tapering tip; sulcus inclination angle very variable, 13 – 23 °; OL: SuL = 1.5 – 1.6. Subcaudal iugum long, narrow, seemingly ingressing cauda ventrally, extending as narrow feature around caudal tip and sometimes also extending anteriorly below ostium. Dorsal depression wide, with distinct crista superior towards sulcus. Ventral furrow distinct, close and parallel to ventral rim of otolith and regularly curved. Outer face evenly convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFEFFFA20236FB1063AAF8B8.taxon	discussion	Discussion Microgobius glaber differs from the other species in the Microgobius camur plexus in the combination of the smooth rims and inner and outer faces and the unusually narrow cauda.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFEDFFA3018CF890601FFED8.taxon	description	Figure 31 q – v Holotype NMB P 15729 (Fig. 31 q), Langhian to Serravallian, upper Brasso FM, Trinidad, KR- 9244. Paratypes 7 specimens: 4 specimens Langhian to Serravallian, same data as holotype (NMB P 15730 - 33). 3 specimens Tortonian, Gatun FM, Atlantic Panama, PPP 2167 (NMB P 15734 - 36). Referred specimen 6 specimens Langhian to Serravallian, same data as holotype. Etymology A combination of prae (Latin) = before, earlier and the species-name glaber of Microgobius glaber indicating its possible relationship to that species and its earlier occurrence. Diagnosis Otolith size up to 0.85 mm in length. OL: OH = 0.79 – 0.83. Otolith shape high-bodied; dorsal rim elevated, with rounded postdorsal angle. Anterior rim slightly curved; posterior rim vertical, straight. Ventral rim curved. Sulcus short, anteriorly widened, posteriorly tapering, eccentrically forward positioned; OL: SuL = 2.0 – 2.2; sulcus inclination angle 12 – 16 °. Subcaudal iugum long, wide, ingressing into rear part of cauda. Ventral furrow close and parallel to ventral rim of otolith, regularly curved. Description Otolith size up to 0.85 mm in length (holotype). OH: OT = 3.3 – 4.0. Otolith shape high-bodied. Dorsal rim high, rounded or with rounded postdorsal angle; predorsal angle indistinct, postdorsal projection indistinct. Anterior rim slightly curved, posterior rim nearly vertical, straight. Ventral rim distinctly curved. Preventral and postventral angles broadly rounded. All rims smooth. Inner face completely flat. Sulcus short, droplet-shaped, eccentrically positioned towards anterior, with rounded, widened anterior tip and rounded ostial lobe, and short cauda, with tapering, often pointed tip; sulcus inclination angle 12 – 16 °; OL: SuL = 2.0 – 2.2. Subcaudal iugum long, wide, broadly extending around caudal tip and ingressing into cauda from ventral and posterior. Dorsal depression distinct. Ventral furrow distinct, close and parallel to ventral rim of otolith and regularly curved. Outer face evenly convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFEDFFA3018CF890601FFED8.taxon	discussion	Discussion Microgobius praeglaber is characterized by small otoliths that at first sight could be taken as stemming from juvenile specimens of some other gobiid species. It does, however, show a number of unique features that exclude such explanation, including the droplet-shaped forward-positioned sulcus and the massive ingression of the wide subcaudal iugum into the cauda. We therefore postulate that M. praeglaber and M. aphioides represented small, possibly hemipelagic gobies similar to the extant Aphia minuta.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFECFFA3018CFBB0669CFCF8.taxon	description	Rüber et al. (2003) and Tornabene et al., (2016, 2022) have undertaken extensive phylogenetic analyses of the Gobiosoma group. Tornabene et al. (2016) recognized two subgroups, the Barbulifer and the Nes subgroups. Tornabene et al. (2022) recognized an additional subgroup, the Robinsichthys subgroup. Rüber et al. (2003) did not formulate subgroups but instead recognized nine clades in the Gobiosomatini, two in the Microgobius group (the Bollmannia and the Microgobius clades) and seven in the Gobiosoma group (the Nes, Aboma, Barbulifer, Gobiosoma, Risor, Tigrigobius, and Elacatinus clades). Te phylogenies of the three publications are generally congruent but differ somewhat in details. With some variations, we have followed here the grouping in clades by Rüber et al. (2003), primarily because it is easier to relate to the otolith findings. We are aware that certain clades or subgroups are not represented by otoliths for the reasons mentioned above, i. e., the Aboma clade of Rüber et al. (2003) and the Robinsichthys subgroup of Tornabene et al. (2022). Only the Barbulifer and Nes clades of Rüber et al. (2003) are comprehensively represented. Most other clades or subgroups are represented by a limited number of taxa. Tis means that species recognition in the fossil record is hampered, except for the Barbulifer and Nes clades and a few other characteristic genera, and therefore many otoliths must remain in open nomenclature for the time being.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFECFFA3018CFE3060D0FC58.taxon	description	Figure 31 w Material 1 specimen Calabrian, Armuelles FM, Pacific Panama, PPP 3236 (NMB P 15737). Discussion A single, 1.45 mm long and relatively well-preserved specimen from the middle Pleistocene of Pacific Panama is characterized by a regularly crenulated otolith rim, a small and narrow sulcus with a very low ostial lobe and a massive subcaudal iugum ingressing into the cauda from ventral and posterior. Te ratio OL: OH is 0.85; the ratio OL: SuL = 2.2. Tere is a good chance that this otolith belongs to one of the four extant Microgobius species from the East Pacific from which otoliths are not yet known. Tis would also mean that the Microgobius camur otolith plexus, which apparently is extinct in the West Atlantic could have survived in the East Pacific.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFECFFAD0236FC50604CF918.taxon	description	Figure 32 c – g Holotype NMB P 15739 (Fig. 32 c), Gelasian, Bastimentos FM, Ground Creek Unit, Atlantic Panama, PPP 1256. Paratypes 7 specimens: 2 specimens Piacenzian, Escudo de Veraguas FM, Atlantic Panama, PPP 1241 (NMB P 15745 - 46). 5 specimens Calabrian, Swan Cay FM, Atlantic Panama, PPP 2221 (NMB P 15740 - 44). Referred specimen 7 specimens Gelasian: 4 specimens Escudo de Veraguas FM, Atlantic Panama, PPP 2170; 3 specimens Bastimentos FM, Ground Creek Unit, Atlantic Panama, 1 specimen PPP 3202, 2 specimens PPP 1256. Etymology Named after its occurrence in the West Atlantic. Diagnosis Otolith size up to 1.65 mm in length. OL: OH = 1.25 – 1.3. Otolith shape nearly rectangular with orthogonal predorsal, preventral and postventral angles and not protruding postdorsal projection. Dorsal rim highest at obtuse postdorsal angle; ventral rim straight to slightly bent, horizontal. Sulcus small, without ostial lobe; OL: SuL = 2.15 – 2.55; sulcus inclination angle 10 – 15 °. No subcaudal iugum. Dorsal depression connected to ventral furrow forming continuous circumsulcal furrow. Description Otolith size up to 1.65 mm in length (holotype). OH: OT = 2.3 – 2.6. Otolith shape nearly rectangular. Dorsal rim low, anteriorly straight and slightly inclined, with broad, obtuse postdorsal angle as highest point, and short, not protruding postdorsal projection. Predorsal and preventral angles nearly orthogonal and anterior rim almost vertical. Postventral angle rounded orthogonal and as strongly developed as postdorsal projection; posterior rim near vertical, with moderate concavity at about level of caudal tip. Ventral rim flat or slightly curved, horizontal to slightly inclined. All rims smooth. Inner face flat but area around sulcus somewhat bulged. Sulcus small, somewhat deepened, evenly wide, without ostial lobe, and rounded caudal tip; sulcus inclination angle 10 – 15 °; OL: SuL = 2.15 – 2.55. No subcaudal iugum. Dorsal depression distinct, narrow, curved, close to dorsal rim of otolith, continuously connected around sulcus with distinct ventral furrow. Resulting circumsulcal furrow almost oval, clipping otolith angles. Outer face evenly and relatively strongly convex, smooth. Discussion Te genus Aruma today is known from a single species, Aruma histrio (Jordan, 1844) (Fig. 32 a – b), a secretive species inhabiting rocky crevices and tide pools in the Gulf of California (Robertson & Allen, 2015). Its otoliths are characterized by the elongate rectangular outline and the small sulcus without clear differentiation of ostium and cauda and no ostial lobe. Aruma otoliths differ from those of the other genus in the clade, Barbulifer, in the absence of a protruding postdorsal projection. It is of interest to note that, from the Late Pliocene to the late Early Pleistocene, a presumably vicariant species existed in the tropical West Atlantic. Aruma atlantica differs from otoliths of the Pacific A. histrio in the shorter sulcus (OL: SuL = 2.15 – 2.55 vs 1.8 – 2.0) and the obtuse postdorsal angle (vs flat dorsal rim). Aruma atlantica seems also to have grown to larger sizes than its extant counterpart in the East Pacific. Aruma atlantica has been found in only a few localities, which may indicate a specific environmental adaptation. Of particular interest is its relatively common occurrence in the Swan Cay FM in locality PPP 2221, which represents a carbonatic sand layer in a cavity in a fossil reef bioherm.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE2FFAE018CF8B0610EFD78.taxon	description	Figure 32 k – n Holotype NMB P 15747 (Fig. 32 m), Calabrian, Swan Cay FM, Atlantic Panama, PPP 2531. Paratypes 4 specimens same data as holotype (NMB P 15748 - 51). Referred specimen 9 specimens: 2 specimens Gelasian, Bastimentos FM, Ground Creek Unit, Atlantic Panama, PPP 3202. 