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Palaeo
Ichthyologica
Systematik
Morphologie
Palökologie
Paläogeographie
Stratigraphie
Verlag Dr. Friedrich Pfeil
Orangel A. AGUILERA, Werner SCHWARZHANS and Philippe BÉAREZ:
Otoliths of the Sciaenidae from the Neogene of tropical America
Werner SCHWARZHANS and Orangel A. AGUILERA:
Otoliths of the Ophidiiformes from the Neogene of tropical America
14
14
Palaeo
Ichthyologica
Systematik
Morphologie
Palökologie
Paläogeographie
Stratigraphie
Orangel A. AGUILERA, Werner SCHWARZHANS and Philippe BÉAREZ:
Otoliths of the Sciaenidae from the Neogene of tropical America
Werner SCHWARZHANS and Orangel A. AGUILERA:
Otoliths of the Ophidiiformes from the Neogene of tropical America
Verlag Dr. Friedrich Pfeil
•
München, April 2016
ISSN 0724-6331
•
ISBN 978-3-89937-207-6
14
Begründet und herausgegeben von Dr. Friedrich H. PFEIL, München
In der Reihe Palaeo Ichthyologica werden Originalarbeiten und Dissertationen zur Systematik, Morphologie, Palökologie,
Paläogeographie und Stratigraphie rezenter und fossiler Fische veröffentlicht.
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Copyright
© 2016 by Verlag Dr. Friedrich Pfeil, München
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Druck: Winterwork, Grimma
Printed in the European Union
ISSN 0724-6331
ISBN 978-3-89937-207-6
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Die Deutsche Nationalbibliothek verzeichnet diese Publikation in der Deutschen Nationalbibliografie;
detaillierte bibliografische Daten sind im Internet über
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Palaeo Ichthyologica
14
1-124 München, April 2016154 figs., 18 pls.
3
Palaeo Ichthyologica 14
Contents
Orangel A. AGUILERA and Werner SCHWARZHANS:
Preface ....................................................................................................................................... 5
Orangel A. AGUILERA, Werner SCHWARZHANS and Philippe BÉAREZ:
Otoliths of the Sciaenidae from the Neogene of tropical America ................................................ 7
Werner SCHWARZHANS and Orangel A. AGUILERA:
Otoliths of the Ophidiiformes from the Neogene of tropical America ........................................... 91
Palaeo Ichthyologica
14
1-124 München, April 2016154 figs., 18 pls.
4
Palaeo Ichthyologica 14
5
Palaeo Ichthyologica 14
Palaeo Ichthyologica 14, p. 5-6
© 2016 Verlag Dr. Friedrich Pfeil, München
Palaeo Ichthyologica
14
5-6 München, April 20161 fig.
Preface
This now is the second compendium of articles dealing with
the fossil otoliths that were collected from Central America
during several decades. Following the initial study of the
otoliths of the mesopelagic lantern fishes (Myctophidae)
published as the first part of the ongoing investigations
in Palaeo Ichthyologica volume 13 (SCHWARZHANS &
AGUILERA, 2013) and a review of the otoliths of the ma-
rine catfish family Ariidae (AGUILERA, MORAES-SANTOS,
COSTA, OHE, JARAMILLO & NOGUIERA, 2013), we are
now covering the drum fishes (Sciaenidae) and cusk eels
(Ophidiiformes).
Most of the otoliths studied for this compendium were
obtained from young Neogene strata, chiefly of late Miocene
(Tortonian) to middle Pleistocene (Calabrian) age. Clearly,
one has to expect many Recent species to be retrieved from
so young strata. Therefore, a good command of the otoliths
of the relevant extant fishes of the region (and beyond) is
a prerequisite for everybody attempting to identify such
otolith assemblages. Consequently we have focused first
on selected groups of fishes for which adequate knowledge
of Recent otoliths is available or could be made available
with reasonable efforts. Fortunately, they are also the most
common and diverse groups of fishes and they represent
the prime environments sampled, as is exemplified in the
graph below.
We plan to continue with our task of describing the
Neogene otoliths of tropical America in further studies.
However, the knowledge base is often much less compre-
Sciaenidae
inner shelf
estuarine
freshwater
common in Cantaure FM,
Pirabas FM, Manzanilla FM,
Urumaco FM
common in Brasso FM,
Cantaure FM, Gurabo and
Mao FMs, Onzole FM
-
dominant in Chagres FM,
Cubagua FM, Bowden FM,
Burica FM,
Boca del Toro FMs
common in Angostura FM,
Onzole FM, Canoa FM,
Cercado FM
common in Pirabas FM
common in Angostura FM,
Cantaure FM, Cercado FM,
Gatun FM, Manzanilla FM,
Canoa FM
-
dominant in Pirabas FM,
Urumaco FM, Rio Banano FM
inner shelf
estuarine
freshwater
mesopelagic
benthopelagic
benthic on shelf
bathybenthic
benthic on shelf
reef associated
Batrachoididae
Ophidiidae
Myctophidae
Ariidae
6
Palaeo Ichthyologica 14
hensive of otoliths of extant species of those fish families
still outstanding and therefore identifications at the species
level may not always be possible in the extent desirable. It
is our aim to try to accomplish an adequate data base of
pertinent Recent species as good as possible in the course
of our quest to extract the maximum possible information
of this extraordinary collection.
Orangel Aguilera* and Werner Schwarzhans**
Niteroi and Hamburg, May 15
th
2015
References
Aguilera, O. A., Heloisa Moraes-Santos, H., Costa, S.,
Ohe, F., Jaramillo, C. & Nogueira, A. 2013. Ariid
sea catfishes from the coeval Pirabas (Northeastern Brazil),
Cantaure, Castillo (Northwestern Venezuela), and Castil-
letes (North Colombia) formations (early Miocene), with
description of three new species. – Swiss Journal of Palae-
ontology, 132: 45-68. doi: 10.1007/s13358-013-0052-4.
Schwarzhans, W. & Aguilera, O. A. 2013. Otoliths of the
Myctophidae from the Neogene of tropical America. –
Palaeo Ichthyologica, 13: 83-150.
* Orangel
aguilera, Universidade Federal Fluminense, Instituto de Biologia, Programa de Pós-graduação em Biología Marinha. Campus
do Valonguinho, Outeiro São João Batista, s/nº. CEP: 24020-141, Niterói, Rio de Janeiro, Brasil; e-mail: orangel.aguilera@gmail.com
** Werner schwarzhans, Ahrensburger Weg 103, D-22359 Hamburg, and Natural History Museum of Denmark, Zoological
Museum, Universitetsparken 15, DK-2100 Copenhagen; e-mail: wwschwarz@aol.com
7
Palaeo Ichthyologica 14
Palaeo Ichthyologica 14, pp. 7-90, 10 figs., 18 pls.
© 2016 Verlag Dr. Friedrich Pfeil, München
Palaeo Ichthyologica
14
7-90 München, April 201610 figs., 18 pls.
Otoliths of the Sciaenidae
from the Neogene of tropical America
Orangel A. Aguilera*, Werner Schwarzhans**
and Philippe Béarez***
Abstract
Fossil otoliths of the family Sciaenidae are described from Neogene strata from a wide array of locations in tropical America rang-
ing from Ecuador to Panama, Costa Rica, Dominican Republic, Colombia, Venezuela, Trinidad and Brazil. A total of 85 species are
recognized by means of otoliths, 20 of which represent Recent species also recorded as fossil. Two otolith-based fossil genera and 30
otolith-based fossil species are described new to science. The new genera are:
†
Amazonasciaena n. gen.,
†
Protonebris n. gen. The new
species are, in the sequence described: Callaus cubaguanus n. sp., Umbrina abbreviata n. sp., Umbrina bananensis n. sp., Umbrina
laxa n. sp., Umbrina opima n. sp., Umbrina sublima n. sp., Umbrina surda n. sp., Polycirrhus jaramilloi n. sp., Polycirrhus mustus n. sp.,
Larimus angosturae n. sp., Larimus humboldti n. sp., Larimus pandus n. sp., Nebris dioneae n. sp., Protonebris sanchezi n. sp., Cynoscion
latiostialis n. sp., Cynoscion prolixus n. sp., Cynoscion scitulus n. sp., Isopisthus acer n. sp., Plagioscion ultimus n. sp., Atractoscion odeai
n. sp., Ophioscion amphiamericanus n. sp., Ophioscion inflaticauda n. sp., Ophioscion transitivus n. sp., Stellifer abbreviatus n. sp.,
Stellifer acerbus n. sp., Stellifer bicornutus n. sp., Stellifer depressifrons n. sp., Stellifer onzole n. sp., Xenotolithus semiostialis n. sp. and
Xenotolithus sineostialis n. sp.
The otolith record shows that the northern shores of South America with their diverse paleoenvironmental setting and the large and
shifting river drainage systems of the paleo-Amazon and paleo-Orinoco offered favorable conditions for the evolution of the Sciaeni-
dae, a group with marked preference for estuarine and other near shore environments. Evidently, tropical America was an evolutionary
center of sciaenids since at least early Miocene times. The presence in the marine Neogene of several species belonging to genera that
nowadays are restricted to freshwater, i. e. Aplodinotus and Plagioscion, indicates that freshwater is a secondary habitat colonized late
in the evolution of sciaenids.
Diversification of otolith-based sciaenid lineages is high since early Miocene representing all relevant extant tribes and dominated
by genera indigenous to the Americas. We observed also, however, that certain lineages became extinct primarily in the Caribbean due
to either shifts and expansions of the large river discharge systems during the late Miocene or to a yet insufficiently understood extinc-
tion event at the end of the late Pliocene, about 1.5 to 2 million years after the closure of the Isthmus of Panama at about 3.5 Ma.
The extinct lineages comprise five fossil otolith-based sciaenid genera, one of which,
†
Xenotolithus, represents an entire extinct group
thought to be related to the Stelliferinae with a highly specialized otolith morphology.
Finally, the presence, diversity and abundance of Neogene sciaenids from both sides of the Isthmus of Panama allowed us to study
the timing and effects of the forced allopatric speciation in this family, which was caused by the gradual emersion of the Isthmus. In
the Recent, there is not a single species of Sciaenidae recorded from both sides of the Isthmus of Panama. We found that the youngest
amphi-American species as well as the earliest occurrence of geminate species point to a separation of eastern Pacific and Caribbean line-
ages in sciaenids predating the final closure of the Pacific-Atlantic seaway and the uplift of the Isthmus of Panama (~3.5 Ma) by 2 to 5 Ma.
Our findings support the exchange of marine biota, represented here by the fish family Sciaenidae, between the Atlantic and Pacific
oceans in tropical America during the late Miocene Tortonian. We also conclude however that the marine biota were already separated,
possibly environmentally induced, to a degree during latest Miocene and early Pliocene that was sufficient to trigger a geminate specia-
tion.
Key words: Otoliths, Miocene, Pliocene, Sciaenidae, Central America, northern South America, tropical West Atlantic
* Orangel A. Aguilera, Universidade Federal Fluminense, Instituto de Biologia, Programa de Pós-graduação em Biología Marinha e
Ambientes Costeiros. Campus do Valonguinho, Outeiro São João Batista, s/nº. CEP: 24020-141, Niterói, Rio de Janeiro, Brasil;
e-mail: orangel.aguilera@gmail.com
** Werner Schwarzhans, Ahrensburger Weg 103, D-22359 Hamburg, and Natural History Museum of Denmark, Zoological Museum,
Universitetsparken 15, DK-2100 Copenhagen; e-mail: wwschwarz@aol.com
*** Philippe Béarez, UMR 7209 CNRS/MNHN, Département « Écologie et gestion de la biodiversité », Muséum national d’histoire na-
turelle, 55 rue Buffon, F-75231 Paris cedex 05, France; e-mail: bearez@mnhn.fr
8
1. Introduction .................................................................... 9
2. Recent comparative material ........................................... 10
3. Otolith terminology ........................................................ 10
4. Depository, conventions and techniques applied .............. 10
5. Acknowledgements ......................................................... 11
6. Regional Geology and Localities ..................................... 11
7. Systematic Part ............................................................... 22
Family Sciaenidae .......................................................... 22
Subfamily Pachyurinae .................................................... 22
Subfamily Protosciaeninae .............................................. 22
Genus Protosciaena ....................................................... 24
Protosciaena brasiliensis ........................p. 24 – Pl. 1, Figs. 1-5
Protosciaena neritica ...........................p. 24 – Pl. 1, Figs. 6-10
Protosciaena trewavasae ....................p. 24 – Pl. 1, Figs. 11-12
“Protosciaena” bathytatos ..................p. 24 – Pl. 1, Figs. 13-15
Subfamily Sciaeninae ..................................................... 24
Tribe Ctenosciaenini ....................................................... 24
Genus Ctenosciaena ...................................................... 25
Ctenosciaena gracilicirrhus ....................p. 25 – Pl. 2, Figs. 1-4
Ctenosciaena peruviana .........................p. 25 – Pl. 2, Figs. 2-5
Tribe Pogoniini new tribe ................................................ 25
Genus
†
Amazonasciaena n. gen .................................... 26
†
Amazonasciaena rossettiae ................. p. 26 – Pl. 3, Figs. 4-6
Genus Aplodinotus ......................................................... 26
Aplodinotus hoffmani ......................... p. 26 – Pl. 2, Figs. 17-20
Aplodinotus santosi ............................p. 28 – Pl. 2, Figs. 14-16
Genus Sciaenops ........................................................... 28
Sciaenops reyesi ....................................p. 28 – Pl. 3, Figs. 2-3
Sciaenops sp. ........................................... p. 28 – Pl. 3, Figs. 7
Tribe Sciaenini ................................................................ 28
Genus Callaus ............................................................... 30
Callaus cubaguanus ..........................p. 30 – Pl. 3, Figs. 12-16
Genus
†
Frizzellithus ........................................................ 30
†
Frizzellithus longecaudatus ...............p. 30 – Pl. 3, Figs. 17-21
Genus Umbrina .............................................................. 31
Umbrina abbreviata n. sp. .....................p. 31 – Pl. 4, Figs. 4-9
Umbrina bananensis n. sp. ................p. 31 – Pl. 4, Figs. 14-17
Umbrina laxa n. sp. ...........................p. 31 – Pl. 4, Figs. 18-21
Umbrina opima n. sp. ............................p. 32 – Pl. 5, Figs. 1-6
Umbrina sublima n. sp. ......................p. 34 – Pl. 4, Figs. 10-13
Umbrina surda n. sp. ...........................p. 34 – Pl. 5, Figs. 8-11
Umbrina sp. ......................................p. 34 – Pl. 5, Figs. 12-13
Tribe Equetini ................................................................. 36
Genus
†
Equetulus ........................................................... 36
†
Equetulus amazonensis .........................p. 36 – Pl. 6, Figs. 1-4
†
Equetulus davidandrewi .....................p. 36 – Pl. 6, Figs. 5-10
†
Equetulus fitchi ................................p. 36 – Pl. 6, Figs. 11-15
Genus Pareques ............................................................. 38
Pareques jungi ...................................p. 38 – Pl. 5, Figs. 14-18
Tribe Menticirrhini new tribe ........................................... 38
Genus Menticirrhus ........................................................ 38
Menticirrhus chaoi ...................................................... 38
Menticirrhus sp. ...................................... p. 38 – Pl. 6, Figs. 16
Tribe Lonchurini .............................................................. 39
Genus Lonchurus ............................................................ 39
Lonchurus lanceolatus ........................p. 39 – Pl. 6, Figs. 18-20
Genus Paralonchurus ...................................................... 39
Paralonchurus rathbuni ........................... p. 39 – Pl. 6, Figs. 17
Genus Polycirrhus ........................................................... 39
Polycirrhus jaramilloi n. sp. ...................p. 39 – Pl. 7, Figs. 9-15
Polycirrhus mustus n. sp. ..........................p. 4 – Pl. 7, Figs. 1-3
Polycirrhus schwarzhansi ......................p. 4 – Pl. 7, Figs. 16-22
Polycirrhus trinidadensis .....................p. 42 – Pl. 7, Figs. 16-22
Subfamily Lariminae ....................................................... 42
Tribe Larimini ................................................................. 42
Genus Larimus ............................................................... 42
Larimus angosturae n. sp. ..................p. 42 – Pl. 8, Figs. 11-17
Larimus argenteus ..................................p. 44 – Pl. 8, Figs. 1-2
Larimus breviceps ..................................p. 44 – Pl. 8, Figs. 3-7
Larimus fasciatus .................................p. 45 – Pl. 9, Figs. 8-10
Larimus gatunensis ................................p. 45 – Pl. 8, Figs. 8-9
Larimus humboldti n. sp. ........................p. 45 – Pl. 9, Figs. 1-
3
Larimus pacificus ...............................p. 46 – Pl. 9, Figs. 11-15
Larimus pandus n. sp. ............................p. 46 – Pl. 9, Figs. 4-5
Larimus sp. 1 ........................................... p. 46 – Pl. 8, Figs. 8
Larimus sp. 2 ........................................p. 46 – Pl. 9, Figs. 6-7
Genus Nebris ................................................................. 48
Nebris dioneae n. sp. ......................p. 48 – Pl. 10, Figs. 14-15
Nebris sp. (juveniles) ...........................p. 48 – Pl. 10, Figs. 6-7
Genus Paranebris ........................................................... 48
Paranebris bauchotae ........................p. 48 – Pl. 10, Figs. 8-11
Paranebris steurbauti ...........................p. 50 – Pl. 10, Figs. 2-5
Genus
†
Protonebris n. gen. ............................................ 50
†
Protonebris sanchezi n. sp. .................... p. 50 – Pl. 10, Figs. 1
Subfamily Cynoscioninae ................................................ 51
Tribe Cynoscionini .......................................................... 51
Genus Cynoscion ........................................................... 51
Cynoscion analis ................................. p. 51 – Pl. 11, Figs. 1-2
Cynoscion arenarius ............................ p. 51 – Pl. 11, Figs. 3-5
Cynoscion latiostialis n. sp. ..............p. 51 – Pl. 11, Figs. 14-17
Cynoscion nothus ................................p. 52 – Pl. 11, Figs. 6-9
Cynoscion prolixus n. sp. .................p. 52 – Pl. 11, Figs. 11-13
Cynoscion scitulus n. sp. ..................p. 54 – Pl. 11, Figs. 18-19
Cynoscion sp. ....................................... p. 54 – Pl. 11, Figs. 20
Genus Isopisthus ............................................................ 55
Isopisthus acer n. sp. ...........................p. 55 – Pl. 12, Figs. 4-6
Isopisthus remifer .................................p. 55 – Pl. 12, Figs. 2-3
Genus Macrodon ........................................................... 55
Macrodon spina ......................................................... 55
Genus Plagioscion .......................................................... 55
Plagioscion marinus ...........................p. 55 – Pl. 12, Figs. 7-10
Plagioscion travassosi ........................... p. 56 – Pl. 12, Figs. 11
Plagioscion urumacoensis ........................ p. 56 – Pl. 13, Figs. 2
Plagioscion ultimus n. sp. .................p. 56 – Pl. 12, Figs. 12-16
Tribe Atractoscionini ....................................................... 58
Genus Atractoscion ........................................................ 58
Atractoscion odeai n. sp. .....................p. 58 – Pl. 13, Figs. 3-6
Subfamily Micropogoniinae ............................................ 60
Tribe Micropogoniini ....................................................... 60
Genus Micropogonias .................................................... 60
Micropogonias altipinnis ......................... p. 60 – Pl. 13, Figs. 9
Micropogonias coatesi .........................p. 60 – Pl. 13, Figs. 7-8
Micropogonias furnieri .......................... p. 60 – Pl. 13, Figs. 10
Subfamily Stelliferinae .................................................... 60
Genus
†
Protolarimus ...................................................... 61
†
Protolarimus henrici ...........................p. 61 – Pl. 14, Figs. 1-3
†
Protolarimus lundbergi .......................... p. 61 – Pl. 14, Figs. 4
†
Protolarimus? mauryae .......................... p. 62 – Pl. 14, Figs. 5
Tribe Odontoscionini ...................................................... 62
Genus Bairdiella ............................................................ 62
Bairdiella ronchus ................................... p. 62 – Pl. 14, Figs. 7
Bairdiella sp. .....................................p. 64 – Pl. 14, Figs. 8-10
Genus Odontoscion ....................................................... 64
Odontoscion dentex ............................... p. 64 – Pl. 14, Figs. 6
Tribe Stelliferini ............................................................... 64
Genus Ophioscion ......................................................... 64
Ophioscion amphiamericanus n. sp. p. 64 – Pl. 16, Figs. 1-7
Ophioscion inflaticauda n. sp. ..............p. 66 – Pl. 15, Figs. 7-9
Ophioscion transitivus n. sp. ............... p. 68 – Pl. 15, Figs. 1-3
Genus Stellifer ................................................................ 68
Stellifer abbreviatus n. sp. ....................p. 68 – Pl. 17, Figs. 1-2
Stellifer acerbus n. sp. ........................... p. 69 – Pl. 13, Figs. 13
Stellifer bicornutus n. sp. ......................p. 69 – Pl. 17, Figs. 3-4
Stellifer depressifrons n. sp. ..................... p. 70 – Pl. 17, Figs. 5
Stellifer onzole n. sp. .............................. p. 70 – Pl. 17, Figs. 6
Lapilli of Stelliferini ......................................................... 72
Ophioscion sp. (lapillus) ......................... p. 72 – Pl. 18, Figs. 3
Stellifer sp. (lapillus) ............................p. 72 – Pl. 18, Figs. 1-2
Tribe indet. of Stelliferinae .............................................. 74
Genus
†
Xenotolithus ....................................................... 74
†
Xenotolithus retrolobatus ..................p. 74 – Pl. 18, Figs. 9-12
†
Xenotolithus sasakii ........................p. 74 – Pl. 18, Figs. 13-16
†
Xenotolithus semiostialis n. sp. ............. p. 74 – Pl. 18, Figs. 17
†
Xenotolithus sineostialis n. sp. .........p. 75 – Pl. 18, Figs. 18-19
8. Faunal reconstruction ..................................................... 75
8.1 Sciaenid otoliths from the Neogene of tropical
America – expression from a center of diversity ........ 75
8.2 Miocene sciaenid otoliths in South American as
indicators of the paleoenvironment .......................... 78
8.3 Tropical American sciaenid lineages that became
extinct during Neogene ............................................ 79
8.4 Amphi-American sciaenid species and geminate
sciaenid species pairs .............................................. 83
9. References ..................................................................... 85
Contents and Index
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
9
Palaeo Ichthyologica 14
The Miocene and Pliocene sediments of tropical America are
rich in otoliths. Otolith assemblages have been described
from Panama (SCHUBERT, 1908), Mexico (WEILER, 1959),
Trinidad (NOLF, 1976), Venezuela (NOLF & AGUILERA,
1998; AGUILERA & RODRIGUES DE AGUILERA, 2001,
2003, 2004), Costa Rica (LAURITO, 1996), Western Ama-
zonia basin (MONSCH, 1998), the Dominican Republic
(NOLF & STRINGER, 1992) and Eastern Amazonia basin
(AGUILERA et al., 2014) and have been mentioned from
Panama in MARTIN & DUNN (2000) and AGUILERA &
RODRIGUES DE AGUILERA, (1999), from Jamaica in FITCH
(1969) and from Ecuador by BIANUCCI et al. (1993, 1997),
LANDINI et al. (1991, 2001, 2002) and CARNEVALE et
al. (2011). Recently, SCHWARZHANS & AGUILERA (2013)
dedicated a volume with similar format to the present one
to the description of the myctophid otoliths from the Neo-
gene of tropical America and announced further volumes
to follow on other teleost groups. This now represents the
second volume dealing with Neogene sciaenid otoliths from
tropical America.
The purpose of this study is to describe and document
the rich Miocene and Pliocene sciaenid otolith assemblages
collected by the senior author, Carlos Jamarillo and other
colleagues mostly under the scope of the Panama Pale-
ontological Project (PPP) in Brazil, Colombia, Costa Rica,
Ecuador, Panama, Trinidad and Venezuela, and to review
previously published material. Previously described sciae-
nid otoliths by NOLF & AGUILERA (1998), AGUILERA &
RODRIGUES DE AGUILERA (2003, 2004) and AGUILERA,
SCHWARZHANS, MORAES-SANTOS & NEPOMUCENO
(2014) have been available for review. The sciaenid otoliths
studied by NOLF (1976) had been reviewed by SCHWAR-
ZHANS (1993). The material studied by NOLF & STRINGER
(1992) was also available for review, but that of weiler
(1959) is not accessible.
All in all, over 1300 identifiable specimens of sciaenid
otoliths have been studied representing 85 species, thereof
30 new, 10 in open nomenclature and 20 Recent species
recorded as fossil. The fossil sciaenid otoliths from South
America have proven to represent a highly diverse assem-
blage covering all major Recent groups of the region plus
one extinct genus group (Xenotolithus group).
Sciaenid otoliths are large, mostly robust and highly
diverse. That has attracted scientists and non-scientists for
a long time, and it is in fact a sciaenid otolith that was
described first from the fossil record (LAWLEY, 1876), even
before KOKEN (1884) really initiated fossil otolith research.
The Sciaenidae were also the first family in which otoliths
were used as an additional character for taxonomic purposes
(CHU, LO & WU, 1963; MOHAN, 1969; TREWAVAS, 1977;
CHAO, 1978, 1986, 2001; CHAO et al., 2001; SASAKI,
1989; BÉAREZ & SCHWARZHANS, 2013). SCHWARZHANS
(1993) wrote an extensive monograph on Recent and fossil
sciaenid otoliths covering almost all extant genera and a
large proportion of extant species as well as all then known
fossil species. This work is used as the main reference for
comparison with Recent sciaenid otoliths, being comple-
mented in certain instances with new data, particularly for
the genera Isopisthus, Nebris, Paranebris, Ophioscion and
Stellifer. Sciaenid otoliths are primarily recognized by their
advanced heterosulcoid sulcus organization composed of
a wide to very wide shallow ostium and a steeply curved
cauda of varying depth. This principal pattern is seen in its
most plesiomorphic expression in such genera as Pachy-
urus or Umbrina and has developed into a wide variety of
morphologies across the entire family, which clearly is an
aspect of the advanced sound and hearing abilities of the
“croakers” (Sciaenidae) and which is also expressed in the
specialization of their gas bladder patterns (TREWAVAS,
1977; CHAO, 1978; SASAKI, 1989), although there is
no readily recognizable link between the morphological
developments of both organs. Many of the specializations
seen in sciaenid otoliths are unique, but owing to their,
unresolved, functional morphology may still have occurred
more than once in the evolution of fishes of this family. For
instance the tadpole shaped, widened caudal tip observed
in the genera Protosciaena, Micropogonias, Macrodon
and the genera of the Otolithes and Collichthys Groups
(of SCHWARZHANS, 1993), many of which are not con-
sidered to be related, and seemingly represent homoplasy
of a highly unique feature. We hope that such examples
will serve as a warning to those postulating phylogenetic
relationships of fishes based on otoliths alone.
According to ESCHMEYER & FONG (2015), Recent Sci-
aenidae comprise 291 valid species world wide, distributed
among 67 genera. More than half of the species (169)
and the genera (36) occur in and around North and South
America, making it the richest region of the world in sciaenid
biodiversity. Sciaenids form an important faunal element in
the warm shallow seas and estuaries of the world. They are
particularly adapted to soft and sandy bottomed near-shore
marine and brackish environments, where they sometimes
occur in large shoals, but are generally missing from oceanic
islands, with a few exceptions, such as the Galapagos and
Juan Fernandez Islands in the Eastern Pacific or Bermuda
Island in the Western Atlantic. Only few species and genera
are known from deeper shelf environments. In addition,
there are four genera recognized from the freshwaters of
the Americas, and one from the freshwaters of Indochina.
The genera and species of the family Sciaenidae show a
high degree of regionalization with three individual centers
of distribution. The by far highest diversity is observed in the
Americas, followed by the Indo-West Pacific and a much
smaller center along the shores of West Africa. Interaction
between these regions apparently is very limited, with all
species confined to their respective regions. Even at the
genus level, there is only one, Umbrina, which is distributed
across all three regions and very few across more than one
region. In the case of the Sciaenidae of the Americas, the
only genus other than Umbrina with a wider distribution
is Atractoscion, known from the Pacific shores of North
America and the Indo-West Pacific. However, the relation-
ships of “Protosciaena” bathytatos in the Caribbean deep
water remains unresolved and there are reliable records
of indigenous American genera in the fossil record of the
Neogene of Europe, such as Cynoscion, Sciaenops, Seriphus
and possibly others.
The fossil record of sciaenid otoliths in the Neogene of
tropical America likewise is very rich and its knowledge base
is rapidly increasing, practically with every new collection
being uncovered. It currently comprises 85 species of 30
genera, including six extinct otolith-based genera. Another
28 species have been recorded from the Neogene of NE-
America by MÜLLER (1999), arriving at a total of 117 species
in the fossil record, which includes 31 extant species also
recorded as fossils. This impressive number of fossil sciaenid
1. Introduction
10
species recorded by otoliths from the American Neogene
is all represented by indigenous American genera, except
Atractoscion and Umbrina. It clearly documents that the
main splitting of the three sciaenid centers of diversity had
occurred prior to the Neogene. It is of interest to note that
North America is also rich with sciaenid otoliths in Eocene
strata. Unfortunately, we have no Eocene or older fossil
records from tropical America to enable a comparison with
North America.
Another aspect emerging from this study is the pres-
ence in the marine Neogene of several species belonging
to genera that nowadays are restricted to freshwater, i. e.
Aplodinotus and Plagioscion. They indicate that freshwater
is a secondary habitat colonized late in the evolution of
sciaenids. Finally, the presence, diversity and abundance
of Neogene sciaenids from both sides of the Isthmus of
Panama allowed the timing and effects of the forced al-
lopatric speciation that was caused by the gradual emersion
of the Isthmus, to be studied in this family. In the Recent,
there is not a single species of Sciaenidae recorded from
both sides of the Isthmus of Panama.
2. Recent comparative material
A reliable identification of fossil otoliths of Miocene and
Pliocene age is only possible through detailed comparison
with relevant Recent otoliths of the species of the region.
This is also important with respect to the assessment of the
interspecific diversity in a genus, ontogenetic changes, or
intraspecific variability. For the purpose of this study, the
description of Recent sciaenid otoliths from the Americas
in the monograph of SCHWARZHANS (1993) required
only minor complementation for certain genera to form an
adequate basis for comparisons.
The Recent fish, from which the otoliths were obtained,
have mostly been obtained from the collections of AMS
(Sydney), BMNH (London), HUMZ (Hakodate), ISH and
ZMH (both Hamburg), LACM (Los Angeles), USNM (Wash-
ington D.C.), ZMUC (Copenhagen) and the collections of
L. CHAO (Taipeh), W. AGUIRRE (Chicago) and the authors
(OA donated to WS, WS and PB at MNHN).
3. Otolith terminology
The otolith terminology is used as shown in text-figure 1.
The morphological nomenclature has been established by
KOKEN (1884) and WEILER (1942).
Otolith measurements (Text-fig. 1) were made with a
camera lucida, following SCHWARZHANS (1993) and as
described in BÉAREZ & SCHWARZHANS (2013): Otolith
length (OL) is the greatest anteroposterior distance; otolith
height (OH) is the greatest dorsoventral distance; otolith
thickness (OT) is the greatest thickness in dorsal or ventral
view; the curvature of the inner face (Z) is the maximum
distance of the ventral rim of the otolith to the arc connecting
anterior and posterior tips of the otolith; sulcus length (SUL)
is the greatest distance from the anterior rim of the ostium
to the posterior-most extension of the cauda; ostium length
(OCL) is the greatest distance from the anterior rim of the
ostium to the posterior rim of the postostial lobe; ostium
height (OCH) is the greatest dorso-ventral height of the
ostium; cauda length (CCL) is measured from the upper
(dorsal) ostial-caudal joint to the posterior-most extension
of the cauda; distance from ostium to downturned cauda
(W) is measured as the minimal horizontal distance of the
posterior-most position of the postostial lobe to the anterior
margin of the downturned part of the cauda; length of
proximal part of cauda (X) is measured from lower (ventral)
ostial-caudal joint to the incursion of the ventral margin
of the cauda at the turning point; height of cauda (Y) is
measured from the incursion of the ventral margin of the
cauda at the turning point to the caudal tip. Calculations
of different ratios follow SCHWARZHANS (1993): OL : OH;
OH : OT; CCL : OCL; OCL : OCH, W as % of SUL (ostial-
caudal interspace), X : Y (the caudal curvature index), and
Z as % of OL (the curvature index of the inner face).
4. Depository, conventions and techniques applied
All photographs of otoliths were made with a Wild M400
photomacroscope. A Canon EOS digital camera was
mounted on a specifically designed adapter on the photo-
tubus of the photomacroscope. Photos were taken remotely
controlled from a computer. Multiple sequentially focused
photos were taken in regular field of depth steps of each
view. The focused parts of each photograph representing a
specific field of depth range were then automatically fused
with the HeliconFocus software of HeliconSoft (Charkov,
Ukraine) in order to produce a continuous field of depth.
Finally, the resulting photographs were digitally retouched
as far as possible without afflicting alterations to the mor-
phology of the shown otolith (removal of sand grains,
incrustations, pigmentations).
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
11
Palaeo Ichthyologica 14
For optimal comparison purposes, all figures show
otoliths from the right side. Photos of left otoliths have been
mirror imaged and are marked with an (r) for reversed.
All holotypes of newly described species are deposited
at Naturhistorisches Museum, Basel, Switzerland (NMB P
catalog), except those from Castilletes, which are deposited
at the Mapuka Museum of Universidad del Norte, Colom-
bia (MUN and STRI catalog). Type-material of previously
described species by the senior author are deposited at the
Universidad Nacional Experimental Francisco de Miranda,
Coro, Venezuela (UNEFM) and the Museu Paraense Emilio
Goeldi (MPEG-V), Brazil. Location numbers refer to the Pan-
ama Paleontology Project (PPP) from Ecuador, Panama and
Venezuela, locations catalogued by the Naturhistorisches
Museum Basel (NMB), locations collected by K. Rohr (K.R.)
and C. J. Campbell (Cb.) both from Trinidad and locations
collected by Orangel Aguilera (OA) in Venezuela.
5. Acknowledgements
We would like to thank the following colleagues for their help in
collecting the material described herein, often under arduous con-
ditions: Jorge Carrillo, Anthony Coates, Carlos de Gracia, Jeremy
Jackson, Carlos Jaramillo, Peter Jung, Aaron O´Dea, Félix Rodri-
guez, Dione Rodrigues de Aguilera, Julio Reyes, Marcelo Sánchez
and Rodolfo Sánchez, and the crew of the R/V Urraca for their
kind assistance. The Smithsonian Tropical Research Institute from
Panama, the Francisco de Miranda University from Venezuela, the
Paleontological Institute and Museum from the University of Zurich,
the Museu Paraense Emilio Goeldi and the Universidade Federal
Fluminense from Brazil provided funds and logistical support. Walter
Etter and Olivier Schmidt (NMB) are thanked for making available
the fossil otoliths from Trinidad. Trevor H. Worthy (Adelaide) is
thanked for improving the English.
In respect to the recent comparative material used for this study,
the authors would like to cordially thank for their support and advice
W. Aguirre (Chicago), L. Chao (Taipeh), O. Crimmen and the late
E. Trewavas (BMNH), A. Gomes and T. Berriel (AGEVAP-ICMBio Rio
de Janeiro), R. Feeney (LACM), P. Moeller and J. Nielsen (ZMUC),
F. Ohe (Seto City, Japan), J. Paxton (AMS), A. Post (formerly ISH),
S. Raredon and J. Williams (USNM), S. Santos (UFPA), K. Sasaki
(Kochi) and R. Thiel (ZMH).
SUL
OCL
CCL
OL
OH
Y
X
OCH
anterior
anterior
outer face
dorsal
ventral
inner face
OT
Z
OL
ostium
cauda
umbo
ostium
cauda
postostial lobe
W
Text-figure 1. Schematic drawing of a sciaenid otolith to visualize terminology and measurements (amended after BÉAREZ & SCHWARZ-
HANS, 2013). Abbreviations: OL = otolith length, OH = otolith height, OT = otolith thickness, OCL = ostium length, OCH = ostium
height, CCL = cauda length, SUL = sulcus length, W = distance rear margin of ostium to downturned portion of cauda, X = length of
horizontal portion of cauda, Y = height of downturned portion of cauda, Z = curvature of inner face.
6. Regional Geology and Localities
The otoliths described in the following were obtained from
a variety of Neogene formations from various locations, to
which the stratigraphic information has been referred on
various occasions in the relevant literature. Many of them
have been mentioned and discussed in detail in SCHWAR-
ZHANS & AGUILERA (2013). For those, description is kept to
a minimum while those not represented in the earlier paper
are more extensively discussed. All stratigraphical data to
the geochronological time scale, ages and names follow the
most recent International Chronostratigraphic Chart (ICS;
COHEN et al. 2013), edition 2015/01, published by the
International Commission on Stratigraphy. This differs from
the version shown in SCHWARZHANS & AGUILERA (2013)
in the recognition of the Gelasian stage as early Pleistocene
12
“above“
Cerro Negro Mb
Cerro Verde Mb
Cumana
FM
Caujarao FM
Castillo FM Cantaure FM
Brasso FM
Pirabas FM
Falcon
Lara
Northeastern
Venezuela
Gros Morne FM
San José Mb
Telemaque Mb
Montserrat Mb
Southern
Range
Bragança
Viseu Basin
Chucunaque FM
Tuira FM
Yaviza
FM
Darien Bowden Cibao Valley
PleistoceneP lioceneM iocene
Atlantic
Caribbean coast
Caribbean coast
Pacific coast
Canoa
FM
Moin
FM
Swan
Cay FM
Cayo Agua
FM
Shark Hole
Point FM
Escudo de
Veraguas FM
Upper Coastal
Group
Bowden
FM
Mao
FM
Cercado
FM
Gurabo FM
Cubagua FM
Chagres FMGatun FM
Nancy Point
FM
Tobabe
FM
Jama
FM
Armuelles
FM
La Peñi-
ta FM
Burica FM
Angostura FM Onzole FM
Manzanilla FM
Springvale
FM
Baitoa FM
Castilletes FM
Jimol
FM
Urumaco FM
Rio Ban-
ano FM
Central
Range
Costa Rica Panama
Jamaica
Dominican
Republic
EcuadorPanama
Ma
Ma
Epoch Stage
Nanno-
plankton
planktonic
Foramini-
fera
Limon
Esmeralda-
Borbon
Bastimentos
Cayo Agua
Valiente
Peninsula
Escudo de
Veraguas
Panama Canal
Basin
Burica
Peninsula
NN 1
NN 21-
NN 20
N 4
N 5
N 7-6
N 8
N 9
N 10
N 12
N 14
N 15
N 16
N 17
N 18
N 19
N 20
/
N 21
N 22
N 11
N 13
P 22
NP 25
NN 2
NN 3
NN 5
NN 6
NN 7
NN 8
NN 9
NN 10
NN 11a
NN 11b
NN 16
NN 19
NN 12
NN 15
-
NN 13
NN 18-
NN 17
NN 4
Aquitanian
Oligocene
Bastimen-
tos FM
Colombia Venezuela Trinidad Brazil
Alta Guajira
Peninsula
Holocene
Ionian
Clabri-
an
Gelasi-
an
Piacen-
zian
ZancleanMessinianTortonianSerravallianLanghianBurdigalian
L
L
L
M
E
M
M
E
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
Text-figure 2. Stratigraphic correlation chart depicting stratigraphic positions of otoliths obtained. Time scale is from ICC; for strati-
graphic correlation see detailed references in chapter 6.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
13
Palaeo Ichthyologica 14
“above“
Cerro Negro Mb
Cerro Verde Mb
Cumana
FM
Caujarao FM
Castillo FM Cantaure FM
Brasso FM
Pirabas FM
Falcon
Lara
Northeastern
Venezuela
Gros Morne FM
San José Mb
Telemaque Mb
Montserrat Mb
Southern
Range
Bragança
Viseu Basin
Chucunaque FM
Tuira FM
Yaviza
FM
Darien Bowden Cibao Valley
PleistoceneP lioceneM iocene
Atlantic
Caribbean coast
Caribbean coast
Pacific coast
Canoa
FM
Moin
FM
Swan
Cay FM
Cayo Agua
FM
Shark Hole
Point FM
Escudo de
Veraguas FM
Upper Coastal
Group
Bowden
FM
Mao
FM
Cercado
FM
Gurabo FM
Cubagua FM
Chagres FMGatun FM
Nancy Point
FM
Tobabe
FM
Jama
FM
Armuelles
FM
La Peñi-
ta FM
Burica FM
Angostura FM Onzole FM
Manzanilla FM
Springvale
FM
Baitoa FM
Castilletes FM
Jimol
FM
Urumaco FM
Rio Ban-
ano FM
Central
Range
Costa Rica Panama
Jamaica
Dominican
Republic
EcuadorPanama
Ma
Ma
Epoch Stage
Nanno-
plankton
planktonic
Foramini-
fera
Limon
Esmeralda-
Borbon
Bastimentos
Cayo Agua
Valiente
Peninsula
Escudo de
Veraguas
Panama Canal
Basin
Burica
Peninsula
NN 1
NN 21-
NN 20
N 4
N 5
N 7-6
N 8
N 9
N 10
N 12
N 14
N 15
N 16
N 17
N 18
N 19
N 20
/
N 21
N 22
N 11
N 13
P 22
NP 25
NN 2
NN 3
NN 5
NN 6
NN 7
NN 8
NN 9
NN 10
NN 11a
NN 11b
NN 16
NN 19
NN 12
NN 15
-
NN 13
NN 18-
NN 17
NN 4
Aquitanian
Oligocene
Bastimen-
tos FM
Colombia Venezuela Trinidad Brazil
Alta Guajira
Peninsula
Holocene
Ionian
Clabri-
an
Gelasi-
an
Piacen-
zian
ZancleanMessinianTortonianSerravallianLanghianBurdigalian
L
L
L
M
E
M
M
E
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
14
instead of late Pliocene. Color-coding does not follow ICC,
and is selected to allow for reasonable visual differentiation
of the time interval depicted. Correlation with nannoplankton
and planktonic foraminifera zones is based on GRADSTEIN
et al. (2004). Text-figure 2 summarizes the local formations
in a regional stratigraphic frame highlighting the intervals
from which otoliths have been obtained for this study.
6.1 Brazil
The northeastern-most Atlantic coast of South America of
Brazil is located on a stable platform not directly affected by
the Cenozoic South American orogenesis (ALMEIDA et al.,
2000; ROSSETTI, 2006; MOHRIAK, 2008) of the Andean
uplift (GREGORY-WODZICKI, 2000; GARZIONE et al.,
2008; SHEPHARD, et al. 2010). The Oligocene-Miocene
deposits along the equatorial margin of Brazil correspond
to the Cametá Sub-Basin and the São Luís-Grajaú Basin
and, along the eastern margin, such deposits are found
in the Cumuruxatiba, Jequitinhonha, Camamu/Almada,
Recôncavo/Tucano, Sergipe/Alagoas, Paraíba and Potiguar
basins, and directly overlay Precambrian crystalline base-
ment or Paleozoic basins (ROSSETTI et al., 2013). The
equatorial margin and part of the eastern coasts of Brazil
form a narrow but extensive belt of more than 5000 km
in length. The Oligocene-Miocene deposits exposed along
the equatorial margins are lithostratigraphically known as
Pirabas and Barreiras formations. The Pirabas Formation
of late Oligocene to early Miocene age is exposed only
along the equatorial margin and consists of mixed carbon-
ate and siliciclastic deposits. The thickness of the Pirabas
Formation at Salinopolis is calculated by FREIMANN (2014)
at about 130 m (120 m bellow the present sea level) and
the paleoenvironment as interpreted by ROSSETTI & GÓES
(2004) and AGUILERA et al. (2013) consisted of inner
shelf, restricted shelf-lagoon, shoreface-foreshore, and
mangrove-mud flat deposits.
Along the equatorial and eastern margins of Brazil, two
main transgressive episodes were recognized by ROSSETTI
et al. (2013), one in the Oligocene-Miocene and the other
in the middle Miocene. The fact that these events on the
passive margin of Brazil show a reasonable correspondence
with sea-level highstands observed elsewhere suggests that
these transgressive deposits might be in part associated with
eustatic fluctuations (ROSSETTI et al., 2013). Estimates of
the Miocene sea-level highstand magnitude however vary
from as high as 100-130 m (HAQ et al., 1987; HAQ &
AL-QAHTANI, 2005) to as low as 50 m (JOHN et al., 2004;
KOMINZ et al. 2008) or even slightly below present-day
sea level (MILLER et al., 2005). The profile sections from
the Pirabas FM at Salinopolis, Primavera and Capanema
and the nature of the fossil assemblages relate best to the
sea level postulated by KOMINZ et al. (2008).
The Pirabas FM (MAURY, 1925) is of late Oligocene
to early Miocene age and consists of carbonate coquina
rocks of an offshore platform environment (grainstone
and consolidated packstone, stratified wackestone to
packstone and laminated mudstone). In addition, littoral
facies (shoreface/foreshore), marginal lagoons, restricted
platform environments (gray to olive mudstone and con-
glomeratic sandstone) and mangrove estuarine lagoons
(dark mudstone, massif or laminated) have been recorded
(GOES et al., 1990; ROSSETTI, 2001; ROSSETTI & GOES,
2004; ROSSETTI et al., 2013; AGUILERA et al., 2013a,b).
The planktonic foraminifera association from the rocks
from which otoliths have been extracted of the Pirabas Fm.
(PETRI, 1957; FERNANDES, 1984, 1988; FERNANDES &
TAVORA, 1990; TAVORA & FERNANDES, 1999) correlates
with the early Miocene, Aquitanian to early Burdigalian,
N4 to N5 plankton foraminiferal biozones (BLOW, 1969).
A palynological investigation of a sample from the Pirabas
Fm. has revealed an age not younger than early Miocene
(com. pers. C. JARAMILLO).
Locations (Text-fig. 3): Atalaia beach, Salinópolis Mu-
nicipality, Pará state (see AGUILERA et al. 2014, fig. 5 for
further details)
6.2 Ecuador
A string of forearc and foreland basins developed during
the Neogene in the areas immediately to the west and
east of the Andean Cordillera, with their subsidence his-
tory related to events of relative sea-level rise in magnitude
and timing which were not necessarily contemporaneous
with the overall global pattern (HERNÁNDEZ et al., 2005;
CUITIÑO et al., 2012). Along the Ecuadorian coast, a
series of sedimentary basin developed which are character-
ized by a narrow platform shelf bordered seawards by the
deep Pacific Ocean margin and landwards by the western
slopes of the Andes, much as today. The fluctuations in
the subsidence of the basin floor were caused by regional
tectonic events and relate to variations in rates and local
punctuation of subduction which resulted in the formation
of marine terraces. The Bordón and Manabí basins are
filled with partially exposed late Cenozoic shallow marine
sediments (EVANS & WHITTAKER, 1982; BRISTOW &
HOFSTETTER, 1986; ROSANÍA, 1989; BIANUCCI et al.,
1997; CATALAMESSA et al., 2007; ALVAREZ et al., 2009).
These basins are located in the east, in forearc positions
of the modern convergent North Andes block-Nazca plate
boundary and are separated from the Manabí Basin to the
east by the early Pliocene coastal range (DI CELMA et al.
2005). The Canoa Basin at Cabo San Lorenzo in central
Ecuador was an island separated from mainland Ecuador
during the early Pleistocene (DI CELMA et al. 2005; PEDOJA
et al., 2006) and then connected to the mainland later in
the Pleistocene by an east-west trending land bridge that
separated the Manta Basin to the north from the Canoa
Basin to the south. During the middle and late Pleistocene
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
15
Palaeo Ichthyologica 14
interglacial periods, flooding of the shelf created a small,
southwesterly facing coastal embayment of the Canoa Basin,
which has been progressively in-filled by cyclically stacked
shallow-marine strata (PEDOJA et al., 2006).
The Angostura FM (STAINFORTH, 1948) is of late Mio-
cene, Tortonian age (NN 10-11), and is found in the western
Borbón Basin, northwestern coast of Ecuador. It includes
middle neritic and shallow outer neritic facies exposed along
the Rio Santiago and Punta Verde, respectively. It consists
of muddy, fine grained sandstones with mollusk shells dis-
persed in the matrix (BALDOCK, 1982; CANTALAMESSA
et al., 2007). The supposed paleoenvironment is shoreface
to inner-shelf.
Locations (Text-fig. 3): PPP 3439, PPP 3440, PPP 3441and
PPP 3444 in Cueva de Angostura; PPP 3456 in Angostura
beach; PPP 3457 in Palo Palo river bank; PPP 3459 in
Playa del Toro, Río Santiago (Playa de Oro, Selva Alegre),
Borbón. PPP 3406 in Telembí, PPP 3431 and PPP 3432 in
Quebrada Telembito, Río Cayapas, Borbón. PPP 3587 in
Favio Alvaro road cut, Manabí.
The Canoa FM, has extensive outcrops along the sea
cliff south of Manta in the Manabí Province (LANDINI et
al., 2001, 2002), and its age has been attributed to the
early Pleistocene (TSUCHI et al., 1988; WHITTAKER, 1988;
BIANUCCI et al., 1997; LANDINI et al., 2001, 2002; DI
CELMA et al., 2005). The lithology of the Canoa FM consists
mainly of sandstone and sandy mudstone, divided into two
sections (PILSBRY & OLSSON, 1941). The lower Canoa FM,
exposed on the sea cliff south of Punta Canoa, is a 36 m
thick, cyclothemic succession of bluish, chiefly fine-grained
strata that rest unconformably on Miocene rocks of the
Tosagua FM. The upper Canoa FM of 38 m thick shore face
and inner-shelf deposits, and all of them represent shallow
water deposits (DI CELMA et al., 2005).
Locations (Text-fig. 3): PPP 3514, PPP 3515, PPP 3516 and
PPP 3519 in Punta Canoa (Punta Canoa waterfall Valley);
PPP 3529 and PPP 3537 in Santa Rosa, Punta Canoa,
Manabí.
The Jama FM, has been assigned to the middle Pleistocene
(PILSBRY & OLSSON, 1941), and has an 180 m thick clastic
succession deposited in an array of terrestrial and shallow-
marine settings along the eastern margin of the Esmeraldas-
Caráquez Basin (ROSANIA, 1989). It is located along the
Bahía Jama in the Manabí Province, and is subdivided into
three main informal units (CANTALAMESSA et al., 2005).
The lower Punta Pasa Borracho Member (~20 m thick) is
exposed between Punta Pasa Borracho and Punta Cabuya;
and the Punta Ballena Member (~100 m thick) is exposed
Text-figure 3. Location plate for Ecuador and Brazil.
16
from Punta Cabuya to Punta Alcatraz; the El Matal Member
(~60 m thick) is exposed between Punta Alcatraz and the
village of El Matal. The age of the Punta Ballena and El
Matal members yielded a weighted mean radioisotopic age
of 1.16 ± 0.06 Ma (CANTALAMESSA et al., 2005).
Locations (Text-fig. 3): PPP 3560, PPP 3561, PPP 3562,
PPP 3563, PPP 3565 and PPP 3566 in Punta La Ceraza,
Jama, Manabí.
The Onzole FM is assigned to the late Miocene
-
early
Pliocene (SCHULMAN et al., 1965; WHITTAKER, 1988;
CARNEVALE et al., 2011). WHITTAKER (1988) subdivided
the Onzole Formation into two main units (informally named
lower Onzole Formation and upper Onzole Formation)
separated by a regionally extensive angular unconformity of
late Miocene
-
early Pliocene age. These outcrops are situ-
ated near Punta Gorda and Esmeraldas, and are composed
mainly of fine-grained muddy turbidites, having regular
vertical sequences of sedimentary structures associated with
fining upward sequences, and bioturbation restricted mostly
to the tops of the beds. The remainder of the beds measured
consists of volcanic ash, mud pelagite, and glauconitic silts
and turbidites. Vertical sequential analyses of stratigraphic
sections for the most part show no pronounced trends in
bed thickness or grain size. Facies relations, paleontologi-
cal data, and regional geologic settings suggest sediment
accumulation on an inner trench slope in a basin situated
ocean-ward of the Pliocene trench-slope break (AALTO &
MILLER, 1999). The otoliths described here were all obtained
from the lower Onzole Formation of late Miocene age.
Locations (Text-fig. 3): PPP 3467, PPP 3470 and PPP 3471
in Las Antonias, Río Santiago; PPP 3484 in Camarones
River, Borbón.
6.3 Costa Rica
The Río Banano FM was described by TAYLOR (1975),
and CASSELL (1986) established the type section west of
the railroad bridge at Bomba on the Banano River, 14 km
south-southwest of Limon. The oldest unit of the Río Banano
FM, from which the otoliths were obtained, correlates bi-
ostratigraphically to the late early to late Pliocene, although
it may be older in the southern Limon Basin. The formation
is ~750 m thick and consists of burrow-mottled, blue-gray
siltstone and volcanic litharenite with frequent shell-rich
lenses and scattered mollusc hash. Molluscs, sand dollars,
and callianassid burrows are common.
Locations (Text-fig. 4): PPP 1726, PPP 1727, PPP 1728, PPP
1731 and PPP 1732 in La Bomba; PPP 1734 and PPP 1735
in Quitaria, Río Banano, Limón (see COATES et al., 1992,
appendix A, section 29 for further details); PPP 1777 and
PPP 1776 in Quebrada Brasso Seco, Limón (out of section).
The Moin FM (CASSELL, 1986; CASSELL & SEM GRUPTA,
1989; COATES et al., 1992) of Late Pliocene to early Pleis-
tocene age (3.0 to 1.9-1.6 Ma) is located near the mouth
of the Moin River, about six km west of Limón in Costa
Rica, and includes two members: the Empalme Mb and
the Lomas del Mar Mb. The Lomas del Mar Mb, where the
otoliths come from, is composed of blue-gray, clayey silt-
stone and calcarenite that flank beds immediately adjacent
to reef patches. Sediments are packed with small, diverse
mollusk, including vermetids, small solitary and oculinid
ahermatypic corals, bryozoans, serpulids and large echinoid
spines (COATES, 1999).
Locations (Text-fig. 4): urban construction site in Loma del
Mar Eastern, Limón.
6.4 Dominican Republic
The Baitoa FM (COOKE, 1920) of late early to early mid-
dle Miocene, Burdigalian age, is located near the Village
López in the Río Yaque del Norte, Dominican Republic. It is
composed of finer-grained silts interbedded with conglomer-
ates (SAUNDERS et al., 1986).
Locations (Text-fig. 5): NMB17287 in Río Yaque del Norte
(see SAUNDERS et al., 1986, Text-fig. 21, 22 for further
details).
The Cercado FM (COOKE, 1920) of late Miocene, late
Tortonian-Messinian age (NN 11) is found in several river
bank sections of the Cibao Valley, Dominican Republic.
It is composed of calcareous silts, which are occasionally
separated by thin, sandy conglomerates in a shallow, near
shore environment in the west (Rio Cana) and gradually
deepening eastwards (Río Gurabo and Río Mao) (SAUN-
DERS et al., 1986).
Locations (Text-fig. 5): Río Cana, locations NMB16817,
16845, Rio Gurabo, locations NMB15900, 15903, 15904,
15914, Río Mao, locations TU1294, NMB16913, 16915,
16917, 16918, 16923, 16927 downstream Maury’s Bluff 3
(see SAUNDERS et al., 1986, Text-figure 29, 33 for further
details).
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
17
Palaeo Ichthyologica 14
6.5 Panama
The Bastimentos FM (COATES et al., 1992) is of late
Pliocene, Gealsian age (NN 16-18), and its stratotype is
located in the Bastimentos Island, Atlantic Panama. It con-
sists of reef deposits sitting directly on the Miocene basalt
of the Valiente Formation, which are well exposed on Wild
Cane Key and along the coast for 2 km to the east, with
reef limestone extensively re-crystallized with many coral
colonies. A particularly well preserved example of the La
Gruta reef, packed with large and diverse coral colonies
is exposed at the base of the cliffs in three small bays at
Fish Hole, at the southern end of Long Beach. The coral
assemblage is represented for more that 50 % by extinct
species (COATES et al., 2005).
Locations (Text-fig. 4): PPP 1256 at Fish Hole promontory,
Fish Hole Bay, Bastimentos Island, Bocas del Toro (see
COATES, 1999, appendix A, section 23; COATES et al.,
2005, fig. 7 for further details).
The Cayo Agua FM (COATES et al., 1992) of early to late
Pliocene, Zanclean to Piacenzian age (NN 13-16) is found
on the Cayo Agua Island that lies about 6 km to the west
of Toro Point, Valiente Peninsula, Atlantic Panama. The
formation is well exposed along the east coast and consists
lithologically of pervasively bioturbated, muddy, silty sand-
stone with common horizons of abundant thick-shelled mol-
lusks, ahermatypic corals and wood fragments. Occasional
horizons of pebble conglomerate and very coarse-grained
volcaniclastic sandstone are common in the middle part of
the formation. A distinctive marker bed of corals occurs near
the top of the formation and is well exposed at Tiburón Point
and the unnamed point to the south. The corals appear to
be mostly free living, hermatypic, grass flat corals, as well as
ahermatypic species, associated with benthic foraminifera,
ostracods and teleostean fish. The depositional environment
is assumed in water depths of 30-50 m (COLLINS et al.,
1999 AGUILERA & RODRIGUES DE AGUILERA, 1999).
The age of the Cayo Agua Formation is dated at the base
~5.0-3.5 Ma and at the top 3.7-3.4 Ma (COATES et al.,
1992, 1999, 2005). Otoliths were collected from the lower
and middle part of the section below the limestone marker.
Locations (Text-fig. 4): PPP 1188 and PPP 2237 at Punta
Piedra Roja, Cayo Agua; PPP 1256 on Fish Hole promon-
tory, Bastimentos Island; PPP 1276 on the northeastern
coast of Isla de Popa; PPP 2225 in Punta Norte, Cayo Agua
Island; PPP 2243 in Cayo Zapatilla; PPP 2248 at Ground
Creek, Colón Island, Bocas del Toro (see COATES, 1999,
appendix A, section 17 and 19; COATES et al., 2005,
fig. 6 for details).
The Chagres FM (MACDONALD, 1915) of late Miocene,
late Tortonian to Messinian age (NN 11) consists at the
base of massive sandstones overlying the Gatun Formation
that occurs west of Colón City between Toro Point and the
Chagres River, Atlantic Panama. It is composed of massive,
homogeneous, fine-grained, silty sandstones with scattered
mollusks. The unit becomes finer grained westward for 50
km along the coast where it becomes a gray-green siltstone
with lithic and quartz grains and scattered mollusks. A distinct
basal unit, the Toro Point Member (WOODRING, 1957)
of late Tortonian age (NN 11a) is exposed west of Colón
City. It is somewhat cemented with calcite, consisting of a
coarse, cross-bedded coquina with many large barnacle
plates, but also mollusk and coral fragments, echinoid
spines, and interbedded coarse calcarenites (COATES et
al., 1992; COLLINS et al., 1996). Previous research of
fossil fish includes AGUILERA & RODRIGUES DE AGUILERA
(1999). The paleo-depth were estimated between 50 to 80 m
(COLLINS et al., 1999), but the Toro Point Mb, which has
yielded the majority of otoliths described from this FM, has
been deposited at upper bathyal depth between 200-500 m
depth and is noted for faunal influence from the East Pacific
(COLLINS et al., 1996).
Locations (Text-fig. 4): PGP 620002, (PPP 1652) (see COL-
LINS et al., 1996, fig. 1 for further details).
The Escudo de Veraguas FM (COATES et al., 1992) ranges
in age from late Pliocene, Piacenzian to early Pleistocene,
Gelasian age (NN 16-18). It is known only from the island
of the same name that lies 27 km east of Nancy Point, At-
lantic Panama. The lower section of this formation (3.5 Ma)
is found along the coast on the east side of the V-shaped
embayment situated in the central part of the north coast,
about 1 km east of Long Bay Point. It is more indurated
with very common and densely packed cemented ichno-
fossils, scattered ahermatypic corals through the middle
part, as well as variably abundant mollusks, and at the
top a coral biostrome dominated by Stylophora and sand
dollars. The paleo-depth based on ahermatypic species,
benthic foraminifera, ostracods and teleostean fish were
estimated between 100 to 150 m (COLLINS et al., 1999;
AGUILERA & RODRIGUES DE AGUILERA, 1999). The upper
section of the formation (1.8 Ma) lies along the west coast
for about one km south of Long Bay Point and consists of
gray-blue, pervasively bioturbated clayey siltstone and silty
claystone, with frequent concretions, and scattered shelly
hash, often with scattered whole and diverse mollusks and
small, cornute, ahermatypic corals (COATES et al., 1992,
1999, 2005). The top and bottom units were deposited in
an inner neritic environment, but the middle part of this
lower succession, from which most of the otoliths were
collected, has abundant benthic foraminifera indicating
deposition in outer neritic to upper bathyal water depths
(COLLINS, 1993).
Locations (Text-fig. 4): PPP 2185 in Escudo de Veraguas,
Southeastern Coast; PPP 2330 in Plantain Cay, Tobabe
Point, Valiente Península, Bocas del Toro (see COATES,
1999, appendix A, section 11; COATES et al., 2005, fig. 6
for further details).
The Gatun FM (HOWE, 1907) has first been described
from Gatun in the course of the construction of the Panama
Canal between Mount Hope and Sabanita at the northern
end of the Panama Canal. The formation is divided into
three members (WOODRING, 1957). The section of the
lower member is exposed along the trans-isthmian highway
about 12 km east of Colón City, from Sabanita to Cativa.
This member consists of burrowed, concretionary, gray-
green, tuffaceous, silty litharenite. The section of the middle
member is located in the region between Cativa and Gatun
and consists of massive, borrow-mottled, silty, tuffaceous
and occasionally calcareous litharenite, ashy siltstone and
thin conglomerate with very abundant mollusks, echinoid
fragments, and wood. The section of the upper part, of late
Miocene, middle Tortonian age (NN 10), is exposed about
2-3 km north of Gatun and to the west of the Panama Canal.
This section has yielded the otoliths described herein and is
18
characterized by clayey, biocalcarenit, tuffaceous siltstone
and fine silty litharenite that is burrow-mottled and rich in
scattered mollusks (COATES et al., 1992) and is assumed
to have been deposited at a paleo-depth between 15 to
40 m (COLLINS et al., 1999). The total thickness of the
Gatun Formation, recorded in a borehole near Colón City,
is about 500 m (WOODRING, 1957).
Locations (Text-fig. 4): PPP 04 and PPP 08 in Sabanita; PPP
223 at Dr. Martin Luther King residences; PPP 2164 and
PPP 2165 at Payardi refinery, Payardi Island; PPP 2167 in
San Judas Tadeo, Colón (see COATES, 1999, appendix B,
section 1 for further details).
Text-figure 4. Location plate for Panama and Costa Rica.
The Chucunaque FM (SHELTON, 1952) from the Tuquesa
river in Darien, consists of gray weathering, greenish blue
to black, blocky to massive, silty claystone and siltstone, with
minor thin horizons and stringers of volcanic sandstone.
Calcareous concretions are common. Calcified thalassinoid
burrows are typical. The age is estimated at about 7.1
(possibly from 8.6) to 5.6 Ma (NN 11a/NN 11b to NN 11/
NN 12 zonal boundary) (COATES et al., 2004).
Locations (Text-fig. 4): PPP 1612 in Tuquesa River, Darien
(see COATES et al. 2004, fig. 4, appendix 1, Section 42
for further details)
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
19
Palaeo Ichthyologica 14
The Tuira FM (TERREY, 1926; Esso Exploration and Pro-
duction Panama, 1970) from the Chucunaque-Tuira basin
consists of thin and regularly bedded alternations of blue
gray greywacke and arkosic sandstone with dark green to
black, silty claystone and siltstone. Abundant plant debris,
scattered small mollusks, particularly pectinids and nucula-
nids. Many units have pervasive bioturbation or thalassinoid
burrow systems. Pebble breccia, shell beds, and stringers of
rip-up clasts may occur occasionally. The lower part of the
Tuira FM cannot be assigned to a calcareous nannofossil
biozone, but represents the planktonic foraminiferal zone
N 15. Its upper part belongs to subzone NN 9b and partly
into zone NN 10 and zone N 16 respectively, ranging from
11.2 to 9.4 Ma (COATES et al., 2004). The otoliths have
been collected from the upper part.
Locations (Text-fig. 4): PPP 1164 in Río Chico; 1139
between Charco Chivo and village Marraganti, Tuquesa
River; PPP 1142 near La Pullida, Tupisa River; PPP 1554
Río Cubilele near Río Chico (see COATES et al., 2004,
fig. 4, appendix 1, sections 42, 43, 44 for further details).
PPP 1593 small tributary of Río Icuanati.
The Yaviza FM (COATES et al., 2004) of middle late Miocene
age of the nannofossil zone NN10 between 9.4 and 8.6 Ma
consists of clayey and silty, shelly sandstone and blue-gray,
volcaniclastics. Whole arcids, Pecten, turretellids and other
mollusks are abundant. Shelly hard beds are common,
especially with Turritella, Anadara, and oysters. Many units
are pervasively bioturbated, with shell filled burrows, with
occasional ahermatypic corals and common, often large
concretions (COATES et al., 2004).
Locations (Text-fig. 4): PPP 1149 in Punta Grande village,
Tupisa River; PPP 1566 near village Corozal, Río Chico,
Darien (See COATES et al. 2004, fig. 4, appendix 1, section
43 and 44 for further details).
6.6 Colombia
The Castilletes FM (RENZ, 1960; ROLLINS, 1965; MORENO
et al., 2015) outcrops in the Alta Guajira Peninsula (IRVING,
1972), northern Colombia, is composed of gray massive
mudstone, fossiliferous mudstone and siltstone, wacke-
stone to packstone biosparite, and medium-grained to
conglomeratic fossiliferous sandstone. Thick successions of
mudstone (~50 m) dominate the unit and form long valleys.
Thin beds of biosparite and sandstone (50 cm to 2 m) are
interbedded and form laterally extensive ridges. Sandstones
often present planar and cross-bedded stratification. Sandy
and silty facies increase toward the top of the formation,
forming prominent hills. Fossiliferous horizons are common
within the unit, with marine invertebrate fossils occurring
in a broad range of facies (calcareous, sandy, silty, and
muddy sediments), whereas terrestrial vertebrates tend to
be restricted to muddy sediments and are often associated
with freshwater invertebrate fossils. A late early Miocene
to early middle Miocene (late Burdigalian
-
Langhian) age
is assigned to the Castilletes Formation on the basis of
macro-invertebrate biostratigraphy and
87
Sr/
86
Sr isotope
chronostratigraphy (MORENO et al. 2015).
Locations (Text-fig. 5): 290448, 290610, 290616, 290632,
290666, 290674, 290840, 290685, 390094, 430202 and
390126 Pataju Valley, Cocinetas Basin (see MORENO et
al., 2014, figs. 3, 4 for further details).
The Jimol FM (RENZ, 1960) is composed of grey calcareous
sandstone, yellowish gray biosparite, and gray to brown
siltstones and mudstone. At the base occur 50 cm to 1 m
thick beds of coarse calcareous sandstone with ripples,
cross and planar bedding, and wackestone to packstone
biosparite dominate the sequence. There are occasional
~5 m thick beds of siltstone and mudstone in this part of
the sequence. At the top mudstone and fine-grained cal-
careous sandstone in 5 m to 20 m thick beds dominate the
sequence interbedded with 50 cm to 2 m thick beds of fine
to medium grained calcareous sandstone, and wackestone
to packstone biosparites. A late early Miocene (Burdigalian)
age is therefore assigned to the Jimol Formation on the basis
of macro-invertebrate biostratigraphy and
87
Sr/
86
Sr isotope
chronostratigraphy (MORENO et al., 2015).
Locations (Text-fig. 5): 290602, 390085 and 390090 in La
Tiendita, Paraguachón Valley (see MORENO et al., 2015,
fig. 3, 4, 5 for further details).
6.7 Trinidad
The Brasso FM (RENZ, 1942) of late early to early middle
Miocene, late Burdigalian to late Langhian age (NN 4-5,
N 7-11) has been described from Central Trinidad. It con-
sists of massive, gray to gray brown claystones and sandy
claystones. Planktonic and benthonic foraminifera indicate
paleo-depths range form ~60 m (middle neritic) to ~470 m
(deep upper bathyal) (WILSON 2003, 2005).
Locations (Text-fig. 5): PPP 2661 beyond bridge over Gorge
River; KR 9244 in Ganteaume River (see KUGLER et al.,
2001, geologic map 20 and section 21a for further details).
The Gros Morne FM (or Morne l’Enfer FM) (MACREADY,
1921) is composed of sand and silt of latest Miocene to
early Pliocene age. It is found in the Moruga Bay along
the south coast of Trinidad and corresponds roughly to the
Telémaque Mb of the Manzanilla FM in the Manzanilla Bay
in north-eastern Trinidad (HUGGINS, 2012).
Location (Text-fig. 5): K 12059 at the Mogue River.
The Manzanilla FM (WALL & SAWKINS, 1860) dates from
upper Miocene to early Pliocene and is located in the north-
ern Trinidad Basin and includes three members: Telémaque,
Montserrat and San José Mbs. The San José Mb is the
20
oldest of late Tortonian age comprising dark gray to black,
calcareous silts which have been burrowed. A rich fauna of
foraminifera and small mollusk indicates an open marine,
inner sublittoral environment of deposition (DONOVAN,
1994). The Montserrat Mb overlies the San José Mb and
is associated with a delta front environment (HUGGINS,
2012). The Telémaque Mb finally is the youngest, primarily
early Pliocene in age and is considered to be delta plain
related (HUGGINS, 2012).
Locations (Text-fig. 5): San José Mb: PPP 2667, PPP 2669,
PPP 2670, K 10842, K 12051, K 12052 and K 12054 in
cliffs along shore of the Manzanilla Bay, Cb 1633, PPP
2675, PPP 2676 at the San José River, between Gran Couva
and Brasso, SM 1399 in Pointe à Pierre (see KUGLER et
al., 2001, stratigraphic column 21b for further details);
Montserrat Mb: PPP 2665 in the Manzanilla Bay, PPP 2673
in Point Paloma beach, PPP 2677 at the San José River,
between Gran Couva and Brasso; Telémaque Mb: K 9845
at the Savaneta River.
The Springvale FM (GUPPY, 1910) is of early to late Plio-
cene age and is composed of sandstones with intercalated
mudstones (KUGLER, 1956).
Location (Text-fig. 5): K 9894 at the Savaneta River.
The Tamana FM dates as middle Miocene and comprises
massive, white to yellow, granular to crystalline limestones
with intercalated sands and clays. Various calcareous units
are included within this formation. The fauna is dominated
by benthic foraminifer and coral alga, with some corals and
mollusks (DONOVAN, 1994).
Location (Text-fig. 5): Cb 2076 at the Cascade River.
Text-figure 5. Location plate for Colombia, Venezuela and Trinidad.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
21
Palaeo Ichthyologica 14
The Cantaure FM (HUNTER & BARTOK, 1974) is of late
early Miocene, late Burdigalian to early Langhian age
(NN 4-5, N 7-8). Its stratotype is located approximately
10 km west of Pueblo Nuevo on the Paraguaná Peninsula,
Falcón State, Venezuela. Outcrops of the formation are
found west of Casa Cantaure and are composed of silty
shales interbedded with thin algal limestones and shell
beds (HUNTER & BARTOK, 1974). There is an unexposed
unit of Cantaure FM that is 48 m thick and was accessed
by a local artisan well. The section consists mainly of silty
to medium sandstone, intercalated with massive mudstone.
Research of planktonic foraminifera and calcareous nan-
nofossil have revealed late Burdigalian to early Langhian
age (DIAZ DE GAMERO, 1974; REY, 1996). A diverse fos-
sil fauna has been described, rich in mollusks, decapods
and fish (JUNG, 1965; THOMAS & MACDONALD, 1970;
NOLF & AGUILERA, 1998; AGUILERA, 2010; AGUILERA &
LUNDBERG, 2010; AGUILERA & MARCENIUK, 2012). The
fossil composition is indicative of a tropical-marine, clear
near shore neritic environment of normal marine salinity,
probably not far from open marine environments (JUNG,
1965; DIAZ DE GAMERO, 1974; NOLF & AGUILERA, 1998;
AGUILERA & RODRIGUES DE AGUILERA, 2001). The age
of the Cantaure FM is dated at the base at about 16.5 Ma
(GRIFFITHS et al., 2012).
Location (Text-fig. 5): San José de Cocodite, pozo Cantaure,
Paraguaná Peninsula (see AGUILERA et al. 2013, fig. 2b
for further details).
The Castillo FM (WHEELER, 1960, 1963) dates as late
Oligocene to early Miocene (LORENTE, 1997) and is situ-
ated in the western Falcón Basin, northwestern Lara State,
Venezuela. The outcrop of the formation is exposed at Cerro
La Cruz near La Mesa Town, on the southern flank of the
Serranía La Baragua, and consists of marls, sandstones, and
claystones. WHEELER (1960) suggested an early Miocene
age for the exposures of the Castillo Formation in the area
studied in this paper. The palaeoenvironment is interpreted
as nearshore marine and includes diverse assemblages of
mollusks, crustaceans, fishes, turtles, crocodiles, and terres-
trial, aquatic and marine mammals (SÁNCHEZ-VILLAGRA et
al. 2000; SÁNCHEZ-VILLAGRA & CLACK, 2004; JOHNSON
et al., 2009).
Location (Text-fig. 5): Cerro La Cruz near La Mesa Town,
on the southern flank of the Serranía La Baragua, Lara
State (see AGUILERA et al. 2013, fig. 3a for further details).
The Caujarao FM (WIEDENMAYER, 1937) of late Miocene
in northwestern Venezuela includes three members: the
Muaco Mb, lower, mainly clayey, organogenic limestones
and interbedded fossiliferous marls and some friable fine-
grained sands; the Mataruca Mb, intermediate, character-
ized by three or more layers of prominent nodular marly and
fossiliferous limestones interbedded with fossiliferous shales
and calcareous marls and sands; and the Taratara Mb,
higher prevalence of clays and micro-fossiliferous shales.
The otoliths have been collected from the middle Mataruca
Mb, considered to be of late Tortonian age.
Location (Text-fig. 5): PPP 2534 in Carrizal Cementery,
village of Carrizal, near La Vela town, Falcón State (see
SMITH et al. 2010, fig. 2 for further details).
The Cubagua FM (DALTON, 1912) of late Miocene to early
Pliocene age (NN 10-17), is found in the north-eastern part
of Venezuela. It is divided in two sections, the lower Cuba-
gua FM consisting of gray shale with glauconite, abundant
pyrite nodules, gray limonite, and some sandy intercalation
with fine clastics, probably emplaced by turbidity currents.