7 specimens Calabrian, Swan Cay FM, Atlantic Panama, 1 specimen PPP 2221, 6 specimens same data as holotype. Etymology From amplus (Latin) = great, wide, referring to the comparatively large size of the otoliths. Diagnosis Otolith size up to 1.9 mm in length. OL: OH = 1.25 – 1.35. Otolith shape parallelogram-like to slightly trapezoidal. Dorsal rim highest at obtuse, rounded postdorsal angle; predorsal angle distinct; postdorsal projection distinctly protruding; ventral rim straight to slightly bent, horizontal. Sulcus long, with very low ostial lobe; OL: SuL = 1.85 – 2.05; sulcus inclination angle 13 – 15 °. No subcaudal iugum. Dorsal depression anteriorly connected to ventral furrow. Description Otolith size up to 1.9 mm in length (holotype). OH: OT = 2.5 – 3.1. Otolith shape parallelogram-like or sometimes trapezoidal (Fig. 32 m). Dorsal rim low, anteriorly straight and slightly inclined, with broadly rounded postdorsal angle as highest point. Predorsal angle distinct, sometimes protruding (Fig. 32 m, n), postdorsal projection more or less strongly protruding. Preventral angle orthogonal or protruding, postventral angle broadly rounded. Anterior rim short, vertical or slightly inclined, with distinct central concavity; posterior rim slightly inclined toward ventral, with moderate concavity at level above caudal tip. Ventral rim flat or slightly curved, horizontal. All rims smooth. Inner face moderately convex in both directions. Sulcus long, somewhat deepened, with very low ostial lobe, and rounded caudal tip; sulcus inclination angle 13 – 15 °; OL: SuL = 1.85 – 2.05. No subcaudal iugum. Dorsal depression distinct, narrow, curved, close to dorsal rim of otolith, anteriorly connected around sulcus with distinct ventral furrow. Ventral furrow distinct, regularly curved, clipping preventral angle. Outer face evenly convex, about as strong as inner face, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE2FFAE018CF8B0610EFD78.taxon	discussion	Discussion Barbulifer amplus has relatively large otoliths for the genus indicating fish sizes in the range of 30 mm SL or more. Te only extant Barbulifer species reaching that size is B. pantherinus (Pellegrin, 1901) (Fig. 32 h – i) from the Gulf of California. Its otoliths have a more pronounced postdorsal projection and a small sulcus. Today, Barbulifer has five species, two in the East Pacific, two in the Caribbean, and one off central Brazil. Te two extant Caribbean Barbulifer species, B. antennatus Böhlke & Robins, 1968, and B. ceuthoecus (Jordan & Gilbert, 1884), seem not to achieve sizes larger than 20 mm SL for B. antennatus (see Froese & Pauly, 2023) and 30 mm SL for B. ceuthoecus (see Robertson & Allen, 2015). Otoliths of B. ceuthoecus (Fig. 32 j) do resemble B. amplus in otolith shape but have a larger sulcus (OL: SuL = 1.65 vs 1.85 – 2.05). Barbulifer amplus has been found in the same environment as Aruma atlantica, primarily in the sand layer in a cavity in a fossil reef bioherm of the Swan Cay FM at Swan Cay (PPP 2221 and 2531), and differs in the longer sulcus (OL: SuL = 1.85 – 2.05 vs 2.15 – 2.55) and the protruding postdorsal projection.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE1FFA80236FD306061FCF8.taxon	description	Figure 33 e – f Holotype NMB P 15752 (Fig. 33 e), Tortonian, Yaviza FM, Pacific Panama, Darien, PPP 1145. Paratypes 2 specimens Tortonian, Tuira FM, Pacific Panama, Darien, PPP 1137 (NMB P 15753 - 54). Etymology From altus (Latin) = high, referring to the comparatively compressed shape of the otoliths. Diagnosis Otolith size up to 1.15 mm in length. OL: OH = 1.05 – 1.1. Otolith shape rectangular to parallelogram-like. Postdorsal projection distinctly protruding; ventral rim straight, horizontal. Sulcus long, with very low ostial lobe; OL: SuL = 1.7 – 1.85; sulcus inclination angle 14 – 16 °. No subcaudal iugum. Ventral furrow close to ventral rim of otolith. Description Otolith size up to 1.15 mm in length (holotype 1.1 mm). OH: OT = 2.5 – 2.8. Otolith shape rectangular to parallelogram-like. Dorsal rim anteriorly depressed, with broadly rounded, shallow postdorsal angle. Predorsal angle distinct, not protruding, postdorsal projection pointed, slightly protruding. Preventral angle orthogonal, not protruding, postventral angle rounded. Anterior rim slightly inclined, with weak or without central concavity; posterior rim slightly inclined toward ventral, with (See figure on next page.) Fig. 33 a – b Chriolepis semisquamata (Rutter, 1904), extant, USNM 322622, 27 ° 57 ’ N 111 ° 22 ’ W. c Chriolepis zebra Ginsburg, 1938, extant, USNM 322602, Gulf of California, SL 26 mm. d Psilotris laurae van Tassell, Tornabene & Baldwin, 2016, extant, USNM 442092, Bonaire, 12 ° 05 ’ N 68 ° 17 ’ W, 134 - 165 m, SL 28 mm. e – f Chriolepis altus n. sp.; e holotype, NMB P 15752, Yaviza FM, Darien, Panama, PPP 1145; f paratype, Tuira FM, Darien, Panama, PPP 1137, NMB P 15753. g Chriolepis sp. 1, Bastimentos FM, Ground Creek Unit, Bocas del Toro, Panama, PPP 3202, NMB P 15755. h – k Chriolepis balboa n. sp.; h holotype, NMB P 15756, Angostura FM, Ecuador, PPP 3301; i – k paratypes, Gatun FM, Colon, Panama, PPP 2168, NMB P 15757 - 59. l Chriolepis sp. 2, Cumaná FM, Venezuela, PPP 3038, NMB P 15761. m – n Chriolepis sp. 3, Moin FM, Limón, Costa Rica, PPP 3259, NMB P 15762 - 63. o Chriolepis sp. 4, Río Banano FM, Limón, Costa Rica, PPP 3249, NMB P 15764. p – q Chriolepis sp. 5, Cubagua FM, Venezuela; p PPP 2557, NMB P 15765; q PPP 3057, NMB P 15766 moderate concavity at level above caudal tip. Ventral rim flat, horizontal. All rims smooth. Inner face moderately convex in both directions; area around sulcus slightly bulged. Sulcus moderately long, slightly deepened, with very low ostial lobe, and rounded caudal tip; sulcus inclination angle 14 – 16 °; OL: SuL = 1.7 – 1.85. No subcaudal iugum. Dorsal depression indistinct; ventral furrow distinct, close and parallel to ventral rim of otolith. Outer face evenly convex, slightly stronger than inner face, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE1FFA80236FD306061FCF8.taxon	discussion	Discussion Chriolepis altus is recognized by its relatively compressed shape and the relatively large sulcus. Te available otoliths of the few extant species show a great variety in sulcus size and development of the preventral angle and the postdorsal projection. A typical feature is also the bulging of the inner face around the sulcus which is often supported by the ventral furrow being connected around the sulcus to the dorsal depression.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE7FFA8018CFCD06751FD78.taxon	description	Figure 33 h – k Holotype NMB P 15756 (Fig. 33 h), Tortonian, Angostura FM, Ecuador, PPP 3301. Paratypes 4 specimens Tortonian: 1 specimen Tuira FM, Pacific Panama, Darien, PPP 1163 (NMB P 15760); 3 specimens Gatun FM, Atlantic Panama, PPP 2168 (NMB P 15757 - 59). Referred specimens 2 specimens Tortonian, Gatun FM, Atlantic Panama, PPP 2168. Etymology Named after Vasco Núñez de Balboa, the first to name the Pacific Ocean after having crossed the Isthmus of Panama. Diagnosis Otolith size up to 1.15 mm in length. OL: OH = 1.1. Otolith shape nearly rectangular but with depressed predorsal section. Postdorsal projection slightly protruding; ventral rim straight, horizontal. Sulcus long, with very low ostial lobe; OL: SuL = 1.55 – 1.7; sulcus inclination angle 8 – 14 °. No subcaudal iugum. Ventral furrow close to ventral rim of otolith, regularly curved. Description Otolith size up to 1.15 mm in length (holotype). OH: OT = 2.4 – 2.6. Otolith shape nearly rectangular with orthogonal preventral and postventral angles but somewhat depressed predorsal region and obtuse predorsal angle. Postdorsal projection rounded, slightly protruding. Anterior rim nearly vertical; posterior rim slightly inclined toward ventral, with weak concavity at level above caudal tip. Ventral rim flat, horizontal. All rims smooth. Inner face moderately convex in both directions; area around sulcus slightly bulged. Sulcus long, slightly deepened, with very low ostial lobe, and rounded caudal tip; sulcus inclination angle 8 – 14 °; OL: SuL = 1.55 – 1.7. No subcaudal iugum. Dorsal depression indistinct; ventral furrow distinct, close to ventral rim of otolith and regularly curved. Outer face evenly convex, slightly stronger than inner face, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE7FFA8018CFCD06751FD78.taxon	discussion	Discussion Chriolepis balboa is a typical otolith of the genus Chriolepis but with an unusually long sulcus.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE7FFA80236FD5061D9FBD8.taxon	description	Figure 33 g Material 3 specimens Gelasian, Bastimentos FM, Ground Creek Unit, Atlantic Panama, PPP 3202 (NMB P 15755). Discussion Unusually large otoliths for the genus Chriolepis up to 1.65 mm in length with a rounded dorsal rim, a short preventral projection, a weak postdorsal projection (OL: OH = 1.25) and a relatively long sulcus (OL: SuL = 1.7).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE7FFA80236FB306748FA58.taxon	description	Figure 33 l Material 1 specimen Calabrian, Cumaná FM, Venezuela, PPP 3038 (NMB P 15761). Discussion A rather compressed, nearly rectangular otolith with a high positioned predorsal angle and a weak postdorsal projection. Te sulcus is short and shows moderate ostial lobe. OL: OH = 1.1; OL: SuL = 2.0.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE7FFA90236F9B060A3FE98.taxon	description	Figure 33 m – n Material 3 specimens Gelasian, Moin FM, Atlantic Costa Rica, 1 specimen PPP 1773, 2 specimens PPP 3259 (NMB P 15762 - 63). Discussion Relatively compact otoliths with weak preventral and postdorsal projections. Characteristic is the straight, inclined dorsal rim, which is anteriorly depressed, and the long sulcus oriented parallel to the dorsal rim. OL: OH = 1.1 – 1.2; OL: SuL = 1.7 – 1.75.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE6FFA9018CFEF06307FD58.taxon	description	Figure 33 o Material 1 specimen Piacenzian, Río Banano FM, Atlantic Costa Rica, PPP 3249 (NMB P 15764). Discussion Te single, relatively large otolith of 2 mm in length is remarkable for its thick appearance (OH: OT = 2.2), the rather large, wide and somewhat deepened sulcus (OL: SuL = 1.65) and the relatively slen- der shape (OL: OH = 1.25) with a rounded postdorsal projection.