Occasionally clastic metamorphic and volcanic components
are present as well. The upper Cubagua FM consists of
bioclastic banks of reef, mollusk and bryozoans, quartz
sandstone, calcareous sandstone, bioclastic limestone, with
interstratified olive gray shale, laminar glauconitic clay and
gray limonite. The sediments of the lower part appear to
have been deposited in deep water, and the upper part in
shallow tropical water. The Cubagua FM is divided into four
formal members (Mbs): the Cerro Verde Mb, exposed on
the Cubagua Island and westernmost Araya Peninsula; the
Cerro Negro Mb, from which most of the otoliths described
here were obtained, exposed along the Western Araya
Peninsula and Cubagua Island; and the La Tejita and Las
Hernandez Mbs, exposed on Margarita Island. The oldest of
these, the Cerro Verde Mb (late Miocene to earliest Pliocene,
NN 10-12), consists of conglomerate with sandy matrix,
vertically grading into fossiliferous conglomeratic sandstone,
bioclastic sandstone, and shale. The Cerro Verde Mb is
45 m thick at the type section and rests unconformably on
metamorphic rocks of the late Jurassic to early Cretaceous
Manicuare FM. Upwards, the Cerro Verde Mb passes
conformably into the Cerro Negro Mb of early Pliocene,
Zanclean (NN 13-15) age, and strata occasionally sam-
pled above the Cerro Negro Mb are considered as of late
Pliocene age. The Cerro Negro Mb consists of a 2 m thick
basal sandy marl, which is highly fossiliferous with Ostrea
haitensis and O. crassissima, followed by thin intercalations
of sandy lime, fossiliferous marl, and limestone marl. The
total thickness of the member is 22 m at the type locality.
The La Tejita Mb (late Miocene to early Pliocene) consists of
a 45 m thick basal conglomerate with quartz pebbles, schist
and igneous rocks, overlain by gypsum clay, calcareous
argillaceous sandy and fossiliferous marl with O. crassissima
(AGUILERA et al., 2001; HEAD et al., 2006). The La Tejita
Mb rests unconformably upon metamorphic rocks and the
Eocene Punta Carnero Group. The Las Hernandez Mb (early
Pliocene) consists of unconsolidated marl, clay, and lime.
Locations (Text-fig. 5): La Tejita Mb: PPP 2573 on bank of
Laguna de las Marites canal, near Margarita Airport, Mar-
garita Island; Cerro Verde Mb: location OA 99-47, Cerro
El Macho, Araya Peninsula; Cerro Negro Mb: PPP 2553,
PPP 2556 and PPP 2557 at Cerro Barrigón, PPP 3087 at
Margarita airport, Margarita Island; “above” Cerro Negro
Mb: PPP 2567, PPP 2569 and PPP 3093 at Cañón de Las
Calderas, Cubagua Island, PPP 2656 and PPP 2572 in
Bahia de Charagato.
The Cumaná FM (RIVERO, 1956; Léxico Estratigráfico de
Venezuela, 1997) of middle Pleistocene, is outcrops in the
Cerro Caigüire, in Cumaná city. It consist mainly of shellfish
beds and bryozoans with remains of corals; micritic, light-
colored limestone; mudstones, light gray clays interbedded
with light gray to bluish gray silts, with abundant fragments
of echinoderms, bivalves and other molluscs; fossiliferous
calcarenite, calcareous and quartz sandstone, gray and
fine-grained to very fine, olive siltstones.
6.8 Venezuela
22
Location (Text-fig. 5): PPP 3032, Cerro Caigüire, Cumaná,
Sucre State.
The Urumaco Formation (GARNER, 1926; Léxico Es-
tratigráfico de Venezuela, 1997; QUIROZ & JARAMILLO,
2010) in northwestern Venezuela includes three members
comprising 2560 m of sedimentary sequence. The upper
member, where the fossils come from, comprises gray to
brown, often limey claystone with thin intercalated and lo-
cally conchiferous sandstones. Several localities and levels
have concentrations of vertebrate fossils. The vertebrate
fauna includes marine, estuarine and freshwater fishes, ter-
restrial, freshwater and marine turtles and crocodilians, and
terrestrial and aquatic/semiaquatic mammals, summarized
in SANCHEZ-VILLAGRA & AGUILERA (2006). The otoliths
have been collected from the middle, marine member
considered to be of middle Tortonian age.
Location (Text-fig. 5): El Hatillo, northeastern of Urumaco
town, Falcón State (see SMITH et al., 2010, fig. 2; QUIROZ
et al., 2010 fig. 8.6; AGUILERA & MARCENIUK, 2012, fig. 2
for further details).
7. Systematic Part
Subfamily categories follow those proposed by SASAKI
(1989) with certain amendments, which are explained in the
context. Tribes are based on definitions by SASAKI (1989),
genus groups as defined by CHAO (1978) and otolith genus
groups as defined by SCHWARZHANS (1993) with certain
amendments that are discussed in the context.
Family Sciaenidae Cuvier, 1829
Subfamily Pachyurinae SASAKI, 1989
Genera:
†
Eokokenia,
†
Jefitchia, Pachypops and Pachyurus.
Discussion: This subfamily comprises otolith morphologies
considered the most basal within Sciaenidae (SCHWARZ-
HANS, 1993), resembling that of the family Haemulidae,
which are considered as the sistergroup to Sciaenidae by
TREWAVAS (1977) and SCHWARZHANS (1993). The fossil
otolith-based genera
†
Eokokenia and
†
Jefitchia are amongst
the earliest fossil records of the Sciaenidae from the middle
Eocene of the US Gulf Coast (MÜLLER, 1999). The extant
genera Pachyurus and Pachypops are restricted to the fresh-
waters of South America. Recent otoliths of Pachypops have
been figured in NOLF (1976) and SCHWARZHANS (1993)
and those of Pachyurus in CHAO (1978), AGUILERA (1983),
SCHWARZHANS (1993), NOLF (2003) and AGUILERA &
RODRIGUES DE AGUILERA (2004)
1
. Fossil otolith-based re-
cords in past literature from marine Neogene strata of South
America have proven not to represent pachyurins: Pachy-
pops fitchi SCHWARZHANS, 1993 has been transferred to
the fossil otolith-based genus
†
Equetulus by AGUILERA et
al., 2014 and Pachyurus jungi AGUILERA & RODRIGUES DE
AGUILERA, 2004 is placed here in Pareques. Both genera
are placed in the Equetes Group of the Sciaeninae. Thus, the
South American freshwater subfamily Pachyurinae remains
the only sciaenid subfamily not represented in the Neogene
of tropical America.
Subfamily Protosciaeninae SASAKI, 1989
Genus: Protosciaena.
Discussion: SASAKI (1989) established the Protosciaeninae
solely to accommodate for the then newly established mono-
specific genus Protosciaena SASAKI, 1989 (type-species
Sciaena trewavasae CHAO & MILLER, 1975), which he
stated as “strikingly primitive, in that no advanced charac-
ters are found”. This is not entirely true when it comes to
otoliths since P. trewavasae is one of those species with a
tadpole-like widened caudal tip. SASAKI did not conclude
on other so-called New World “Sciaena” species, which
he recorded as incertae sedis, but argued that those are
unlikely to represent Sciaena species. In 2003, CHAO also
placed Sciaena bathytatos CHAO & MILLER, 1975 into
Protosciaena. Otoliths however suggest that this species
represents some other, probably unrelated genus, and
very likely an undefined genus. SCHWARZHANS (1993)
tentatively proposed a relationship with the Pseudotolithi-
nae, an endemic subfamily of the tropical Eastern Atlantic
based on the similarity of the otoliths of Sciaena bathytatos
and Miracorvina angolensis. “Protosciaena” bathytatos is
provisionally left in Protosciaeninae until a conclusive review
has been done.
1 An unresolved discrepancy exists in the cited literature as to the
identification of otoliths of the species Pachyurus schomburgki
GÜNTHER, 1860, which requires review.
Plate 1
Figs. 1-5. Protosciaena brasiliensis AGUILERA & SCHWARZHANS,
2014. 1, 3-5, paratypes, MPEG-1817-V; 2, holotype, MPEG-
1816-V, Brazil, Atalaia beach, Pirabas FM, early Miocene.
Figs. 6-10. Protosciaena neritica AGUILERA & RODRIGUES DE
AGUILERA, 2004. 6-8, 9-10 (r),Venezuela, San José de Cocodite,
Cantaure FM, early Miocene.
Figs. 11-12. Protosciaena trewavasae (CHAO & MILLER, 1975).
11 (r), AFORO database, fish ID 9309, Recent; 12 (r), paratype of
Ctenosciaena latecaudata NOLF & STRINGER, 1992, NMB P128,
Dominican Republic, Rio Gurabo, Cercado FM, late Miocene.
Figs. 13-15. “Protosciaena” bathytatos (CHAO & MILLER, 1975).
13 (r), coll. WS, Venezuela off Cumana, Recent; 14, 15 (r), Ven-
ezuela, Araya Peninsula, Cubagua FM, late Pliocene.
/
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
23
Palaeo Ichthyologica 14
Protosciaena brasiliensis
Protosciaena neritica
Protosciaena trewavasae
“Protosciaena” bathytatos
1a
1b
2
3
4
5
6a
6b
6c
6d
7
8
9a
9b
10
11
12
13a
13b
14
15
2 mm
2 mm
2 mm
2 mm
24
Genus Protosciaena SASAKI, 1989
Protosciaena brasiliensis
AGUILERA & SCHWARZHANS, 2014
(Plate 1, Figs. 1-5)
2014 Protosciaena brasiliensis – AGUILERA & SCHWARZHANS:
figs. 11.1-11.7
Material: 37 specimens, Atalaia beach, Brazil, early Miocene,
MPEG-1816-1818-V.
Diagnosis: OL : OH 1.05-1.1. Dorsal rim shallow, ventral
rim deeply curved. Rear margin of ostium deep. Ostium
higher than long, anteriorly narrowed. Caudal tip rounded
but not or only weakly widened.
Discussion: Protosciaena brasiliensis is the earliest known
species of the genus. It shows the rounded triangular out-
line, the high, compressed ostium, anteriorly narrowed in
this case and a rather short and less than 90° downward-
curved rear part of the cauda, all of which is typical for
the genus, but it lacks the tadpole-type widened caudal tip
observed in the Recent P. trewavasae. The younger P. neritica
is intermediate in the latter aspect in that shows a widened
caudal tip, but the caudal colliculum not widened distally
as it is the case with the Recent species. This lineage thus
confirms clearly how the characteristic tadpole-shape caudal
tip has developed independently in this genus.
Distribution: Aquitanian to early Burdigalian of Brazil.
Protosciaena neritica
AGUILERA & RODRIGUES DE AGUILERA, 2004
(Plate 1, Figs. 6-10)
1998 Protosciaena aff. trewavasae (CHAO & MILLER, 1975)
– NOLF & AGUILERA: part: pl. 12, figs. 5-9
2004 Protosciaena neritica – AGUILERA & RODRIGUES DE
AGUILERA: text-figs. 2A-D
2013 Protosciaena aff. trewavasae (CHAO & MILLER, 1975)
– NOLF: pl. 284
Material: 13 specimens; 11 specimens location OA-95-2, San
José de Cocodite, Paraguaná Peninsula,Venezuela, Cantaure
FM, figured specimens NMB P970-971, and 2 specimens
location 290632, Castilletes, Colombia, Castilletes FM, STRI
34770, both early Miocene.
Diagnosis: OL : OH 1.1-1.2. Dorsal rim shallow, ventral
rim deeply but regularly curved. No postostial lobe. Ostium
nearly round. Caudal tip slightly widened distally, but caudal
colliculum not widened.
Discussion: Protosciaena neritica represents the perfect
intermediate form between the older P. brasiliensis and the
Recent P. trewavasae as far as the development of the caudal
tip is concerned. The round shape of the ostium and the
relatively short downturned part of the cauda however are
additional distinctive characters of P. neritica.
Distribution: Late Burdigalian to early Langhian of Ven-
ezuela and Colombia.
Protosciaena trewavasae (CHAO & MILLER, 1975)
(Plate 1, Figs. 11-12)
1992 Ctenosciaena latecaudata – NOLF & STRINGER: pl.16,
figs. 6-7
1998 Protosciaena aff. trewavasae (CHAO & MILLER, 1975)
– NOLF & AGUILERA: part: synonymization of Ctenos-
ciaena latecaudata, no figures
Material: 2 specimens; 1 specimen, Río Gurabo, NMB 15907,
Dominican Republic, Cercado FM, late Tortonian-Messinian,
NMB P128 (paratype of Ctenosciaena latecaudata); 1 speci-
men, Bastimentos Island, PPP 1256, Atlantic Panama, late
Pliocene, NMB P972.
Diagnosis: OL : OH 1.2-1.3. Dorsal rim shallow, ventral
rim deeply but regularly curved. Weak postostial lobe. Os-
tium about as high as long, triangular in shape, anteriorly
narrowing. Caudal tip distinctly widened inclusive of the
caudal colliculum, typically tadpole-shaped.
Remark: After reviewing of the type specimens of Ctenos-
ciaena latecaudata (paratype Fig. 12) we follow NOLF &
AGUILERA (1998) in recognizing it as junior synonym of the
Recent T. trewavasae. The diagnostic widening of the caudal
tip was not shown in the drawings of NOLF & STRINGER
(1992). The single fossil specimen from Bastimentos is poorly
preserved, but nevertheless identifiable because of is char-
acteristic shape of the ostium and the caudal tip. A Recent
specimen (Fig. 11) is figured for comparison reasons from
the AFORO online otolith catalog.
Distribution: Late Miocene of the Dominican Republic and
late Pliocene of Atlantic Panama and Recent at 70-220 m
water depth along the Atlantic shore of Colombia and off
Venezuela (CHAO, 2002).
“Protosciaena” bathytatos (CHAO & MILLER, 1975)
(Plate 1, Figs. 13-15)
Material: 5 specimens, Araya Peninsula, PPP 2656, Venezuela,
Cubagua FM, ‘above Cerro Negro Member’, late Pliocene,
figured specimens NMB P973-974.
Diagnosis: OL : OH 1.4-1.5. Oval outline, anteriorly nar-
rowed, posteriorly broad. Outer face with massive post-
central umbo. Moderate postostial lobe. Ostium narrow
(OCL : OCH = 1.5) and short (SUL : OCL = 1.8-2.0). Cauda
downward curved to level much below lower ostium mar-
gin; caudal tip tapering. Interspace between rear margin
of ostium and downturned portion of cauda moderately
wide; caudal curvature index = 0.8; ostial-caudal inter-
space = 25-30 %.
Remark: A Recent specimen from WS’ collection (Fig. 13)
is figured for comparison reasons.
Distribution: Late Pliocene of Venezuela and Recent at 70-
300 m water depth along the Atlantic shore of Colombia
and off Venezuela (CHAO, 2002).
Subfamily Sciaeninae GILL, 1861
Definition: In the limits as defined by SASAKI (1989), except
inclusion of the Lonchurini SASAKI, 1989, which he placed
in his newly established subfamily Micropogoniinae.
Tribe Ctenosciaenini SASAKI, 1989
Definition and Discussion: The tribe Ctenosciaenini was
established by SASAKI (1989) to accommodate only the
name giving genus Ctenosciaena based on a single autapo-
morphy, the presence of the single, median, mental barbel.
The otoliths of Ctenosciaena resemble very much those of
the tribe Sciaenini, particularly of the genus Umbrina and
SCHWARZHANS (1993) therefore included Ctenosciaena in
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
25
Palaeo Ichthyologica 14
the Sciaena Group, similar to CHAO (1978). However, we
here follow SASAKI based on the observation that Ctenos-
ciaena otoliths are distinguished from those of Sciaenini
by the wide interspace between rear end of ostium and
downturned portion of cauda expressed in a high caudal
curvature index of about 1.0 or more (vs. <0.8), a character
which it shares with the Pogonini in the following.
Genus Ctenosciaena
FOWLER & BEAN, 1923
Ctenosciaena gracilicirrhus (METZELAAR, 1919)
(Plate 2, Figs. 1-4)
Material: 14 specimens; 3 specimens PPP 2553, 7 specimens
PPP 2557, 1 specimen PPP 2572, 3 specimens OA 99-47,
Araya Peninsula, Cubagua FM, ‘above Cerro Verde Member’,
Venezuela, Zanclean, early Pliocene, figured specimens NMB
P975-978.
Description (of fossil specimens): Moderately compressed,
oval otoliths up to about 7 mm length (up to 9 mm in the
Recent). OL : OH = 1.3-1.45; OH : OT about 3.0. Dorsal rim
shallow, anteriorly slightly inclined, predorsal angle broad,
postdorsal portion broadly rounded, smooth or slightly
undulating; ventral rim moderately deep, smooth, regularly
curved; anterior and posterior rims broadly rounded.
Inner face moderately convex with slightly supramedian
sulcus. Ostium large, wide, spatulate, anteriorly somewhat
narrowing, markedly shorter than cauda, with moder-
ate postostial lobe. Cauda anteriorly straight, posteriorly
bent downward at 90° angle, with slightly widened tip.
OCL : OCH = 0.95-1.05; CCL : OCL = 1.1-1.2; caudal cur-
vature index = 1.0-1.1; ostial-caudal interspace 27-33 %.
Narrow, indistinct dorsal depression; no ventral furrow.
Outer face slightly convex, with broad, rather flat postcentral
umbo, smooth.
Discussion: Otoliths of the genus Ctenosciaena are difficult
to distinguish from those of Umbrina. The best characters for
distinction are probably the rather flat to only moderately
convex inner face and the high caudal curvature index
(1.0-2.0 vs. always smaller than 1.0).
A small, eroded specimen described as C. aff. gracilicir-
rhus by NOLF (1976) from the early Miocene of Trinidad
represents an unidentifiable species of the genus Umbrina.
Specimens described as Ctenosciaena latecaudata NOLF &
STRINGER, 1992 from the late Miocene of the Dominican
Republic have subsequently been synonymized with Proto-
sciaena trewavasae (see above) by NOLF & AGUILERA
(1998). Otolithus (Sciaenidarum) mexicanus LERICHE, 1938
was described from unspecified Miocene rocks of Coatza-
coalcos, Veracruz Province, Atlantic Mexico (erroneously also
mentioned as stemming from Colombia in LERICHE, 1938)
and was subsequently placed in Ctenosciaena by NOLF
(1985). We follow the generic interpretation judging from
LERICHE’s photos, but a specific assessment is not possible
without review of the type-specimen.
Distribution: Early Pliocene of Venezuela and Recent along
the South American Atlantic coast from Nicaragua to south-
ern Brazil in the Recent.
Ctenosciaena peruviana CHIRICHIGNO, 1969
(Plate 2, Figs. 5-12)
Material: 24 specimens; 2 specimens PPP 3514, 11 specimens
PPP 3515, 3 specimens PPP 3516, 1 specimen PPP 3519, 7
specimens PPP 3529, Canoa FM, Ecuador, Gelasian, early
Pleistocene, figured specimens NMB P979-985.
Description (of fossil specimens): Moderately elongate, oval
otoliths up to about 8 mm length (up to 10 mm in the Recent).
OL : OH = 1.25-1.45, increasing with size; OH : OT about
2.5-3.0. Dorsal rim anteriorly and posteriorly inclined,
mediodorsal angle broad, prominent, postdorsal portion
broadly rounded, smooth or slightly undulating; ventral rim
moderately deep, smooth, regularly curved; anterior and
posterior rims broadly rounded.
Inner face moderately convex, only slightly bent in vertical
direction, with slightly supramedian sulcus. Ostium rather
narrow for a sciaenid, slightly upward oriented, sometimes
with distinct diagonal furrows, markedly shorter than cauda,
without postostial lobe. Cauda anteriorly straight, posteri-
orly bent downward at 90° angle. OCL : OCH = 1.1-1.25;
CCL : OCL = 1.15-1.25; caudal curvature index = 1.8-2.1;
ostial-caudal interspace 37-42 %. Broad dorsal depression
with distinct crista superior towards sulcus; no ventral furrow.
Outer face slightly convex, with broad, rather flat postcentral
umbo, smooth.
Discussion: Otoliths of C. peruviana are readily recognized
by the small ostium, the very high caudal curvature index and
the relatively weakly bent inner face. A Recent otolith (Fig. 5)
is figured for comparison from the collection of MNHN.
Very similar otoliths are found in Genyonemus lineatus (see
SCHWARZHANS, 1993 for figures) from the Pacific coast
of North America, differing primarily in the more elongate
shape. Two fossil Genyonemus species had been described
from the middle Miocene Langhian of NE-America – G. cal-
vertensis MÜLLER, 1999 and G.? pertenuis MÜLLER, 1999
– which are also both more elongate in outline.
Distribution: Late Pliocene of Ecuador and Recent along the
coasts (40-200 m) of southern Ecuador and northern Peru.
Tribe Pogoniini new tribe
(= Pogonias and Sciaenops Groups
sensu CHAO, 1978)
Diagnosis: A tribe of the Sciaeninae characterized by a
swimbladder with complex lateral diverticulae or numerous
labyrinthic chambers and thin otoliths showing a cauda with
a high caudal curvature index of about 1.0. It comprises
the genera Aplodinotus, Genyonemus, Pogonias, Roncador,
Sciaenops and the fossil otolith-based genera
†
Trewasciaena
and
†
Amazonasciaena n.gen.
Discussion: CHAO (1978) established the Pogonias Group
to comprise Aplodinotus and Pogonias and the monoge-
neric Sciaenops Group. He did not study the Pacific genera
Genyonemus and Roncador. SASAKI (1989) showed all five
genera clustered closely together in his cladogram and sepa-
rated from other Sciaeninae, but fell short of defining them
as a tribe of their own. SCHWARZHANS (1993) showed all
five genera plus the two fossil otolith-based genera
†
Frizzel-
lithus and
†
Trewasciaena in a Pogonias Group (
†
Frizzellithus
is now removed to Sciaenini). This group seems to entertain
a broad and rather long fossil otolith-based record into Oli-
26
gocene times indicating that it has played a more important
role in sciaenids in the past. In a recent multi-gene mollecular
study LO et al. (2015) confirmed the position of the genera
Genyonemus, Roncador and Seriphus in a distinct lineage,
but it should also be noted, however, that they associated
Sciaenops with Micropogonias and showed the monotypic
genera Aplodinotus and Pogonias widely separated and at
isolated positions in the molecular phylogeny.
Genus
†
Amazonasciaena n. gen.
Type species: Sciaenops rossettiae Aguilera & Schwarzhans,
2014.
Etymology: Referring to the occurrence of the type-species
in the Miocene close to the present day Amazonas river
mouth in combination with the genus name Sciaena Lin-
naeus, 1758.
Diagnosis: A fossil otolith-based genus of the family
Sciaenidae with the following combination of characters.
Moderately elongate otoliths with a ratio OL : OH of
1.45-1.6, moderately thin, without umbo on outer face.
Dorsal rim slightly curved; ventral rim deeply and regularly
curved; anterior and posterior rims inclined resulting in a
parallelogram-like outline. Ostium short, OCL : CCL = 0.8.
Cauda bent in less than 90°, namely 50-60°; caudal cur-
vature index about 1.5.
Discussion: The specific outline and the rear part of the
cauda being bent downwards at only 50 to 60° distinguish
these otoliths from the living genus Sciaenops GILL, 1863,
with which it is considered to be most closely related. The
low curvature of the cauda resembles the status found in
some of the earliest and putative most primitive, extinct,
otolith-based genera
†
Jefitchia FRIZZELL & DANTE, 1965
and
†
Eokokenia FRIZZEL & DANTE, 1965, and here too are
taken as an indication of a plesiomorphic character status.
Species: A single species, A. rossettiae, from the early
Miocene Pirabas FM of Brazil.
†
Amazonasciaena rossettiae
(AGUILERA & SCHWARZHANS, 2014)
(Plate 3, Figs. 4-6)
2014 Sciaenops rossettiae – AGUILERA & SCHWARZHANS:
figs. 10.4-10.8.
Material: 14 specimens, Pirabas FM, Brazil, early Miocene, MPEG-
1808-1810-V.
Description: Moderately elongate otoliths up to about
10 mm length. OL : OH = 1.45-1.6; OH : OT about 3.0. Dor-
sal rim shallow, slightly curved without angles except for ob-
tuse, rounded postdorsal angle located slightly behind bent
of cauda; ventral rim deep, very regularly curved, smooth;
anterior rim distinctly anterior-ventrally inclined, slightly pro-
jecting above upper ostial margin; posterior rim posterior-
dorsally inclined with obtuse tip slightly above tip of cauda.
Inner face moderately convex with distinctly supramedian
sulcus. Ostium moderately large, not inclined, with moder-
ate postostial lobe. Cauda curved at about 50° to 60° with
relatively short inclined section and low caudal curvature
index of about 1.5; caudal tip pointed. OCL : OCH = 1.05-
1.3; CCL : OCL = 0.8. Dorsal depression narrow, small; no
ventral furrow. Outer face flat to slightly convex, less convex
than inner face, smooth.
Discussion: See discussion to genus.
Distribution: Aquitanian to early Burdigalian of Brazil.
Genus Aplodinotus RAFINESQUE, 1819
Aplodinotus hoffmani NOLF & AGUILERA, 1998
(Plate 2, Figs. 17-20)
1998 Aplodinotus hoffmani – NOLF & AGUILERA: pl. 9,
figs. 1-4
Material: 9 specimens; 5 specimens, San José de Cocodite, Para-
guaná Peninsula, Venezuela, Cantaure FM, late Burdigalian
to Langhian, 4 specimens coll. Schwarzhans (leg. Aguilera), 1
specimen NMB P986; 2 specimens, location 290666, Castil-
letes, Colombia, Castilletes FM, late Burdigalian to Langhian,
STRI 34778; 2 tentatively assigned specimens, Cerro la Cruz,
Venezuela, Castillo FM, Aquitanian to early Burdigalian, figured
specimen NMB P987.
Description: Moderately elongate, oval otoliths up to
about 10 mm length. OL : OH = 1.3-1.5; OH : OT about
2.5. Predorsal lobe moderately developed, shallow; dorsal
rim nearly flat, horizontal or slightly inclined backwards,
smooth or indented at its middle, with oblique postdorsal
angle located close to posterior tip of otolith; ventral rim
moderately deep, smooth, regularly curved; anterior and
posterior rims blunt, almost vertically cut.
Inner face markedly convex with distinctly supramedian
sulcus. Ostium large, wide, spatulate, slightly shorter than
cauda, straight, moderate postostial lobe. Cauda anteri-
orly straight, posteriorly bent downward at 90° angle, with
tapering tip. OCL : OCH = 1.1-1.2; CCL : OCL = 0.8-1.0;
caudal curvature index = 1.25-1.6; ostial-caudal interspace
30-35 %. Narrow dorsal depression, often with sharp edge
close to dorsal rim of otolith; no ventral furrow. Outer face
flat to moderately convex in large specimens, with broad,
rather flat postcentral umbo, smooth.
Discussion: The genus Aplodinotus is known in the Recent
from a single freshwater species in North America. Aplodi-
notus hoffmani represents one of the fossil marine species of
the genus. The specimens figured originate from the same
formation as the type-specimens, but appear to be generally
more elongate than those figured by NOLF & AGUILERA,
while the one figured from the slightly older Castillo FM
as A. aff. hoffmani is somewhat more compressed. Until
more specimens have become available for a statistically
more meaningful assessment of the intraspecific variability,
Plate 2
Figs. 1- 4. Ctenosciaena gracilicirrhus (METZELAAR, 1919).
1-3, 4 (r), Venezuela, Araya Peninsula, Cubagua FM, early Pliocene.
Figs. 5-12. Ctenosciaena peruviana CHIRICHIGNO, 1969. 5, coll.
MNHN, off Peru, Recent; 6 (r), 7-11, 12 (r), Ecuador, Punta Canoa,
Canoa FM, early Pleistocene.
Fig. 13. Aplodinotus grunniens RAFINESQUE, 1819; coll. WS,
Texas, Recent, freshwater.
Figs. 14-16. Aplodinotus santosi AGUILERA & SCHWARZHANS,
2014. 14, holotype, MPEG-1792-V; 15-16, paratypes, MPEG-
1793-V, Brazil, Atalaia beach, Pirabas FM, early Miocene.
Fig. 17. Aplodinotus aff. hoffmani NOLF & AGUILERA, 1998;
Venezuela, Cerro la Cruz, Castillo FM, early Miocene.
Figs. 18-20. Aplodinotus hoffmani NOLF & AGUILERA, 1998.
18 (r), 19-20, Venezuela, San José de Cocodite, Cantaure FM,
early Miocene.
Fig. 21. Pogonias chromis (LINNAEUS, 1766); coll. WS, off
Brazil, Recent.
/
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
27
Palaeo Ichthyologica 14
Ctenosciaena gracilicirrhus
Ctenosciaena peruviana
Aplodinotus santosi
Aplodinotus hoffmani
Pogonias chromis
Aplodinotus aff. hoffmani
Aplodinotus grunniens
1a
1c
1b
2
3
4a
4b
5a
5b
5c
6a
6b
6c
6d
7
8
9a
9b
10
11a
11b
12
13a
13b
14a
14b
15
16
17a
17b
17c
18a
18b
18c
18d
19a
19b
20a
20b
21
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
28
we leave these apparent variations within a single species.
Recent otoliths figured for comparison reasons are
Aplodinotus grunniens RAFINESQUE, 1819 (Fig. 13) and
Pogonias chromis (LINNAEUS, 1766) (Fig. 21). Aplodinotus
hoffmani clearly differs from both Recent species in the more
elongate otolith shape.
Distribution: Late Burdigalian to early Langhian of Ven-
ezuela and Colombia.
Aplodinotus santosi
AGUILERA & SCHWARZHANS, 2014
(Plate 2, Figs. 14-16)
2014 Aplodinotus santosi – AGUILERA & SCHWARZHANS:
figs. 9.1-9.5
Material: 44 specimens, Pirabas FM, Brazil, early Miocene, MPEG-
1792-1794-V.
Diagnosis: OL : OH = 1.05-1.15. Ostium very large, about
as wide as long. Cauda strongly bent downwards; posterior
tip pointed with dorsally expanded margin reaching very
close to posterior rim of otolith. Caudal curvature index
about 0.7. Ostial-caudal interspace 22-25 %. Outer face
mildly convex without umbo.
Discussion: Aplodinotus santosi represents a second fossil
marine species of the genus. It differs from A. hoffmani in
the more compressed shape, the wider ostium, the shape of
the caudal tip and the low caudal curvature index. A further
fossil species has been recorded from the early Miocene
of NE-America – Aplodinotus primigenius MÜLLER, 1999.
Distribution: Aquitanian to early Burdigalian of Brazil.
Genus Sciaenops GILL, 1863
Sciaenops reyesi
AGUILERA & RODRIGUES DE AGUILERA, 2004
(Plate 3, Figs. 2-3)
1998 Sciaenops sp. – NOLF & AGUILERA: pl. 13, figs. 1-2
2004 Sciaenops reyesi – AGUILERA & RODRIGUES DE AGUI-
LERA: text-fig. 2G-J
Material: 7 specimens; 3 specimens San José de Cocodite, Para-
guaná Peninsula, Venezuela, Cantaure FM, late Burdigalian
to Langhian, figured specimen paratype UNEFM-PF-02; 3
specimens, Castilletes, Colombia, Castilletes FM, late Burdi-
galian to Langhian, locations 290610, 290632 (STRI 34759,
34894, 34770); 1 specimen Castilletes, Colombia, Jimol FM,
late Burdigalian, location 290602 (STRI 37940).
Diagnosis (of large, adult specimen of >15 mm length):
Anterior rim broadly rounded; posterior rim vertically cut,
massive; dorsal and ventral rims nearly flat, horizontal.
OL : OH = 1.6-1.7. Otolith thin with concave outer face
and without umbo. Ostium very large, almost entirely oc-
cupying anterior half of inner face. Cauda deeply curved.
CCL : OCL = 0.75-0.85; caudal curvature index = 0.9.
Ostial-caudal interspace 22-25 %.
Ontogeny: Otoliths of the genus Sciaenops show a re-
markable ontogenetic change with specimens up to about
10 mm length exhibiting a convex ventral rim and a dorsally
pronounced anterior tip, while in very large specimens
such as the figured paratype of about 16.5 mm length
both dorsal and ventral rims are flat, the posterior tip is
massive and vertically cut and the anterior rim is almost
symmetrically developed.
Discussion: Otoliths of S. reyesi from the early Miocene
of Venezuela and Colombia are very similar to the Recent
S. ocellatus LINNAEUS, 1766 (Fig. 1), distinguished mainly
by the less pronounced postdorsal angle, which in S. ocel-
latus is projecting, and the wider ostium and the slightly
smaller caudal curvature index (0.9 vs. 1.1). Smaller otoliths
of S. reyesi of less than 10 mm length resemble those of
Atractoscion odeai n. sp., described later, differing in the
longer distance of the downcurved cauda from the posterior
rim of the otolith, the more cut shape of the posterior rim
and the flatter dorsal rim.
Distribution: Late Burdigalian to early Langhian of Ven-
ezuela and Colombia.
Sciaenops sp.
(Plate 3, Fig. 7)
Material: 1 specimen Araya Peninsula, Cerro Barrigón, Venezuela,
Cubagua FM, Cerro Negro Member, Zanclean, NMB P988.
Discussion: A single otolith of about 7 mm length and
moderately elongate shape; OL : OH = 1.5. It differs from
similar sized otoliths of the early Miocene S. reyesi in the
deeper, more convex ventral rim and in the slightly smaller
ratio OL : OH (1.5 vs. 1.7-1.7). From the Recent S. ocel-
latus (LINNAEUS, 1766) it differs in the absence of the
postdorsal projection.
Tribe Sciaenini TREWAVAS, 1962
Definition and Discussion: The tribe Sciaenini was defined
by TREWAVAS (1962) and modified by SASAKI (1989).
We have here reduced this group to the genera Callaus,
Cheilotrema, Cilus, Robaloscion, Sciaena, Umbrina and
the fossil otolith-based genus
†
Frizzellithus. The extent cor-
responds to definition of SASAKI (1989), but excluding the
Ctenosciaenini SASAKI, 1989 and excluding the Pogoniini
as above and the Menticirrhini (Menticirrhus Group sensu
CHAO, 1978). The tribe Sciaenini as used here corresponds
to the Umbrina and part of the Sciaena Groups as defined
by CHAO, 1978 and the Sciaena Group of SCHWARZHANS,
Plate 3
Fig. 1. Sciaenops ocellatus LINNAEUS, 1766; coll. WS, Gulf of
Mexico, Recent.
Figs. 2-3. Sciaenops reyesi AGUILERA & RODRIGUES DE AGUILE-
RA, 2004; 2 (r), paratype, UNEFM-PF-02, Venezuela, San José de
Cocodite, Cantaure FM, early Miocene; 3, Colombia, La Guajira,
Castilletes FM, early Miocene.
Figs. 4-6. Amazonasciaena rossettiae (AGUILERA & SCHWARZ-
HANS, 2014); 4, holotype, MPEG-1808-V; 5- 6, paratypes,
MPEG-1809-V; Brazil, Atalaia beach, Pirabas FM, early Miocene.
Fig. 7. Sciaenops sp.; Venezuela, Araya Peninsula, Cubagua FM,
early Pliocene.
Fig. 8. Umbrina dorsalis GIL, 1862; coll. MNHN, off Peru, Recent.
Fig. 9. Cheilotrema saturnum (GIRARD, 1858); coll. WS, off
California, Recent.
Fig. 10. Cheilotrema fasciatum TSCHUDI, 1846; coll. WS, off
Peru, Recent.
Fig. 11. Callaus deliciosa (TSCHUDI, 1846); coll. WS, off Peru,
Recent.
Figs. 12-16. Callaus cubaguanus n. sp. 12, holotype, NMB
P989; 13-16, paratypes, NMB P990-994; Venezuela, Araya
Peninsula, Cubagua FM, late Pliocene.
Figs. 17-21. Frizzellithus longecaudatus (NOLF & AGUILERA,
1998). 17- 20, 21 (r), Colombia, La Guajira, Castilletes FM,
early Miocene.
/
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
29
Palaeo Ichthyologica 14
2 mm
2 mm
2 mm
2 mm
2 mm
8a
8b
9a
9b
10a
10b
11a
11b
Sciaenops ocellatus
Sciaenops reyesi
Amazonasciaena rossettiae
Sciaenops sp.
1a
2a
2b
2c
3
4a
4b
5
6
7
Cheilotrema fasciatum
Callaus
deliciosus
2 mm
2 mm
17
Callaus cubaguanus
Umbrina dorsalis
Cheilotrema
saturnum
1b
12a
12b
12c
12d
13
14
15a
15b
15c
16
Frizzellithus
longecaudatus
18a
18b
19a
19b
20
21
30
1993. LO et al. (2015) expressed the view that the genera
Umbrina and Sciaena are polyphyletic and require re-
definition. In their molecular phylogenetic analysis various
species of the genus Umbrina were associated with widely
separated lineages. The old world species of Umbrina and
Sciaena umbra would form a distinct clade, while several of
the new world species of Umbrina, Callaus and Cheilotrema
would be associated with the Equetini. Otolith morphology
could be used to support such interpretation. We conclude
that both Sciaenini and Equetini may require re-definition.
Genus Callaus JORDAN, 1889
Remarks: Callaus was introduced by JORDAN (1886)
as a subgenus of Sciaena LINNAEUS, 1758 for Corvina
deliciosa TSCHUDI, 1846. We consider the genus Sciaena
a monotypic genus with S. umbra in the NE-Atlantic and
the Mediterranean. The subgenus Callaus was elevated
to genus level by JORDAN et al. (1930), and despite this
decision was not followed by successive authors, we here
consider it as valid and closely related to the genera Umbri-
na CUVIER, 1816 and Cheilotrema TSCHUDI, 1846. It is
provisionally considered a monotypic genus in the Recent,
with the status of “Sciaena” callaensis HILDEBRAND, 1946
being unresolved.
Callaus cubaguanus n. sp.
(Plate 3, Figs. 12-16)
Holotype: Pl. 3, Fig. 12, NMB P989, PPP 2553, N. Cerro Barri-
gón, Araya Peninsula, Venezuela, Cubagua FM, Cerro Negro
Member, Zanclean, early Pliocene.
Paratypes: 8 specimens; 3 specimens same data as holotype,
NMB P990-992; 3 specimens, NMB P993-995, PPP 2556,
Cerro Barrigón, Araya Peninsula, Venezuela, Cubagua FM,
Cerro Negro Member, Zanclean, early Pliocene; 2 specimens,
NMB P996-997, location OA 99-47, Cerro Barrigón, Araya
Peninsula, Venezuela, Cubagua FM, Cerro Verde Member, late
Tortonian to Messinian, late Miocene.
Further material: 2 specimens; 1 specimen, PPP 2572, Cañón de
Charagato, Cubagua Island, Venezuela, Cubagua FM, above
Cerro Negro Member, Piacenzian, late Pliocene; 1 specimen,
PPP 3087, Cerro Barrigón, Araya Peninsula, Venezuela, Cu-
bagua FM, Cerro Negro Member, Zanclean, early Pliocene.
Name: Referring to the Cubagua Formation.
Diagnosis: OL : OH = 1.6-1.7 (1.5 in juveniles). Ante-
rior and posterior tips shifted dorsally. Ventral rim gently
curved; dorsal rim anteriorly depressed. Ostium narrow
for a sciaenid, its anterior-dorsal rim depressed; cauda
slightly forward-bent at its tip. CCL : OCL = 0.9-1.0;
OCL : OCH = 1.4-1.5; caudal curvature index = 0.8; ostial-
caudal interspace 25-27 %.
Description: Elongate otoliths with their longest axis distinct-
ly shifted dorsally, up to about 8 mm length. OL : OH = 1.5-
1.7, increasing with size; OH : OT = 1.7-2.0. Dorsal rim
shallow, with broadly rounded, slightly expanded section
above rear part of ostium, anteriorly depressed. Ventral
rim moderately deeply and very regularly curved, smooth.
Anterior and posterior rims rounded, with their tips distinctly
dorsally shifted.
Inner face markedly convex with distinctly supramed-
ian sulcus. Ostium rather narrow for a sciaenid otolith,
somewhat narrowing anteriorly with downward flexed dor-
sal margin, moderate postostial lobe. Cauda moderately
deepened and rather regularly curved, with tapering tip
slightly curving forward. Indistinct and very small dorsal
depression, only above ostial-caudal joint; no ventral furrow.
Outer face moderately, with broad, rather flat postcentral
umbo, smooth.
Discussion: The otoliths of C. cubaguanus resemble a
number of otoliths of Recent fishes of the genera Callaus,
Cheilotrema and Umbrina with the peculiar shape of the
ostium, the outline of the otolith with the dorsally shifted
posterior tip and the anteriorly depressed dorsal rim. Those
species of the genus Umbrina that resemble C. cubaguanus,
for instance of Umbrina dorsalis GILL, 1862 (Pl. 3, Fig. 8),
all have a much wider ostium (OCL : OCH 1.2 vs. 1.4-1.5)
and differ also in the much narrower ostial-caudal inter-
space (20 % vs. 25-30 %). Likewise the otoliths of the genus
Cheilotrema, Cheilotrema fasciatum TSCHUDI, 1846 (Pl. 3,
Fig. 10) and Cheilotrema saturnum (GIRARD, 1858) (Pl. 3,
Fig. 9) have a narrower ostial-caudal interspace (20 % vs.
25-30 %) and a more downward inclined ostium than those
of Callaus like the Recent C. deliciosa (Pl. 3, Fig. 11) and
the fossil C. cubaguanus. The Recent, Pacific C. deliciosa dif-
fers from the fossil, Caribbean C. cubaguanus in the more
compressed shape (OL : OH = 1.5 vs. 1.6-1.7) and the
slightly larger ostial-caudal interspace (30 % vs. 25-27 %).
Callaus cubaguanus probably represents the Caribbean
geminate counterpart of the Pacific C. deliciosa, but became
extinct during or past late Pliocene.
Distribution: Late Miocene (late Tortonian to Messinian) to
late Pliocene of Venezuela.
Genus
†
Frizzellithus
SCHWARZHANS, 1993
†
Frizzellithus longecaudatus
(NOLF & AGUILERA, 1998)
(Plate 3, Figs. 17-21)
1998 Aplodinotus longecaudatus – NOLF & AGUILERA: pl. 9,
figs. 8-12
Material: 66 specimens; 2 specimens San José de Cocodite,
Venezuela, Cantaure FM, late Burdigalian to Langhian; 27
specimens Castilletes, Colombia, Castilletes FM, late Bur-
digalian to Langhian, locations 16555, 290616, 290632,
290666, 290674, 290685, 290840, 340041, 390090,
390126, 430202 (STRI 34798, 34765, 34770, 34778, 34895,
34792, 37371, 37408, 38223, 34755, 37617, 37403); 37
specimens Castilletes, Colombia, Jimol FM, Burdigalian, loca-
tions 290602, 390085, 390090 (STRI 34773, 34755, 34780,
34786, 34794).
Description: Compressed, high-bodied otoliths up to about
8 mm length. OL : OH = 1.05-1.15; OH : OT about 2.5.
Outline with shallow dorsal rim without angles except for
a rounded postdorsal angle at junction with posterior rim;
ventral rim very deeply and regularly curved; anterior tip
blunt, rounded, dorsally pronounced; posterior tip likewise
blunt, almost vertically cut. All rims smooth.
Inner face markedly convex with supramedian sulcus.
Ostium large, very wide, slightly longer than cauda, with
distinct and broad postostial lobe, its dorsal margin slightly
inclined. Cauda with very short anterior portion, steeply
and regularly bent downward, its tapering tip slightly bent
forward and reaching distinctly deeper than lower margin of
ostium. OCL : OCH = 0.9-1.0; CCL : OCL = 0.7-0.8; cau-
dal curvature index = 0.55-0.65; ostial-caudal interspace
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
31
Palaeo Ichthyologica 14
17-22 %. No distinct dorsal depression, but occasionally
edge close to dorsal rim of otolith; no ventral furrow. Outer
face flat to slightly convex in the largest specimens, without
umbo, slightly and irregularly ornamented.
Discussion: These otoliths were originally placed in the
genus Aplodinotus, but the very narrow ostial-caudal inter-
space, the high caudal curvature index and the long cauda
extending below the level of the ostium all are characters
that distinguish these otoliths from those of Aplodinotus
and instead relate them to the extinct otolith-based genus
Frizzellithus. In fact F. longecaudatus represents the youngest
known representative of the genus.
Distribution: Burdigalian to early Langhian of Venezuela
and Colombia.
Genus Umbrina CUVIER, 1816
Umbrina abbreviata n. sp.
(Plate 4, Figs. 4-9)
1998 Equetes aff. punctatus (BLOCH & SCHNEIDER, 1801) –
NOLF & AGUILERA: pl. 10, figs. 6-9
Holotype: Pl. 4, Fig. 5, MUN2001, Castilletes, location 390126,
Colombia, Castilletes FM, late Burdigalian to Langhian.
Paratypes: 5 specimens, all Castilletes, Colombia, Castilletes FM,
late Burdigalian to Langhian; 4 specimens location 290840,
MUN2002; 1 specimen location 430202, MUN2003; 3
specimens Castilletes, Colombia, Jimol FM, late Burdigalian; 1
specimen location 290602, MUN2004; 2 specimens location
390085, MUN2005.
Name: From abbreviatus (Latin = shortened, abbreviated),
referring to the compressed shape of the otoliths compared
to supposed related Recent species.
Diagnosis: OL : OH = 1.2-1.3. Anterior-dorsal rim some-
what expanded, projecting. Posterior tip nearly vertically
cut. Ventral rim regularly curved, posteriorly bent upwards
to about
1
/
3
before meeting posterior rim. Ostium mod-
erately wide, its anterior-dorsal rim slightly depressed;
cauda slightly forward-bent at its tip. CCL : OCL = 0.8-0.9;
OCL : OCH = 1.05-1.15; caudal curvature index = 0.55-
0.7; ostial-caudal interspace 22-26 %.
Description: Compressed, rather thin otoliths up to about
7.5 mm length. OH : OT = 2.0-2.2. Dorsal rim shallow,
slightly backward inclined towards angle with posterior rim,
anteriorly projecting and expanded in large specimens.
Ventral rim deeply and regularly curved, smooth. Anterior
rim rounded, with its tip shifted way dorsally. Posterior tip
vertically cut between postdorsal angle and angle with ven-
tral rim at level of about
1
/
3
up from deepest ventral point.
Inner face markedly convex with supramedian sulcus.
Ostium moderately wide, anteriorly with downward flexed
dorsal margin, moderate postostial lobe. Cauda moderately
deepened and rather regularly curved, with tapering tip
slightly curving forward. No distinct dorsal depression, but
occasionally edge close to dorsal rim of otolith; no ventral
furrow. Outer face flat, without umbo, slightly and irregularly
ornamented. All rims sharp.
Discussion: Umbrina abbreviata resembles the parallel
occurring Pareques jungi, but is thinner, with a flat outer
face, a slightly wider and less inclined ostium and with the
characteristic predorsal expansion and projection. These
characters are in line with several of the Recent Umbrina
species known from the American shores, such as the inter-
related Umbrina broussonnetii CUVIER, 1830 (Pl. 4, Fig. 2),
Umbrina coroides CUVIER, 1830 (Pl. 4, Fig. 3) and Umbrina
xanti GILL, 1862 (Pl. 4, Fig. 1). The otoliths of these species
all show a larger ostium, are somewhat more elongate
(OL : OH = 1.45-1.6 vs. 1.2-1.3) and have a dorsally
pronounced posterior tip (vs. vertically cut). Umbrina brous-
sonnetii from the Caribbean and U. xanti from the tropical
Eastern Pacific are here considered geminate species.
Distribution: Late Burdigalian to early Langhian of Co-
lombia.
Umbrina bananensis n. sp.
(Plate 4, Figs. 14-17)
Holotype: Pl. 4, Fig. 14, NMB P998, PPP 1727, Costa Rica, Río
Banano FM, Piacenzian, late Pliocene.
Paratypes: 4 specimens; 1 specimen, NMB P999, same data as
holotype; 1 specimen, NMB P1000, PPP 1732, Costa Rica,
Río Banano FM, Piacenzian, late Pliocene; 1 specimen, NMB
P1001, PPP 223, Colón Island, Atlantic Panama, upper Cayo
Agua FM, Zanclean, late early Pliocene.
Name: Referring to the Río Banano Formation.
Diagnosis: OL : OH = 1.1-1.2. Outline nearly round with
blunt posterior tip and regularly rounded anterior rim. Os-
tium wide, its anterior-dorsal rim inclined, but not depressed;
cauda not forward-bent at its tip. CCL : OCL = 0.85-0.95;
OCL : OCH = 1.0-1.1; caudal curvature index = 0.7-0.8;
ostial-caudal interspace 22-27 %.
Description: Compressed, nearly round and rather thin
otoliths up to about 5 mm length. OH : OT about 3.0. All rims
rather regularly curved, smooth, without prominent angles
except occasional obtuse postdorsal angle. Posterior rim
often rather blunt; dorsal rim sometimes less strongly bent.
Inner face moderately convex with supramedian sulcus.
Ostium wide, anteriorly with slightly downward inclined but
not flexed dorsal margin, moderate postostial lobe. Cauda
moderately deepened and rather regularly curved, with
tapering tip not or very slightly curving forward. Narrow
and indistinct dorsal depression above horizontal portion
of cauda; ventral furrow indistinct, sometimes faintly visible
close to caudal tip. Outer face flat, without umbo, smooth.
Discussion: Umbrina bananensis is a relatively small species
characterized by the compressed shape and nearly round
outline combined with a moderately wide ostium without
depressed dorsal margin.
Distribution: Late early Pliocene of Atlantic Panama and
Piacenzian of Costa Rica.
Umbrina laxa n. sp.
(Plate 4, Figs. 18-21)
Holotype: Pl. 4, Fig. 18, MUN2006, Castilletes, location 290840,
Colombia, Castilletes FM, late Burdigalian to Langhian.
Paratypes: 29 specimens, Castilletes, Colombia, Castilletes FM,
late Burdigalian to Langhian; 8 specimens location 290840,
MUN2007; 1 specimen location 290685, MUN2008; 1
specimen location 340041, MUN2009; 2 specimens location
290674, MUN2010; 2 specimens location 16677, MUN2011;
15 specimens location 290632, MUN2012; 1 specimen
Castilletes, Colombia, Jimol FM, late Burdigalian, location
290602, MUN2013.
Name: From laxus (Latin = broad, wide, capacious), refer-
ring to the very wide ostium.
32
Diagnosis: OL : OH = 1.2-1.3. Outline regularly oval with
slightly less bent dorsal rim. Anterior tip shifted far dorsally.
Ostium very wide, spacious, occupying most of anterior
half of inner face, its anterior-dorsal rim not depressed,
broad postostial lobe; cauda slightly forward-bent at its tip.
CCL : OCL = 0.65-0.75; OCL : OCH = 1.0-1.1; caudal cur-
vature index = 0.7-0.9; ostial-caudal interspace 16-22 %.
Description: Moderately compressed, rather massive oto-
liths up to about 7.5 mm length. OH : OT about 2.0. Dorsal
rim less strongly bent than ventral rim, without prominent
angles. Ventral rim deeply and regularly curved, smooth.
Anterior rim rounded, with its tip shifted way dorsally. Pos-
terior rim regularly rounded.
Inner face markedly convex with supramedian sulcus.
Ostium very wide, occupying most of anterior part of inner
face, anterior-dorsally not depressed, strong and broad
postostial lobe. Cauda moderately deepened and rather
regularly curved, with tapering tip slightly curving forward.
Dorsal depression very indistinct and small, above anterior
part of cauda only; no ventral furrow. Outer face uniformly
thickened, but without distinct umbo.
Discussion: Umbrina laxa is readily recognized by its very
wide ostium that shows no deflection or depression of its
anterior-dorsal margin. With its regular oval outline and
the absence of a depressed anterior-dorsal margin of the
ostium, it resembles otoliths of certain Old World Umbrina
species such as Umbrina canariensis VALENCIENNES, 1843,
Umbrina ronchus VALENCIENNES, 1843, and Umbrina
sinuata DAY, 1876 (see SCHWARZHANS, 1993 and SMALE
et al., 1995 for otolith figures), but none of those show a
similarly wide ostium.
Distribution: Late Burdigalian to early Langhian of Co-
lombia.
Umbrina opima n. sp.
(Plate 5, Figs. 1-6)
Holotype: Pl. 5, Fig. 1, NMB P1002, PPP 2572, Cubagua Island,
Cañón de Charagato, Venezuela, Cubagua FM, above Cerro
Negro Member, late Pliocene.
Paratypes: 5 specimens; 1 specimen NMB P1003, same location
as holotype; 4 specimens NMB P1004-1007, PPP 2164, Gatún,
Panama, Gatún FM, Tortonian.
Further material: 7 specimens; 2 specimens PPP 04, 1 specimen
PPP 08, both Gatún FM, Panama, Tortonian; 3 specimens PPP
1139, Río Tuquesa, Darien, Panama, Tuira FM, Tortonian;
1 specimen PPP 1612, Río Tuquesa, Darien, Panama, Chucu-
naque FM, late Miocene.
Tentatively assigned specimens: 1 specimen NMB P1008, San
José de Cocodite, Paraguaná Peninsula, Venezuela, Cantaure
FM, late Burdigalian to Langhian.
Name: From opimus (Latin = abundant, copious), referring
to the widespread occurrence of the species.
Diagnosis: OL : OH = 1.25-1.4; OH : OT = 1.7-2.0. Ante-
rior-dorsal rim and dorsal rim of ostium slightly inclined.
Posterior tip broadly rounded. Ventral rim regularly curved.
Ostium wide, with broad postostial lobe; cauda not or only
very slightly forward-bent at its tip. CCL : OCL = 0.7-0.85;
OCL : OCH = 1.05-1.15; caudal curvature index = 0.65-
0.75; ostial-caudal interspace 20-24 %. Broad umbo on
outer face.
Description: Moderately compressed, oval, thick otoliths up
to about 12.5 mm length. OH : OT = 1.7-2.0 in specimens
larger than 5 mm length and decreasing with size. Dorsal
rim rather regularly curved, anteriorly slightly inclined, pos-
teriorly with a broad, rounded postdorsal angle. Ventral rim
deeply and regularly curved, smooth. Anterior rim regularly
rounded. Posterior tip blunt, broadly rounded.
Inner face markedly convex with median to slightly
supramedian sulcus. Ostium wide, anteriorly downward
inclined but not depressed, strong and broad postostial lobe.
Cauda moderately deepened and rather regularly curved,
with tapering tip not or very slightly curving forward. Small
dorsal depression above ostial-caudal joint only; no ventral
furrow. Outer face broadly convex, with massive postcentral
umbo in large specimens, smooth.
Discussion: Umbrina opima is a rather inconspicuous
species exhibiting a notable variability in its proportions
(OL : OH = 1.25-1.4 and CCL : OCL = 0.7-0.85) A speci-
men from the early Miocene of the Cantaure FM (Fig. 2)
is only tentatively associated with the species differing
somewhat in the less regularly, more flattish ventral rim.
Umbrina opima resembles U. bananensis and U. laxa as
well as the Recent species U. milliae MILLER, 1971 (Pl. 5,
Fig. 7) and Umbrina otoliths from the Old World (see above)
in the wide ostium without depressed dorsal margin. Otoliths
of the deep water Caribbean U. milliae resemble most, dif-
fering primarily in the narrower and less voluminous ostium
and, visible in the lateral view in the thinner anterior part
of the otolith and the thicker postcentral umbo. Smaller
specimens of U. opima (Figs. 3, 4, 6) have a similarly narrow
ostium indicating that small specimens of the two species
could be difficult to distinguish. Umbrina laxa has a still more
voluminous ostium than both U. opima and U. milliae and
shows no inclination of the anterior-dorsal rim. Umbrina
opima differs from U. bananensis in the wider ostium and
the more elongate shape (OL : OH = 1.25-1.4 vs. 1.1-1.2).
Distribution: Widely distributed in the late Miocene and
early Pliocene of Panama and Venezuela and possibly also
in the late Burdigalian to Langhian of Venezuela.
Plate 4
Fig. 1. Umbrina xanti GILL, 1862; coll. Aguirre, off Ecuador, Recent.
Fig. 2. Umbrina broussonnetii CUVIER, 1830; BMNH 1920.12.
22.77-79, off Tobago, Recent.
Fig. 3. Umbrina coroides CUVIER, 1830; coll. OA, off Coro,
Venezuela, Recent.
Figs. 4-9. Umbrina abbreviata n. sp. 5 (r), holotype, MUN2001;
4, 9, 6-8 (r), paratypes, MUN2002-2005; Colombia, La Guajira,
Castilletes FM, early Miocene.
Figs. 10-13. Umbrina sublima n. sp. 10 (r), holotype, NMB P1009,
Venezuela, Araya Peninsula, Cubagua FM, late Miocene; 11, para-
type, NMB P1010, Venezuela, Araya Peninsula, Cubagua FM, late
Miocene; 12, Umbrina aff. sublima, NMB P1012, Panama, Cayo
Agua Island, Cayo Agua FM, early Pliocene; 13, paratype, NMB
P1011, Panama, Colón Island, Cayo Agua FM, early Pliocene.
Figs. 14-17. Umbrina bananensis n. sp. 14, holotype, NMB P998,
Costa Rica, Río Banano FM, late Pliocene; 15-16, paratypes,
NMB P999-1000, Costa Rica, Río Banano FM, late Pliocene;
17, paratype, NMB P1001, Panama, Colón Island, Cayo Agua
FM, early Pliocene.
Figs. 18-21. Umbrina laxa n. sp. 18 (r), holotype, MUN2006;
19-21, paratypes, MUN 2007-2009; Colombia, La Guajira,
Castilletes FM, early Miocene.
/
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
33
Palaeo Ichthyologica 14
Umbrina xanti
Umbrina broussonnetii
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
18a
18b
18c
18d
19
20
21
4a
4b
5a
5b
6
7
8a
8b
9
5c
1a
1b
2a
2b
3a
3b
Umbrina abbreviata
Umbrina sublima
10a
10b
10c
11
12a
12b
13a
13b
13c
Umbrina coroides
Umbrina bananensis
2 mm
14a
14b
15
16
17
Umbrina laxa
34
Umbrina sublima n. sp.
(Plate 4, Figs. 10-13)
Holotype: Pl. 4, Fig. 10, NMB P1009, location OA 99-47, Araya
Peninsula, Venezuela, Cubagua FM, above Cerro Verde Mem-
ber, late Tortonian to Messinian.
Paratypes: 2 specimens; 1 specimen NMB P1010, same location
as holotype; 1 specimen NMB P1011, PPP 2243, Colón Island,
Atlantic Panama, Cayo Agua FM, Zanclean.
Tentatively assigned specimens: 1 specimen NMB P1012, PPP
2237, Cayo Agua Island, Atlantic Panama, Cayo Agua FM,
Zanclean.
Name: From sublimus (Latin = sublime), referring to the
outline of the otolith characterized by the distinctive post-
dorsal projection.
Diagnosis: OL : OH = 1.35-1.45. Dorsal rim shallow, with
sharp, projecting postdorsal angle. Ventral rim regularly
curved. Ostium wide, its dorsal margin downward deflected
close to ostial opening, broad postostial lobe; cauda slightly
forward-bent at its tip. CCL : OCL = 0.8; OCL : OCH = 1.15-
1.25; caudal curvature index = 0.75-0.85; ostial-caudal
interspace 18-22 %.
Description: Moderately elongate, rather thin otoliths up to
about 10 mm length. OH : OT = 2.0-2.5. Dorsal rim shal-
low, anteriorly level or slightly inclined, posteriorly with sharp,
projecting postdorsal angle. Ventral rim deeply and regularly
curved, smooth. Anterior rim regularly rounded. Posterior
rim rounded, leading up to prominent postdorsal angle.
Inner face markedly convex with slightly supramedian
sulcus. Ostium wide, its dorsal margin downward deflected
close to anterior rim of otolith; strong and broad postostial
lobe. Cauda moderately deepened and rather regularly
curved, with tapering tip slightly curving forward. Narrow
dorsal depression extending above entire cauda, with fine
margin backwards towards posterior rim; faint ventral furrow
occasionally visible close to ventral rim in space between
ostium and caudal tip. Outer face flat, only slightly convex
at rear part, little ornamented.
Discussion: Umbrina sublima is readily recognized by the
sharp, projecting postdorsal angle and the abruptly deflected
dorsal margin of the ostium with that deflection located
close to the anterior rim of the otolith. A smaller specimen
of about 5 mm length (Fig. 13) shows a less pronounced,
more rounded postdorsal angle. A specimen from Cayo de
Agua (Fig. 12) exhibits a rather blunt anterior rim and a
deeply curved preventral part followed by less intensely bent
middle part of the ventral rim. This specimen is tentatively
associated wit U. sublima.
Distribution: Tortonian to Messinian of Venezuela and
Zanclean of Atlantic Panama.
Umbrina surda n. sp.
(Plate 5, Figs. 8-11)
Holotype: Pl. 5, Fig. 8, NMB P1013, PPP 3459, Río Santiago,
Ecuador, Angostura FM, Tortonian.
Paratypes: 7 specimens, all Angostura FM, Tortonian, Ecuador; 1
specimen NMB P1014, same location as holotype; 2 specimens
NMB P1015-1016, PPP 3587; 2 specimens NMB P 1017-
1018, PPP 3444; 1 specimen NMB P 1019, PPP 3439; 1
specimen NMB P 1020, PPP 3406.
Name: From surdus (Latin = indifferent, mute), referring to
the inconspicuous appearance of the otoliths.
Diagnosis: OL : OH = 1.25-1.3. Dorsal rim shallow, almost
flat, rounded angles at junction with anterior and posterior
rims. Ventral rim deep, regularly curved. Ostium moder-
ately wide, its dorsal margin inclined, but not deflected,
moderate postostial lobe; cauda not forward-bent at its
tip. CCL : OCL = 0.85-0.95; OCL : OCH = 1.1-1.3; caudal
curvature index = 0.75-0.85; ostial-caudal interspace 30 %.
Description: Moderately elongate, oval, rather thin otoliths
up to about 6.5 mm length. OH : OT = 2.3-3.0. Dorsal rim
shallow, almost flat, anteriorly and posteriorly with rounded
angles at junction with anterior and posterior rims respec-
tively, occasionally shallow mediodorsal expansion. Ventral
rim deeply and regularly curved. Anterior and posterior rims
regularly rounded, dorsally pronounced. All rims smooth,
or occasionally intensely and delicately crenulated.
Inner face mildly convex with distinctly supramedian
sulcus. Ostium moderately wide, its dorsal margin slightly
downward inclined but not deflected, moderate postostial
lobe. Cauda moderately deepened and rather regularly
curved, with tapering tip not curving forward. Indistinct
dorsal depression only above anterior part of cauda; no
ventral furrow. Outer face flat except for shallow postcentral
umbo, ornamented at variable intensity.
Discussion: Umbrina surda is a rather inconspicuous spe-
cies that combines characters of the genera Umbrina and
Ctenosciaena. With otoliths of Ctenosciaena it shares the
wide ostial-cauda interspace, but differs in the caudal cur-
vature index being less than 1.0 and also the CCL : OCL
index being less than 1.0. With its Umbrina congeners it
resembles most U. opima from which it mainly differs in
the wide ostial-caudal interspace (30 % vs. 20-24 %), the
smaller ostium and the specific shape of the dorsal rim.
Distribution: Tortonian of Ecuador.
Umbrina sp.
(Plate 5, Figs. 12-13)
Material: 3 specimens, Castilletes, Colombia, Castilletes FM, late
Burdigalian to Langhian, locations 290448, 290632, 290840
(STRI 34791, 34770, 37408).
Discussion: Three specimens of 5 to 5.5 mm length dif-
fer from the co-occurring U. abbreviata and U. laxa in the
wide ostial-caudal interspace of 26-28 % (vs. 22-26 % and
16-22 % respectively). From U. abbreviata they differ further
in the outline of the otolith and the lack of a deflection of
Plate 5
Figs. 1-6. Umbrina opima n. sp.; 1, holotype, NMB P1002, Vene-
zuela, Cubagua Island, Cubagua FM, late Pliocene; 2 (r), tentatively
assigned specimen, NMB P1008, Venezuela, San José de Cocodite,
Cantaure FM, early Miocene; 3-5, paratypes, NMB P1004-1007,
Panama, Gatún FM, late Miocene; 6 (r), paratype, NMB P1003,
Venezuela, Cubagua Island, Cubagua FM, late Pliocene.
Fig. 7. Umbrina milliae MILLER, 1971; USNM 407628, Caribbean
off Colombia, 250-290 m.
Figs. 8-11. Umbrina surda n. sp. 8, holotype, NMB P1013,
9-11, paratypes, NMB P1014-1016; Ecuador, Angostura FM,
late Miocene.
Figs. 12-13. Umbrina sp.; Colombia, La Guajira, Castilletes FM,
early Miocene.
Figs. 14- 18. Pareques jungi AGUILERA & RODRIGUES DE
AGUILERA, 2004. 14, 15 (r), Venezuela, San José de Cocodite,
Cantaure FM, early Miocene; 16-17, 18 (r), Colombia, La Guajira,
Castilletes FM, early Miocene.
Fig. 19. Pareques lanfeari (BARTON, 1947); coll. MNHN, off
Peru, Recent.
/
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
35
Palaeo Ichthyologica 14
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
1a
1b
2a
2b
2c
3
4
5a
5b
6
8a
8b
9a
9b
9c
9d
10a
10b
11
12
13a
13b
13c
14
15a
15b
16a
16b
16c
17
18
19a
19b
7a
7b
7c
Umbrina opima
Umbrina milliae
Umbrina surda
Pareques jungi
Pareques lanfeari
Umbrina sp.
36
the dorsal margin of the ostium, and from U. laxa in the
much smaller ostium. These otoliths probably represent yet
another species of the genus Umbrina, but the few specimens
available are rated as not sufficiently adult to be suited for
specific definition.
Tribe Equetini SASAKI, 1989
Definition and Discussion: The tribe Equetini contains the
two extant genera Equetes and Pareques following the defi-
nition of SASAKI (1989) and the fossil otolith-based genus
†
Equetulus. CHAO (1978) had also included Sciaena in this
group, which was then consequently named Sciaena Group.
LO et al. (2015) place Callaus, Cheilotrema and certain
species of the new world species of Umbrina in the same
lineage as Equetes and Pareques following their molecular
analysis. We conclude that the limits and definition of the
Equetini may require re-defintion.
Genus
†
Equetulus
AGUILERA & SCHWARZHANS, 2014
†
Equetulus amazonensis
AGUILERA & SCHWARZHANS, 2014
(Plate 6, Figs. 1-4)
2014 Equetulus amazonensis – AGUILERA & SCHWARZHANS,
in: AGUILERA, SCHWARZHANS, MORAES-SANTOS &
NEPOMUCENO: figs. 9.6-9.11
Material: 733 specimens Atalaia beach, Brazil, Pirabas FM, early
Miocene, MPEG-1796-V-1798-V.
Description: Compressed, moderately thick otoliths up
to 5.5 mm length, with somewhat parallelogram-shaped
outline due to strongly projecting anterior-dorsal lobe and
presence of distinct inferior posterior tip. OL : OH = 1.05-
1.15; OH : OT about 2.5. Dorsal rim straight, inclined
backwards from anterior-dorsal projection; anterior and
ventral rims regularly and continuously curving; posterior
rim upward-anteriorly inclined above inferior tip and joint
to dorsal rim in gentle curve.
Inner face moderately convex with distinctly supramed-
ian sulcus. Ostium extremely small as compared to cauda,
shallow, sometimes with reduced opening, slightly bent
downwards. Cauda long, steeply curving in a regular bent;
its vertical section reaching close to posterior-ventral rim
with pointed tip. Cauda somewhat deepened and reach-
ing its greatest width within the maximally bent region.
OCL : OCH about 1.0; CCL : OCL = 2.5-3.0; caudal cur-
vature index = 0.8-1.0; ostial caudal interspace 50-60 %.
Dorsal depression minute, indistinct; ventral furrow feeble,
recognizable along narrow stretch close to mid-ventral rim
and connecting to tip of cauda. Outer face slightly convex
posteriorly and flat anteriorly, smooth.
Discussion: Equetulus amazonensis is readily recognized
by the combination of a strong predorsal lobe and the
small ostium. In comparison, E. davidandrewi sometimes
has a similarly strong predorsal lobe, but a larger ostium
(CCL : OCL = 2.5-3.0 vs. 1.8-2.2), while E. fitchi shares
the small ostium, but has a depressed predorsal region.
Distribution: Aquitanian to early Burdigalian of Brazil.
†
Equetulus davidandrewi
(NOLF & AGUILERA, 1998)
(Plate 6, Figs. 5-10)
1998 Equetus davidandrewi – NOLF & AGUILERA: pl. 10,
figs. 1-5
Material: 147 specimens; 63 specimens Castilletes, Colombia,
Castilletes FM, late Burdigalian to Langhian, locations 16344,
16555, 290448, 290610, 290616, 290632, 290666, 290674
(figured specimen STRI 34895), 290685, 290840, 390126,
430202. GAB059; 14 specimens Castilletes, Colombia, Ji-
mol FM, Burdigalian, locations 290602, 390085, 390090;
4 specimens San José de Cocodite, Paraguaná Peninsula,
(figured specimen NMB P1021), Venezuela, Cantaure FM, late
Burdigalian to Langhian; 5 specimens, PPP 1554, Rio Chico
and PPP1593, Rio Icuanati, both Darien, Panama, Tuira FM,
early Tortonian; 6 specimens location OA 01-1 (figured speci-
men NMB P1022), El Hatillo, Venezuela, Urumaco FM middle
member, middle Tortonian; 30 specimens, Sabanita, PPP 04
(figured specimens NMB P1023-1024), 08, 2165 and San
Judas Tadeo, PPP 2167, Atlantic Panama, Gatun FM middle
member, middle Tortonian; 25 specimens, Rio Cayapas, PPP
3406, 3431, 3432, Rio Santiago, PPP 3444, 3456, 3457, and
Favio Alvaro, PPP 3587, Ecuador, Angostura FM, Tortonian.
Discussion: Equetulus davidandrewi is readily recognized
by the combination of a moderately to strongly developed
predorsal lobe and a comparatively large ostium (se also
above: E. amazonensis).
Distribution: Equetulus davidandrewi is one of the most
widely distributed species in the early to late Miocene of
tropical America from Ecuador to Panama and Venezuela
thus exhibiting a clear tropical amphiamerican pattern.
†
Equetulus fitchi (SCHWARZHANS, 1993)
(Plate 6, Figs. 11-15)
1976 “genus aff. Pachypops” sp. – NOLF: pl. 8, figs. 4-5
1993 Pachypops fitchi – SCHWARZHANS: figs. 15-18
2014 Equetulus fitchi (SCHWARZHANS, 1993) – AGUILERA,
SCHWARZHANS, MORAES-SANTOS & NEPOMUCENO:
figs. 9.12-9.14
Material: 297 specimens; 240 specimens Atalaia beach, Brazil,
Pirabas FM, early Miocene, MPEG-1799-V-1800-V; 25 speci-
mens, PPP 2667, 2669 (figured specimen NMB P1025), Man-
zanilla Bay, 2673 (figured specimens NMB P1026-1027), Point
Paloma, 2675, 2676, 2677, San José river, Trinidad, Manzanilla
FM, San José and Monserrat Mbs, late Tortonian-Messinian;
32 specimens, NMB, KR 9244, Ganteaume river, Trinidad,
Brasso FM, early
-
middle Miocene and KR 25959, Mayo river,
Plate 6
Figs. 1-4. Equetulus amazonensis AGUILERA & SCHWARZHANS,
2014. 6, holotype, MPEG-1796-V; 2-4, paratypes, MPEG-1797-V;
Brazil, Atalaia beach, Pirabas FM, early Miocene.
Figs. 5-10. Equetulus davidandrewi (NOLF & AGUILERA, 1998).
5 (r), Venezuela, San José de Cocodite, Cantaure FM, early Mio-
cene; 6, 8, Ecuador, Angostura FM, late Miocene; 7, Panama,
Gatun FM, late Miocene; 9, Venzuela, El Hatillo, Urumaco FM,
late Miocene; 10 (r), Colombia, La Guajira, Castilletes FM, early
Miocene.
Figs. 11-15. Equetulus fitchi (SCHWARZHANS, 1993). 11-12,
Brazil, Atalaia beach, Pirabas FM, early Miocene; 13-15, Trinidad,
Manzanilla FM, late Miocene.
Fig. 16. Menticirrhus sp.; NMB P1028, Panama, Darien, Tuira
FM, late Miocene.
Fig. 17. Paralonchurus rathbuni (JORDAN & BOLLMAN, 1890);
NMB P1032, Ecuador, Jama FM, middle Pleistocene.
Figs. 18-20. Lonchurus lanceolatus (BLOCH, 1788); NMB P1029-
1031, Trinidad, Manzanilla FM, late Miocene.
/
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
37
Palaeo Ichthyologica 14
Equetulus amazonensis
Equetulus davidandrewi
Equetulus fitchi
Menticirrhus sp.
Paralonchurus rathbuni
Lonchurus lanceolatus
1a
1b
2
3
4
5a
5b
5c
6a
6b
7
8
9
10
11
12
13
14a
14b
15
16a
16b
17a
18a
18b
19
20a
20b
20c
17b
17c
1 mm
1 mm
1 mm
1 mm
1 mm
1 mm
38
KR 25971, Guaracarito river, K 10842, K 12051, K 12052,
K 12054, Manzanilla coast, SM 1399, Pointe-à-Pierre, Cb
1633, San José river, Trinidad, Manzanilla FM, late Tortonian.
Discussion: Equetulus fitchi is recognized by the combina-
tion of the depressed predorsal rim and a very small ostium
(se also above: E. amazonensis).
Distribution: Equetulus fitchi replaces E. davidandrewi
geographically eastwards of Venezuela, in Trinidad and
Brazil during early to late Miocene.
Genus Pareques GILL, 1876
Pareques jungi
(AGUILERA & RODRIGUES DE AGUILERA, 2004)
(Plate 5, Figs. 14-18)
2004 Pachyurus jungi – AGUILERA & RODRIGUES DE AGUI-
LERA: pl. 1, figs. 13-16
2014 Pachyurus jungi AGUILERA & RODRIGUES DE AGUILERA,
2004 – AGUILERA, SCHWARZHANS, MORAES-SANTOS
& NEPOMUCENO: fig. 10.3
Material: 87 specimens; 26 specimens Atalaia beach, Brazil,
Pirabas FM, early Miocene, MPEG-1807-V; 31 specimens
Castilletes, Colombia, Castilletes FM, late Burdigalian to Lang-
hian, locations 16555, 290448, 290616, 290632, 290666,
290674, 290685, 290840 (STRI 34791, 34773, 34770,
34778, 34895, 34792, 37408, 37371, 34780); 7 specimens
Castilletes, Colombia, Jimol FM, late Burdigalian, locations
290602, 390085, 390090 (STRI 34755, 34798, 34786);
23 specimens, paratypes, location OA-95-02, San José de
Cocodite, Paraguaná Península, Cantaure FM, Venezuela, late
Burdigalian to Langhian.
Description: Moderately compressed, oval otoliths up to
about 7.5 mm length. OL : OH = 1.4; OH : OT about 1.5.
Predorsal lobe moderately developed, somewhat project-
ing anteriorly; dorsal rim rather flat, smooth; ventral rim
shallow, smooth, regularly curved; anterior and posterior
rims broadly rounded.
Inner face markedly convex with distinctly supramedian
sulcus. Ostium large, equal in length to cauda, somewhat
bent downwards towards anterior. Cauda very regularly
half-moon shaped with its pointed tip distinctly bent for-
ward, leaving little space on inner face towards posterior
rim. OCL : OCH about 1.0; CCL : OCL = 0.7-0.8; caudal
curvature index = 0.6; ostial-caudal interspace 22-26 %.