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE6FFA9018CFCB062EAFA38.taxon	description	Figure 33 p – q Material 9 specimens Zanclean, Cubagua FM, Venezuela, 1 specimen PPP 2549, 1 specimen PPP 2555, 2 specimens PPP 2556, 1 specimen PPP 2557 (NMB P 15765), 2 specimens PPP 2563, 2 specimens PPP 3057 (NMB P 15766). Discussion Tis is a rather characteristic species with a highly elevated postdorsal region and a strongly depressed predorsal angle and with a steeply inclined predorsal rim (25 – 33 °). Anterior and posterior rims are nearly vertical and the postdorsal projection is very week. OL: OH = 1.05 – 1.15; OL: SuL = 1.7 – 1.8; sulcus inclination angle = 13 °. Tese otoliths may represent an undescribed species but because of their low stratigraphic age and the lack of knowledge of West Atlantic Chriolepis, Pinnichthys and Psilotris species we have refrained from a formal action.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE6FFA9018CFA1061CFFD58.taxon	description	(Fig. 34 d) — and Nes Ginsburg, 1933 — N. longus (Nichols, 1914) (Fig. 34 e). Te otoliths of Gobulus and Eleotrica are recognized by their relatively elongate shape with a distinct postdorsal projection (OL: OH = 1.2 – 1.35), the inner face being markedly bent in the horizontal direction, and the sulcus being relatively large, distinctly deepened and with a low ostial lob and relatively wide cauda (OL: SuL = 1.7 – 1.85); the sulcus inclination angle is relatively low at 5 – 15 °. Te otoliths of Gymneleotris differ in being more compact without a pronounced postdorsal projection and with a small sulcus (OL: SuL = 2.2). Te otoliths of Nes finally differ in being more high-bodied (OL: OH about 0.95) and showing a narrow and rather steeply inclined sulcus (> 20 °).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE6FFAA0236FCB066A9F9A6.taxon	description	Figure 34 f – g Holotype NMB P 15767 (Fig. 34 f), Gelasian, Moin FM, Atlantic Costa Rica, PPP 1773. Paratype 1 specimen Gelasian, Moin FM, Atlantic Costa Rica, PPP 1772 (NMB P 15768). Etymology Named after the province Limón, Costa Rica, where the specimens were collected. Diagnosis Otolith size up to 2.5 mm in length. OL: OH = 1.2 – 1.25. Otolith moderately elongate but with depressed predorsal section. Postdorsal projection and postventral angle equally protruding with broad concavity of posterior rim in between; ventral rim curved, horizontal. Sulcus moderately long, wide, with very low ostial lobe; OL: SuL = 1.85 – 1.95; sulcus inclination angle 5 – 6 °. No subcaudal iugum. Ventral furrow regularly curved. Description Otolith size up to 2.5 mm in length (holotype). OH: OT = 2.8 – 3.2. Otolith moderately elongate; dorsal rim shallow, predorsal section depressed, regularly declining towards low predorsal angle; postdorsal angle highest point, broadly rounded. Postdorsal projection distinct, moderately protruding, about as strongly as projecting postventral angle; posterior rim with broad, distinct concavity between postdorsal and postventral projections. Anterior rim short, nearly vertical. Ventral rim curved, horizontal. All rims smooth. Inner face moderately convex in both directions; area around sulcus slightly bulged. Sulcus relatively short but wide, deepened, with very low ostial lobe, and cauda about equally wide as ostium; sulcus inclination angle 5 – 6 °; OL: SuL = 1.85 – 1.95. No subcaudal iugum. Dorsal depression indistinct, joint to distinct ventral furrow around sulcus; ventral furrow half-moon shaped, close to ventral rim of otolith only at central part. Outer face nearly flat, smooth. Discussion Otoliths of Gobulus limonensis are exceptionally large for otoliths of the Nes clade or the genus Gobulus. An otolith of the extant G. crescentalis of 61 mm SL, which is the largest recorded specimen of the species (Robertson & Allen, 2015) and represents the largest extant species of the genus, is 1.8 mm in length. Te fishes from which the fossil G. limonensis otoliths are described may have surpassed 80 mm SL. In a redescription of the only extant Atlantic species, Gobulus myersi Ginsburg, 1939, Hastings (1983) found no specimen larger than 29.3 mm SL among all known specimens. Te size correlation therefore rules out G. limonensis being synonymous with the extant G. myersi, even though no otoliths are known from the latter.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE5FFAB0236F9FE60B2FE3F.taxon	description	(Fig. 35 b), and V. cephalocellatus (Fig. 35 c). Te otoliths show a parallelogram-like shape, as is also typical for many Atlanto-Mediterranean goby otoliths, with a relatively short sulcus. Te expression of the ostial lobe appears to be highly variable and could represent an important characteristic for differentiation of species in this genus.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE4FFB4018CFE1060AAF9F8.taxon	description	Figure 35 d – f Holotype NMB P 15769 (Fig. 35 d), Zanclean, Cubagua FM, Venezuela, PPP 2568. Paratypes 2 specimens same data as holotype (NMB P 15770 - 71). Referred specimen 1 specimen Zanclean, Cubagua FM, Venezuela, PPP 3093. Etymology Named after the occurrence of the species in the (early) Pliocene. Diagnosis Otolith size up to 1.9 mm in length. OL: OH = 1.2 – 1.3. Otolith shape parallelogram-like with pronounced preventral angle and expanded postdorsal projection. Dorsal rim regularly curved. Sulcus moderately long, relatively narrow, with pointed ostial tip and tapering caudal tip; ostial lobe rounded with colliculum not extending into lobe; OL: SuL = 1.75 – 1.9; sulcus inclination angle 12 – 15 °. No subcaudal iugum. Ventral furrow regularly curved. Description Otolith size up to 1.9 mm in length (holotype). OH: OT = 2.7 – 2.9. Otolith parallelogram-like in shape with pronounced preventral angle, broad and expanded postdorsal projection and obtuse, rounded predorsal and postventral angles. Dorsal rim regularly curved with broad, rounded postdorsal angle. Anterior rim short, nearly vertical to slightly inclined towards dorsal. Posterior rim slightly inclined towards ventral, with narrow concavity at level of tip of cauda. Ventral rim straight, horizontal. All rims smooth. Inner face moderately convex in both directions. Sulcus moderately long, narrow, slightly deepened, its ostial tip pointed, caudal tip tapering, narrow; ostial lobe low with rounded expansion without extension of colliculum; sulcus inclination angle 12 – 15 °; OL: SuL = 1.75 – 1.9. No subcaudal iugum. Dorsal depression indistinct; ventral furrow regularly curved, closest to ventral rim of otolith at central part. Outer face convex, usually more strongly than inner face, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFE4FFB4018CFE1060AAF9F8.taxon	discussion	Discussion Varicus pliocenicus is characterized by the parallelogram-like shape that resembles the extant V. veliguttatus; the unusual expansion of the ostial lobe without extension of the colliculum, a feature also seen in a slightly different form in the extant V. decorum; and the pointed ostial tip and tapering, narrow caudal tip of the sulcus. Varicus pliocenicus is thus a well-defined and easily recognized otolith of this genus. None of the 12 extant Varicus species are known from off Venezuela, but three are known off Curacao: V. decorum, V. lacerta, and V. veliguttatus. Otoliths of V. decorum (Fig. 35 a) resemble V. pliocenicus in the unusual development of the ostial lobe but otherwise show little similarity. Otoliths of V. veliguttatus resemble V. pliocenicus in otolith proportions and the pointed ostial tip of the sulcus, but differ in the rounded caudal tip, the lack of the expansion of the ostial lobe, and the less strongly developed postdorsal projection. Otoliths of V. lacerta are not known.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFBFFB4018CF9D0619FFED8.taxon	description	Figure 35 g – h Material 3 specimens Calabrian, Cumaná FM, Venezuela, PPP 3038 (figured specimens NMB P 15772 - 73). Discussion Tese are relatively small and compressed specimens with a small and simple-shaped sulcus that resemble in habitus the extant V. cephalocellatus (Fig. 35 c).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFBFFB40236FE3061B0FD78.taxon	description	Figure 35 i – j Material 2 specimens Zanclean, Cubagua FM, Venezuela, 1 specimen PPP 3057 (NMB P 15774), 1 specimen PPP 3055 (NMB P 15775). Discussion Like Varicus sp. 1 mentioned above these are compressed otoliths but with a relatively long and structured sulcus.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFBFFB50236FD50631AFE38.taxon	description	Te fishes of the genera Gobiosoma and Enypnias occur mostly in shallow near-shore environments, for instance in mangrove and tide pools or rocky environments. Several species inhabit freshwater and brackish waters or are euryhaline.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFAFFB5018CFC3E660FFC18.taxon	description	Figure 36 l Holotype NMB P 15777 (Fig. 36 l), Messinian, Chucunaque FM, Pacific Panama, Darien, PPP 1612. Etymology Named after the indigenous Indian tribe Embera inhabiting the Darien province of Panama. Diagnosis Otolith size 0.9 mm in length. OL: OH = 0.95. Otolith shape more or less rectangular with pronounced predorsal, preventral and postventral angles and angular postdorsal projection. Anterior, posterior and predorsal rims slightly concave. Sulcus long, relatively narrow, deepened, without ostial lobe; OL: SuL = 1.55; sulcus inclination angle 20 °. No subcaudal iugum. Ventral furrow regularly curved. Description A small otolith of 0.9 mm in length (holotype). OH: OT = 2.8. Otolith more or less rectangular. Dorsal rim shallow, with nearly orthogonal, high positioned predorsal angle, slightly concave ascending predorsal rim, obtuse but distinct postdorsal angle and angular, not protruding postdorsal projection. Ventral rim nearly flat, horizontal, with orthogonal preventral angle and rounded postventral angle. Anterior rim vertical, broadly concave; posterior rim vertical, very slightly concave. All rims smooth. Inner face moderately convex in both directions; area around sulcus markedly bulged. Sulcus long, relatively narrow, distinctly deepened, its ostial tip more pointed than caudal tip; no ostial lobe discernable; sulcus inclination angle 20 °; OL: SuL = 1.55. No subcaudal iugum. Dorsal depression marked, long and narrow, with wide crista superior toward sulcus, connected to evenly curving ventral furrow around sulcus. Outer face convex, smooth. Discussion Despite its small size, G. emberae is well characterized by its shape, particularly the sharply angled, high predorsal angle and the concave predorsal and anterior rims. Tese characters and the long sulcus set G. emberae apart from coeval otoliths and the known extant Gobiosoma otoliths.