Indistinct, narrow dorsal depression; no ventral furrow.
Outer face convex, with postcentral umbo and sometimes
with broad tubercles.
Discussion: Pareques jungi is removed here from its origi-
nal position with Pachyurus in the Pachyurinae to Pareques
following the close resemblance with otoliths of the Recent
Pareques lanfeari (BARTON, 1947) (Fig. 19) from the Pa-
cific coast of Ecuador and Peru and Pareques fuscovittatus
(KENDALL & RADCLIFFE, 1912) from the Pacific coast of
Mexico (otolith studied from specimen CAS 214483, but
too poorly preserved for figuring). It differs from P. lanfeari
in the more compressed shape (OL : OH = 1.2-1.4 vs. 1.6),
the somewhat wider ostium and narrower downward bent
portion of the cauda. Similar otoliths have been figured
by BAREMORE & BETHEA (2010) from the Atlantic spe-
cies Pareques iwamotoi MILLER & WOODS, 1988 and
Pareques umbrosus (JORDAN & EIGENMANN, 1889), the
latter probably the geminate counterpart of P. lanfeari.
Both differ from the fossil P. jungi primarily in the larger
ostial-caudal interspace (33-35 % vs. 22-26 %) and the
narrower ostium. Pareques iwamotoi differs further in the
strongly oblique anterior rim.
Other fossil otoliths possibly representing a further fossil
species have been described as Pareques sp. by NOLF &
STRINGER (1992) from the early Pliocene Gurabo FM of
the Dominican Republic.
Distribution: Aquitanian to early Burdigalian of Brazil
and late Burdigalian to early Langhian of Colombia and
Venezuela.
Tribe Menticirrhini new tribe
(= Menticirrhus Group sensu CHAO,
1978)
Diagnosis (following CHAO, 1978): A tribe of the subfam-
ily Sciaeninae characterized by a swimbladder atrophied
in adults, no drumming muscles, and a thin and elongate
otolith. It comprises the genus Menticirrhus. The genus
Leiostomus, which has a simple, carrot-shaped swimblad-
der, is tentatively placed here on the basis of similarities
of the otoliths.
Discussion: CHAO (1978) established the Menticirrhus
Group mainly based on its distinct otolith pattern. SASAKI
(1989) considered Umbrina and Menticirrhus as sister
genera, but then listed 6 autapomorphies for Menticirrhus
without considering otoliths. In his cladogram he placed the
genus Leiostomus next to his Umbrina-Menticirrhus branch
and characterized by a set of 8 autapomorphies exclud-
ing otoliths. A recent molecular study by LO et al. (2015)
confirmed the separation of Meniticirrhus, but showed
Leiostomus as the in-group sister-taxon of all remaining
Sciaenidae.
Genus Menticirrhus GILL, 1861
Menticirrhus chaoi NOLF & STRINGER, 1992
1992 Menticirrhus chaoi – NOLF & STRINGER: pl. 16, fig. 9
Remarks: Otoliths of M. chaoi from the late Miocene Cer-
cado FM of the Dominican Republic are characterized by
their compressed shape with an OL : OH of less than 2.0.
Menticirrhus sp.
(Plate 6, Fig. 16)
Material: 1 specimen, PPP 1142 (NMB P1028), Rio Tupisa, Darien,
Panama, Tuira FM, early Tortonian.
Discussion: A single, eroded specimen with the typical
outline and sulcus of the otoliths of the genus Menticir-
rhus. It differs from M. chaoi NOLF & STRINGER, 1992,
the only fossil otolith-based species of the genus known to
date, from the late Miocene, Messinian of the Dominican
Republic primarily in being more elongate (OL : OH = 2.4
vs. 1.95). At a size of about 6.5 mm length the specimen
is reasonably large, but too poorly preserved for specific
identification.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
39
Palaeo Ichthyologica 14
Tribe Lonchurini SASAKI, 1989
Definition and Discussion: The tribe Lonchurini contains
the two extant genera Lonchurus and Paralonchurus follow-
ing the definition of SASAKI (1989) and of the Lonchurus
Group of CHAO (1978) and of SCHWARZHANS (1993).
The definition of the genera appears now resolved for the
genus Lonch urus containing two extant species (see dis-
cussion in SASAKI, 1989), but we still consider Paralonch-
urus as containing different lineages as indicated by the
divergence of otolith morphology. We therefore maintain
Polycirrhus BOCOURT, 1869 as a valid genus following
SCHWARZHANS (1993). We have relocated the Lonchurini
from their position within the Micropogoniinae in SASAKI
(1989) to a placement within Sciaeninae primarily because
otolith morphology strongly contradicts a relationship of
Lonchurini with Micropogoniini. Recent molecular studies
(SANTOS et al., 2013 and VINSON et al., 2004) support a
more distant relation of the two groups and in fact points to
Lonchurini representing a clade separated sufficiently early
to potentially warrant placement of a subfamily of its own.
Genus Lonchurus BLOCH, 1793
Lonchurus lanceolatus (BLOCH, 1788)
(Plate 6, Figs. 18-20)
Material: 3 specimens PPP 2673 (NMB P1029-1031), Point
Paloma, Trinidad, Manzanilla FM, Montserrat Mb, Messinian.
Description (of fossil specimens): Very elongate, thin oto-
liths up to about 6 mm length (to 7.5 mm in the Recent).
OL : OH = 2.6-2.9; OH : OT = 1.6- 1.7. Predorsal lobe
broadly expanded, forward projecting; mid- and postdorsal
rims depressed, nearly straight, horizontal; ventral rim flat,
straight, horizontal, smooth; anterior tip broadly rounded;
posterior tip with moderately developed angle at junction
with dorsal rim.
Inner face slightly convex, nearly flat in anterior view.
Sulcus occupying almost entire inner face except for area
of predorsal lobe, ostial-caudal interspace and small area
inside of postdorsal angle. Ostium large, downward bent,
with flat ventral and anteriorly inclined dorsal margin. Cauda
with relatively long anterior section and strongly down-
ward curved, slightly widened posterior section, running
close to dorsal and posterior rims. OCL : OCH about 2.0;
CCL : OCL = 0.7-0.8; caudal curvature index = 1.3-1.4;
ostial-caudal interspace about 25 %. No dorsal depres-
sion or ventral furrow. Outer face flat except for moderate
postcentral umbo, smooth.
Discussion: Lonchurus lanceolatus is the most derived
otolith pattern within the Lonchurini, characterized by a
strong predorsal lobe, but otherwise depressed dorsal rim
and flat ventral rim.
Distribution: Late Miocene, Tortonian of Trinidad and
Recent in the Caribbean from Colombia to the Amazon
river mouth in Brazil.
Genus Paralonchurus BOCOURT, 1869
Paralonchurus rathbuni
(JORDAN & BOLLMAN, 1890)
(Plate 6, Fig. 17)
Material: 1 specimens PPP 3563 (NMB P1032), Jama, Ecuador,
Jama FM, middle Pleistocene.
Description (of fossil specimens): A single very elongate,
thin otolith of about 5.5 mm length (to about 11 mm in the
Recent). OL : OH = 2.1; OH : OT = 1.6. Predorsal lobe mod-
erately expanded, forward projecting; mid- and postdorsal
rims inclined, moderately depressed; ventral rim shallow,
nearly flat except for middle portion, smooth; anterior tip
broadly rounded; posterior tip with weakly developed angle
at junction with dorsal rim.
Inner face slightly convex, nearly flat in anterior view.
Sulcus occupying much of inner face except for area of
predorsal lobe, ostial-caudal interspace and narrow strip
between downward bent part of cauda and posterior rim.
Ostium large, downward bent, with flat ventral and anteriorly
inclined dorsal margin. Cauda with relatively long anterior
section and strongly downward curved, slightly widened
posterior section. OCL : OCH about 1.7; CCL : OCL = 0.8;
caudal curvature index = 1.0; ostial-caudal interspace about
22 %. No dorsal depression or ventral furrow. Outer face
flat except for moderate postcentral umbo, smooth.
Discussion: This otolith differs from Lonchurus lanceola-
tus described above in being less elongate and the less
strongly developed predorsal lobe. It differs from the only
known Recent otolith (SCHWARZHANS, 1993, fig. 213) in
the slightly convex ventral rim (vs. completely flat). Due to
the difference in size of both specimens, this is regarded
as an ontogenetic change.
Distribution: Middle Pleistocene from Ecuador and Recent
along the Pacific shores of America from Panama to Peru.
Genus Polycirrhus BOCOURT, 1869
Remarks: According to ESCHMEYER (2014), Polycirrhus is
not preoccupied by Polycirrus GRUBE, 1850 in Polychaeta
and therefore Polyclemus BERG, 1895 is an unneeded
replacement. We regard Polycirrhus as a valid genus fol-
lowing SCHWARZHANS (1993) with a single Recent spe-
cies – Polycirrhus dumerilii BOCOURT, 1869, known from
the Pacific coast of America from El Salvador to Peru. Its
otoliths represent the most plesiomorphic pattern in the
Lonchurini with the least reduced dorsal rim, the least
developed predorsal lobe and the least expanded sulcus
over the inner face.
Polycirrhus jaramilloi n. sp.
(Plate 7, Figs. 9-15)
Holotype: Pl. 7, Fig. 9, MUN2014, Castilletes, location 430202,
Colombia, Castilletes FM, late Burdigalian to Langhian.
Paratypes: 6 specimens; 2 specimens Castilletes, Colombia,
location 290616 (MUN2015), location 390094 (MUN2017),
Castilletes FM, late Burdigalian to Langhian; 1 specimen
Castilletes, Colombia, Jimol FM, Burdigalian, location 390090
(MUN2016); 3 specimens NMB P1033-1035, San José de
Cocodite, Paraguaná Península, Venezuela, Cantaure FM, late
Burdigalian to Langhian.
40
Further material: 23 specimens; 19 specimens Castilletes, loca-
tions 290448, 290616, 290632 290840, 430202, 390094,
Colombia, Castilletes FM, late Burdigalian to Langhian; 2
specimens Castilletes, Colombia, Jimol FM, Burdigalian,
locations 390090; 2 specimens, San José de Cocodite, Para-
guaná Península, Venezuela, Cantaure FM, late Burdigalian
to Langhian.
Name: Named in honor of Carlos Jaramillo (STRI), who
explored the prolific Castilletes section in Colombia.
Diagnosis: OL : OH = 1.6-1.75. Ostium large, OCL : OCH
= 1.3-1.5, slightly downward bent with parallel dorsal and
ventral margins. Otolith anteriorly thin, posteriorly with
broad umbo on outer face.
Description: Moderately elongate and moderately thick
otoliths to about 12 mm length. OL : OH = 1.6-1.75, in-
creasing with size; OH : OT = 1.7-1.8. Predorsal lobe not
expanded upwards, but slightly forward projecting; mid- and
postdorsal rims shallow, somewhat undulating; ventral rim
shallow, slightly and regularly bent, smooth; anterior rim
very broadly rounded, centrally or dorsally pronounced;
posterior tip blunt, sometimes with very weak angle at
junction with dorsal rim.
Inner face moderately convex, with median, large sulcus.
Ostium large, wide, downward bent, with parallel dorsal
and ventral margins. Cauda with moderately long anterior
section and strongly downward curved posterior section and
tapering tip slightly curving forward. OCL : OCH = 1.3-1.5;
CCL : OCL = 0.65-0.8; caudal curvature index = 0.7-0.8;
ostial-caudal interspace 20-23 %. No dorsal depression or
ventral furrow. Outer face anteriorly thin, posteriorly with
broad, moderately strong postcentral umbo, smooth.
Discussion: Polycirrhus jaramilloi occurs parallel to
P. schwarzhansi and differs primarily in the somewhat more
elongate form, which is best expressed in specimens of more
than 8 mm length (OL : OH = 1.7-1,75 vs. 1.4-1.55), and
the anteriorly relatively thin otoliths (vs. thickset throughout).
Distribution: Burdigalian to Langhian of Colombia and
Venezuela.
Polycirrhus mustus n. sp.
(Plate 7, Figs. 1-3)
Holotype: Pl. 7, Fig. 2, MUN2018, Castilletes, location 390090,
Colombia, Jimol FM, Burdigalian.
Paratypes: 3 specimens; 2 specimens, Castilletes, Colombia, loca-
tion 290616 (MUN2019) and location 290632 (MUN2020),
Castilletes FM, late Burdigalian to Langhian; 1 specimen NMB
P1036, San José de Cocodite, Paraguaná Península, Venezuela,
Cantaure FM, late Burdigalian to Langhian.
Name: From mustus (Latin = fresh, young), referring to
the modern morphological otolith pattern, which appears
transitional between Polycirrhus and Paralonchurus.
Diagnosis: OL : OH = 1.95-2.1. Ostium large, OCL : OCH
= 1.5-1.6, slightly downward bent with deflected dorsal
margin. Downward bent portion of cauda leaving moder-
ately wide posterior field to posterior rim of otolith. Otolith
anteriorly thin, posteriorly with massive, tuberculate umbo
on outer face.
Description: Elongate, thick otoliths to about 14.5 mm
length. OL : OH = 1.95-2.1; OH : OT = 1.1-1.3. Predorsal
lobe slightly forward expanded; dorsal rim flat, somewhat
undulating, without prominent angles; ventral rim shallow,
almost flat, smooth; anterior rim very broadly and regularly
rounded; posterior tip vertically cut, with moderate angles
at junctions with dorsal and ventral rims.
Inner face distinctly convex, with median, large sulcus.
Ostium large, wide, downward bent, with deflected dorsal
margin at about mid-section and gently anteriorly inclined
ventral margin. Cauda with moderately long anterior
section and strongly downward curved posterior section
and tapering tip slightly curving forward. Downward bent
portion of cauda reaching moderately close to posterior
rim of otolith leaving a moderately wide posterior field in
between. OCL : OCH = 1.5-1.6; CCL : OCL = 0.6-0.75;
caudal curvature index = 0.7-0.8; ostial-caudal interspace
14-20 %. Occasionally narrow dorsal depression visible;
no ventral furrow. Outer face anteriorly thin, posteriorly with
massive, thick and strongly ornamented postcentral umbo.
Discussion: Polycirrhus mustus is the most elongate of the
three Polycirrhus species occurring simultaneously in the
early Miocene of Venezuela and Colombia and also the one
with the most strongly developed umbo on the outer face.
Further, it is characterized by a marked deflection of the
dorsal margin of the ostium. The overall shape resembles
otoliths of the genus Paralonchurus, but the strongly bent in-
ner face and the moderately wide dorsal and posterior fields
of the inner face are more in agreement with Polycirrhus.
Distribution: Burdigalian to Langhian of Colombia and
Venezuela.
Polycirrhus schwarzhansi
(AGUILERA & RODRIGUES DE AGUILERA, 2004)
(Plate 7, Figs. 16-22)
2004 Paralonchurus schwarzhansi – AGUILERA & RORDIGUES
DE AGUILERA: pl. 1, figs. 19-22
Material: 26 specimens; 10 specimens, Castilletes, Colombia,
locations 16344, 16519,16555, 138110, 290674, 390094,
390126, GABo35, Castilletes FM, late Burdigalian to Langhian;
6 specimens Castilletes, Colombia, Jimol FM, late Burdigalian,
locations 290602, 390085; 10 specimens, figured specimens
NMB P1037-1038, San José de Cocodite, Paraguaná Penín-
sula, Venezuela, Cantaure FM, late Burdigalian to Langhian.
Description: Moderately compressed and moderately
thick otoliths to about 10 mm length. OL : OH = 1.4-1.55;
OH : OT = 1.9-2.0. Predorsal lobe indistinct or weak, then
slightly forward expanded; dorsal rim flat, somewhat un-
Plate 7
Figs. 1-3. Polycirrhus mustus n. sp. 2 (r), holotype, MUN2018,
Colombia, La Guajira, Castilletes FM, early Miocene; 1, paratype,
NMB P1036, Venezuela, San José de Cocodite, Cantaure FM,
early Miocene; 3, paratype, MUN2019, Colombia, La Guajira,
Castilletes FM, early Miocene.
Figs. 4-5. Polycirrhus dumerilii BOCOURT, 1869; coll. Aguirre,
off Ecuador, Recent.
Figs. 6- 8. Polycirrhus trinidadensis (SCHWARZHANS, 1993).
6, Venezuela, El Hatillo, Urumaco FM, late Miocene; 7-8, Trinidad,
Manzanilla FM, late Miocene.
Figs. 9-15. Polycirrhus jaramilloi n. sp. 9, holotype MUN2014,
Colombia, La Guajira, Castilletes FM, early Miocene; 12-13 (r),
15 (r), paratypes, MUN2015-2017, Colombia, La Guajira, Cas-
tilletes FM, early Miocene; 10-11, 14, paratypes, NMB P1033-
1035, Venezuela, San José de Cocodite, Cantaure FM, early
Miocene.
Figs. 16-22. Polycirrhus schwarzhansi (AGUILERA & SCHWARZ-
HANS, 2014). 16-17 (r), 19-20 (r), Colombia, La Guajira, Cas-
tilletes FM, early Miocene; 18 (r), 21-22 (r), Venezuela, San José
de Cocodite, Cantaure FM, early Miocene.
/
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
41
Palaeo Ichthyologica 14
2 mm
2 mm
2 mm
2 mm
2 mm
Polycirrhus mustus
Polycirrhus dumerilii
Polycirrhus jaramilloi
Polycirrhus schwarzhansi
Polycirrhus trinidadensis
1a
1b
2a
2b
2c
3
4a
4b
5a
5b
6a
6b
7
8a
8b
9a
9b
10a
10b
10c
11
12
13a
13b
14
15
16a
16b
16c
17a
17b
17c
18
19
20
21
22
42
dulating, without prominent angles; ventral rim shallow,
rather regularly curved, smooth; anterior rim very broadly
rounded, dorsally pronounced; posterior tip vertically cut,
with moderate angle at junction with dorsal rim and broadly
rounded at junction with ventral rim.
Inner face distinctly convex, with slightly supramedian,
large sulcus. Ostium large, wide, downward bent, its dor-
sal margin more strongly inclined than its ventral margin.
Cauda with short anterior section and strongly downward
curved posterior section and tapering tip slightly curving
forward. Area behind downward bent portion of cauda to
posterior rim moderately narrow. OCL : OCH = 1.5-1.6;
CCL : OCL = 0.75-0.8; caudal curvature index = 0.6-0.7;
ostial-caudal interspace 18- 22 %. Occasionally narrow
dorsal depression visible; no ventral furrow. Outer face
convex throughout, but with no distinct postcentral umbo.
Discussion: Polycirrhus schwarzhansi is the most com-
pressed of the three Polycirrhus species occurring simulta-
neously in the early Miocene of Venezuela and Colombia
and resembles most P. jaramilloi. It represents the most
plesiomorphic otolith morphology of the Lonchurini known
to date. The compressed outline reminds of otoliths of the
Sciaenini and it is mainly the shape of the ostium and
predorsal rim that suggests a placement in the Lonchurini.
Distribution: Late Burdigalian to Langhian of Colombia
and Venezuela.
Polycirrhus trinidadensis (SCHWARZHANS, 1993)
(Plate 7, Figs. 16-22)
1976 Polyclemus sp. – NOLF: pl. 8, fig. 9
1993 Polyclemus trinidadensis – SCHWARZHANS: figs. 227-
228
Material: 20 specimens; 4 specimens (holotype and paratypes), Cb
1633, San José River, Trinidad, Manzanilla FM, late Tortonian,
8 specimens, PPP 2667, 2669, Manzanilla Bay, PPP 2674,
2675, 2677, figured specimens NMB P1039-1040, San José
River, Trinidad, Manzanilla FM, San José and Montserrat Mbs,
late Tortonian-Messinian; 6 specimens, figured specimen NMB
P1041, El Hatillo, Venezuela, Urumaco FM middle member,
middle Tortonian; 1 specimen, PPP 08, Gatun, Panama, Gatun
FM, Tortonian; 1 specimen PPP 3459, Rio Santiago, Ecuador,
Angostura FM, Tortonian.
Description: Moderately compressed and moderately
thick otoliths to about 7 mm length. OL : OH = 1.8-1.9;
OH : OT = 1.3-1.8. Predorsal lobe indistinct, sometimes
very slightly forward expanded; dorsal rim flat, somewhat
undulating, without prominent angles; ventral rim shallow,
nearly flat, smooth; anterior rim broadly rounded, some-
times dorsally pronounced; posterior tip vertically cut, with
moderate angle at junctions with dorsal and ventral rims.
Inner face moderately convex, with slightly supramed-
ian, large sulcus. Ostium large, wide, downward bent,
its dorsal margin more strongly inclined than its ventral
margin. Cauda with short anterior section and strongly
downward curved posterior section and tapering tip slightly
curving forward. Area behind downward bent portion of
cauda to posterior rim very narrow. OCL : OCH = 1.3-1.4;
CCL : OCL = 0.8-0.9; caudal curvature index = 0.75-0.85;
ostial-caudal interspace 20- 23 %. Occasionally narrow
dorsal depression visible; no ventral furrow. Outer face
convex throughout, but with no distinct postcentral umbo.
Discussion: Polycirrhus trinidadensis so far is only known
from relatively small otoliths. Comparable otoliths of the Re-
cent P. dumerilii (pl. 7, figs. 4-5) are larger, 10-13 mm long.
Thus, the observed differences of the Recent otoliths such as
the massive umbo on the outer face, the sharply developed
postdorsal angle and the longer ostium (OCL : OCH = 0.65-
0.75 vs. 0.8-0.9) could be ontogenetically induced. We
maintain P. trinidadensis as a valid species, but consider the
distinction from P. dumerilii not satisfactory resolved without
larger fossil specimens having become available.
Distribution: Polycirrus trinidadensis is widely distributed
in the late Miocene, Tortonian of tropical America, from
Ecuador to Panama, Venezuela and Trinidad, much wider
in fact as the Recent P. dumerilii, which is confined to the
Pacific shores of America from El Salvador to northern Peru.
Subfamily Lariminae GILL, 1861
Tribe Larimini SASAKI, 1989
Definition and Discussion: The tribe Larimini contains the
three extant genera Larimus, Nebris, Paranebris and the
newly established
†
Protonebris n. gen. We follow the defi-
nition of SASAKI (1989), while CHAO (1978) had Larimus
and Nebris in two separate groups.
Genus Larimus CUVIER, 1830
Larimus angosturae n. sp.
(Plate 8, Figs. 11-17)
Holotype: Pl. 8, Fig. 11, NMB P1044, PPP 3406, Río Cayapas,
Ecuador, Angostura FM, Tortonian.
Paratypes: 6 specimens; 3 specimens NMB P1045-1047, PPP
3432, Río Cayapas, and PPP 3441, Río Santiago, Ecuador,
Angostura FM, Tortonian; 3 specimens NMB P1048-1050,
PPP 1726, 1727, 1728, Río Banano, Costa Rica, Río Banano
FM, late Pliocene.
Further material: 8 specimens; 1 specimen, PPP 3440, Río
Santiago, Ecuador, Angostura FM, Tortonian; 7 specimens,
PPP 1726, 1727, 1732, 1734, Río Banano, Costa Rica, Río
Banano FM, late Pliocene.
Name: Named after the stratum typicum, the Angostura
Formation of late Miocene, Tortonian age in Equador.
Diagnosis: OL : OH = 1.35-1.45. Predorsal lobe well
developed; ventral rim moderately deep; posterior tip nar-
rowed. Ostium relatively small, anteriorly narrowed. Ventral
furrow well developed, running oblique from below middle
of ostium to caudal tip.
Plate 8
Figs. 1-2. Larimus argenteus (GILL, 1863); Ecuador, Jama FM,
middle Pleistocene.
Figs. 3-7. Larimus breviceps CUVIER, 1830. 3, 6-7, Trinidad,
Manzanilla FM, late Miocene; 4-5 (r), Dominican Republic, Cer-
cado FM, late Miocene.
Figs. 8-9. Larimus gatunensis (SCHUBERT, 1909). 8, Panama,
Gatun FM, late Miocene; 9 (r), Ecuador, Onzole FM, late Miocene.
Fig. 10. Larimus sp. 1; Venezuela, El Hatillo, Urumaco FM, late
Miocene.
Figs. 11-17. Larimus angosturae n. sp. 11, holotype, NMB P1044;
Ecuador, Angostura FM, late Miocene; 12 (r), 14, 17, paratypes,
NMB P1045-1047, Ecuador, Angostura FM, late Miocene; 13, 15-
16, paratypes, NMB P1048-1050, Costa Rica, Rio Banano FM,
late Pliocene.
/
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
43
Palaeo Ichthyologica 14
Larimus argenteus
Larimus breviceps
Larimus angosturae
1a
1b
1c
2
3a
3b
4a
4b
6
7a
7b
7c
5b
5a
8a
8b
8c
9a
9b
11a
11c
10a
10b
11b
12a
12b
13a
13b
14
15
16b
16a
17a
17b
2 mm
2 mm
2 mm
2 mm
Larimus gatunensis
2 mm
Larimus sp. 1
44
Description: Compressed and moderately thick otoliths up
to 7.5 mm length. OL : OH = 1.35-1.45; OH : OT = 1.5-
1.8. Predorsal lobe well developed above anterior part
of ostium; dorsal rim straight, slightly inclined backwards,
slightly undulating at times; ventral rim moderately deep,
deepest slightly in front of its middle below rear part of
ostium, posteriorly rather straight and turning upwards,
smooth; anterior rim broadly rounded; posterior rim blunt,
with distinct angle at junctions with ventral rim.
Inner face convex, with slightly supramedian, moderately
large sulcus. Ostium moderately large, anteriorly narrowed
and closed, its dorsal margin slightly deflected anteriorly, its
ventral margin with notch at about middle and curving up-
wards towards anterior. Cauda deep, with downward curved
posterior section not or only slightly widened and tapering
tip not bend forward. Area behind downward bent portion
of cauda to posterior rim narrow. OCL : OCH = 1.3-1.4;
CCL : OCL = 0.7-0.8; caudal curvature index = 0.7-0.8;
ostial-caudal interspace 18-22 %. Very narrow and indistinct
dorsal depression, but sometimes marked dorsally by edge
close to dorsal rim of otolith; distinct, sharp ventral furrow
beginning at ventral rim below middle of ostium and run-
ning inwards to meet caudal tip. Outer face strongly convex
throughout, with broad postcentral umbo, rather smooth.
Ontogeny: Larimus otoliths show an unusual ontogeny in
that small specimens have a much smaller ostium than large
specimens from fully adult fishes. We do see this trend within
the specimens of L. angosturae from specimens of about
3.5-5.0 mm length (figs. 12-15), which show a distinctly
narrower ostium with a shallower ventral margin than those
of the largest known specimens between 7.0 and 7.5 mm
length. Since the ontogenetic change seems to be more or
less completed, we assume that L. angusturae actually rep-
resented a smaller species than most other Larimus species.
Discussion: Larimus angosturae represents the most com-
pressed otolith known to date in the genus. It is further
characterized by a relatively small ostium, a deep anterior
ventral rim, a well developed predorsal lobe and a sharp
ventral furrow. In terms of general appearance, the closest
resemblance is with the Recent Larimus effulgens GILBERT,
1898 from the American Pacific coast from Baja Califor-
nia to northern Peru, which however differs in being less
compressed (OL : OH = 1.5-1.6 vs. 1.35-1.45), showing a
broad instead of a sharp ventral furrow, a much less deeply
curved anterior-ventral rim and a larger ostium.
Larimus angosturae is known from two locations of rather
different geographical origin and stratigraphical age: late
Miocene of Ecuador and late Pliocene of Costa Rica, Atlan-
tic side. The Costa Rican specimens appear to be slightly
more compressed at average than the older Ecuadorian
ones, but, as we felt, not sufficient with this knowledge
base for taxonomic distinction. In the Caribbean Pliocene,
L. angosturae probably represented the extinct geminate
sister species to the Pacific Larimus effulgens.
Distribution: Amphiamerican, in the Tortonian of Ecuador
and the Piacenzian of the Atlantic Costa Rica.
Larimus argenteus (GILL, 1863)
(Plate 8, Figs. 1-2)
Material: 5 specimens, PPP 3562, 3562, 3565, 3566, figured
specimens NMB P1051-1052, Punta la Cereza, Jama, Ecuador,
Jama FM, middle Pleistocene.
Discussion: Recent otoliths of L. argenteus are readily rec-
ognized by their projecting, expanded preventral angle set
off from the rest of the inner face by a broad, groove-like
ventral furrow (see SCHWARZHANS, 1993, figs. 330-331).
This character safely distinguishes otoliths of L. argenteus
from the geminate Caribbean Larimus breviceps CUVIER,
1830. However, all otoliths studied from Recent specimens
so far were 10 mm long or larger, while those from the
Pleistocene of Ecuador are 8 mm or shorter. The Pleistocene
otoliths exhibit a less angular and deep development of
the preventral rim than those larger Recent otolith, and we
interpret this subtle difference as an expression of a late
stage ontogenetic change.
Distribution: Middle Pleistocene of Ecuador and in the
Recent from Baja California to Peru.
Larimus breviceps CUVIER, 1830
(Plate 8, Figs. 3-7)
1976 Larimus breviceps CUVIER, 1830 – NOLF: pl. 7, fig. 9
(non fig. 10)
1992 Larimus breviceps CUVIER, 1830 – NOLF & STRINGER:
pl. 16, fig. 12
1993 Larimus gatunensis (SCHUBERT, 1909) – SCHWARZHANS:
figs. 338, 339 (non fig. 337)
Material: 52 specimens; 4 specimens, HF 98-16-1, PPP 2667,
Manzanilla Bay, PPP 2675, 2676, San José River, figured
specimens NMB P1053-1055, Trinidad, Manzanilla FM, late
Tortonian; 48 specimens, NMB 15903, 15904, Río Gurabo,
Dominican Republic, Cercado FM, late Tortonian-Messinian,
NMB P326, 340.
Discussion: Otoliths of L. breviceps share with those of its
geminate counterpart from the Pacific shores of tropical
America, L. argenteus, the anteriorly reduced ostium, the
expanded predorsal lobe and the strong preventral angle
set off from the rest of the inner face by a rather straight,
tangential groove-like ventral furrow (see SCHWARZHANS,
1993, figs. 332-333 for Recent otoliths of L. breviceps). It
differs from L. argenteus in the much less strongly devel-
oped preventral angle, which is well depicted in the large
specimen of about 10 mm length of figure 3. The smaller
specimens of the figures 4 and 5 (5 and 6 mm length)
show the small and even more anteriorly reduced ostium
typical for many subadult and juvenile otoliths of Larimus.
Larimus otoliths have been found in the late Miocene of
Trinidad in the Manzanilla FM (planktonic foraminifera zone
N 16) and the Tamana FM (planktonic foraminifera zone
N 15). NOLF (1976) placed all specimens in L. breviceps
while SCHWARZHANS (1993) placed them in the fossil
L. gatunensis. Newly collected specimens now reveal that
the younger specimens from the Manzanilla FM belong to
L. breviceps, while the older specimens from the Tamana
FM belong to L. gatunensis. Otoliths of L. gatunensis are
distinguished from those of L. breviceps by the shallower
anterior-ventral rim, the absence of a predorsal projection
and a larger ostium reaching closer to the anterior tip of
the otolith.
Distribution: Late Tortonian of Trinidad and in the Recent
throughout the Caribbean and southwards to Rio de Ja-
neiro in Brazil.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
45
Palaeo Ichthyologica 14
Larimus fasciatus HOLBROOK, 1855
(Plate 9, Figs. 8-10)
Material: 8 specimens PPP 2567, 2572, 3093, figured specimens
NMB P1058-1060, Margarita Island and Cubagua Island,
Cañón de las Calderas and Bahia de Charagato, Cubagua
FM, above Cerro Negro member, late Pliocene.
Description (of the fossil specimens): Moderately elon-
gate and thick otoliths up to about 10 mm length.
OL : OH = 1.55-1.7; OH : OT about 1.2. Predorsal lobe low,
somewhat forward projecting; dorsal rim shallow, without
prominent angles, highest above rear part of dorsal mar-
gin of ostium; ventral rim shallow, gently curving, without
preventral projection, smooth; anterior and posterior rims
broadly rounded; posterior rim with rounded angle at junc-
tions with ventral rim.
Inner face strongly convex, with median, large sulcus.
Ostium very large, wide, closed anteriorly, but reaching
close to anterior and anterior-ventral rims of otolith, its
dorsal margin turned downward anteriorly, its ventral margin
with feeble notch at mid-section. Cauda very deep, with
widened downward curved posterior section and tapering
tip not bend forward. Area behind downward bent portion
of cauda to posterior rim very narrow. OCL : OCH = 1.2-
1.35; CCL : OCL = 0.7-0.8; caudal curvature index = 0.9;
ostial-caudal interspace 15-20 %. Very narrow and indistinct
dorsal depression; no ventral furrow. Outer face strongly
convex throughout, with massive postcentral umbo, mildly
tuberculate.
Discussion: Otoliths of L. fasciatus and its putative Pacific
geminate counterpart, Larimus pacificus JORDAN & BOLL-
MAN, 1890, differ from all other Recent Larimus otoliths in
the narrow ventral field below the ostium and the absence
of any indication of a ventral furrow. The otoliths of both
species differ mainly in those of L. pacificus being more
elongate than those of L. fasciatus (OL : OH = 1.75-1.95
vs. 1.55-1.7).
Distribution: Early Pliocene of Venezuela and in the Recent
along the Northeast American shores from Massachusetts
to the Gulf of Mexico, except southern Florida, but missing
from the Caribbean.
Larimus gatunensis (SCHUBERT, 1909)
(Plate 8, Figs. 8-9)
1909 Eques gatunensis – SCHUBERT: fig. 4
1976 Larimus breviceps CUVIER, 1830 – NOLF: pl. 7, fig. 10
(non fig. 9)
1992 Larimus breviceps CUVIER, 1830 – NOLF & STRINGER:
pl. 16, fig. 12
1993 Larimus gatunensis (SCHUBERT, 1909) – SCHWARZHANS:
fig. 337 (non figs. 338, 339)
Material: 5 specimens; 4 specimens, At-14-19-1 and At-14-21-1
(figured specimen NMB P1056), San Judas, Panama, Gatun
FM, Tortonian; 1 specimen NMB P1042, PPP 3470, Río San-
tiago, Ecuador, Onzole FM, Messinian.
Description: Moderately elongate and thick otolith up to
8.5 mm length. OL : OH = 1.7; OH : OT = 1.4. Predorsal
lobe moderately developed above anterior part of ostium;
dorsal rim slightly depressed at mid-section, without promi-
nent angles; ventral rim shallow, anteriorly gently curving,
posteriorly straight and turning upwards at low angle,
smooth; anterior rim broadly rounded; posterior rim blunt,
with distinct angle at junctions with ventral rim.
Inner face convex, with slightly supramedian, large
sulcus. Ostium very large, wide, closed anteriorly, its dorsal
margin slightly deflected anteriorly, its ventral margin with
notch at anterior third. Cauda very deep, with widened
downward curved posterior section and tapering tip not
bend forward. Area behind downward bent portion of
cauda to posterior rim narrow. OCL : OCH = 1.3-1.4;
CCL : OCL = 0.6-0.7; caudal curvature index = 0.7-0.8;
ostial-caudal interspace 20 % (14 % in the Ecuadorian
specimen). Very narrow and indistinct dorsal depression;
broad, indistinct ventral furrow below ostium and up to
caudal tip. Outer face strongly convex throughout, with
massive postcentral umbo, rather smooth.
Discussion: Distinction of these otoliths from the various
living species and other fossil otolith-based species is not
easy and often depends on subtle morphometric characters
and sulcus and otolith relations. Otoliths of L. gatunensis
closely resemble the Pacific Larimus acclivis JORDAN &
BRISTOL, 1898, differing mainly in the narrower sulcus
(OCL : OCH = 1.3-1.4 vs. 1.15) and the less prominently
expanded predorsal lobe. Larimus effulgens GILBERT, 1898,
also from the Pacific, is slightly more compressed than L. ga-
tunensis (OL : OH = 1.5-1.6 vs. 1.7). NOLF (1976) placed
these specimens in L. breviceps, extant in the West Atlantic,
which however shows an expanded preventral rim marked
by a straight, tangential ventral furrow on the inner face
(see SCHWARZHANS, 1993 for figures of Recent otoliths).
The single specimen from the Onzole FM of Ecuador
(pl. 8, fig. 9) differs from the slightly older specimens from
Panama and Trinidad in the oblique anterior/vertical margin
of the cauda resulting in a smaller ostial caudal interspace
(14 % vs. 20 %). If supported by further specimens in the
future, this character could indicate the presence of a further
species.
Distribution: Late Miocene, Tortonian of Panama and
Trinidad and tentatively Messinian of Ecuador.
Larimus humboldti n. sp.
(Plate 9, Figs. 1-3)
Holotype: Pl. 9, Fig. 1, NMB P1061, OA 95-2, San José de
Cocodite, Paraguaná Peninsula, Venezuela, Cantaure FM, late
Burdigalian to early Langhian.
Paratypes: 4 specimens; 1 specimen NMB P1062, same data as
holotype; 3 specimens Jimol FM, Castilletes, Colombia, Burdi-
galian, 1 specimen MUN2022, location 290602, 2 specimens
MUN2021, location 390090.
Name: A homage to Alexander von Humboldt (1769-
1859) in recognition of his early pioneering contribution to
the natural history of South America, and of whom Simon
Bolivar wrote that he was the real discoverer of South
America, “since his work was more useful for our people
than that of all the conquerors”.
Diagnosis: OL : OH = 1.45-1.55. Predorsal lobe low, an-
teriorly projecting; ventral rim gently and regularly curved.
Ostium large, with only incipient indention on ventral
margin and reaching close to anterior rim of otolith. No
ventral furrow.
Description: Moderately compressed, thick otoliths up to
about 9 mm length. OH : OT = 1.1-1.2. Predorsal lobe low,
somewhat forward projecting; dorsal rim shallow, rather
regularly curved, without prominent angles, highest above
anterior part of cauda; ventral rim very gently and regularly
curved, without preventral projection, smooth; anterior and
posterior rims broadly rounded; posterior rim with rounded
angle at junctions with ventral rim.
46
Inner face strongly convex, with median, large sulcus.
Ostium large, wide, reaching very close to anterior rim of
otolith, its dorsal margin somewhat deflected anteriorly,
its ventral margin with incipient indention at mid-section.
Cauda very deep, with steeply downward curved posterior
section and tapering tip slightly bend forward. Area behind
downward bent portion of cauda to posterior rim narrow.
OCL : OCH = 1.15-1.2; CCL : OCL = 0.7-0.8; caudal cur-
vature index = 0.65-0.7; ostial-caudal interspace about
18 %. Very narrow and indistinct dorsal depression; no
ventral furrow. Outer face strongly convex throughout, with
massive postcentral umbo, mildly tuberculate.
Discussion: Larimus humboldti represents the earliest spe-
cies of the genus Larimus so far and shows certain characters
considered here as plesiomorphic for the genus, such as
the gently curved dorsal and ventral rims, the shape of the
ostium and its close reach towards the anterior rim of the
otolith and the lack of a ventral furrow. The latter character
it shares with the Recent species L. fasciatus and L. pacificus.
Distribution: Burdigalian to early Langhian of Venezuela
and Colombia.
Larimus pacificus JORDAN & BOLLMAN, 1890
(Plate 9, Figs. 11-15)
Material: 16 specimens PPP 3515, 3529, 3537 and one unre-
corded location, figured specimens NMB P1063-1067, Punta
Canoa, Ecuador, Canoa FM, early Pleistocene.
Discussion: Larimus pacificus is here considered the gemi-
nate species of the Atlantic L. fasciatus. Its otoliths differ
mainly in being more elongate (OL : OH = 1.75-1.95 vs.
1.55-1.7), but small specimens of 7 mm length or less may
be more compressed and then distinction of both species
by means of otoliths may become improbable.
Distribution: Early Pleistocene of Ecuador and in the Recent
in the East Pacific from Mexico to Peru.
Larimus pandus n. sp.
(Plate 9, Figs. 4-5)
Holotype: Pl. 9, Fig. 4, NMB P1068, PPP 2164, Payardi Island,
Colón, Atlantic Panama, Gatun FM, middle member, middle
Tortonian.
Paratypes: 2 specimens NMB P1069-1070, PPP 3587, Favio
Alvaro road cut, Ecuador, Angostura FM, Tortonian.
Name: From pandus (Latin = bowed, curved) referring to
the regularly curved outline of the otoliths.
Diagnosis: OL : OH = 1.45-1.55. Predorsal lobe low, an-
teriorly projecting; ventral rim gently and regularly curved.
Ostium large, with only incipient indention on ventral margin
and reaching close to anterior rim of otolith. Ostial-caudal
interspace = 13-15 %. No ventral furrow.
Description: Moderately compressed, thick otoliths up to
about 9 mm length. OH : OT = 1.1-1.2. Predorsal lobe low,
somewhat forward projecting; dorsal rim shallow, rather
regularly curved, without prominent angles, highest above
anterior part of cauda; ventral rim very gently and regularly
curved, without preventral projection, smooth; anterior and
posterior rims broadly rounded; posterior rim with rounded
angle at junctions with ventral rim.
Inner face strongly convex, with median, large sulcus.
Ostium large, wide, reaching very close to anterior rim of
otolith, its dorsal margin somewhat deflected anteriorly,
its ventral margin with incipient indention at mid-section.
Cauda very deep, with steeply downward curved posterior
section and tapering tip not bent forward. Area behind
downward bent portion of cauda to posterior rim narrow.
OCL : OCH = 1.25-1.3; CCL : OCL = 0.7; caudal curvature
index = 0.7; ostial-caudal interspace = 13-15 %. Very nar-
row and indistinct dorsal depression; no ventral furrow.
Outer face anteriorly flat, posteriorly strongly convex, with
massive postcentral umbo, mildly tuberculate.
Discussion: Larimus pandus belongs to the Larimus species
without expanded anterior-ventral rim and without ventral
furrow, like the Recent species L. fasciatus and L. pacifi-
cus. Otoliths of L. pandus are less elongate than those of
L. pacificus and slightly less elongate than those of L. fas-
ciatus (OL : OH = 1.45-1.55 vs. 1.75-1.95 and 1.55-1.7
respectively) and the ostial-caudal interspace is narrower
than in L. fasciatus (13-15 % vs. 15-20 %). Larimus pandus
likely represents the common ancestor of the two geminate
Recent species.
Distribution: Tortonian of Panama and Ecuador.
Larimus sp. 1
(Plate 8, Fig. 8)
Material: 8 specimens; 7 specimens location OA 98-9, 99-19,
00-3 (figured specimen NMB P1057), El Hatillo, Venezuela,
Urumaco FM middle member, middle Tortonian; 1 eroded
specimen PPP 2225, east coast of Cayo Agua Island, Atlantic
Panama, Caya Agua FM, early Pliocene.
Discussion: Unfortunately, all otoliths currently available
from Urumaco and Cayo Agua are too much eroded for
a reliable definition. They resemble the extant L. breviceps
in the medioventral expansion associated with a ventral
furrow on the inner face, but this expansion is much less
strongly developed than in the Recent species. They differ
also in the low, almost depressed predorsal rim and the
very large ostium that reaches close to the anterior rim
of the otolith. They probably are an undescribed species
that represents the ancestor to the geminate extant species
L. argenteus and L. breviceps.
Larimus sp. 2
(Plate 9, Figs. 6-7)
Material: 2 juvenile to subadult specimens, 1 specimen San José de
Cocodite, Paraguaná Peninsula, Venezuela, Cantaure FM, late
Burdigalian to early Langhian; 1 specimen location 390090,
Castilletes, Colombia, Jinol FM, Burdigalian.
Discussion: The two small specimens of 6 and 3 mm length
differ from the contemporaneous L. humboldti in the more
/
Plate 9
Figs. 1-3. Larimus humboldti n. sp. 1 (r), holotype, NMB P1061,
Venezuela, San José de Cocodite, Cantaure FM, early Miocene;
2-3 (r), paratypes, MUN2021, Colombia, La Guajira, Castilletes
FM, early Miocene.
Figs. 4- 5. Larimus pandus n. sp. 4, holotype, NMB P1068,
Panama, Gatun FM, late Miocene; 5, paratype, Ecuador, Angos-
tura FM, late Miocene.
Figs. 6-7. Larimus sp. 2. 6 (r), Colombia, La Guajira, Castilletes
FM, early Miocene; 7, Venezuela, San José de Cocodite, Cantaure
FM, early Miocene.
Figs. 8-10. Larimus fasciatus HOLBROOK, 1855; Venezuela,
Margarita Island, Cubagua FM, late Pliocene.
Figs. 11-15. Larimus pacificus JORDAN & BOLLMAN, 1890;
Ecuador, Canoa FM, early Pleistocene.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
47
Palaeo Ichthyologica 14
Larimus fasciatus
Larimus pacificus
2 mm
2 mm
Larimus humboldti
Larimus sp. 2
11a
11b
11c
11d
12
10
13
14
15a
15b
Larimus pandus
4a
4b
4c
5a
5b
5c
6a
6b
7
8a
8b
8c
9
1a
1b
1c
1d
2
3a
3b
3c
2 mm
2 mm
2 mm
48
compressed and thinner appearance (OL : OH = 1.25-1.35;
OH : OT of the larger specimen = 1.4). Its outline with the
tapered posterior tip resembles Umbrina abbreviata, but the
anteriorly narrowed ostium and the massive umbo on the
outer face is characteristic for Larimus otoliths. The specimens
likely represent yet another species of Larimus, but non of
the otoliths now available is considered large enough to
warrant full display of all pertinent diagnostic characters.
Genus Nebris CUVIER, 1830
Nebris dioneae n. sp.
(Plate 10, Figs. 14-15)
Holotype: Pl. 10, Fig. 14, NMB P1071, El Hatillo, Venezuela,
Urumaco FM middle member, middle Tortonian.
Paratypes: 3 specimens NMB P1072-1074, same data as holo-
type.
Name: In honor of Dione Rodrigues de Aguilera, the wife
of the senior author, for her relentless support of their joint
scientific work.
Diagnosis: OL : OH = 2.0; OH : OT = 1.8. Predorsal rim
depressed, straight inclined or concave; ventral rim shallow.
Posterior-ventral lobe more projecting than posterior-dorsal
lobe. Ostium very large, anteriorly closed, but reaching close
to anterior, predorsal and preventral rims. No ventral furrow.
Description: Elongate, rather thin otoliths up to about
12 mm length. Dorsal rim shallow, anteriorly inclined down-
wards straight or concave, its mid and posterior sections
almost horizontal, straight; ventral rim shallow, very gently
curved, smooth; anterior rim pointed, its ventral margin
convex; posterior rim with broad central concavity typical
for Nebris otoliths and posterior-ventral and posterior-dorsal
lobes, the former being longer than the latter.
Inner face convex, with median, large sulcus. Ostium
very large, wide, anteriorly closed except for narrow ostial
channel connecting to tip of otolith, but nevertheless reach-
ing very close to anterior-ventral and anterior-dorsal rims of
otolith. Cauda very deep, very narrow at joint with ostium,
with short anterior portion and widened, kink-bent poste-
rior portion and marked ventral incision at junction. Area
behind downward bent portion of cauda to posterior rim
very narrow except for expanded lobes. OCL : OCH = 1.7-
2.0; CCL : OCL = 0.5; caudal curvature index = 0.5; ostial-
caudal interspace extremely narrow, less than 5 %. Narrow
dorsal depression only above anterior part of cauda; no
ventral furrow. Outer face flat to slightly concave, without
marked umbo.
Discussion: Otoliths of the genus Nebris are readily rec-
ognized by the specific outline with its two posterior lobes
and the highly diagnostic shape of the sulcus that looks like
a Larimus sulcus pattern taken to the extremes. Otoliths of
Nebris (and Paranebris) are the ones with the narrowest
ostial-caudal interspace found in all sciaenids. In the Recent,
two closely related geminate species occur on either side
of the Isthmus of Panama: Nebris microps CUVIER, 1830
(pl. 10, fig. 13) (Atlantic from Colombia to SE-Brazil) and
Nebris occidentalis VAILLANT, 1897 (pl. 10, fig. 12) (Pacific
from Guatemala to Peru). The differences of the otoliths of
the two Recent species are subtle: those of N. microps are
thicker and more compressed than those of N. occiden-
talis (OL : OH = 1.75 vs. 1.85 and OH : OT = 1.3 vs. 1.5
respectively) and the posterior-ventral lobe is longer than
the posterior-dorsal lobe (vs. equally long). The distinction
of N. dioneae from both Recent species likewise is subtle,
albeit to a lesser degree. Otoliths of N. dioneae are more
elongate than both Recent species (OL : OH = 2.0 vs. 1.85
and 1.75 respectively) and thinner (OH : OT = 1.8 vs. 1.5
and 1.3 respectively), the anterior dorsal margin is straight
to concave (vs. convex), the dorsal rim of the otolith is not
reduced above the rear part of the ostium (vs. reduced
with a distinct step-change behind) and the ostium reaches
closer to the anterior-ventral rim of the otolith. We assume
that N. dioneae represents the common ancestor of the
two recent species.
Distribution: Tortonian of Venezuela.
Nebris sp. (juveniles)
(Plate 10, Figs. 6-7)
Remarks: Juvenile otoliths of Nebris not exceeding 5 mm
length have been found in various locations such as the
Manzanilla FM (late Tortonian of Trinidad) (pl. 10, fig. 6),
the Rio Banano FM (late Pliocene of Costa Rica) (pl. 10,
fig. 7) and the Jama FM (middle Pleistocene of Ecuador).
Ontogenetical differences in Nebris otoliths are profound
with juveniles being much more compressed and thinner
than adults, showing a strongly reduced ostium similar to
the observations made in the assumed related genus Larimus
and a still wider ostial-caudal interspace in the order of
9-12 %, prohibiting specific identification of these otoliths
until large, adult specimens have become available from
the respective formations.
Genus Paranebris
CHAO, BÉAREZ & ROBERTSON, 2001
Paranebris bauchotae
CHAO, BÉAREZ & ROBERTSON, 2001
(Plate 10, Figs. 8-11)
Material: 3 specimens; 1 specimen NMB P1075, PPP 1149, Río
Tupisa, Darien, Panama, Yaviza FM, middle Tortonian; 2 speci-
mens NMB P1076-1077, PPP 3471, Río Santiago, Ecuador,
Onzole FM, Messinian.
/
Plate 10
Fig. 1. Protonebris sanchezi n. sp.; holotype, NMB P1078, Ven-
ezuela, Cerro la Cruz, Castillo FM, early Miocene.
Figs. 2-5. Paranebris steurbauti (NOLF & AGUILERA, 1998).
2 (r), 5 (r), Venezuela, San José de Cocodite, Cantaure FM, early
Miocene; 3, 4 (r), Colombia, La Guajira, Castilletes FM, early
Miocene.
Figs. 6-7. Nebris sp. juv. 6, Trinidad, Manzanilla FM, late Mio-
cene; 7, Costa Rica, Rio Banano FM, late Pliocene.
Figs. 8-10. Paranebris bauchotae CHAO, BÉAREZ & ROBERTSON,
2001; 8, 9 (r), Ecuador, Onzole FM, late Miocene; 10 (r), Panama,
Darien, Tuira FM, late Miocene.
Fig. 11. Paranebris bauchotae CHAO, BÉAREZ & ROBERTSON,
2001; coll. MNHN, Pacific off Panama, Recent.
Fig. 12. Nebris occidentalis VAILLANT, 1897; coll. WS, Pacific
off Panama, Recent.
Fig. 13. Nebris microps CUVIER, 1830; coll. OA, off Venezuela,
Recent.
Figs. 14-15. Nebris dioneae n. sp. 14, holotype, NMB P1071;
15, paratype, NMB P1072; Venezuela, El Hatillo, Urumaco FM,
late Miocene.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
49
Palaeo Ichthyologica 14
Protonebris sanchezi
Paranebris steurbauti
Nebris sp. juv.
Nebris dioneae
Paranebris bauchotae
Nebris occidentalis
Nebris microps
2 mm
2 mm
2 mm
2 mm
2 mm
1a
1b
1c
2a
2b
2c
3a
3b
3c
4
5
6a
6b
7
11a
11b
12a
12b
13a
13b
14a
14b
14c
15
Paranebris bauchotae
2 mm
8a
8b
8c
8d
8e
9
10
50
Description (of the fossil specimens): Elongate, thick
otoliths up to about 13.5 mm length. OL : OH = 1.8-1.9;
OH : OT = 1.2-1.3. Dorsal rim irregularly curved, anteriorly
inclined downwards straight or concave, highest above rear
dorsal margin of ostium; ventral rim shallow, very gently
curved, smooth; anterior rim rounded, its ventral margin
convex; posterior rim with markedly projecting posterior-
ventral lobe and rounded posterior-dorsal angle.
Inner face convex, with median, large sulcus. Ostium
very large, wide, anteriorly closed except for narrow ostial
channel connecting to tip of otolith, reaching very close to
anterior-ventral and anterior-dorsal rims of otolith. Cauda
very deep, narrowed just behind connection to ostium, with
short anterior portion and widened, kink-bent posterior
portion and marked by dorsal and ventral incisions at
junction. Area behind downward bent portion of cauda
to posterior rim very narrow except for expanded lobes.
OCL : OCH = 1.35-1.45; CCL : OCL = 0.55-0.6; caudal
curvature index = 0.55; ostial-caudal interspace extremely
narrow, less than 5 %. Remnant of dorsal depression only
above incised part of cauda at bent; no ventral furrow.
Outer face markedly convex, but without distinct umbo,
mildly tuberculate.
Discussion: Otoliths of Paranebris resemble closely those of
Nebris, differing mainly in the shape of the dorsal rim and
the ventrally and dorsally incised cauda at the location of it
kink-bent. We consider the otolith morphology of Paranebris
plesiomorphic as compared to that of Nebris, but advanced
apomorphic when compared to Larimus.
Distribution: Paranebris bauchotae is known from the
eastern Central Pacific off Panama from the Tortonian to
the Recent and apparently extended further southwards to
Ecuador during late Miocene.
Paranebris steurbauti (NOLF & AGUILERA, 1998)
(Plate 10, Figs. 2-5)
1998 Larimus steurbauti – NOLF & AGUILERA: pl. 11, figs. 5-8
Material: 28 specimens; 5 specimens San José de Cocodite, Para-
guaná Peninsula, Venezuela, Cantaure FM, Late Burdigalian
to early Langhian; 13 specimens locations 290448, 290610,
290632, 290636, 290840, 390094 Castilletes, Colombia,
Castilletes FM, late Burdigalian to early Langhian.
Discussion: Otoliths of P. steurbauti are readily distinguished
from those of the late Miocene to Recent P. bauchotae by
the slightly more compressed shape (OL : OH = 1.6-1.7
vs. 1.8-1.9), thinner appearance (OH : OT = 1.6-2.0 vs.
1.2-1.3), the wider ostium (OCL : OCH = 1.2-1.3 vs. 1.35-
1.45), the somewhat less widened downturned part of the
cauda and the lack of the incisions of the anterior part of
the cauda as well an ostial-caudal interspace of 4-8 % (vs.
always smaller than 5 %).
Most specimens show a slightly expanded predorsal
margin, while in others it is somewhat depressed (pl. 10,
fig. 4). This subtle difference however does not appear to
be underpinned by any further character and therefore is
considered to represent variability.
Distribution: Late Burdigalian to early Langhian of Colom-
bia and Venezuela.
Genus
†
Protonebris n. gen.
Type species:
†
Protonebris sanchezi n. sp.
Etymology: Referring to the primitive, plesiomorphic ap-
pearance of this putative early member of the Larimini.
Diagnosis: A fossil otolith-based genus of the family Sciae-
nidae with the following combination of characters. Com-
pressed otolith with a ratio OL : OH of 1.4, moderately thin,
without umbo on outer face. Dorsal rim regularly curved,
predorsally depressed; ventral rim moderately deeply and
regularly curved; anterior rim broadly rounded, posterior
rim with inferior, pointed tip. Ostium anteriorly narrowed
with deflection of dorsal margin and incipient notch at
mid-ventral margin; OCL : OCH = 1.3; CCL : OCL = 0.65.
Downturned part of cauda not widened; caudal curvature
index = 0.55; ostial-caudal interspace 17 %.
Discussion: Protonebris is interpreted as a plesiomorphic
genus within the Larimini close to the branch leading to
Paranebris and Nebris. It differs from the two genera in the
smaller ostium, the larger ostial-caudal interspace and the
not widened downturned part of the cauda and it shares
with them characters considered as early apomorphic such
as the shape of the anteriorly narrowed ostium with its notch
at the ventral margin and the depressed predorsal rim
Species: A single species, P. sanchezi, from the early Mio-
cene Castilllo FM of Venezuela.
†
Protonebris sanchezi n. sp.
(Plate 10, Fig. 1)
Holotype (and unique specimen): NMB P1078, sloth locality, Cerro
La Cruz near La Mesa, Venezuela, niveaux with Encina, Castillo
FM, Aquitanian to early Burdigalian.
Name: In honor of Marcelo Sánchez (Zürich) in recognition
of his support for our studies.
Diagnosis: See genus diagnosis (monospecific genus).
Description: Compressed otolith of 10.5 mm length.
OL : OH = 1.4; OH : OT = 1.8. Dorsal and ventral rims
regularly rounded; predorsal rim depressed, postdorsal
angle broadly rounded; anterior rim broadly rounded,
slightly projecting dorsally; posterior rim with pointed tip
located slightly below median line.
Inner face moderately convex with slightly supramedian
sulcus. Ostium moderately large, inclined, anteriorly nar-
rowed with reduced opening, its dorsal margin anteriorly
deflected and its ventral margin with shallow notch at
center. Cauda with short anterior and steeply curved, not
widened posterior part; tapering caudal tip slightly bent
forward. OCL : OCH = 1.3; CCL : OCL = 0.65; caudal
curvature = 0.55; ostial-caudal interspace 17 %. Dorsal
depression narrow, indistinct, only above anterior part of
cauda; no ventral furrow. Outer face moderately convex,
without distinct umbo, rather smooth.
Discussion: Protonebris sanchezi superficially resembles
Pareques jungi in outline and general character of the sulcus,
but clearly differs in the ostium being anteriorly narrowed
with a dorsal deflection and a ventral notch and the nar-
rower ostial-caudal interspace (17 % vs. 22-26 %), which
we interpret as indication for a relationship with Larimini
(see discussion to genus).
Distribution: Aquitanian to early Burdigalian of Venezuela.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
51
Palaeo Ichthyologica 14
Subfamily Cynoscioninae
TREWAVAS, 1977
Definition: In the limits as defined by SASAKI (1989), but
excluding the old world Kathalini.
Tribe Cynoscionini TREWAVAS, 1962
Definition and Discussion: In the limits as defined by
SASAKI (1989) including the genera Cynoscion, Isopisthus,
Macrodon and the freshwater genus Plagioscion. Recent
molecular-based studies suggest that Plagioscion may be
more closely related to another South American freshwater
sciaenid subfamily, the Pachyurinae, and hence question the
validity of the placement of the genus with the Cynoscioninae
(BOEGER et al., 2014). SCHWARZHANS (1993) also related
Plagioscion with the genera Pachyurus and Pachypops con-
stituting the Pachyurinae on the basis of their otolith analysis.
Nevertheless, we have here selected to provisionally keep
Plagioscion within the Cynoscionini, although SASAKI’s
character analysis (1989) indeed suggests Plagioscion may
be more distantly related to the remaining three marine
genera of the clade.
Genus Cynoscion GILL, 1861
Remarks: Cynoscion is one of the richest sciaenid genera
with 25 valid extant species in the waters around the Ameri-
cas (CHAO, 2002). Otoliths are known from all species (see
AGUIRRE, 2003; CHAO, 1978; CORREA & VIANNA, 1992;
LOMBARTE et al., 2006; ROSSI-WONGTSCHOWSKI et al.,
2014; SCHWARZHANS, 1993). It represents one of the few
indigenous American sciaenid genera, which also has been
recorded from the NE-Atlantic in the geological past, i. e.
the late Miocene of the North Sea Basin (SCHWARZHANS,
2010). Cynoscion otoliths show a rather strong ontogenetic
change and their thin otolith rims break or erode easily.
Because of these two effects only few specimens allow
identification to the species level and many juvenile or
eroded or fragmented otoliths have been omitted from the
following descriptions.
Cynoscion analis (JENYNS, 1842)
(Plate 11, Figs. 1-2)
Material: 3 specimens PPP 3514, 3515, 3529, figured speci-
men NMB P1079, Punta Canoa, Ecuador, Canoa FM, early
Pleistocene.
Description: Elongate, thin otoliths to at least 12.5 mm
length. OL : OH = 2.1-2.3; OH : OT about 3.0. Dorsal rim
shallow, anteriorly inclined and slightly depressed, with
obtuse, distinct middorsal angle above ostial-caudal joint
and postdorsal angle above bend of cauda; section between
middorsal and postdorsal angles broadly concave; ventral
rim shallow, anteriorly regularly curved, with concavity at
about
2
/
3
from anterior tip; anterior rim broadly rounded,
dorsally pronounced; posterior rim with inferior angular tip
slightly above caudal tip; postdorsal rim between postdorsal
angle and posterior tip inclined, straight.
Inner face slightly convex, with slightly supramedian,
moderately large sulcus. Ostium shallow rather small and
narrow, with weak postostial lobe. Cauda slightly deepened,
with long anterior section and short downward curved
posterior section and broadened tip. OCL : OCH = 1.7-
1.8; CCL : OCL = 1.05-1.3; caudal curvature index = 2.7-
3.4; ostial-caudal interspace 35-40 %. Dorsal depression
present above ostial-caudal joint and straight section of
cauda; ventral furrow running close to ventral rim, joining
it anteriorly below ostium and slightly upward trending
posteriorly extending below tip of cauda. Outer face flat to
slightly concave, smooth, without umbo. All rims very thin.
Discussion: Otoliths of C. analis are very characteristic
and resemble with their thin appearance, the low caudal
curvature and the high ostial-caudal interspace those of
Cynoscion othonopterus JORDAN & GILBERT, 1882 from
the Gulf of California and the related genus Isopisthus.
A Recent specimen figured for comparison (pl. 11, fig. 1)
shows excellent comparison with the fossil otolith from the
Canoa FM (pl. 11, fig. 2).
Distribution: Early Pleistocene of Ecuador and Recent in
the East Pacific from Ecuador to 30° S.
Cynoscion arenarius GINSBURG, 1930
(Plate 11, Figs. 3-5)
1976 Cynoscion sp. – NOLF: pl. 7, fig. 3
Material: 10 specimens PPP 2667, 2670, Manzanilla Bay and PPP
2675, San José River, figured specimens NMB P1080-1081,
Trinidad, Manzanilla FM, San José Mb, late Tortonian.
Discussion: None of the otoliths from the Manzanilla FM
is well preserved, but we consider the relative thickness, low
curvature of the inner face, the comparatively small ostium
and the broad mediodorsal expansion of the dorsal rim
sufficiently diagnostic for identification. A Recent otolith is
figured for comparison (pl. 11, fig. 3).
Distribution: Late Tortonian of Trinidad and Recent in the
Gulf of Mexico and off Florida.
Cynoscion latiostialis n. sp.
(Plate 11, Figs. 14-17)
Holotype: Pl. 11, Fig. 15, NMB P1082, At-14-19-1, San Judas,
Panama, Gatun FM, lower member, middle Tortonian.
Paratypes: 3 specimens; 2 specimens NMB P1083-1084, AT-14-
21-1, same data as holotype; 1 specimen NMB P1085, PPP
223, Sabanita, Colón, Panama, Gatun FM, lower member,
middle Tortonian.
Further material: 8 specimens; 7 specimens, At-14-19-1 and 14-
21-1, same data as holotype;1 specimen, El Hatillo, Venezuela,
Urumaco FM middle member, middle Tortonian.
Name: From latus (Latin = large) and the term ostium of
the sulcus of an otolith referring to the very large size of
the ostium.
Diagnosis: OL : OH = 1.9-2.1. Otolith anteriorly distinctly
wider than posteriorly. Dorsal rim with marked, obtuse
postdorsal angle. Posterior tip angular, slightly inferior, not
projecting. Ostium very large and wide, reaching close to
anterior, predorsal and preventral rims of otolith. Ostial-
caudal interspace 15-20 %, rarely 25 %. No ventral furrow.
Outer face convex, with broad postcentral umbo.
Description: Elongate, moderately thick otoliths up to
17 mm length. OH : OT = 1.5-1.8. Dorsal rim shallow, an-
terior two-thirds horizontal with broadly rounded predorsal
and obtuse postdorsal angles, thereafter straight inclined
52
towards inferior, angular, not projecting posterior tip; ventral
rim most deeply curved anteriorly below ostium, with slight
concavity just before caudal tip and thereafter moderately
curved; anterior rim broadly rounded, dorsally pronounced.
Inner face markedly convex, with very large sulcus.
Ostium shallow, very large and wide, making up for about
half of area of inner face in large specimens, with moderate
postostial lobe. Cauda slightly deepened, with about equally
long anterior and downturned posterior sections, caudal tip
broadened. OCL : OCH = 1.25-1.3; CCL : OCL = 0.6-0.8,
decreasing with size; caudal curvature index = 0.9-1.2,
decreasing with size; ostial-caudal interspace 15-25 %,
decreasing with size. Dorsal depression very small, present
only above ostial-caudal joint; no ventral furrow. Outer face
convex, with broad postcentral umbo, smooth. Anterior rim
thin.
Ontogeny: Otoliths of C. latiostialis show the following
ontogenetic changes from specimens of 9 mm length
(fig. 20) to 17 mm length (fig. 18): decreasing OL : OH
from 2.1 to 1.9, decreasing CCL : OCL from 0.8 to 0.6,
decreasing caudal curvature index from 1.2 to 0.9 and
strongly decreasing ostial-caudal interspace from 25 % to
15 %. In connection with the latter, the size of the ostium
also increases with the size of the otolith.
Discussion: Otoliths of C. latiostialis are characterized by
the large ostium, the low ostial-caudal interspace and the
thick outer face when compared with Recent otoliths of
Cynoscion. The most resembling otoliths are from the extant
Cynoscion acoupa (LACEPÈDE, 1802) from the Atlantic
shores of South America and its geminate Pacific counterpart
Cynoscion albus (GÜNTHER, 1864), both of which differ in
the thinner appearance, the larger ostial-caudal interspace
(27-30 % vs. 15-20 %, rarely up to 25 %) and the smaller
ostium. Other similar Recent otoliths are Cynoscion stein-
dachneri (JORDAN, 1889), which has a still smaller ostium
and extensive ornamentation, and Cynoscion microlepidotus
(CUVIER, 1830), which is thinner and shows a much nar-
rowed posterior part of the otolith (see SCHWARZHANS,
1993 for figures of Recent otoliths). The low ostial-caudal
interspace of C. latiostialis is probably the most obvious
distinctive character, but only so in specimens exceeding
12 mm length.
Distribution: Middle Tortonian of Panama and Venezuela.
Cynoscion nothus (HOLBROOK, 1848)
(Plate 11, Figs. 6-9)
Material: 5 specimens; 1 specimen location 17468, NMB P1086,
Manzanilla FM, San José Mb, late Tortonian; 4 specimens PPP
1727, 1728, 1734, NMB P1087-1090, Río Banano, Costa
Rica, Río Banano FM, late Pliocene.
Description: Moderately compressed and moderately
thin otoliths to at least 12 mm length. OL : OH = 1.9-2.0
in specimens of 9 to 12 mm length, and down to 1.6 in
small specimens of 5 mm length (fig. 9); OH : OT = 1.9-
2.5, decreasing with size. Dorsal rim with broad, obtuse
predorsal angle located at above termination of dorsal
margin of ostium, decreasing in strength with size, and
lower, obtuse postdorsal angle; ventral rim deeply curved,
deepest below postostial lobe; anterior rim broadly rounded,
distinctly dorsally pronounced; posterior rim with slightly
projecting inferior tip positioned at mid-level of downturned
part of cauda.
Inner face moderately convex, with slightly supramedian,
moderately large sulcus. Ostium shallow, rather short, with
weak postostial lobe. Cauda slightly deepened, with long
anterior section and short downward curved posterior section
and rounded tip. OCL : OCH = 1.0-1.1; CCL : OCL = 1.05-
1.15; caudal curvature index = 1.2-1.6; ostial-caudal
interspace 30-35 %. Dorsal depression present above
ostial-caudal joint and straight section of cauda; no ventral
furrow. Outer face moderately convex, smooth, with broad,
indistinct postcentral umbo. Anterior rim thin.
Discussion: Otoliths of C. nothus are readily recognized by
their rather compressed shape with the low OL : OH ratio
in combination with the short and wide ostium.
Distribution: In the late Tortonian of Trinidad, late Plio-
cene of Costa Rica and Recent in the western Atlantic from
Maryland to Florida and throughout the Gulf of Mexico,
but not in the Caribbean.
Cynoscion prolixus n. sp.
(Plate 11, Figs. 11-13)
Holotype: Pl. 11, Fig. 11, NMB P1091, PPP 2573, Margarita
airport, Margarita Island, Venezuela, Cubagua FM, La Tejita
Mb, late Tortonian.
Paratypes: 4 specimens; 1 specimen NMB P1092, PPP 1652,
Atlantic Panama, Chagres FM, Toro Mb, late Tortonian; 1
specimen NMB P1093, PPP 2572, Bahia de Charagato, Cu-
bagua Island, Venezuela, Cubagua FM, above Cerro Negro
Mb, late Pliocene; 1 specimen NMB P1094, PPP 2567, Ca-
ñon de las Calderas, Cubagua Island, Venezuela, Cubagua
FM, above Cerro Negro Mb, late Pliocene; 1 specimen NMB
P1095, El Hatillo, Venezuela, Urumaco FM middle member,
middle Tortonian.
Further material: 17 mostly eroded or fragmented specimens; 11
specimens, El Hatillo, Venezuela, Urumaco FM middle mem-
ber, middle Tortonian; 5 specimens, PPP 2567, Cañon de las
Calderas, PPP 2572, Bahia de Charagato, Cubagua Island,
Venezuela, Cubagua FM, above Cerro Negro Mb, late Pliocene.
Name: From prolixus (Latin = lengthy, stretching) referring
to the elongate shape of the otoliths.
Diagnosis: OL : OH = 2.15-2.3. Otolith anteriorly distinctly
wider than posteriorly. Dorsal rim anteriorly shallow, often
concave at mid-section and with broad, somewhat elevated,
obtuse postdorsal angle. Anterior tip distinctly dorsally pro-
/
Plate 11
Figs. 1-2. Cynoscion analis (JENYNS, 1842). 1, coll. WS, off Peru,
Recent; 2, Ecuador, Canoa FM, early Pleistocene.
Figs. 3-5. Cynoscion arenarius GINSBURG, 1930. 3, coll. WS,
Gulf of Mexico, Recent; 4-5, Trinidad, Manzanilla FM, late
Miocene.
Figs. 6-9. Cynoscion nothus (HOLBROOK, 1848). 6, 8-9, Costa
Rica, Rio Banano FM, late Pliocene; 7, Trinidad, Manzanilla FM,
late Miocene.
Fig. 10. Cynoscion jamaicensis (VAILLANT & BOCOURT, 1883);
coll. OA, off Venezuela, Recent.
Figs. 11-13. Cynoscion prolixus n. sp. 11 (r), holotype, NMB
P1091, Venezuela, Margarita Island, Cubagua FM, late Mio-
cene; 12, paratype, NMB P1093, Cubagua Island, Cubagua
FM, late Pliocene; 13, paratype, NMB P1092, Panama, Chagres
FM, late Miocene.
Figs. 14-17. Cynoscion latiostialis n. sp. 15, holotype, NMB
P1082; 14 (r), 16 (r), 17, paratypes, NMB P1083-1085; Panama,
Gatun FM, late Miocene.
Figs. 18-19. Cynoscion scitulus n. sp. 18 (r), holotype, NMB
P1096; 19 (r), paratype, NMB P 1097; Panama, Gatun FM, late
Miocene.
Fig. 20. Cynoscion sp.; Panama, Escudo de Veraguas FM, late
Pliocene.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
53
Palaeo Ichthyologica 14
Cynoscion arenarius
Cynoscion nothusCynoscion analis
Cynoscion jamaicensis
Cynoscion prolixus
Cynoscion latiostialis
Cynoscion scitulus
Cynoscion sp.
18a
18b
18c
18d
19a
19b
20
19c
1a
1b
2a
2b
3a
3b
3c
4
5a
5b
6a
6b
7a
7b
8
9
10a
10b
11a
11b
11c
11d
12
13a
13b
15a
15b
15c
15d
16
17a
17b
14a
14b
17c
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
54
nounced; posterior tip slightly inferior, projecting. Ostium
rather short, its ventral margin distinctly turned upwards.
Ostial-caudal interspace 30 %. No ventral furrow.
Description: Elongate, thin otoliths up to at least 14 mm
length. OH : OT = 2.3-2.5. Dorsal rim shallow, anteriorly
almost flat with weak, broad predorsal angle becoming
minute in large specimens, slightly concave middorsal sec-
tion, broad, somewhat elevated, rounded or obtuse post-
dorsal angle; ventral rim shallow, deepest below postostial
lobe, with small concavities in front of and behind caudal
tip resulting in caudal tip appearing like extending on a
projection; anterior rim ventrally backward inclined leading
to pronounced dorsally positioned tip; posterior tip inferior,
positioned at about level of mid section of downturned part
of cauda, somewhat projecting, postdorsal rim between
postdorsal angle and posterior tip usually slightly concave.
Inner face mildly convex, with moderately large sulcus.
Ostium shallow, short and wide, its upper margin horizontal
and its ventral margin steeply inclined, with moderate posto-
stial lobe. Cauda slightly deepened, with long anterior and
short downturned posterior sections, caudal tip widened,
rounded. OCL : OCH = 1.1-1.2; CCL : OCL = 1.05-1.15;
caudal curvature index = 1.5-1.8; ostial-caudal interspace
30 %. Dorsal depression small, present above ostial-caudal
joint and anterior part of cauda; no ventral furrow. Outer
face flat to slightly concave, without umbo, smooth. All rims
thin.
Ontogeny: Most otoliths of C. prolixus have an incomplete
anterior tip due to its thin nature that easily resulted in ero-
sion or fragmentation. The smallest otoliths available thus
lack the anterior portion, but nevertheless it appears clear
that they would be more compressed than the large speci-
mens of 9 and more mm length, which is in line with the
usual observations about ontogenetic changes in Cynoscion
otoliths (AGUIRRE, 2003).
Discussion: Otoliths of C. prolixus resemble those of Cy-
noscion jamaicensis (VAILLANT & BOCOURT, 1883), known
in the western Atlantic from Jamaica and Puerto Rico to
southern Brazil, in most aspects, but differ in the more
elongate shape (OL : OH = 2.15-2.3 vs. 1.95-2.05), the
posteriorly reduced height and a marked inferior projection
of the posterior tip. We have figured a Recent specimen of
C. jamaicensis for comparison (pl. 11, fig. 10) and refer to
ROSSI-WONGTSCHOWSKI et al., 2014 for further figures.
Cynoscion jamaicensis does not appear to have an obvious
allopatric counterpart on the Pacific side, although otoliths
of Cynoscion phoxocephalus JORDAN & GILBERT, 1882,
Cynoscion praedatorius (JORDAN & GILBERT, 1889) and
Cynoscion squamipinnis (GÜNTHER, 1867) (see SCHWARZ-
HANS, 1993 for figures), show some similarity, but differ
in the flat, nearly straight dorsal rim and in the case of the
latter two also in being thicker and showing a wider ostium.