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFAFFB50236FC706742FA38.taxon	description	Figure 36 p – q Material 3 specimens Zanclean, Cubagua FM, Venezuela, PPP 3057 (figured specimens NMB P 15778 - 79). Discussion Tese otoliths are in many ways similar to G. emberae from the Tortonian but differ in the posterior rim being inclined toward dorsal, the more rounded angles and the ventral furrow running further away from the ventral rim of the otolith. It also resembles the extant G. robustum (Fig. 36 m – o), but is thicker and has a shorter sulcus with a lower, almost flat ostial lobe.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFAFFB70236FA106310FE18.taxon	description	Figure 36 r (See figure on next page.) Fig. 36 a Enypnias aceras Ginsburg, 1939, extant, BMNH 1975.4.25.11 - 12, Panama canal, Miraflores. b – d Enypnias seminudus (GÜnther, 1861), extant, USNM 407784, 13 ° 22 ’ N 87 ° 52 ’ W. e Garmannia sp., Springvale FM, Trinidad, KR- 1216, NMB P 15776. f Garmannia schultzi Ginsburg, 1944, extant, USNM 121550, Venezuela, Lago Maracaibo. g Gobiosoma chiquita (Jenkyns & Evermann, 1889), extant, LACM coll Fitch, Gulf of California. h – k Gobiosoma bosc (Lacepède, 1800), extant; h, k TCWC- 20131.04 Texas, Oso Bay SL 36, 54 mm; i LACM coll Fitch, Rappahonnock; j USNM 420904, 28 ° 41 ’ N 96 ° 13 ’ W. l Gobiosoma emberae n. sp., holotype, NMB P 15777, Tuira FM, Darien, Panama, PPP 1612. m – o Gobiosoma robustum Ginsburg, 1933, extant; m, o TCWC 20141.03, off Texas, SL 23,5 and 25, 5 mm; n CAS 36379, Florida, Sanibel Island. p – q Gobiosoma sp. 1, Cubagua FM, Venezuela, PPP 3057, NMB P 15778 - 79. r Gobiosoma sp. 2, Tuira FM, Darien, PPP 1593, NMB P 15780. s Gobiosoma sp. 3, Swan Cay FM, Atlantic Panama, PPP 2221, NMB P 15781. t – v Gobiosoma sp. 4, Cumaná FM, Venezuela, PPP 3038, NMB P 15782 - 84. W Gobiosoma sp. 5, Moin FM, Limón, Costa Rica, PPP 2038, NMB P 15785 Material 1 specimen Tortonian, Tuira FM, Pacific Panama, Darien, PPP 1593 (NMB P 15780). Discussion A relatively high-bodied otolith (OL: OH = 0.93) that also resembles otoliths of Elacatinus (see below).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF8FFB7018CF9D061D0FB58.taxon	description	Te aforementioned genera of this clade contain nearly 50 valid species according to Fricke et al. (2023): 24 species in Elacatinus, 16 in Tigrigobius, five in Evermannichthys, and one each in Risor, Ginsburgellus, Ophiogobius, and Pariah. Otoliths are known from only a few of these taxa: Elacatinus chancei (Beebe & Hollister, 1933) (Fig. 37 a – b), E. horstii (Metzelaar, 1922) (Fig. 37 c – d), E. lori Colin, 2002 (Fig. 37 e), E. oceanops Jordan, 1904 (Fig. 37 f), E. puncticulatus (Ginsburg, 1938) (Fig. 37 g), E. xanthipora (Böhlke & Robins, 1968) (Fig. 37 h), Ginsburgellus novemlineatus (Fowler, 1950) (Fig. 38 a), Risor ruber (Rosén, 1911) (Fig. 38 c), and Tigrigobius macrodon (Beebe & Tee Van, 1928) (Fig. 38 d). Because of this low coverage, the fossil otoliths described here have been left in open nomenclature, except for one otolith that can be attributed to an extant species (Ginsburgellus novemlineatus).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF8FFB70236FAF061FEF8D8.taxon	description	Figure 37 i – j Material 4 specimens: 1 specimen Zanclean, Cubagua FM, Venezuela, PPP 2568; 2 specimens Gelasian, Bastimentos FM, Ground Creek Unit, Atlantic Panama, PPP 2247 (figured specimen NMB P 15787); 1 specimen Calabrian, Bastimentos FM, Ground Creek Unit, Atlantic Panama, PPP 2250 (NMB P 15786). Discussion Tese otoliths are characterized by their high-bodied appearance (OL: OH = 0.88 – 0.93), the strong postdorsal angle and the relatively large sulcus without ostial lobe. Tey resemble otoliths of the extant species E. chancei (Fig. 37 a – b) and E. lori (Fig. 37 e, formalin-eroded specimen).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF8FFB7018CFE70606EFCD8.taxon	description	Figure 36 s Material 1 specimen Calabrian, Swan Cay FM, Atlantic Panama, PPP 2221 (NMB P 15781). Discussion A relatively large otolith for a Gobiosoma species of 1.3 mm length. It resembles G. emberae in shape but has a sharper preventral projection and a more forward positioned postdorsal angle. Tis otolith likely pertains to an extant species that may have derived from such a Late Miocene form like G. emberae.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF8FFB7018CFC30605AFB78.taxon	description	Figure 36 t – v Material 20 specimens Calabrian, Cumaná FM, Venezuela, PPP 3038 (figured specimens NMB P 15782 - 84). Discussion Tese otoliths resemble Gobiosoma sp. 1 from the Zanclean of Venezuela and differ mainly in the more rounded dorsal rim and in being thinner.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF8FFB7018CFB5063A5F9F8.taxon	description	Figure 36 w Material 1 specimen Gelasian, Moin FM, Atlantic Costa Rica, PPP 2038 (NMB P 15785). Discussion Te single, slightly eroded otolith resembles otoliths of the extant G. robustum but is remarkable for its rounded triangular shape with the relatively small dorsal field on the inner face.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFEFFB1018CFF5060A8FDF8.taxon	description	Figure 37 k Material 1 specimen Zanclean, Shark Hole FM, Atlantic Panama, PPP 2218 (NMB P 15788). Discussion Te single, small specimen of 0.7 mm length is remarkable for its rather evenly rounded shape, which could be an indication for a juvenile specimen, and the very small kidney-shaped sulcus (OL: SuL = 2.3).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFEFFB1018CFDD0603EFBB8.taxon	description	Figure 37 l Material 1 specimen Zanclean, Cubagua FM, Venezuela, PPP 2549 (NMB P 15789). Discussion Tis small otolith resembles Elacatinus sp. 2 but has an even smaller sulcus (OL: SuL = 2.65) which is also somewhat forward positioned. It further differs in the more strongly inclined sulcus (22 ° vs 10 °). It thus resembles the extant E. puncticulatus (Fig. 37 g) which, however, has a distinctly larger sulcus.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFEFFB1018CFB9062E4F9D8.taxon	description	Figure 37 m – o Material 3 specimens Zanclean, Cubagua FM, Venezuela, 1 specimen PPP 2557 (NMB P 15790), 1 specimen PPP 3057 (NMB P 15791), 1 specimen PPP 3055 (NMB P 15792). Discussion Tese are relatively distinctive otoliths characterized by a nearly quadrate shape (OL: OH = 1.0 – 1.05), and only slightly inclined oval sulcus (7 – 10 °) and the dorsal depression and ventral furrow joint to a nearly circular furrow surrounding the sulcus. Te area between the circular furrow and the sulcus is bulged. Elacatinus sp. 4 is very similar to the extant E. oceanops (Fig. 37 f) and could possibly represent the same or a closely related species.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFEFFB1018CF9706603FE58.taxon	description	Figure 38 a – b Material 1 specimen late Zanclean, upper Cayo Agua FM, Atlantic Panama, PPP 1188 (NMB P 15793). Discussion Te single fossil specimen perfectly matches the available extant specimen although the latter is slightly eroded (Fig. 38 a), but is slightly larger. Characteristic features are the rounded orthogonal preventral and postventral angles, the rounded middorsal angle, the otolith proportion (OL: OH = 1.0 in the extant specimen vs 1.03 in the fossil) and the small, oval sulcus which is only slightly inclined (7 °).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFEFFB10236FDF066D4FB18.taxon	description	Figure 38 e Material 1 specimen Tortonian, Tuira FM, Pacific Panama, Darien, PPP 1132 (NMB P 15794). Discussion Only two extant specimens have been available from the many species in the genus: T. macrodon (Fig. 38 d) and T. saucrus (Robins, 1960) kindly made available as photograph by Chien-Hsiang Lin. Both are characterized by a high-bodied shape (OL: OH <1.0), a rounded ventral rim and a very small, oval sulcus. An otolith of Risor ruber (Fig. 38 c) is figured for further comparison. Te single fossil otolith from the Tortonian shows the same small, oval sulcus and a reasonable comparison of the otolith shape to interpret it as an early representative of Tigrigobius. It is, however, too poorly preserved for any further identification.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFEFFB10236FB7061CCF998.taxon	description	Figure 38 f – h Material 4 specimens Calabrian, Cumaná FM, Venezuela, PPP 3038 (figured specimens NMB P 15795 - 97). Discussion Tese otoliths show a high oval shape with a small kidney-shaped sulcus that reminds of Tigrigobius. However, their notable variability hinders a further identification.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFDFFB30236F9E26358FE38.taxon	description	Figure 38 k – l Material 2 specimens: 1 specimen Zanclean, Cubagua FM, Venezuela, PPP 2549 (NMB P 15799). 1 specimen Calabrian, Swan Cay FM, Atlantic Panama, PPP 2221 (NMB P 15800). Discussion Two slightly eroded small otoliths (0.95 and 0.8 mm long) could possibly represent specimens pertaining to Lythrypnus. Particularly the specimen from Swan Cay (Fig. 38 l) bears the typical characters of the Priolepis lineage.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFCFFB3018CFC9061D1FDF8.taxon	description	(Fig. 39 c – d), C. thrix Böhlke & Robins, 1960 (Fig. 39 e), C. urospilus Ginsburg, 1938 (the only East Pacific species in the genus) (Fig. 39 f), C. venezuelae Cervigón, 1966 (Fig. 39 g – h), C. eidolon Böhlke & Robins, 1960 (Fig. 39 i), C. tortugae (Jordan, 1904) (Fig. 39 j), C. lipernes Böhlke & Robins, 1962 (Fig. 40 a), and C. personatus (Jordan & Tompson, 1905) (Fig. 40 c – d). Lophogobius contains two species, L. cristulatus Ginsburg, 1939, in the East Pacific and L. cyprinoides (Pallas, 1770) (Fig. 40 i – n) in the West Atlantic. Rhinogobiops is monospecific with R. nicholsi (Bean, 1882) (Fig. 40 o – p) in the East Pacific. Rhinogobiops nicholsii is a large-growing species (up to 150 mm TL according to Froese & Pauly, 2023), and the two otoliths figured from a large adult specimen (OL = 2.75 mm; Fig. 40 o) and a small juvenile specimen (OL = 1.15 mm; Fig. 40 p) show a significant degree of allometric growth. Te still relatively large size of the otolith from the juvenile specimen exemplifies the problem that can arise when identifying small goby otoliths in the range of about 1 mm in length.