Distribution: Tortonian to late Pliocene of Venezuela.
Cynoscion scitulus n. sp.
(Plate 11, Figs. 18-19)
Holotype: Pl. 11, Fig. 18, NMB P1096, At-14-19-1, San Judas,
Panama, Gatun FM, lower member, middle Tortonian.
Paratypes: 1 specimen, NMB P1097, same data as holotype.
Further material: 4 fragmented specimens, same data as holotype
Name: From scitulus (Latin = elegant) referring to the deli-
cate shape of the otoliths.
Diagnosis: OL : OH = 2.45. Otolith anteriorly slightly wider
than posteriorly, with anterior tip pronounced dorsally.
Dorsal rim with very broad and gently rounded postdorsal
angle. Posterior tip angular, distinctly inferior, slightly pro-
jecting. Ostium large and wide, reaching close to anterior,
predorsal and preventral rims of otolith. Ostial-caudal
interspace 25 %. No ventral furrow. Outer face nearly flat,
with shallow, ventrally shifted tuberculate ridge.
Description: Very elongate, thin otoliths up to 15 mm
length. OH : OT = 2.0-2.5. Dorsal rim shallow, anterior third
slightly inclined up to very broad, rounded predorsal lobe
situated above rear part of ostium; middorsal section very
slightly inclined and slightly concave until very broad and
gently rounded postdorsal angle, thereafter inclined towards
distinctly inferior, angular, slightly projecting posterior tip.
Ventral rim shallow, deepest below postostial lobe, with
slight concavity just before caudal tip and thereafter almost
straight; anterior rim gently ascending towards distinctly
dorsally pronounced tip.
Inner face slightly convex, with moderately large sulcus.
Ostium shallow, large, moderately wide, with moderate pos-
tostial lobe. Cauda slightly deepened, with distinctly longer
anterior and shorter downturned posterior sections, caudal
tip slightly broadened. OCL : OCH = 1.7; CCL : OCL = 0.8;
caudal curvature index = 1.7; ostial-caudal interspace 25 %.
Dorsal depression indistinct; no ventral furrow. Outer face
flat to slightly convex posteriorly, with ventrally shifted tuber-
culate ridge on rear part, anteriorly smooth. All rims thin.
Discussion: Otoliths of C. scitulus are readily distinguished
from other Miocene congeners by the thin, slender appear-
ance, its distinctive outline and the ventrally shifted ridge
on the rear part of the outer face. The slender shape, the
ventrally projecting posterior tip and the thin appearance
resembles otoliths of the Recent Atlantic species Cynoscion
nebulosus (CUVIER & VALENCIENNES, 1830), Cnoscion
regalis (BLOCH & SCHNEIDER, 1801) and Cynoscion thalas-
sinus (HOLBROOK, 1855) (see SCHWARZHANS, 1993 for
figures), but they show a smaller ostium and a lesser caudal
curvature index of 2.1-3.0 (vs. 1.7). Cynoscion reticulatus
(GÜNTHER, 1864), Cynoscion parvipinnis (AYRES, 1862)
and Cynoscion xanthulus JORDAN & GILBERT, 1881 (see
SCHWARZHANS, 1993 for figures) from the Pacific shores of
Tropical America also resemble in general appearance, but
all show a more strongly projecting posterior tip. Cynoscion
reticulatus is further distinguished by its smaller ostium,
C. parvipinnis by the lesser caudal curvature index (2.0 vs.
1.7) and the very slender shape (OL : OH = 3.0 vs. 2.45)
and C. xanthulus by the higher caudal curvature index (1.4
vs. 1.7). In conclusion, C. scitulus represents a third major
lineage of the genus Cynoscion parallel to C. latiostialis
and C. prolixus.
Distribution: Middle Tortonian of Panama.
Cynoscion sp.
(Plate 11, Fig. 20)
Material: 1 specimen NMB P1098, PPP 2185, Escudo de Vera-
guas, Atlantic Panama, Escudo de Veraguas FM, late Pliocene.
Discussion: A single, large, well preserved specimen
lacking the posterior part. It is characterized by a shallow
preventral rim, a rather narrow ostium and a regularly
rounded anterior tip.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
55
Palaeo Ichthyologica 14
Genus Isopisthus GILL, 1862
Isopisthus acer n. sp.
(Plate 12, Figs. 4-6)
Holotype: Pl. 12, Fig. 4, NMB P1099, PPP 1727, Río Banano,
Costa Rica, Río Banano FM, late Pliocene.
Paratypes: 3 specimens NMB P1100-1102, same data as holotype.
Name: From acer (Latin = sharp, acute), referring to the
sharp, projecting postdorsal angle.
Diagnosis: OL : OH = 1.35-1.45 in otoliths to 6 mm length.
Dorsal rim flat, with distinct, projecting postdorsal angle.
Ventral rim deepest in front of middle. Anterior tip broadly
rounded; posterior tip short, cut. Ostium short, wide.
Description: Compressed, moderately thin otoliths up to
at least 6 mm length [specimens of Recent species are
known up to 10.5 mm length; see I. remifer pl. 12, fig. 2].
OH : OT about 3.0. Dorsal rim nearly flat, with massive,
slightly backward projecting postdorsal angle; ventral rim
deep, deepest well in front of middle, with broad, shallow
concavity across most of postventral rim; anterior rim broadly
rounded, blunt; posterior tip median, blunt, postdorsal rim
between postdorsal angle and posterior tip very steep.
Inner face mildly convex, with moderately large, slightly
supramedian sulcus. Ostium shallow, short and wide,
no postostial lobe. Cauda slightly deepened, with long
anterior and short downturned posterior sections, caudal
tip rounded. OCL : OCH = 0.95-1.05; CCL : OCL = 1.15-
1.25; caudal curvature index = 2.15-2.25; ostial-caudal
interspace 35-40 %. Dorsal depression moderately wide,
present above ostial-caudal joint and anterior part of cauda;
no ventral furrow. Outer face flat to slightly convex, without
or with very shallow umbo, smooth. All rims thin.
Ontogeny: Otoliths of Isopisthus show a strong allomet-
ric ontogenetic growth. ROSSI-WONGTSCHOWSKI et
al. (2014) figured two specimens of Isopisthus parvipin-
nis (CUVIER, 1830), one at about 8.5 mm long with an
OL : OH of 1.8, the other about 4.5 mm long with an
OL : OH of 1.45. A specimen of I. parvipinnis of 9 mm
length figured here (pl. 12, fig. 1) has an OL : OH of 1.8,
and two specimens of about 5 and 8 mm length figured
in SCHWARZHANS (1993) show a ratio OL : OH of about
1.5 and 1.7 respectively. This indicates that the change of
the otolith proportions from compressed to more slender
takes place at a rather late ontogenetic stage between 5
and 8 mm length. The specimens of I. acer hence probably
represent the compressed ontogenetic stage.
Discussion: Otoliths of I. acer readily differ from the
two geminate Recent species I. parvipinnis from the At-
lantic (pl. 12, fig. 1) and I. remifer JORDAN & GILBERT,
1882 from the Pacific (pl. 12, fig. 2) in the shorter ostium
(OCL : OCH = 0.95-1.05 vs. 1.2- 1.45), the distinctly
anteriorly positioned deepest point at the ventral rim (vs.
mid-ventral) and the strongly developed and projecting
postdorsal angle, which are all evident from comparison
of similar sized specimens.
Distribution: Late Pliocene of Costa Rica, indicating the
presence of a extinct lineage of the genus in the Pliocene
of the Caribbean, separate from the lineage containing the
two extant geminate species.
Isopisthus remifer JORDAN & GILBERT, 1881
(Plate 12, Figs. 2-3)
Material: 2 specimens PPP 3560, 3562, figured specimen NMB
P1103 Punta La Cereza, Jama, Ecuador, Jama FM, middle
Pleistocene.
Discussion: Otoliths of I. remifer differ from their allopatric
counterpart in the West Atlantic, I. parvipinnis, primarily
in the more elongate shape (OL : OH = 1.9 vs. 1.7-1.8),
although smaller specimens below 6 mm length may not
be possible to be distinguished. A Recent otolith is figured
for comparison (pl. 12, fig. 2).
Distribution: Middle Pleistocene of Ecuador and Recent in
the East Pacific from Mexico to Peru.
Genus Macrodon SCHINTZ, 1822
Macrodon spina SCHWARZHANS, 1993
1993 Macrodon spina – SCHWARZHANS: fig. 84
Remarks: This species is based on a single, anteriorly
eroded otolith of about 6 mm length from the Morne l’Enfer
FM (late Miocene to early Pliocene) of Trinidad and char-
acterized by the unique development of a spinous, sharply
pointed posterior tip. Otherwise Macrodon otoliths are
easily recognized by their oblong shape, the rather nar-
row, downward inclined spatulate ostium and the drop-like
widened downturned part of the cauda. For Recent otoliths
see SCHWARZHANS (1993), CARDOSO et al. (2012) and
ROSSI-WONGTSCHOWSKI et al. (2014).
Genus Plagioscion GILL, 1861
Remarks: Following a recent revision by CASATTI (2005),
the genus Plagioscion comprises five Recent species in the
freshwater systems of South America from Colombia to
Uruguay. Their otoliths are characterized by a spatulate,
trapezoidal ostium narrowing towards the caudal joint,
showing a weak and ventrally reduced postostial lobe and
a caudal tip being curved forward for a considerable length.
NOLF & AGUILERA (1998) and AGUILERA & RODRIGUES
DE AGUILERA (2004) were the first to prove the presence
of Plagioscion otoliths in the marine strata of the Miocene
of Venezuela thereby showing that the genus may have
had a marine origin.
Plagioscion marinus
AGUILERA & RODRIGUES DE AGUILERA, 2003
(Plate 12, Figs. 7-10)
1998 Plagioscion sp. – NOLF & AGUILERA: pl. 12, figs. 1-4
2003 Plagioscion marinus – AGUILERA & RODRIGUES DE
AGUILERA: figs. 4.1-4.4
2014 Plagioscion marinus AGUILERA & RODRIGUES DE AGU-
ILERA, 2003 – AGUILERA, SCHWARZHANS, MORAES-
SANTOS & NEPOMUCENO: figs. 10.9-10.11
Material: 81 specimens; 50 specimens Atalaia beach, Brazil,
Aquitanian to early Burdigalian, MPEG-1801-1802-V; 22
specimens locations 16344, 16555, 16677, 290602, 290616,
290632, 290666, 290685, 390085, 390090, 390094,
430202 Castilletes, Colombia, Castilletes FM, late Burdigalian
to early Langhian; 2 specimens Castilletes, Colombia, Jimol
FM, Burdigalian, location 390090; 7 specimens, San José de
Cocodite, Paraguaná Península, Venezuela, Cantaure FM, late
Burdigalian to Langhian.
56
Description: Compressed, moderately thick otoliths to
about 12.5 mm length. OL : OH = 1.4-1.5; OH : OT about
1.8. Dorsal rim shallow, with postdorsal angle positioned
behind termination of cauda; ventral rim moderately deep,
middle section less bent than anterior and posterior sections,
posterior most section below posterior tip concave; anterior
rim very broad, cut, inclined backward-downwards; poste-
rior tip slightly inframedian, angular, postdorsal rim between
postdorsal angle and posterior tip straight, steeply inclined.
Inner face mildly convex, with moderately large, slightly
supramedian sulcus. Ostium shallow, short and wide, anteri-
orly widened, moderately expanding postostial lobe. Cauda
slightly deepened, with short anterior and long downturned
posterior sections, with caudal tip turning forward for a
long section until about middle of horizontal part of cauda,
tapering. OCL : OCH = 0.95-1.05; CCL : OCL = 0.9-0.95;
caudal curvature index = 0.65-0.7; ostial-caudal interspace
25-30 %. Dorsal depression very narrow, present above
ostial-caudal joint and anterior part of cauda; no ventral
furrow. Outer face distinctly convex, with diffuse, broad
postcentral umbo, smooth. Rims thick except for anterior
part of otolith.
Discussion: Palgioscion marinus differs from the otoliths of
the Recent species (otoliths known from four of five Recent
species; see SCHWARZHANS, 1993) in the wide ostium,
which is not much ventrally reduced towards the caudal
joint. It resembles Plagioscion auratus (CASTELNAU, 1855)
most in the shallow postventral rim, but differs in the expres-
sion of the posterior tip (see AGUILERA & RODRIGUES DE
AGUILERA, 2004, for figures).
Distribution: Widely distributed and rather common in the
early Miocene, Aquitanian to early Langhian of Colombia,
Venezuela and Brazil.
Plagioscion travassosi
AGUILERA & SCHWARZHANS, 2014
(Plate 12, Fig. 11)
2014 Plagioscion travassosi – AGUILERA & SCHWARZHANS:
figs. 10.12-10.13
Material: 3 specimens (holotype and 2 paratypes), Atalaia beach,
Brazil, early Miocene, MPEG-1803-1804-V.
Discussion: Plagioscion travassosi is known from few large
specimens, which are readily recognized by their elongate
shape (OL : OH = 1.7-1.8), the wide and large ostium and
the upward oriented shape of the anterior ventral rim. The
holotype of about 24 mm length is here refigured.
Distribution: Aquitanian to early Burdigalian of Brazil.
Plagioscion urumacoensis
AGUILERA & RODRIGUES DE AGUILERA, 2003
(Plate 13, Fig. 2)
2003 Plagioscion urumacoensis – AGUILERA & RODRIGUES
DE AGUILERA: figs. 4.5-4.8
Material: 2 specimens (holotype and 1 paratype), El Hatillo, Uru-
maco, Venezuela, Urumaco FM, middle Mb, middle Tortonian,
UNEFM-PF-049-050.
Discussion: Plagioscion urumacoensis is known from two
well preserved and very large specimens of about 25 mm
length, a size otherwise only reached by the giant Recent
Plagioscion squamosissimus (HECKEL, 1840). They differ
in the lack of an umbo on the outer face (vs. massive
postcentral umbo), the postventral rim being deeper than
in P. squamosissimus and a pronounced, somewhat pointed
posterior tip. The holotype is here refigured together with
a large specimen of P. squamosissimus (pl. 13, fig. 1) for
comparison reasons.
Distribution: Middle Tortonian of Venezuela.
Plagioscion ultimus n. sp.
(Plate 12, Figs. 12-16)
1976 Plagioscion sp. – NOLF: pl. 8, fig. 7
2004 Plagioscion sp. – AGUILERA & RODRIGUES DE AGUIL-
ERA: figs. 4.9-4.10
Holotype: Pl. 12, Fig. 12, NMB P1043, PPP 3093, Cañon de las
Calderas, Cubagua Island, Venezuela, Cubagua FM, above
Cerro Negro Mb, late Pliocene.
Paratypes: 9 specimens; 1 specimen NMB P1201, same data as
holotype; 1 specimen NMB P661, K 9894, Savaneta River,
Trinidad, Springvale FM, early to late Pliocene; 6 specimens
NMB P662, K 9845, Savaneta River, Trinidad, Manzanilla
FM, Telémaque Mb, early Pliocene; 1 specimen NMB P531, K
12059, Mogue River, Trinidad, Gros Morne FM, early Pliocene.
Name: From ultimus (Latin = last), referring to the species
representing the last / youngest record of Plagioscion in a
marine environment.
Diagnosis: OL : OH = 1.5-1.7. Anterior tip dorsally pro-
nounced; posterior tip narrowed, slightly projecting, angular.
Ostium short, triangular shaped, without postostial lobe.
Cauda bent downward and forward at sharp flexure, its tip
reaching almost the level of ostial-caudal joint.
Description: Moderately compressed, thick otoliths of
relatively small size not reaching beyond 8-9 mm length.
OH : OT = 1.3-1.6, much less in specimens smaller than
5 mm (2.0-2.7). Dorsal rim shallow, rather regularly curved
except for broad, obtuse postdorsal angle behind curvature
point of cauda; ventral rim moderately deep, deepest at
tip of cauda, with obtuse angle at deepest point of ostium;
rear part of ventral rim almost straight ascending to angular
posterior tip; anterior rim obliquely cut, dorsally pronounced;
posterior tip distinctly supramedian, angular, narrowed,
slightly projecting.
Inner face only slightly convex, with moderately large,
slightly supramedian sulcus. Ostium shallow, short, trian-
gular in shape with its dorsal margin straight, horizontal
and its ventral margin straight, downward inclined leading
/
Plate 12
Fig. 1. Isopisthus parvipinnis (CUVIER, 1830); coll. OA, Lake
Maracaibo, Venezuela, Recent.
Figs. 2-3. Isopisthus remifer JORDAN & GILBERT, 1881. 2 (r),
coll. Aguirre, off Ecuador, Recent; 3, Ecuador, Jama FM, middle
Pleistocene.
Figs. 4-6. Isopisthus acer n. sp. 4, holotype, NMB P1099; 5-6,
paratypes, NMB P1100-1101; Costa Rica, Rio Banano FM, late
Pliocene.
Figs. 7-10. Plagioscion marinus AGUILERA & RODRIGUES DE
AGUILERA, 2003. 7 (r), Venezuela, San José de Cocodite, Cantaure
FM, early Miocene; 8-9 (r), 10, Colombia, La Guajira, Castilletes
FM, early Miocene.
Fig. 11. Plagioscion travasossi AGUILERA & SCHWARZHANS,
2014; holotype, MPEG-1803-V, Brazil, Atalaia beach, Pirabas
FM, early Miocene.
Figs. 12- 16. Plagioscion ultimus n. sp. 12, holotype, NMB
P1043, Venezuela, Cubagua Island, Cubagua FM, late Pliocene;
13, 15-16, paratypes, NMB P662, Trinidad, Manzanilla FM, early
Pliocene; 14, paratype, NMB P531, Trinidad, Gros Morne FM,
early Pliocene.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
57
Palaeo Ichthyologica 14
Isopisthus parvipinnis
Isopisthus remifer
Isopisthus acer
Plagioscion marinus
Plagioscion travasossi Plagioscion ultimus
7a
7b
7c
7d
8
9a
9b
9c
10
11a
12a
12b
12c
13a
13b
13c
15
16a
16b
11b
14a
14b
1 mm
2 mm
2 mm
1a
1b
2a
2b
3a
3b
3c
4a
4b
4c
5
6a
6b
5 mm
2 mm
2 mm
58
to wide opening towards anterior-ventral rim of otolith;
no postostial lobe. Cauda slightly deepened, with short
anterior and long downturned posterior sections; caudal
flexure very sharp, v-shaped; caudal tip turning forward
for a long section almost reaching to level of ostial-caudal
joint, tapering. OCL : OCH = 1.0-1.2; CCL : OCL = 0.8-1.0;
caudal curvature index = 0.75-0.9; ostial-caudal interspace
27-35 %. Dorsal depression indistinct; no ventral furrow.
Outer face distinctly convex, with broad ventrally shifted
umbo, smooth; slightly convex only in small specimens.
Ventral and posterior rims thick, anterior and dorsal rims
thin.
Discussion: When NOLF (1976) and AGUILERA & ROD-
RIGUES DE AGUILERA (2004) first recorded these otoliths
they left them in open nomenclature because of them being
somewhat eroded and considered to represent juveniles in
the light of Recent and other known fossil otoliths of the
genus reaching much larger sizes. In fact the combination
of both records shows the presence of a continuous ontoge-
netic succession with the largest specimens at 8 mm length
exhibiting all the attributes of otoliths of adult specimens of
the genus such as the massive ventrally shifted umbo on
the outer face. We therefore now conclude that P. ultimus
represents a small, “dwarfed” species and its distinctive
diagnostic characters warrant definition of a new species
despite all specimens showing some degree of erosion.
Otoliths of P. ultimus are readily distinguished from all
known fossil and Recent Plagioscion species by the sharp,
v-shaped caudal flexure, the much forward extended caudal
tip almost to the level of the ostial-caudal joint and the tri-
angular shape of the ostium without exhibiting a postostial
lobe.
Distribution: Plagioscion ultimus represents the last ma-
rine species known of Plagioscion, which in the Recent
is exclusively known from South American river schemes.
Plagioscion ultimus has been recorded from the early to late
Pliocene of Venezuela and Trinidad. In Trinidad juveniles
are commonly found associated with the sediments of the
Telémaque Mb that deposited in the delta estuary of a
paleo-Orinoco branch.
Tribe Atractoscionini SASAKI, 1989
Definition and Discussion: SASAKI (1989) established the
tribe Atractoscionini solely to accommodate for the genus
Atractoscion, while it was included into Cynoscionini by
TREWAVAS (1977) and SCHWARZHANS (1993). Atracto-
scion is one of the few genera with a broad geographic
distribution with one Recent species – Atractoscion aequidens
(CUVIER, 1830) – known from the East Atlantic and the
western Indian Ocean to Australia and a second – Atracto-
scion nobilis (AYRES, 1862) – along the northwestern shores
of America from Alaska to the Gulf of California. There is
also fossil otolith-based evidence from Europe – Atractoscion
elongatissimus SCHWARZHANS, 1993 – from the Central
Paratethys and the North Sea Basin (SCHWARZHANS,
2010).
Genus Atractoscion GILL, 1862
Atractoscion odeai n. sp.
(Plate 13, Figs. 3-6)
Holotype: Pl. 13, Fig. 3, NMB P1104, San José de Cocodite, Para-
guaná Peninsula, Venezuela, Cantaure FM, late Burdigalian
to early Langhian.
Paratypes: 3 specimens; 2 specimens NMB P1105-1106, same
data as holotype; 1 specimen MUN2023, location 290632,
Castilletes, Colombia, Castilletes FM, late Burdigalian to early
Langhian.
Further material: 7 specimens; 1 specimen San José de Cocodite,
Paraguaná Peninsula, Venezuela, Cantaure FM, late Burdiga-
lian to early Langhian; 6 specimens locations 290610, 290632,
290840, GAB035, Castilletes, Colombia, Castilletes FM, late
Burdigalian to early Langhian.
Name: In honor of Aaron O’Dea (STRI, Panama) in recogni-
tion of his many contributions to the knowledge of the fossil
record from Panama and the Caribbean and his support
with otoliths from Gatun.
Diagnosis: OL : OH = 1.7-1.8. OH : OT = 2.2-2.5. Dorsal
rim with distinct, obtuse postdorsal angle. Anterior tip dis-
tinctly dorsally pronounced; posterior tip blunt, with obtuse
median angle, not extending much behind cauda. Ostium
long, wide, its dorsal margin horizontal and its ventral
margin slightly turned upwards. Ostial-caudal interspace
23-26 %. No ventral furrow. No umbo on outer face.
Description: Elongate, thin otoliths up to about 12.5 mm
length. Dorsal rim shallow, anteriorly almost flat with weak,
broad predorsal angle positioned above rear end of upper
margin of ostium, flat middorsal section, distinct, obtuse
postdorsal angle; ventral rim moderately deep, deepest
below postostial lobe, very regularly curved; anterior rim
rounded, dorsally pronounced, ventrally inclined; posterior
tip broad, with obtuse postdorsal and postventral angles
and obtuse median angle.
Inner face moderately convex, with moderately large
sulcus. Ostium shallow, long and wide, its upper margin
horizontal and its ventral margin slightly inclined, with dis-
tinct postostial lobe. Cauda slightly deepened, with about
equally long anterior and downturned posterior sections,
caudal tip tapering, slightly bent forward. OCL : OCH = 1.2-
1.3; CCL : OCL = 0.75-0.9; caudal curvature index = 1.0-
1.1; ostial-caudal interspace 23-26 %. Dorsal depression
narrow, present above ostial-caudal joint and anterior part
/
Plate 13
Fig. 1. Plagioscion squamosissimus (HECKEL, 1840); coll. WS,
Amazon River, Recent.
Fig. 2. Plagioscion urumacoensis AGUILERA & RODRIGUES DE
AGUILERA, 2003; holotype, UNEFM-PF-049, Venezuela, El Hatillo,
Urumaco FM, late Miocene.
Figs. 3-6. Atractoscion odeai n. sp. 3 (r), holotype, NMB P1104,
Venezuela, San José de Cocodite, Cantaure FM, early Miocene;
4-5 (r), paratypes, NMB P1105-1106, Venezuela, San José de
Cocodite, Cantaure FM, early Miocene; 6 (r), paratype, MUN2023,
Colombia, La Guajira, Castilletes FM, early Miocene.
Figs. 7-8. Micropogonias coatesi AGUILERA & RODRIGUES DE
AGUILERA, 2004. 7, holotype, UNEFM-PF-053, Venezuela, El
Hatillo, Urumaco FM, late Miocene; 8 (r), Venezuela, Carrizal,
Caujarao FM, late Miocene.
Fig. 9. Micropogonias altipinnis (GÜNTHER, 1864); Ecuador,
Canoa FM, early Pleistocene.
Fig. 10. Micropogonias furnieri (DESMAREST, 1823); (r) Venezuela,
Cumana, Cumana FM, middle Pleistocene.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
59
Palaeo Ichthyologica 14
5 mm
5 mm
5 mm
Plagioscion squamisissimus
Atractoscion odeai
Micropogonias coatesi
Micropogonias
altipinnis
Micropogonias furnieri
2 mm
2 mm
2 mm
Plagioscion urumacoensis
1a
1b
1c
1d
2a
2b
2c
2d
3a
3b
3c
3d
4
5a
5b
6
7a
7b
7c
7d
8a
8b
9a
9b
9c
10a
10b
10c
10d
60
of cauda; no ventral furrow. Outer face almost flat, without
umbo, little ornamentation. All rims thin.
Discussion: Otoliths of A. odeai are readily distinguished
from the two Recent species and A. elongatissimus from
the Miocene of Europe by the complete absence of a
postcentral umbo on the outer face resulting in a much
thinner appearance (OH : OT = 2.2-2.5 vs. 1.1-1.3). Other
differences are the low index OL : OH of 1.8-1.8 (vs. 2.0-
2.8), the shape of the ostium and the dorsally pronounced
anterior rim. Otoliths of A. odeai can be confused with
smaller otoliths of the contemporaneous S. reyesi of less
than 10 mm length, but differ in the slightly more elongate
shape (OL : OH = 1.7-1.8 vs. 1.6-1.7), the longer distance
of the downcurved cauda from the posterior rim of the
otolith, the shape of the posterior rim with its three obtuse
angles (vs. vertically cut) and the presence of an obtuse
predorsal angle (vs. flat dorsal rim).
Distribution: Late Burdigalian to early Langhian of Ven-
ezuela and Colombia.
Subfamily Micropogoniinae SASAKI, 1989
Definition: We have reduced the Micropogoniinae of SA-
SAKI (1989) to incorporate only the tribe Micropogoniini,
while we moved Lonchurini to the Sciaeninae.
Tribe Micropogoniini SASAKI, 1989
Definition and Discussion: In the limits as defined by SA-
SAKI (1989) to accommodate for the genus Micropogonias.
This is also in agreement with the Micropogonias group of
CHAO (1978) and LO et al. (2015) (who however further
included Sciaenops in this lineage), while SCHWARZHANS
(1993) combined Micropogonias with Protosciaena and the
Otolithini sensu MOHAN (1972), including the Pennahiini
and Atrobuccini of SASAKI (1989) based on the spectacular
and unique tadpole-like or buffer-shaped caudal tip. New
data on fossil Protosciaena otoliths (see earlier), however,
suggest that despite its uniqueness this otolith character may
not necessarily represent a synapomorphy of the genera
where it occurs, but might have been originated in two or
three sciaenid lineages independently. We have therefore
selected to maintain separate positions for the three tribes
Protosciaenini, Micropogoniini and Otolithini (not present
in America).
Genus Micropogonias
BONAPARTE, 1831
Micropogonias altipinnis (GÜNTHER, 1864)
(Plate 13, Fig. 9)
Material: One very large specimen NMB P1107, PPP 3514, Punta
Canoa, Ecuador, Canoa FM, early Pleistocene.
Discussion: The single well preserved and very large otolith
of nearly 28 mm length is identified as a representative of
the extant East Pacific species based on its low index OL : OH
of 1.2 and its high dorsal field, both typical for otoliths
of M. altipinnis (see SCHWARZHANS, 1993 for figures of
Recent specimens).
Distribution: Early Pleistocene of Ecuador and Recent from
the Gulf of California to Peru.
Micropogonias coatesi
AGUILERA & RODRIGUES DE AGUILERA, 2004
(Plate 13, Figs. 7-8)
2004 Micropogonias coatesi – AGUILERA & RODRIGUES DE
AGUILERA: pl. 1, figs. 1-4
Material: 3 specimens (holotype and two paratypes); 2 specimens,
El Hatillo, Urumaco, Venezuela, Urumaco FM, middle Mb, mid-
dle Tortonian, UNEFM-PF-053-054; 1 specimen, PPP 2534,
Carrizal, Venezuela, Caujarao FM, Mataruca Mb, late Tortonian.
Description: Compressed, thick otoliths up to about 10 mm
length. OL : OH = 1.1-1.2. OH : OT about 1.6. Dorsal rim
relatively regularly curved with obtuse mediodorsal angle;
ventral rim very deep, deepest below postostial lobe, with
flat or slightly concave stretch below caudal tip, steeply
ascending to posterior tip; anterior rim very high, broadly
rounded; posterior tip rounded, positioned above caudal tip.
Inner face strongly convex, with very large sulcus. Ostium
extremely large and wide, occupying about half of inner
face, with distinct postostial lobe. Cauda shallow, short,
with about equally anterior and downturned posterior sec-
tions, caudal tip tadpole-like widened. OCL : OCH = 1.0;
CCL : OCL = 0.5-0.6; caudal curvature index = 0.9; ostial-
caudal interspace 7-10 %. Dorsal depression indistinct,
narrow, only above anterior part of cauda; no ventral
furrow. Outer face as strongly convex as inner face, with
broad, supramedian umbo, little ornamentation.
Discussion: Otoliths of M. coatesi are easily recognized
and distinguished from all Recent species by the strongly
biconvex, thick appearance and the very large ostium, which
is best expressed in the narrow index CCL : OCL (0.5-0.6
vs. 0.9-1.25) and the very low ostial-caudal interspace
(7-10 % vs. 23-33 %). It is probably not related to any of
the Recent species, and due to the apomorphic status of
the main distinguishing characters it is likely to represent a
fossil lineage within the genus.
Distribution: Tortonian of Venezuela.
Micropogonias furnieri (DESMAREST, 1823)
(Plate 13, Fig. 10)
Material: 1 specimen NMB P1108, Cerro Caiguire, Cumaná,
Venezuela, Cumaná FM, middle Pleistocene.
Discussion: A single well preserved otolith of about
8.5 mm length characterized by a rather elongate shape
(OL : OH = 1.35), a flat and smooth dorsal rim and a
strongly indented concavity at the postventral rim, which
allow identification as M. furnieri (see SCHWARZHANS,
1993 for figures of Recent specimens).
Distribution: Middle Pleistocene of Venezuela and Recent
from the Caribbean to Argentina.
Subfamily Stelliferinae SASAKI, 1989
Definition and Discussion: In the limits as defined by
SASAKI (1989) plus the fossil otolith-based genera
†
Proto-
larimus AGUILERA & SCHWARZHANS, 2014 and
†
Xeno-
tolithus SCHWARZ HANS, 1993. SASAKI (1989) listed nine
synapomorphies for the subfamily. CHAO (2001) notes that
the species of the Stelliferinae “are unique in having a two
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
61
Palaeo Ichthyologica 14
chambered gas-bladder, the anterior one yoke-shaped and
the posterior main chamber carrot-shaped, and two large
pairs of otoliths (sagitta and lapillus)”.
In fact, the Stelliferinae show one of the most aberrant
otolith morphologies within Sciaenidae, (and entire Teleostei)
characterized by a number of putative synapomorphies
which support well their monophyly. The most striking
character is the reduction of the anterior tip of the sagitta
otolith, which is also bent outwards, often resulting in a
strong convexity of the anterior part of the inner face. This
reduction of the anterior part of the sagitta goes in hand
with an enlargement of the lapillus otolith located in front
of the sagitta (CHAO, 1978). The lapilli appear to have
developed some advanced morphological characters of
their own in the course of this specialization, which may
prove to be also of taxonomic value in this group, subject
however to detailed investigations, which we were unable
to perform in the scope of this study. In some species of
the genus Stellifer the lapillus has achieved almost the size
of the sagitta, which in turn is reduced in relative size as
compared to most other otoliths of the Sciaenidae. – Other
highly apomorphic characters of the sagitta otoliths of the
Stelliferinae are thorns, spikes and projections of the anterior
rim of the otolith as a result of the anterior reduction, the
reduction and bending of the ostium and the deep cauda,
which is often bent downwards to less than 90°.
The specialization of the sagitta otoliths (‘otoliths’ in
the following, if not noted otherwise) also allows for a well
supported recognition of three groups referred to in the
following as Odontoscionini, Stelliferini and the Xenotolithus
otolith group, the latter considered a fossil tribe incertae
sedis within Stelliferinae. The fossil otolith-based genus
Protolarimus is considered a plesiomorphic representative
of the stem group of the Stelliferinae and is therefore not
associated with a specific tribe.
The Stelliferinae are a species rich subfamily and
their Recent otoliths have been known rather incompletely
in the past, particularly for the genera Ophioscion and
Stellifer. We have therefore undertaken specific efforts to
complement the coverage of otoliths to those figured by
CHAO (1978, 2001, 2002), CORREA & VIANNA (1992),
SCHWARZHANS (1993) and ROSSI-WONGTSCHOWSKI et
al. (2014) now leaving only five species unaccounted for:
Bairdiella armata GILL, 1863, Corvula batabana (POEY,
1860), Ophioscion simulus GILBERT, 1898, Stellifer magoi
AGUILERA, 1983 and Stellifer melanocheir EIGENMANN,
1924. Ophioscion adustus (AGASSIZ, 1831) is considered
a junior synonym of Micropogonias furnieri (DESMAREST,
1823) in agreement with the presumption of CHAO (1978)
and after having investigated several specimens identified
as O. adustus (BMNH 1903.6.9.62, BMNH 1931.12.5.164,
BMNH 1935.8.29.19), which all turned out to represent
M. furnieri and which are easily recognized by the very
distinctive otolith patterns found in both genera.
The many Recent Stelliferinae species generally show
only rather subtle differences in their otolith morphology,
particularly so in the Odontoscionini, where not all species
may actually be distinguishable by otoliths. In contrast the
degree of ontogenetic changes is very low. On the higher
taxonomic level, otolith morphologies are mostly useful
for distinguishing genera and tribes, but at the same time
demonstrate that the distinction of both levels within Stel-
liferinae requires review. Recent phylogenetic studies by
BARBOSA et al. (2014), LO et al. (2015) and SANTOS et
al. (2013) all came to similar conclusions and are largely
supported by our otolith analysis.
Genus
†
Protolarimus
AGUILERA & SCHWARZHANS, 2014
Remarks: This genus was originally established with the
assumption that it would represent a plesiomorphic genus
close to Larimus. More detailed comparison with various
extant stelliferin otoliths has led us to move this genus to
the Stelliferinae where we assume it occupies a very basal
position in the stem group of the subfamily.
†Protolarimus henrici (NOLF & AGUILERA, 1998)
(Plate 14, Figs. 1-3)
1998 Larimus henrici – NOLF & AGUILERA: pl. 11, figs. 1, 3, 4
(non fig. 2)
Material: 8 specimens; 4 specimens, Castilletes, Colombia, lo-
cations 290840 (STRI 37408), 390094, Castilletes FM, late
Burdigalian to Langhian; 4 specimens, figured specimens coll.
Schwarzhans, San José de Cocodite, Paraguaná Peninsula,
Venezuela, Cantaure FM, late Burdigalian to Langhian.
Description: Elongate and moderately thick otoliths to
about 11 mm length. OL : OH = 2.0-2.25; OH : OT = 1.3.
Dorsal rim flat, sometimes with middorsal notch and step
and predorsal projection; ventral rim shallow, nearly flat,
smooth; anterior rim very broadly rounded to slightly inclined
with dorsal projection; posterior tip with distinct inferior
angle about
1
/
3
up from ventral rim; section immediately
below slightly concave.
Inner face strongly bent along the horizontal axis with
a curvature index of about 22 %. Ostium shallow, smooth,
long, moderately wide, its dorsal margin slightly inclined
downwards, margins with broad indention slightly anterior of
their middle, moderate postostial lobe. Cauda deep, poste-
rior portion of cauda bent downward at about 90°, slightly
widened. OCL : OCH = 1.7-1.95; CCL : OCL = 0.65-0.75;
caudal curvature index = 0.8-0.9; ostial-caudal interspace
17-20 %. No dorsal depression, but variable edge running
from tip of predorsal projection to postdorsal bent along
dorsal rim of otolith, slightly departing inwards anteriorly;
no ventral furrow. Outer face nearly flat, irregular, somewhat
tuberculate.
Discussion: Protolarimus henrici is considered one of the
most plesiomorphic otolith morphologies to be expected in
the Stelliferinae, close to their basal dichotomy. Obviously,
the anterior part of the otolith is not (yet) reduced, but a
mild predorsal projection and the strong bent of the inner
face in the horizontal axis are already indicative, as well as
the incipient postdorsal projection still situated rather low
on the posterior rim. The ostium is much larger than in
modern Stelliferinae, presumably also owing to the lack of
a anterior reduction of the otolith, but does show an outline
that resembles the more advanced forms. Other similarities
are with the Larimini, where the species has originally been
placed, and even more though with the Lonchurini. Of the
latter, the parallel occurring Polycirrhus mustus is the one
probably resembling closest. Protolarimus henrici differs
in the expression of an incipient predorsal projection (vs.
broad lobe) and posterior projection (vs. vertically cut), in
a flat outer face (vs. with strong postcentral umbo) and a
narrower ostium (OCL : OCH 1.7-1.95 vs. 1.5-1.6). We
consider the mild predorsal and posterior projections as
incipient developments indicating a basal position in Stellif-
erinae and possibly also indicating an outgroup relationship
to Lonchurini or related groups.