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFCFFB3018CFE1063AEFCB8.taxon	description	Figure 38 j Material 1 specimen Calabrian, Swan Cay FM, Atlantic Panama, PPP 2221 (NMB P 15798). Discussion Tis otolith is very similar to the East Pacific Lythrypnus zebra (Fig. 38 i). It is also interesting that it was found in a closely reef-associated setting, i. e., in a fossil reef cave sediment.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFCFFBD0236FA906307FA58.taxon	description	Figure 39 m Holotype NMB P 15803 (Fig. 39 m), Tortonian, Tuira FM, Pacific Panama, Darien, PPP 1137. (See figure on next page.) Fig. 39 a Coryphopterus dicrus Böhlke & Robins, 1960, extant, USNM 413296, 11 ° 09 ’ N 60 ° 50 ’ W, 5 - 9 m. b Coryphopterus curasub Baldwin & Robertson, 2015, extant, USNM 442134, Bonaire SL 31 mm. c – d Coryphopterus glaucofraenum Gill, 1863, extant, USNM 388062, 17 ° 21 ’ N 63 ° 15 ’ W, 15 - 18 m. e Coryphopterus thrix Böhlke & Robins, 1960, extant, USNM 413803, Curacao. f Coryphopterus urospilus Ginsburg, 1938, extant, coll. Schwarzhans leg. Fitch, Baja California. g – h Coryphopterus venezuelae Cervigón, 1966, extant; g USNM 413044, S Caicos; h USNM 413804, Curacao. i Coryphopterus eidolon Böhlke & Robins, 1960, USNM 397696, 17 ° 30 ’ N 63 ° 13 ’ W. j – l Coryphopterus tortugae (Jordan, 1904); j extant, USNM 411902, 21 ° 29 ’ N 71 ° 31 ’ W; k – l Cumaná FM, Venezuela, PPP 3038, NMB P 15801 - 02. m Coryphopterus cuevae n. sp., holotype, NMB P 15803, Tuira FM, Darien, Panama, PPP 1137. n – q Coryphopterus rodriguezi n. sp., Tuira FM, Darien, Panama; n holotype, NMB P 15804, PPP 1612; o – q paratypes; o – p PPP 1612, NMB P 15805 - 06; q PPP 1137, NMB P 15807. r – t Coryphopterus xenosus n. sp.; r holotype, NMB P 15808, Cubagua FM, Venezuela, PPP 2556; s – t paratypes; s Cubagua FM, Venezuela, PPP 2556, NMB P 15809; t Manzanilla FM, Trinidad, PPP 2673, NMB P 15810 Etymology Named after the Cueva, indigenous Indians living in Panama. Diagnosis Otolith size 1.5 mm in length. OL: OH = 1.0. Otolith with and regularly curved dorsal rim. Preventral projection pointed; postdorsal projection rounded, slightly protruding. Sulcus relatively short and narrow, with strong ostial lobe and narrow cauda; OL: SuL = 1.8; sulcus inclination angle 13 °. No subcaudal iugum. Ventral furrow regularly curved. Description Otolith size 1.5 mm in length (unique holotype). OH: OT = 3.6. Otolith with high and regularly curved dorsal rim; predorsal angle rounded, relatively low; postdorsal projection broadly rounded, slightly protruding. Anterior rim slightly inclined towards dorsal, with prominent and pointed preventral projection and broad concavity at level of ostial tip. Posterior rim slightly inclined towards ventral, with broad, shallow concavity at level of tip of cauda and rounded orthogonal postventral angle. Ventral rim straight, horizontal. All rims smooth, except dorsal rim slightly undulating. Inner face moderately convex in both directions. Sulcus relatively short, narrow, slightly deepened, its ostial and caudal tips tapering; ostial lobe strong, as much expanded as sulcus height; sulcus inclination angle 13 °; OL: SuL = 1.8. No subcaudal iugum. Dorsal depression indistinct, several weak but long radial furrows ingressing dorsal field from dorsal rim of otolith; ventral furrow regularly curved, closest to ventral rim of otolith at central part. Outer face slightly convex, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFFCFFBD0236FA906307FA58.taxon	discussion	Discussion Coryphopterus cuevae is easily recognized by its high and regularly curved dorsal rim resulting in a relatively low ratio OL: OH of about 1.0 and the strong ostial lobe. With the latter it resembles such otoliths as from the extant C. dicrus (Fig. 39 a), C. glaucofraenum (Fig. 39 c – d) or C. venezuelae (Fig. 39 g – h) but is further distinguished by the less strongly protruding postdorsal projection.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF2FFBD018CF9B06757F8F8.taxon	description	Figure 39 n – q Holotype NMB P 15804 (Fig. 39 n), Messinian, Chucunaque FM, Pacific Panama, Darien, PPP 1612. Paratypes 3 specimens: 2 specimens Messinian: same data as holotype (NMB P 15805 - 06). 1 specimen Tortonian, Tuira FM, Pacific Panama, Darien, PPP 1137 (NMB P 15807). Referred specimens 3 specimens: 2 specimens Messinian same data as holotype, PPP 1612. 1 specimen Tortonian, Tuira FM, Pacific Panama, Darien, PPP 1137. Etymology Named in honor of Felix Rodriguez (Panama City, Republic of Panama) for his support during field trips and laboratory work. Diagnosis Otolith size up to 1.6 mm in length. OL: OH = 1.1 – 1.15. Otolith rectangular to parallelogramshaped. Preventral projection blunt, short; postdorsal projection relatively sharp, moderately protruding. Sulcus relatively short and narrow, with low ostial lobe and narrow cauda; OL: SuL = 1.7 – 1.8; sulcus inclination angle 10 – 17 °. No subcaudal iugum. Ventral furrow relatively indistinct, regularly curved. Description Otolith size up to 1.6 mm in length (holotype). OH: OT = 2.8 – 3.0. Otolith rectangular to parallelogram-shaped; dorsal rim anteriorly depressed and its predorsal part straight, downward inclined; postdorsal angle obtuse, angular; predorsal angle rounded, relatively low; postdorsal projection relatively sharp, somewhat protruding. Anterior rim slightly inclined towards dorsal, with short, blunt preventral projection, straight or with weak concavity at level of ostial tip. Posterior rim slightly inclined towards ventral, with broad, relatively distinct concavity at level of tip of cauda. Ventral rim straight, horizontal, with rounded orthogonal postventral angle. All rims smooth or slightly undulating. Inner face moderately convex in vertical direction and distinctly bent in horizontal direction. Sulcus relatively short, narrow, slightly deepened, its ostial and caudal tips tapering; ostial lobe low; sulcus inclination angle 10 – 17 °; OL: SuL = 1.7 – 1.8. No subcaudal iugum. Dorsal depression relatively distinct, narrow, with broad crista inferior towards sulcus; ventral furrow relatively indistinct, close and parallel to ventral rim of otolith. Outer face slightly convex, smooth. Discussion Because of the low ostial lobe, otoliths of C. rodriguezi resemble those of the extant species C, eidolon (Fig. 39 i) or C. thrix (Fig. 39 e) but differ in the relatively low dorsal rim and the blunt preventral projection. Also the angular postdorsal angle and the relatively straight inclined predorsal rim distinguish C. rodriguezi from the two extant species. From C. eidolon it further differs in the more slender shape (OL: OH = 1.1 – 1.15 vs 1.05).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF1FFBE0236FC1066FCFA58.taxon	description	Figure 40 a – b Material 1 specimen Zanclean, Springvale FM, Trinidad, KR- 1216 (NMB P 15811).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF1FFBE0236FC1066FCFA58.taxon	discussion	Discussion Coryphopterus lipernes and C. personatus belong to the high-bodied otolith morphotype found in the genus, which is further characterized by high positioned predorsal angle and thus is easily distinguished from the known otoliths of the other extant Coryphopterus species. Te otoliths of C. lipernes are slightly less high-bodied than those of C. personatus (OL: OH = 0.9 – 0.95 vs 0.85 – 0.9) and show a narrower sulcus. Te only available extant specimen of C. lipernes (Fig. 40 a) is somewhat eroded by formalin but nevertheless allows a reliable correlation with the fossil specimen (Fig. 40 b).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF1FFBE018CFF5060C7FD18.taxon	description	Figure 39 j – l Material 8 specimens Calabrian, Cumaná FM, Venezuela, PPP 3038 (figured specimens NMB P 15801 - 02). Discussion Te specimens retrieved from the Calabrian of the Cumaná FM (Fig. 39 k – l) match well its extant counterpart (Fig. 39 j) in otolith proportions and the low sulcus inclination angle (11 – 12 °). Smaller specimens differ in a relatively small and less structured sulcus (Fig. 39 l). Today, C. tortugae does not seem to occur so far south in the Caribbean at the coast of Venezuela.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF1FFBE018CFD70663CFC38.taxon	description	Figure 39 r – t Holotype NMB P 15808 (Fig. 39 r), Zanclean, Cubagua FM, Venezuela, PPP 2556. Paratypes 2 specimens: 1 specimen same data as holotype (NMB P 15809); 1 specimen Messinian, Manzanilla FM, Montserrat Member, Trinidad, PPP 2673 (NMB P 15810). Referred specimens 1 specimen Zanclean, Cubagua FM, Venezuela, PPP 25565. Etymology From xenos (Greek) = foreign, referring to the unusual pattern of the otolith. Diagnosis Otolith size up to 1.35 mm in length. OL: OH = 1.15. Otolith skewed rectangular with relatively high predorsal angle and slightly protruding postdorsal projection. Preventral projection blunt, short. Sulcus short and narrow, with moderately developed low ostial lobe and narrow cauda; OL: SuL = 1.9; sulcus inclination angle 12 – 14 °. No subcaudal iugum. Ventral furrow distinct, regularly curved. Description Otolith size up to 1.35 mm in length (holotype). OH: OT = 2.5 – 2.8. Otolith skewed rectangular; dorsal with straight, downward inclined predorsal part, shallow postdorsal angle and high, distinct, rounded predorsal angle; postdorsal projection rounded, slightly protruding beyond broadly rounded postventral angle. Anterior rim nearly vertical, with weak concavity at level of ostial tip. Posterior rim slightly inclined towards ventral, with weak concavity at level of tip of cauda. Ventral rim straight, horizontal, with rounded slightly protruding preventral projection. All rims smooth. Inner face slightly convex. Sulcus relatively short, narrow, distinctly deepened, its ostial and caudal tips tapering; ostial lobe moderately developed; sulcus inclination angle 12 – 14 °; OL: SuL = 1.9. No subcaudal iugum but sometimes weak indications (Fig. 39 s, t) similar to certain extant otoliths of the genus. Dorsal depression relatively distinct, narrow, short, with broad crista inferior towards sulcus; ventral furrow distinct, close and to ventral rim of otolith. Outer face distinctly convex, much stronger than inner face, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF1FFBE018CFD70663CFC38.taxon	discussion	Discussion Coryphopterus xenosus differs from the otoliths of the known extant species in the more rectangular otolith shape and the relatively thick habitus of the otolith. Also untypical is the high positioned predorsal angle. It is possible that this species rather represents an extinct genus but we would consider such allocation premature at the moment considering the limited knowledge of fossil Coryphopterus today.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF7FFB80236FAD0666FF898.taxon	description	Figure 41 a – c Material 1 specimen, Calabrian, Armuelles FM, Pacific Panama, PPP 3235 (NMB P 15819). Discussion Te single, slightly eroded otolith from the Armuelles FM (Fig. 41 c) resembles extant otoliths of C. andrei well but show a postdorsal projection that protrudes as much as the postventral projection, i. e., more strongly than observed in the two extant specimens (Fig. 41 a – b). We consider these small differences to reflect a degree of variability.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF7FFB8018CF9906644FAF8.taxon	description	Te overall otolith pattern is similar to the characterization above, but certain differences exist in the expression of features or proportions. Some species show a more pronounced albeit rounded postdorsal projection, which, however, never expands further than the postventral projection (C. andrei, C. casamancus, C. lacertus); other species are characterized by a more compressed otolith shape and a high, expanded postdorsal angle (C. geminatus, C. mystacium).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF7FFB8018CFE106097FBD8.taxon	description	Figure 40 i – n Material 11 specimens: 3 specimens Zanclean, Cubagua FM, Venezuela, 1 specimen PPP 2549 (NMB P 15818), 2 specimens PPP 2556. 8 specimens Calabrian, Cumaná FM, Venzuela, PPP 3038 (figured specimens NMB P 15814 - 17). Discussion Otoliths of Lophogobius cyptinoides show pronounced preventral and postdorsal projections and do not differ significantly from certain species in the genus Coryphopterus such as C. thrix (Fig. 39 e) or C. urospilus (Fig. 39 f). Lophogobius otoliths tend to have a smaller sulcus and lower ostial lobe than species of Coryphopterus with the exception of C. curasub (Fig. 39 b). Te fossil specimens (Fig. 40 j – n) appear to have a higher postdorsal rim than the figured extant specimen (Fig. 40 i). However, the only available extant specimen has a slightly damaged dorsal rim because of formalin erosional effects and therefore this could be a misleading impression.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF7FFB8018CFB306370FA78.taxon	description	Figure 40 g – h Material 2 specimens, late Burdigalian, Baitoa FM, Dominican Republic, NMB 17290 (NMB P 15849). Discussion Te two small and eroded specimens likely represent a species of either the genus Lophogobius to which it is tentatively associated, or another genus of the Lophogobius lineage.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF5FFBA0236FDD0671FFB98.taxon	description	Figure 42 d – e Material 2 specimens Gelasian, Moin FM, Atlantic Costa Rica, PPP 1772 (NMB P 15820 - 21). Discussion Two otoliths found in the Moin FM of Costa Rica resemble extant otoliths of C. boleosoma (Fig. 42 a – c) but differ slightly in showing a deepened sulcus (vs shallow). Tey also do not show the widened pre- and postdorsal margins of the otolith, which appears to be typical for some Ctenogobius species. Terefore, and because of the inconspicuous overall appearance the fossil otoliths are only tentatively assigned to C. boleosoma.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF5FFBA018CFE106718FDF8.taxon	description	(Fig. 42 k – m), and C. stigmaticus (Poey, 1860) (Fig. 42 p) from the West Atlantic; and C. manglicola (Jordan & Starks, 1895) (Fig. 42 h) and C. sagittula (Günther, 1862) (Fig. 43 a – c) from the East Pacific. For further comparison, otoliths are figured from the extant Evorthodus lyricus (Girard, 1858) (Fig. 42 v – x), E. minutus Meek & Hildebrand, 1928 (Fig. 42 y), Gobioides peruanus (Steindachner, 1880) (Fig. 43 f), and G. broussonnetii Lacepède, 1800 (Fig. 43 g). For the genera Gobionellus and Gnatholepis, see below. Otoliths of the genus Ctenogobius show considerable variation, but are mostly compressed, usually higher than long (OL: OH ≤ 1.0), with a sole-shaped sulcus. Te details of the otolith outline, particularly the dorsal rim, vary from one species to another, but the otoliths usually show a moderate or regressed postdorsal projection. Te otoliths of one species, however, differ considerably from those known of other species. Ctenogobius sagittula has more elongate otoliths (OL: OH = 1.05 – 1.15), a massively developed postdorsal projection that expands toward dorsally and posteriorly with a flat to slightly concave middorsal section and the presence of a distinct subcaudal iugum. Te species also keys out first in the identification key published by Pezold (2022), who confirmed in a personal message (2023) that C. sagittula is the most divergent species in the genus. Otoliths of Evorthodus have a reduced sulcus morphology. Tose of Gobioides also show a somewhat simplified sulcus morphology, a nearly horizontal sulcus, and a distinct circumsulcal furrow composed of the ventral furrow and the furrow-like dorsal depression. Species of Ctenogobius inhabit shallow sand and mud bottoms in bays and estuaries, often in tidal or mangrove environments. Many species are euryhaline and enter brackish water; some inhabit fresh and brackish water (Pezold, 2004, 2022). Most species of Ctenogobius are facultative air-breathing (Froese & Pauly, 2023).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFF5FFBA0236FBF063BCF83B.taxon	description	Figure 42 k – o Material 13 specimens Gelasian, Moin FM, Atlantic Costa Rica, PPP 3256 (figured specimens NMB P 15822 - 23). Discussion Te fossil otoliths (Fig. 42 n – o) placed here in C. smaragdus are smaller than the extant ones (Fig. 42 km) studied. However, they show a very similar otolith shape, the ventral furrow closely approaching the anterior and posterior rims and a deepened sulcus with a well-developed ostial lobe and a rounded caudal tip, which we consider to be typical for the species. (See figure on next page.) Fig. 42 a – c Ctenogobius boleosoma (Jordan & Gilbert, 1882), extant, TCWC 18306.13, Texas, Brazoria, SL 25, 27 and 29 mm. d – e Ctenogobius aff. boleosoma (Jordan & Gilbert, 1882), Moin FM, Limón, Costa Rica, PPP 1772, NMB P 15820 - 21. f – g Ctenogobius fasciatus Gill, 1858, extant; f CAS 18574, 09 ° 07 ’ N 77 ° 57 ’ W, Panama, San Blas, Mosquito Bay; g TCWC 20534.10 Trinidad and Tobago, SL 24 mm. h Ctenogobius manglicola (Jordan & Starks, 1895), extant, USNM 407631, 13 ° 16 ’ N 87 ° 44 ’ W. i – j Ctenogobius shufeldti (Jordan & Eigenmann, 1887), extant, BMNH 1985.6.30.37 – 39, Baia de Sepetiba, Brazil. k – o Ctenogobius smaragdus (Valenciennes, 1837); k – m extant, USNM 371772, 23 ° 02 ’ N 80 ° 03 ’ W, mangrove swamp; n – o Moin FM, Limón, Costa Rica, PPP 3256, NMB P 15822 - 23. p Ctenogobius stigmaticus (Poey, 1860), extant, USNM 357714, Rio de Janeiro, SL 50 mm. q – t Ctenogobius sp. 1, Springvale FM, Trinidad, KR- 1216, NMB P 15824 - 27. u Ctenogobius sp. 2, Moin FM, Limón, Costa Rica, PPP 1772, NMB P 15828. v – X Evorthodus lyricus (Girard, 1858), extant, BMNH 1980.9.22.48 - 51, Florida, Sebastian Inlet. y Evorthodus minutus Meek & Hildebrand, 1928, extant, USNM 408804, 13 ° 13 ’ N 87 ° 31 ’ W	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCBFF84018CFF50603DFCD8.taxon	description	Figure 42 q – t Material 38 specimens Zanclean: 34 specimens Springvale FM, Trinidad, KR- 1216 (figured specimens NMB P 15824 - 27); 4 specimens Cayo Agua FM, Atlantic Panama, PPP 1276. Discussion Tese are otoliths characterized by a high-bodied shape (OL: OH = 0.83 – 0.9) and the straight, slightly inclined posterior rim, and a distinct ventral furrow curving around the cauda to near the narrow dorsal depression. Tese otoliths resemble the extant C. stigmaticus (Fig. 42 p) except for the development of the ventral furrow. We have refrained from a species description because of the relatively small size of the otoliths, the presence of only a few well-preserved specimens in the otherwise large assemblage, and the limited knowledge of otoliths of extant Ctenogobius species.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCBFF84018CFC3066D5FDB8.taxon	description	Figure 42 u Material 1 specimen Gelasian, Moin FM, Atlantic Costa Rica, PPP 1772 (NMB P 15828). Discussion Te single, somewhat eroded otolith is remarkable for its depressed, shallow dorsal rim. Otherwise it shows the expanded pre- and postdorsal projections which are typical for several extant species of the genus like C. boleosoma (Fig. 42 a – c), C. fasciatus (Fig. 42 f – g) and C. shufeldti (Fig. 42 i – j).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCBFF850236FD9061C3FE78.taxon	description	Figure 43 d – e Holotype NMB P 15829 (Fig. 43 d), Tortonian, Yaviza FM, Pacific Panama, Darien, PPP 1607. Paratypes 2 specimens Tortonian, Tuira FM, Pacific Panama, Darien, 1 specimen PPP 1554 (NMB P 15830), 1 specimen PPP 1171 (NMB P 15831). Etymology Named after the province Darien of Panama where the otoliths of this species have been found. Diagnosis Otolith size up to 1.95 mm in length. OL: OH = 0.95. Otolith ventrally box-shaped and dorsally with strongly expanded postdorsal region. Preventral and postventral angles orthogonal; postdorsal projection broad, protruding. Sulcus relatively short, with moderate ostial lobe, narrowed at ostial-caudal joint and rounded caudal tip; OL: SuL = 1.9 – 2.0; sulcus inclination angle 6 – 10 °. Small subcaudal iugum located within ventral collum-like ingression. Ventral furrow distinct, regularly curved and relatively distant from ventral rim of otolith. Description Otolith size up to 1.95 mm in length (holotype). OH: OT = 3.7. Otolith high-bodied with rectangular, box-shaped ventral part and high expanded postdorsal part; dorsal rim with relatively high, rounded predorsal angle, ascending to massive postdorsal region; postdorsal projection broad, high, distinctly protruding. Anterior rim slightly inclined towards dorsal, straight. Posterior rim slightly inclined towards ventral, with weak concavity below postdorsal projection. Ventral rim perfectly straight and horizontal, with orthogonal pre- and postventral angles. All rims smooth or slightly undulating. Inner face distinctly bent in horizontal direction. Sulcus relatively short, relatively shallow, sole-shaped with moderate ostial lobe, its ostial tip tapering; sulcus margins at ostial-caudal joint broadly but not strongly ingressing, thereby narrowing sulcus; cauda slightly widened and with rounded tip; sulcus inclination angle 6 – 10 °; OL: SuL = 1.9 – 2.0. Small subcaudal iugum in ventral ingression of sulcus margin at collum (ventral ingression of sulcus margin). Dorsal depression relatively indistinct; ventral furrow distinct, relatively distant from ventral rim of otolith, anteriorly approaching anterior rim of otolith, posteriorly curving behind cauda. Outer face slightly concave, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCBFF850236FD9061C3FE78.taxon	discussion	Discussion Ctenogobius darienensis closely resembles otoliths of the extant C. sagittula (Fig. 43 a – c) in the specific development of the postdorsal region, the very specific shape of the sulcus and even the expression of the ventral furrow. Te main differences are the more compressed shape (OL: OH = 0.95 vs 1.05 – 1.15), the straight predorsal rim (vs middorsal concavity), the non-protruding rounded predorsal angle and the smaller subcaudal iugum. Ctenogobius darienensis looks like one might envisage the special features of C. sagittula have developed from more plesiomorphic otolith morphologies.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCAFF850236FE506167FADF.taxon	description	(Fig. 44 g) from the East Pacific; G. oceanicus (Pallas, 1770) (Fig. 44 d) from the West Atlantic; and G. occidentalis (Boulenger, 1909) from the East Atlantic. Otoliths are not known from the two remaining West Atlantic species, i. e., G. munizi Vergara, 1978, endemic to Cuba, and G. stomatus Starks, 1913, endemic to Brazil. Te otoliths of Gobionellus are characterized by a high-bodied shape (OL: OH = 0.8 – 1.0) with a much-expanded postdorsal region, a commonly strongly sculptured dorsal rim, a relatively short and somewhat simplified sole-shaped sulcus (OL: SuL = 1.8 – 2.2) with a low inclination angle (3 – 10 °), and the ventral furrow running relatively distant from the ventral rim of the otolith and commonly opening to the anterior-ventral and posterior-ventral rims of the otolith. Te otoliths are usually thin with a concave to flat outer face. A subcaudal iugum is usually not present, except in G. microdon, where it is large and well developed. Specimens of Gobionellus are collected in shallow estuarine, tide pool, and mangrove environments (Pezold, 2004).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFC8FF87018CFA906171F898.taxon	description	Figure 44 k – k Holotype NMB P 15833 (Fig. 44 k), Tortonian, Gatun FM, Atlantic Panama, PPP 2168. Paratypes 6 specimens Tortonian: 4 specimens same data as holotype (NMB P 15834 - 37); 2 specimens Tuira FM, Pacific Panama, Darien, PPP 1137 (NMB P 15838 - 39). Referred specimens 12 specimens Tortonian: 2 specimens Tuira FM, Pacific Panama, Darien, PPP 1137; 1 specimen Yaviza FM, Pacific Panama, Darien, PPP 1553; 9 specimens same data as holotype. Etymology Named after the Guna, the indigenous Indians inhabiting the region of the type location. Diagnosis Otolith size up to 1.05 mm in length. OL: OH = 0.85 – 0.92. Otolith high-bodied, with expanded, angular middorsal angle. Preventral angle pointed, postventral angle orthogonal; postdorsal projection rounded, slightly protruding. Sulcus large, with moderate ostial lobe; OL: SuL = 1.65 – 1.8; sulcus inclination angle 9 – 14 °. No subcaudal iugum. Ventral furrow distinct, regularly curved in specimens below 1 mm in length; anteriorly and posteriorly open in larger specimens. Description Otolith size up to 1.05 mm in length (holotype). OH: OT = 3.3. Otolith high-bodied; dorsal rim high, with angular, relatively high predorsal angle, per- and postdorsal rims ascending to elevated to relatively sharp middorsal angle in straight lines; postdorsal projection broad, high, slightly protruding. Anterior rim slightly inclined towards dorsal, with weak concavity at level of ostium. Posterior rim inclined towards ventral, with weak concavity below postdorsal projection. Ventral rim straight to slightly bent and horizontal, with sharp preventral and orthogonal postventral angles. All rims smooth. Inner face only bent in horizontal direction. Sulcus wide, relatively large, slightly deepened, sole-shaped with moderate ostial lobe, its ostial tip tapering; sulcus margins at ostial-caudal joint broadly but not strongly ingressing, thereby narrowing sulcus; cauda with rounded tip; sulcus inclination angle 9 – 14 °; OL: SuL = 1.65 – 1.8. No subcaudal iugum. Dorsal depression wide, with indistinct margins; ventral furrow distinct, relatively close to ventral rim of otolith, anteriorly and posteriorly opening to ventral portions of anterior and posterior rims in large specimens (Fig. 44 k), curving upward behind cauda in smaller specimens (Fig. 44 l – q). Outer face slightly convex, smooth. Discussion Otoliths of G. gunae are smaller than those of the extant G. thompsoni, but the development of the ventral furrow touching the anterior and posterior rims in morphologically mature specimens indicates that specimens larger than 1 mm in length are mature, and hence G. gunae is indeed a relatively small species. Gnatholepis gunae resembles the extant G. thompsoni in many aspects but differs in the smaller size, the lower OL: OH ratio (0.85 – 0.92 vs 0.8 – 0.88), and the smooth otolith rims. Te occurrence of G. gunae in the Tortonian of tropical America confirms the long presence of this genus in the region.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCFFF800236FDD061BDFA38.taxon	description	Figure 45 c – d Material 2 specimens Zanclean, Springvale FM, Trinidad, KR- 1216 (NMB P 15840 - 41). Discussion Tese two specimens from the Early Pliocene of Trinidad resemble otoliths of the extant Clevelandia ios (Fig. 45 a – b) in the shape of the shallow sulcus, the smooth inner face, and the course of the thin ventral furrow, which tapers anteriorly below the ostium and curves posteriorly upward behind the cauda to connect to the shallow and vague dorsal depression. Te otoliths differ from those of Clevelandia ios in being more compressed (OL: OH = 0.85 – 0.9 vs 0.95 – 1.0) and showing an elevated dorsal rim. It is, however, also possible that these otoliths represent juvenile forms of a species of which otoliths have not yet been found. Te allocation to Clevelandia is therefore tentative until more specimens have been found. Clevelandia ios is the only known species of the genus; it is an euryhaline species inhabiting estuaries, lagoons, and tidal environments on sand and mud bottoms from British Columbia to Baja California (Froese & Pauly, 2023).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCFFF820236FA50666CFE98.taxon	description	Figure 45 n Holotype NMB P 15842 (Fig. 45 n), upper Zanclean, upper Cayo Agua FM, Atlantic Panama, PPP 1188. Etymology Named after the Laguna de Chiriqui located in the back of the Cayo Agua Island, the type locality. Diagnosis Otolith size 1.65 mm in length. OL: OH = 1.0. Otolith high-bodied, with strongly expanded postdorsal region. Preventral angle orthogonal, postventral angle recessed; postdorsal projection angular, not protruding beyond middle part of posterior rim. Sulcus large, shallow, with nivellated ostial lobe and broad indentation of ventral sulcus margin at collum; OL: SuL = 1.65; sulcus inclination angle 20 °. No subcaudal iugum. Ventral furrow narrow, curving around sulcus to join narrow, furrow-like dorsal depression. Description Otolith size 1.65 mm in length (holotype). OH: OT = 3.6. Otolith high-bodied; dorsal rim high, with broad, strongly expanded postdorsal region, slightly depressed rounded predorsal angle, and straight, relatively steeply inclined predorsal rim; postdorsal projection angular, slightly protruding but not beyond tip of posterior rim. Anterior rim nearly vertical, straight. Posterior rim with concavity below postdorsal projection followed towards ventral by distinct bulge. Ventral rim straight, horizontal, with orthogonal preventral angle and deeply recessed postventral angle. All rims smooth. Inner face convex, particularly bent in horizontal direction, relatively smooth. Sulcus moderately wide, shallow, sole-shaped with low, rounded ostial lobe, its ostial tip tapering; sulcus margins at ostial-caudal joint broadly ingressing ventrally; cauda with rounded tip; sulcus inclination angle 20 °; OL: SuL = 1.65. No subcaudal iugum. Dorsal depression very narrow, close to dorsal rim of otolith, relatively indistinct; ventral furrow narrow, relatively close to ventral rim of otolith, turning around sulcus and joining up to dorsal depression. Outer face slightly concave, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCFFF820236FA50666CFE98.taxon	discussion	Discussion Evermannia chiriquiensis is based on a single, well-preserved, and large otolith that matches extant otoliths of Evermannia in many diagnostic features, i. e., the shape of the sulcus with the low ostial lobe and the ventral indentation, the sulcus being shallow, the shape of the posterior rim, the convex and smooth inner face, and the pattern of the ventral furrow joining up with the narrow, furrow-like dorsal depression. Otoliths are known of three of the four extant species of Evermannia, plus a species identified as Evermannia sp. that originates from near the type location of E. longipinnis and may represent that species (Fig. 45 i). Te otoliths of the extant species of Evermannia show a variety of diagnostic features. Tus, the shape of the dorsal rim of E. chiriquiensis resembles the shape in E. sp. (Fig. 45 i) and E. zosterura (Fig. 45 m) but is more strongly expanded. Te specific shape of the posterior rim most resembles that of Evermannia sp. Te sulcus shape resembles that of E. erici (Fig. 45 j) and E. zosterura, while that of Evermannia sp. shows a rounded ostial lobe, and the sulcus of E. panamensis (Fig. 45 k – l) is very small. Evermannia chiriquiensis differs from the otoliths of all extant species in the strongly inclined sulcus (20 ° vs ≤ 15 °).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCDFF830236FEF063E0FE98.taxon	description	Figure 45 o – p Holotype NMB P 15843 (Fig. 45 o), Burdigalian, Cantaure FM, Venezuela, PPP 2545. Paratypes 2 specimens same data as holotype (NMB P 15844 - 45). Referred specimen 1 tentatively assigned specimen, Tortonian, Tuira FM, Pacific Panama, Darien, PPP 1154. Etymology From problematicus (Latin) = problematical, owing to tentative nature of the generic assignment of the species. Diagnosis Otolith size up to 1.5 mm in length. OL: OH = 1.05 – 1.1. Otolith shape nearly quadratic, with flat dorsal rim and pointed, short postdorsal projection. Preventral angle orthogonal, high. Sulcus large, moderately shallow, with rhombic ostium; OL: SuL = 1.7 – 1.8; sulcus inclination angle 8 – 12 °. No subcaudal iugum. Ventral furrow narrow, curving around sulcus to join narrow dorsal depression. Description Otolith size up to 1.5 mm in length (holotype 1.45 mm). OH: OT = 3.4. Otolith nearly quadratic in shape with flat dorsal rim; postdorsal projection narrow, pointed, slightly protruding. Anterior and posterior rims nearly vertical, straight, without concavities. Ventral rim slightly bent, horizontal, with rounded preventral and postventral angles. All rims slightly undulating. Inner face slightly convex. Sulcus relatively large and wide, moderately shallow, ostium diamond-shaped with angular ostial lobe and angular ventral expansion and angular tip; cauda with rounded tip; sulcus inclination angle 8 – 12 °; OL: SuL = 1.7 – 1.8. No subcaudal iugum. Dorsal depression narrow, close to dorsal rim of otolith, relatively indistinct; ventral furrow narrow, relatively close to ventral rim of otolith, turning around sulcus and joining up to dorsal depression. Outer face flat, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCDFF830236FEF063E0FE98.taxon	discussion	Discussion Evermannia? problematica resembles in outline the smooth otolith shape found in the extant E. erici (Fig. 45 j) and E. panamensis (Fig. 45 k – l). Te sulcus shape is similar to that of the otolith of Evermannia sp. (Fig. 45 i) from the northern Gulf of California. However, we are not entirely certain whether this species really represents the genus Evermannia or possibly an extinct lineage in the Gillichthys clade.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCCFF83018CFE30665FFB3F.taxon	description	Figure 46 e Holotype NMB P 15846 (Fig. 46 e), Tortonian, Manzanilla FM, San José Member, Trinidad, PPP 2670. Etymology Named after the occurrence of this species in the Caribbean while the genus today is strictly East Pacific. Diagnosis Otolith size 1.65 mm in length. OL: OH = 1.15. Otolith oval, with regularly curved dorsal rim and expanded postdorsal region. Preventral and postventral angles rounded, recessed; postdorsal projection broad, rounded, protruding. Sulcus large, wide, shallow, with low ostial lobe; OL: SuL = 1.8; sulcus inclination angle 15 °. No subcaudal iugum. Ventral furrow narrow, distant from ventral rim of otolith, curving around sulcus to join narrow, furrow-like dorsal depression. Description Otolith size 1.65 mm in length (holotype). OH: OT = 3.5. Otolith with a generally rounded, more or less oval shape. Dorsal rim high, regularly curved, broadly expanded postdorsal region, slightly depressed rounded and slightly protruding predorsal angle; postdorsal projection broad, rounded, protruding. Anterior rim slightly curved, slightly inclined toward ventral, without concavity. Posterior rim with mild concavity below postdorsal projection followed towards ventral by slight bulge. Ventral rim straight, horizontal, with broadly rounded and recessed preventral and postventral angles. All rims relatively sharp in lateral view, slightly undulating. Inner face convex, particularly bent in horizontal direction, relatively smooth but area outside of central field defined by circular ventral furrow and dorsal depression somewhat undulating. Sulcus large, wide, shallow, sole-shaped with low, rounded ostial lobe, its ostial tip rounded; cauda with rounded tip; sulcus inclination angle 15 °; OL: SuL = 1.8. No subcaudal iugum. Dorsal depression narrow, close to sulcus and distant from dorsal rim of otolith, relatively distinct but not deep; ventral furrow narrow, relatively close to sulcus and distant from ventral rim of otolith, turning around sulcus posteriorly and joining up to dorsal depression; curvature anteriorly interrupted by tip of sulcus. Outer face slightly concave, relatively smooth. Discussion Te single, superbly preserved but thin and fragile otolith bears all the typical hallmarks of otoliths of the extant Gillichthys mirabilis (Fig. 46 a – b), including the general shape of the otolith and the sulcus and the pattern of the ventral furrow and its connection to the dorsal depression, which is distant from the dorsal rim of the otolith. Gillichthys caribbaeus differs from G. mirabilis in being slightly more compressed (OL: OH = 1.15 vs 1.2 – 1.25), the relatively large sulcus (OL: SuL = 1.8 vs 2.0 – 2.2), and the ventral furrow running more distant from the ventral rim of the otolith. Clearly, G. caribbaeus represents an extinct species of the lineage that today is strictly East Pacific but was in the Atlantic during the Late Miocene. In fact the otolith suggests that G. caribbaeus is more closely related to G. mirabilis than the latter is to the second extant species, G. seta (Fig. 46 d). Most of the fishes in the Gillichthys clade inhabit shallow coastal to estuarine environments. Gillichthys mirabilis inhabits shallow coastal environments such as mudflats and bays and is well known for its facultative air-breathing (Todd & Ebeling, 1966).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCCFF8D0236FB5060C0FA38.taxon	description	Figure 46 j Holotype NMB P 15847 (Fig. 46 j), Zanclean, Cubagua FM, Venezuela, PPP 3055. Etymology Named after the type locality on the Araya Peninsula in Venezuela. Diagnosis Otolith size 1.15 mm in length. OL: OH = 1.05. Otolith rectangular with rounded angles. Postdorsal (See figure on next page.) Fig. 46 a – c Gillichthys mirabilis Cooper, 1864, extant; a, c LACM coll. Fitch; b LACM 57250 - 3, California, Marina del Rey, SL 108 mm. d Gillichthys seta (Ginsburg, 1938), extant, LACM coll. Fitch, LTF 701207 - 2, Gulf of California. e Gillichthys caribbaeus n. sp., holotype, NMB P 15846, Manzanilla FM, Trinidad, PPP 2670. f Lepidogobius lepidus (Girard, 1858), extant, coll. Schwarzhans leg. Fitch. g Typhlogobius californiensis Steindachner, 1879, extant, coll. Schwarzhans leg. Fitch. h – i Ilypnus gilberti (Eigenmann & Eigenmann, 1889), extant, LACM coll. Fitch. j Ilypnus arayanensis n. sp., holotype, NMB P 15847, Cubagua FM, Venezuela, PPP 3055. k Quietula ycauda (Jenkins & Evermann, 1889), extant, LACM coll. Fitch. m – n Quietula guamasiae (Jenkins & Evermann, 1889), extant, LACM 680204 - 1, coll. Fitch, Cabo Tepopa, Sonora, SL 52 and 70 mm. o – p Quietula rueberi n. sp., Manzanilla FM, Trinidad, Cb. 1633; o holotype, NMB P 15848; p paratype, NMB P 678 projection broad, rounded, not protruding. Sulcus small, shallow, slightly forward positioned, with rounded, widened ostium and short, narrow cauda; OL: SuL = 2.5; sulcus inclination angle 12 °. No subcaudal iugum. Ventral furrow narrow, close to ventral rim of otolith, curving around sulcus to join narrow dorsal depression. Description Otolith size 1.15 mm in length (holotype). OH: OT = 2.5. Otolith rectangular with regularly rounded predorsal, postdorsal, preventral and postventral angles. Dorsal rim shallow, with weak, broad, rounded middorsal angle. Postdorsal projection broad, rounded, not protruding. Anterior rim straight, vertical, without concavity. Posterior rim with very small concavity at its middle. Ventral rim slightly curved, horizontal, with broadly rounded preventral angle and rounded, recessed postventral angle. All rims thick in lateral view, slightly undulating. Inner face convex, smooth. Sulcus small, shallow, droplet-shaped with rounded ostium and narrow, short cauda; sulcus inclination angle 12 °; OL: SuL = 2.5. No subcaudal iugum. Dorsal depression narrow, shallow, positioned across middle of dorsal field; ventral furrow narrow, relatively close to ventral rim of otolith, turning around sulcus anteriorly and posteriorly and joining up to dorsal depression. Outer face about as convex as inner face, smooth.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFCCFF8D0236FB5060C0FA38.taxon	discussion	Discussion Ilypnus arayanensis resembles the extant I. gilberti (Fig. 46 h – i) in otolith shape, the small, shallow sulcus, the smooth inner face, and the narrow ventral furrow close to the ventral rim of the otolith and joined to the dorsal depression. It differs primarily from the extant species in the lower dorsal rim and the slightly eccentrically forward-positioned sulcus. Otoliths of the second extant species of the genus, Ilypnus luculentus (Ginsburg, 1938) from the Gulf of California, are not known. Ilypnus gilberti is found on mudflats and shallow bays, and males are known to guard the eggs (Froese & Pauly, 2023).	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
492D87AAFFC2FF8D018CFA50606CF918.taxon	description	Figure 46 o – p Holotype NMB P 15848 (Fig. 46 o), Tortonian, Manzanilla FM, San José Member, Trinidad, Cb. 1633.	en	Schwarzhans, Werner W., Aguilera, Orangel A. (2024): Otoliths of the Gobiidae from the Neogene of tropical America. Swiss Journal of Palaeontology (13) 143 (1): 1-129, DOI: 10.1186/s13358-023-00302-5, URL: https://doi.org/10.1186/s13358-023-00302-5