62
Distribution: Late Burdigalian to early Langhian of Ven-
ezuela and Colombia.
†
Protolarimus lundbergi
(AGUILERA & RODRIGUES DE AGUILERA, 2004)
(Plate 14, Fig. 4)
2004 Ophioscion lundbergi – AGUILERA & RODRIGUES DE
AGUILERA: pl. 1, figs. 7-10
Material: 2 specimens, figured specimen NMB P1118, El Ha-
tillo, Urumaco, Venezuela, Urumaco FM, middle Mb, middle
Tortonian.
Description: Elongate and moderately thick otolith of about
8.5 mm length. OL : OH = 1.85; OH : OT = 1.5. Dorsal rim
flat, median section slightly concave, anteriorly with broad,
short predorsal projection (bent upwards in the type speci-
mens figured by AGUILERA & RODRIGUES DE AGUILERA,
2004), posteriorly with broadly rounded postdorsal angle;
ventral rim shallow, nearly flat (or gently curved in the
type specimens figured by AGUILERA & RODRIGUES DE
AGUILERA, 2004), smooth; anterior rim nearly vertically cut
with dorsal projection marked, but very faintly projecting
forward; posterior tip with distinct inferior projection almost
at junction with ventral rim.
Inner face strongly bent along the horizontal axis with
a curvature index of about 25 %. Ostium shallow, smooth,
moderately long, narrow, its dorsal margin slightly inclined
downwards, margins with broad indention slightly anterior
of their middle, no postostial lobe, posterior rim of ostium
box-shaped. Cauda deep, anteriorly slightly narrowed
at ostial-caudal joint, posterior portion of cauda bent
downward at about 90°, not widened. OCL : OCH = 1.5;
CCL : OCL = 0.95; caudal curvature index = 0.8; ostial-
caudal interspace 30 %. No clear dorsal depression, but
sharp edge running very close to dorsal rim of otolith, slightly
departing inwards anteriorly; no ventral furrow. Outer face
nearly flat, irregular.
Discussion: Protolarimus lundbergi is further advanced
than P. henrici and intermediate in appearance to the most
primitive Ophioscion – O. transitivus – but not in a strict
evolutionary sense since it is younger than the latter. The
reduction of the anterior rim is slightly more advanced
in P. lundbergi when compared to P. henrici which is also
evidenced by the strong bent of the inner face along the
horizontal axis and the predorsal and posterior projections
being further developed so that recognition of a Stelliferini
otolith has never been questioned.
Distribution: Middle Tortonian of Venezuela.
†Protolarimus? mauryae
AGUILERA & SCHWARZHANS, 2014
(Plate 14, Fig. 5)
2014 Protolarimus mauryae – AGUILERA & SCHWARZHANS:
figs. 10.1-10.2
Material: 4 specimens (holotype and paratypes), Atalaia beach,
Brazil, early Miocene, MPEG-1805-1806-V.
Discussion: Protolarimus mauryae differs from the two other
species of this extinct genus in the considerably compressed
outline, expressed in the low OL : OH ratio of 1.4-1.5
(vs. 1.65-2.25), the lacking postdorsal projection and
the rounded anterior rim. Its ostium shows the downward
bent and inflection of the dorsal margin that is typical for
many species of the Larimini or Sciaenini, but may also be
considered as a very primitive status in Stelliferinae. The
allocation of this species therefore is regarded as tentative
and with more and better knowledge of early Miocene and
older sciaenid otoliths from South America it may well turn
out to represent a lineage separate from Protolarimus and
with a different relationship.
Distribution: Aquitanian to early Burdigalian of Brazil.
Tribe Odontoscionini SASAKI, 1989
Definition and Discussion: In the limits as defined by
SASAKI (1989) with the genera Corvula, Elattarchus and
Odontoscion, but also Bairdiella, which is moved from
Sasaki’s Stelliferini to Odontoscionini on the basis of their
highly similar otolith morphology with several clear syna-
pomorphies, in line with the definition and explanation of
the Bairdiella otolith group as defined by SCHWARZHANS
(1993). Otoliths of the Odontoscionini are distinguished
from those of the Stelliferini by the development of a
long and sharp predorsal spine, the transformation of
the postdorsal projection into nearly as sharp and long a
spine, and the remarkable reduction of the ventral portion
of the inner face which becomes very short and extremely
bent. CHAO (1978) also related Bairdiella (then including
Corvula as a synonym) and Odontoscion and commented
that “at present, the generic boundaries (of the two genera)
are not clear”.
Genus Bairdiella GILL, 1861
Bairdiella ronchus (CUVIER, 1830)
(Plate 14, Fig. 7)
Material: 2 specimens PPP 1727, 1734, figured specimen NMB
P1109, Río Banano, Costa Rica, Río Banano FM, late Pliocene.
Discussion: Two Bairdiella species occur in the West Atlantic
– B. chrysoura (LACEPÈDE, 1802) from off New York State
to Florida and in the Gulf of Mexico and B. ronchus in the
Caribbean and southwards to southern Brazil. The otoliths of
both species are extremely similar. SCHWARZHANS (1993)
noted three inconspicuous features for distinction, of which
the lack of an upward bent of the ventral margin at the
caudal joint with the ostium is probably the most reliable
and allows identification of these otoliths as B. ronchus.
Distribution: Late Tortonian to Messinian of the Dominican
Republic and late Pliocene of Costa Rica and Recent in the
Caribbean and along the shores of Brazil.
Plate 14
Figs. 1- 3. Protolarimus henrici (NOLF & AGUILERA, 1998).
1-2, Venezuela, San José de Cocodite, Cantaure FM, early Mio-
cene; 3, Colombia, La Guajira, Castilletes FM, early Miocene.
Fig. 4. Protolarimus lundbergi (AGUILERA & RODRIGUES DE AGUI-
LERA, 2004); Venezuela, El Hatillo, Urumaco FM, late Miocene.
Fig. 5. Protolarimus? mauryae AGUILERA & SCHWARZHANS,
2014; holotype, MPEG-1805-V, Brazil, Atalaia beach, Pirabas
FM, early Miocene.
Fig. 6. Odontoscion dentex (CUVIER, 1830); Venezuela, Cumana,
Cumana FM, middle Pleistocene.
Fig. 7. Bairdiella ronchus (CUVIER, 1830); Costa Rica, Rio Banano
FM, late Pliocene.
Figs. 8-10. Bairdiella sp. 8, Ecuador, Onzole FM, late Miocene;
9 (r), 10, Dominican Republic, Cercado FM, late Miocene.
/
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
63
Palaeo Ichthyologica 14
2 mm
2 mm
Protolarimus henrici
Protolarimus lundbergi
Odontoscion dentex
1 mm
1 mm
Protolarimus ? mauryae
5a
5b
6a
6b
6c
7a
7b
7c
7d
7e
8a
9
10a
10b
10c
8b
Bairdiella ronchus
1a
1b
1c
1d
1 mm
2
4a
4b
4c
4d
4e
3
Bairdiella sp.
1 mm
64
Bairdiella sp.
(Plate 14, Figs. 8-10)
1992 Bairdiella ronchus (CUVIER, 1830) – NOLF & STRINGER:
pl. 16, fig. 2
Material: 4 eroded or fragmented specimens; 1 specimen
PPP 3484, NMB P1110, Río Camarones, Ecuador, Onzole
FM, Messinian; 1 specimen PPP 1566, Río Chico, Darien,
Panama, Yaviza FM, early Tortonian; 1 specimen TU 1294,
NMB P123, Río Mao, Dominican Republic, Cercado FM, late
Tortonian-Messinian; 1 specimen NMB 16817, NMB P377,
Cañada de Zamba, Dominican Republic, Cercado FM, late
Tortonian-Messinian.
Discussion: These otoliths are characterized by a long
and rather broad posterior projection and a relatively wide
ostial-caudal interspace, but none of the specimens is well
enough preserved to warrant a differential diagnosis at the
species level in this genus known for its subtle differences
in the otolith morphology.
Genus Odontoscion GILL, 1862
Odontoscion dentex (CUVIER, 1830)
(Plate 14, Fig. 6)
Material: 1 specimen NMB P1111, Cerro Caiguire, Cumaná,
Venezuela, Cumaná FM, middle Pleistocene.
Discussion: Otoliths of Odontoscion are distinguished from
Bairdiella mainly by the shorter predorsal spine, which is
also located higher up relative to the highest point of the
dorsal rim of the otolith. The genus contains two geminate
species on either side of the Isthmus of Panama (and one
endemic species at the Galapagos Islands).
Distribution: Pleistocene of Venezuela and Recent from
Florida to northeastern Brazil and in the Caribbean.
Tribe Stelliferini SASAKI, 1989
Definition and Discussion: In the limits as defined by SA-
SAKI (1989) with the genera Ophioscion and Stellifer, but
excluding Bairdiella, which we moved to Odontoscionini on
the basis of synapomorphies in the otolith morphology (see
above). Otoliths of the Stelliferini are distinguished from
those of the Odontoscionini by the more elongate shape,
the more strongly reduced anterior rim of the otolith with
only a weak or no predorsal spine and a broad, not spiny
postdorsal projection. The generic boundaries of the two
genera contained are not clear and definitely subject to
future revision, as already indicated in the molecular stud-
ies by BARBOSA et al. (2014) and SANTOS et al. (2013).
SCHWARZHANS (1993) resurrected the nominal genera
Nector JORDAN & EVERMANN, 1898, Sigmurus GILBERT,
1898 and Zestidium GILBERT, 1898 solely based on otolith
analyses. While we expect some of this might be validated
in the future, we feel that at present more in depth review
work is required of the many Recent species, particularly
in the genus Stellifer, until such formal action would be
warranted.
Genus Ophioscion GILL, 1863
Remarks: Otoliths of the following Recent species of Ophio-
scion are figured for comparison and as complementation
for CHAO (1978, 2002) and SCHWARZHANS (1993):
Ophio scion imiceps (JORDAN & GILBERT, 1882) (pl. 16,
fig. 8), Ophioscion scierus (JORDAN & GILBERt, 1884)
(pl. 16, fig. 9), Ophioscion strabo GILBERT, 1898 (pl. 15,
fig. 6), Ophioscion typicus GILL, 1863 (pl. 15, fig. 5),
Ophioscion vermicularis (GÜNTHER, 1867) (pl. 15, fig. 4);
as well as two species of the genus Stellifer, which bear
close resemblance to Ophioscion otoliths: Stellifer chryso-
leuca (GÜNTHER, 1867) (pl. 16, fig. 11) and Stellifer minor
(TSCHUDI, 1846) (pl. 16, fig. 10). Otoliths studied from
Ophioscion punctatissimus MEEK & HILDEBRAND, 1925
(BMNH 1993.6.30.41-42) were not well enough preserved
for figuring. Otoliths of O. imiceps resemble Stellifer otoliths,
those of O. vermicularis are characterized by a relatively
long and straight predorsal spine and those of O. typicus
are remarkable for their depressed predorsal rim. The latter
character is also shared with several species of the genus
Stellifer, i. e. Stellifer brasiliensis (SCHULTZ, 1945) (see
ROSSI-WONGTSCHOWSKI et al., 2014), Stellifer microps
(STEINDACHNER, 1864) (pl. 16, fig. 12), Stellifer naso
(JORDAN, 1889) (otolith studied from CAS-SU 69078 not
well enough preserved for figuring) and Stellifer venezuelae
(SCHULTZ, 1945) (see CHAO, 1978; otoliths studied from
USNM 218347 and CAS-SU 52234 are not well enough
preserved for figuring).
Ophioscion amphiamericanus n. sp.
(Plate 16, Figs. 1-7)
1992 Ophioscion species 1 – NOLF & STRINGER: pl. 16, fig. 3
(non fig. 5)
Holotype: Pl. 16, Fig. 1, NMB P1112, PPP 3432, Río Cayapas,
Ecuador, Angostura FM, Tortonian.
Paratypes: 25 specimens; 1 specimen NMB P124, location
NMB 16913, Río Mao, Dominican Republic, Cercado FM,
late Tortonian
-
Messinian; 12 specimens NMB P358, location
NMB 15904, Río Gurabo, Dominican Republic, Cercado FM,
late Tortonian
-
Messinian; 3 specimens, NMB P355, location
NMB 15914, Río Gurabo, Dominican Republic, Cercado FM,
late Tortonian
-
Messinian; 7 specimens, NMB P366, location
NMB 16923, Río Mao, Dominican Republic, Cercado FM, late
Tortonian
-
Messinian; 1 specimen NMB P1113, PPP 3406, Río
Cayapas, Ecuador, Angostura FM, Tortonian; 1 specimen NMB
P1114, same data as holotype.
Further material: 20 specimens; 18 specimens, locations NMB
16913 (NMB P124), 16915 (NMB P371), 16917 (NMB P367),
16918 (NMB P370), 16927 (NMB P375), Rio Mao, NMB 15900
(NMB P362), Río Gurabo, Dominican Republic, Cercado FM,
Plate 15
Figs. 1-3. Ophioscion transitivus n. sp. 3, holotype, NMB P1117,
Trinidad, Brasso FM, early-middle Miocene; 1, 2 (r), paratypes,
NMB P125 and 378, Dominican Republic, Baitoa FM, early-
middle Miocene.
Fig. 4. Ophioscion vermicularis (GÜNTHER, 1867); coll. OA,
Pacific off Panama, Recent.
Fig. 5. Ophioscion typicus GILL, 1863; coll. OA, Pacific off
Panama, Recent.
Fig. 6. Ophioscion strabo GILBERT, 1898; coll. MNHN, off Peru,
Recent.
Figs. 7- 9. Ophioscion inflaticauda n. sp. 7, holotype, NMB
P1115; 8-9, paratypes, NMB P1116-1117; Trinidad, Manzanilla
FM, late Miocene.
/
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65
Palaeo Ichthyologica 14
Ophioscion typicus
Ophioscion strabo
1a
1b
1c
1d
1e
2a
2b
3a
3b
3c
3d
4a
4b
4c
1f
1 mm
1 mm
2 mm
2 mm
2 mm
Ophioscion vermicularis
Ophioscion transitivus
Ophioscion inflaticauda
6a
5a
5b
6b
6c
7a
7b
7c
7d
7e
7f
8
9
66
late Tortonian
-
Messinian; 2 specimens PPP 3432 Río Cayapas
and PPP 3440, Río Santiago, Ecuador, Angostura FM, Tortonian.
Name: Referring to the occurrence of the species on the
Pacific as well as the Atlantic side of northern South America.
Diagnosis: OL : OH = 1.65-1.8. Anterior rim inclined, with
strong predorsal spine. Dorsal rim undulating, with rounded
predorsal and mediodorsal angles and incision in between.
Posterior tip massive, projecting. Ostium short, narrow,
downward inclined, its ventral margin narrowing towards
ostial-caudal joint. Curvature index of inner face 15-20 %.
Description: Moderately elongate, moderately thick otoliths
up to about 5.5 mm length. OH : OT = 1.5-1.8. Dorsal
rim moderately curved, irregular, undulating, with broadly
rounded elevated predorsal angle above ostial-caudal joint
followed by indentation and thereafter broad, shallow medi-
odorsal angle above anterior part of cauda; posterior part
of dorsal rim inclined, straight. Ventral rim anteriorly shallow
and regularly curved, posteriorly strongly bend upwards with
broad and distinct concavity behind caudal tip and prior to
reaching posterior tip. Anterior rim inclined at about 30-35°,
ventrally straight up to a notch marking upper margin of
ostium and above expanding into long and sharp spine.
Posterior rim with broad, long, asymmetrical projection with
curved, convex dorsal and concave ventral rims.
Inner face strongly bent along the horizontal axis with
a curvature index of 15-20 %. Ostium shallow, short, mod-
erately wide, its upper margin bent downwards at about
20°, its ventral margin narrowing towards ostial-caudal
joint instead of a postostial lobe. Narrow furrow present
above ostial colliculum. Cauda very deep, slightly narrow-
ing towards ostial-caudal joint, with sharp edge at ventral
turning point of cauda to about middle of caudal height;
posterior portion of cauda bent downward at nearly 90°.
OCL : OCH = 1.0-1.2; CCL : OCL = 1.5-2.0; caudal curva-
ture index = 0.7-0.75; ostial-caudal interspace 36-42 %.
No dorsal depression, but distinct edge running from tip
of predorsal spine to posterior tip close to dorsal rim of
otolith, slightly departing inwards anteriorly and posteriorly;
no ventral furrow. Outer face flat, anteriorly slightly concave,
posteriorly slightly convex, with distinct, backward oriented
projection on dorsal field underpinning mediodorsal angle.
Discussion: The prominent predorsal spine characterizes
this otolith immediately as a relative of O. vermicularis
(pl. 15, fig. 4) and thus sets it apart from all other Recent
species of the Stelliferinae. Ophioscion amphiamericanus
differs from Ophioscion vermicularis (GÜNTHER, 1867) in
the shape of the dorsal rim, particularly its irregular mid-
dorsal section (vs. nearly flat and regularly formed), the less
pointed posterior tip and the posterior-ventral reduction of
the ostium. Otoliths of the Odontoscionini resemble as well
in general appearance, but are always considerably less
elongate and show a still more pronounced and longer and
sharper predorsal spine and a narrower posterior projection.
Distribution: Tortonian and Messinian of Ecuador and the
Dominican Republic.
Ophioscion inflaticauda n. sp.
(Plate 15, Figs. 7-9)
Holotype: Pl. 15, Fig. 7, NMB P1115, PPP 2667, Manzanilla Bay,
Trinidad, Manzanilla FM, San José Mb, late Tortonian.
Paratypes: 4 specimens; 1 specimen NMB P1116, same data as
holotype; 1 specimen NMB P535, Cb. 1633 San José River,
Trinidad, Manzanilla FM, San José Mb, late Tortonian; 2 speci-
mens, NMB P533-534, Cb. 2076, Cascade River, Trinidad,
Tamana FM, middle Tortonian.
Further material: 5 incomplete specimens, PPP 2667, Manzanilla
Bay and PPP 2675, San José River, Trinidad, Manzanilla FM,
San José Mb, late Tortonian.
Name: From inflatus (Latin = inflated) and cauda (rear part
of sulcus), referring to the widened cauda.
Diagnosis: OL : OH = 1.8. Anterior rim inclined, with strong
predorsal spine. Dorsal rim undulating, with rounded
predorsal and postdorsal angles and middorsal concavity.
Posterior tip massive, projecting. Ostium short, narrow,
downward inclined. Upper section of downturned portion
of cauda bulged on anterior/ventral rim. Curvature index
of inner face 12 %.
Description: Moderately elongate, moderately thick otoliths
up to 6.3 mm length. OH : OT about 1.7. Dorsal rim mod-
erately curved, undulating, with broadly rounded, slightly
elevated predorsal angle above ostial-caudal joint, very
broad, shallow postdorsal angle above rear part of cauda
and broad middorsal concavity in between; posterior part
of dorsal rim regularly curved. Ventral rim anteriorly shallow
and regularly curved, posteriorly strongly bend upwards with
broad and distinct concavity behind caudal tip and prior
to reaching posterior tip. Anterior rim inclined at about
35-40°, straight without notable notch and expanding into
long and sharp predorsal spine. Posterior rim with broad,
long, asymmetrical projection with curved, convex dorsal
and concave ventral rims.
Inner face strongly bent along the horizontal axis with
a curvature index of about 12 %. Ostium shallow, smooth,
short, moderately wide, its upper margin bent downwards at
about 20°, sometimes indented, its ventral margin narrow-
ing towards ostial-caudal joint instead of a postostial lobe.
No distinct narrow furrow above ostial colliculum. Cauda
very deep, distinctly narrowing towards ostial-caudal joint,
with short edge at ventral turning point of cauda; posterior
portion of cauda bent downward at slightly less than 90°,
strongly widened, particularly on its upper half and there
marked by a distinct bulge of anterior/ventral margin of
cauda. OCL : OCH = 1.0; CCL : OCL = 1.35-1.7; caudal
curvature index = 0.6-0.65; ostial-caudal interspace 24-
30 %. No dorsal depression, but distinct edge running from
tip of predorsal spine to posterior tip close to dorsal rim
of otolith, slightly departing inwards anteriorly; no ventral
furrow. Outer face flat, with distinct, backward oriented
projection on dorsal field underpinning mediodorsal angle.
Plate 16
Figs. 1-7. Ophioscion amphiamericanus n. sp. 1 (r), holotype,
NMB P1112, Ecuador, Angostura FM, late Miocene; 3 (r), paratype,
NMB P1113, Ecuador, Angostura FM, late Miocene; 2 (r), 4 (r), 5-7,
paratypes, NMB P124, 355, 358, 366, Dominican Republic, Cer-
cado FM, late Miocene.
Fig. 8. Ophioscion imiceps (JORDAN & GILBERT, 1882); coll. OA,
Pacific off Panama, Recent.
Fig. 9. Ophioscion scierus (JORDAN & GILBERT, 1884); coll. WS,
Pacific off Mexico, Recent.
Fig. 10. Stellifer minor (TSCHUDI, 1846); coll. MNHN, off Peru,
Recent.
Fig. 11. Stellifer chrysoleuca (GÜNTHER, 1867); coll. Aguirre,
off Ecuador, Recent.
Fig. 12. Stellifer microps (STEINDACHNER, 1864); coll. MNHN,
Atlantic off Colombia, Recent.
Fig. 13. Stellifer acerbus n. sp.; holotype, NMB P1122, Trinidad,
Manzanilla FM, late Miocene.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
/
67
Palaeo Ichthyologica 14
1 mm
2 mm
2 mm
2 mm
2 mm
Ophioscion scierus
Stellifer minor
Ophioscion imiceps
Ophioscion amphiamericanus
1a
1b
1c
1d
1e
1f
2a
2b
3a
3b
4
5
6
8a
8b
9a
9b
9c
10a
10b
11a
11b
11c
Stellifer chrysoleuca
Stellifer microps
Stellifer acerbus
2 mm
1 mm
12a
12b
13a
13b
13c
13d
13e
13f
7
68
Discussion: Ophioscion inflaticauda is readily recognized
by the widened upper section of the downturned part of the
cauda, which is particularly well expressed in the bulged,
convex stretch of the anterior (ventral) margin of this part of
the cauda. This is a unique character not seen in any other
Stelliferinae otoliths. This character is so unique that even
poorly preserved specimens or fragments with preserved
cauda can be identified.
The specimens described by NOLF (1976; pl. 7, figs. 11-
13) as Sigmurus sp. show the same characteristic bulge of
the anterior/ventral margin of the downturned section of
the cauda, although somewhat variable, with the slightly
older ones from the Tamana FM depicting a slightly weaker
development.
Distribution: Late Tortonian of Trinidad.
Ophioscion transitivus n. sp.
(Plate 15, Figs. 1-3)
1992 Ophioscion species 2 – NOLF & STRINGER: pl. 16, fig. 4
Holotype: Pl. 15, Fig. 3, NMB P1117, PPP 2661, beyond bridge
over Gorge River, Trinidad, Brasso FM, late Burdigalian to
Langhian.
Paratypes: 2 specimens NMB P125 and NMB P378, location NMB
17287, Rio Yaque del Norte, Dominican Republic, Baitoa FM,
late Brudigalian to Langhian.
Name: From transitivus (Latin = transitional), referring to the
transitional morphology between the extinct
†
Protolarimus
and Ophioscion otolith morphology.
Diagnosis: OL : OH = 1.8. Anterior rim inclined, with
short, robust predorsal spine. Dorsal rim smooth, almost
flat. Posterior tip massive, broad, projecting. Ostium short,
wide, slightly inclined, with small indentions on upper and
lower margins close to opening. Curvature index of inner
face 12 %.
Description: Moderately elongate, moderately thick otoliths
up to 7 mm length. OH : OT = 1.5-2.0, decreasing with
size. Dorsal rim smooth, very shallow and regularly curved,
almost flat; posterior part of dorsal rim curved more strongly
only in front of posterior tip. Ventral rim anteriorly very
shallow, posteriorly strongly bent upwards with broad and
distinct concavity behind caudal tip and prior to reaching
posterior tip. Anterior rim inclined at about 30°, straight with
faint concavity at upper margin of ostium and expanding
into short, robust predorsal spine. Posterior rim with broad,
high, long, asymmetrical, ventrally reduced projection with
curved, convex dorsal and concave ventral rims.
Inner face strongly bent along the horizontal axis with
a curvature index of about 12 %. Ostium shallow, smooth,
short, wide, its dorsal margin bent downwards at about
10°, margins with small indention at anterior part, its
ventral margin not narrowing backwards, box-shaped. No
distinct narrow furrow above ostial colliculum. Cauda very
deep, slightly narrowing towards ostial-caudal joint, without
edge at ventral turning point of cauda; posterior portion
of cauda bent downward at slightly less than 90°, slightly
widened. OCL : OCH = 1.0- 1.3; CCL : OCL = 1.2- 1.3;
caudal curvature index = 0.7-0.9; ostial-caudal interspace
30 %. No dorsal depression, but distinct edge running from
tip of predorsal spine to posterior tip close to dorsal rim of
otolith, distinctly departing inwards anteriorly; no ventral
furrow. Outer face nearly flat, irregular, with moderate,
backward oriented projection on dorsal field at midsection.
Discussion: Ophioscion transitivus is easily recognized by
the smooth, nearly flat dorsal rim and the rather wide,
box-shaped ostium as well as the very broad posterior pro-
jection. These are all considered plesiomorphic characters
and resemble certain species of the extinct
†
Protolarimus,
particularly
†
P. lundbergi, from which it differs in the shorter
ostium (OCL : OCH = 1.0-1.3 vs. 1.5-1.6), the inclined
anterior rim (at 20° vs. nearly vertical), the presence of an,
albeit, short and robust predorsal spine and the ventrally
reduced posterior projection. The Recent O. vermicularis
(pl. 15, fig. 4) resembles in the flat dorsal rim and the
short predorsal spine, which is however still longer than in
O. transitivus, and which differs further in the blunt poste-
rior projection (vs. pointed and much expanded) and the
longer ostium (OCL : OCH = 1.0-1.3 vs. 0.7). We consider
O. transitivus a very plesiomorphic member of the genus
positioned close to its basal dichotomy.
Distribution: Late Brudigalian to Langhian of Trinidad.
Genus Stellifer CUVIER, 1817
Remarks: Otoliths of the following Recent species of Stel-
lifer are figured for comparison and as complementation
for CHAO (1978, 2001, 2002), LOMBARTE et al. (2006),
ROSSI-WONGTSHOWSKI et al. (2014) and SCHWAR-
ZHANS (1993): Stellifer chaoi AGUILERA, SOLANO & VAL-
DEZ, 1983 (pl. 17, fig. 9), Stellifer chrysoleuca (GÜNTHER,
1867) (pl. 16, fig. 11), Stellifer ephelis CHIRICHIGNO, 1974
(pl. 17, fig. 8), Stellifer ericymba (JORDAN & GILBERT, 1882)
(pl. 17, fig. 10, from LOMBARTE et al. (2006)), Stellifer
fuerthii (STEINDACHNER, 1876) (pl. 17, fig. 15), Stellifer il-
lecebrosus GILBERT, 1898 (pl. 17, fig. 7), Stellifer lanceolatus
(HOLBROOK, 1855) (pl. 17, fig. 11, from LOMBARTE et al.
(2006)), Stellifer mancorensis CHIRICHIGNO, 1962 (pl. 17,
fig. 13), Stellifer microps (STEINDACHNER, 1864) (pl. 16,
fig. 12), Stellifer minor (TSCHUDi, 1846) (pl. 16, fig. 10),
Stellifer oscitans (JORDAN & GILBERT, 1882) (pl. 17, fig. 14),
Stellifer pizarroensis HILDEBRAND, 1946 (pl. 17, fig. 16),
Stellifer rastrifer (JORDAN, 1889) (pl. 17, fig. 17), Stellifer
stellifer (BLOCH, 1790) (pl. 17, fig. 12) and Stellifer winter-
steenorum CHAO, 2001 (pl. 17, fig. 18). Key characters for
distinction of species by means otoliths are the ratio OL : OH,
development of the much reduced anterior rim including
the predorsal spine and a spine or angle at the middle
of the anterior rim, and the shape and proportions of the
ostium. The species Stellifer brasiliensis (SCHULTZ, 1945)
(see ROSSI-WONGTSCHOWSKI et al., 2014), Stellifer mi-
crops (STEINDACHNER, 1864) (pl. 16, fig. 12), Stellifer naso
(JORDAN, 1889) and Stellifer venezuelae (Schultz, 1945)
(see CHAO, 1978) represent a different otolith pattern with
a strongly reduced and deflected predorsal area and ostium,
which would warrant their allocation in a separate genus as
suggested by SCHWARZHANS (1993). Similar conclusions
were reached in the molecular phylogenetic investigations
by BARBOSA et al. (2014) and SANTOS et al. (2013).
Stellifer abbreviatus n. sp.
(Plate 17, Figs. 1-2)
Holotype: Pl. 17, Fig. 1, NMB P1119, PPP 1727, Costa Rica, Río
Banano FM, Piacenzian, late Pliocene.
Paratypes: 2 specimens NMB P1120-1121, PPP 1727 and 1728,
Costa Rica, Río Banano FM, Piacenzian, late Pliocene.
Name: From abbreviatus (Latin = abbreviated, truncated)
referring to the nearly vertically cut anterior rim.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
69
Palaeo Ichthyologica 14
Diagnosis: OL : OH = 1.35-1.4. Anterior rim nearly verti-
cally cut, with sharp predorsal spine and broader, similarly
long spine at middle of anterior rim. Ventral rim with con-
cavity behind ostium. Ostium small, short. Cauda curved
downwards posteriorly at about 50-60°.
Description: Compact, moderately thick otoliths up to 5 mm
length. OH : OT = 1.9. Dorsal rim shallow, regularly curved,
with distinct step behind predorsal spine where a dorsal
furrow meets dorsal rim. Ventral rim flat except for steeply
upturned posterior section, anteriorly slightly concave just
behind ostium. Anterior rim near vertically cut with sharp
predorsal spine, broad spine at middle section of anterior
rim and rounded ventral portion with ostium all reach-
ing same level of protrusion. Predorsal spine slightly bent
outwards, section with spine at middle section of anterior
rim and ventral portion with ostium strongly bent outwards.
Posterior rim with broad, angular postdorsal projection.
Inner face strongly bent along the horizontal axis with
a curvature index of 25 %. Ostium shallow, very small and
short, positioned entirely below ostial-caudal joint. Distinct
furrow present above ostial colliculum leading up to cauda.
Cauda very deep, not narrowing towards ostial-caudal joint,
posterior portion of cauda bent downward at about 50-
60°. [The following sulcus proportions measured on views
from inner face, resulting in short values for ostium due to
its strongly outward bent position.] OCL : OCH about 0.5;
CCL : OCL about 3.5; caudal curvature index = 0.45-0.55;
ostial-caudal interspace 47-53 %. No dorsal depression, but
distinct edge or terrace and furrow running close to dorsal
rim of otolith from step above ostium where it meets dorsal
rim to near posterior tip of otoliths where it fades, slightly
less curved than dorsal rim and as result departing inwards
below middorsal rim; no ventral furrow. Outer face flat in
view from ventral but somewhat bulged posteriorly in view
from posterior and with moderate projection on dorsal field
underpinning mediodorsal angle.
Discussion: Stellifer abbreviatus resembles otoliths of
the Recent S. ericymba (pl. 16, fig. 10) and S. illecebrosus
(pl. 17, fig. 7), both from the East Pacific, and S. colonensis
(figures in CHAO, 1978) from the Caribbean in the nearly
vertically cut anterior rim, the equally long predorsal spine
and spine at the middle section of the anterior rim and
the slight concavity of the ventral rim behind the ostium,
but differs from all of them in the more compressed shape
expressed in a low OL : OH ratio of 1.35-1.4 (vs. 1.45-
1.65). Other, similarly compressed Stellifer otoliths have no
distinct spine at the midsection of the anterior rim, like for
instance S. ephelis or S. rastrifer or have a short posterior
projection, like S. chaoi.
Distribution: Late Pliocene of Costa Rica.
Stellifer acerbus n. sp.
(Plate 16, Fig. 13)
Holotype (and unique specimen): Pl. 16, Fig. 13, NMB P1122,
PPP 2675, San José River, Trinidad, Manzanilla FM, San José
Mb, late Tortonian.
Name: From acerbus (Latin = painful, harsh) referring to
the extremely downward bent and small ostium.
Diagnosis: OL : OH = 1.4. Anterior-dorsal rim deeply
depressed, with very small predorsal spine and angular
projection at middle of anterior rim. Ventral rim regularly
curved, without concavity. Ostium extremely small, short,
positioned about half way below ostial-caudal joint, opening
to anterior-ventral rim. Cauda curved downwards posteriorly
at about 60°.
Description: Compact, moderately thin, small otolith
of 3.9 mm length. OH : OT = 2.2. Dorsal rim anteriorly
reduced, compressed, highest at mid-section, posteriorly
regularly curved, with concavity in front of mediodorsal lobe
where a dorsal furrow meets dorsal rim. Ventral rim flat at
mid-section, with steeply upturned anterior and posterior
sections. Anterior rim projecting strongest at position of
angular protrusion at middle of section, predorsal spine
very short, broad, indistinct. Ventral part of anterior rim
moderately bent outwards. Posterior rim with broad, angular
postdorsal projection.
Inner face strongly bent along the horizontal axis with
a curvature index of 15 %. Ostium shallow, extremely
small, short and inferior, positioned nearly 50 % below
ostial-caudal joint. Distinct and wide furrow present above
ostial colliculum leading upwards to cauda. Cauda very
deep, downward bent anteriorly to meet ostium and slightly
narrowing towards ostial-caudal joint, posterior portion of
cauda widened, bent downward at about 60°. [The follow-
ing sulcus proportions measured on views from inner face,
resulting in short values for ostium due to its strongly outward
bent position.] OCL : OCH = 1.25; CCL : OCL = 2.7; caudal
curvature index = 0.7; ostial-caudal interspace 45 %. No
dorsal depression, but distinct edge or terrace and furrow
running close to dorsal rim of otolith from in front of medi-
odorsal lobe where it meets dorsal rim to near posterior tip
of otoliths where it fades, slightly less curved than dorsal
rim and as result departing inwards below middorsal rim;
no ventral furrow. Outer face slightly concave anteriorly
and slightly convex posteriorly, smooth.
Discussion: Stellifer acerbus belongs to the group of Stel-
lifer species with depressed predorsal area including the
following Recent species: S. brasiliensis, S. microps (pl. 16,
fig. 12), S. naso and S. venezuelae. It differs from all of them
in the extremely depressed predorsal area and the extremely
downward shifted, very small ostium and is also the most
compressed of the four. We thus assume that S. acerbus
represents an extinct, highly derived lineage in this group.
Distribution: Late Tortonian of Trinidad.
Stellifer bicornutus n. sp.
(Plate 17, Figs. 3-4)
Holotype: Pl. 17, Fig. 3, NMB P1123, PPP 1727, Costa Rica, Río
Banano FM, Piacenzian, late Pliocene.
Paratypes: 1 specimen NMB P1124, same data as holotype.
Name: From bicornutus (Latin = with two horns) referring
to the two sharp spines at the anterior rim, one predorsal,
the other at middle section of anterior rim.
Diagnosis: OL : OH = 1.5-1.55. Anterior rim projecting
strongest at spine on mid-section above ostium; with equally
sharp, more backward positioned predorsal spine. Ventral
rim flat at mid-section. Ostium small, short, depressed,
positioned about
1
/
3
below ostial-caudal joint. Cauda curved
downwards posteriorly at about 70-80°.
Description: Moderately elongate, moderately thick otoliths
up to about 4.5 mm length. OH : OT = 1.7. Dorsal rim
shallow, regularly curved, with concavity behind predorsal
spine where a dorsal furrow meets dorsal rim. Ventral rim
flat except for steeply upturned posterior section. Anterior
rim near with two sharp spines, one predorsal spine situ-
70
ated somewhat backward and one below mid-section and
above ostium and marking maximum protrusion of anterior
tip; rounded ventral portion with ostium not protruding.
Predorsal spine slightly bent outwards, section with spine
at middle section of anterior rim and ventral portion with
ostium strongly bent outwards. Posterior rim with broad,
angular postdorsal projection.
Inner face strongly bent along the horizontal axis with
a curvature index of 25 %. Ostium shallow, very small and
short, positioned about
1
/
3
below ostial-caudal joint. Distinct
furrow present above ostial colliculum leading up to cauda.
Cauda very deep, not narrowing towards ostial-caudal
joint, posterior portion of cauda bent downward at about
70-80°. [The following sulcus proportions measured on
views from inner face, resulting in short values for ostium
due to its strongly outward bent position.] OCL : OCH about
0.9; CCL : OCL about 2.0; caudal curvature index = 0.5;
ostial-caudal interspace 43 %. No dorsal depression, but
distinct edge or terrace and furrow running close to dorsal
rim of otolith from concavity above ostium where it meets
dorsal rim to near posterior tip of otoliths where it fades,
slightly less curved than dorsal rim and as result departing
inwards below middorsal rim; no ventral furrow. Outer face
flat in view from ventral but somewhat bulged posteriorly
in view from posterior and with small projection on dorsal
field underpinning mediodorsal angle.
Discussion: Stellifer bicornutus resembles otoliths of the
Recent S. lanceolatus (pl. 17, fig. 11) found along the shores
from Virgina to Texas in the Gulf of Mexico in the ventral
portion of the anterior rim projecting more than the predor-
sal spine, the presence of two sharp spines at the anterior
rim and the inferior position of the ostium, but differs in
being more elongate (OL : OH 1.5-1.55 vs. 1.45) and the
reduced anterior-ventral angle (vs. equally projecting as
spine at middle section of anterior rim).
Distribution: Late Pliocene of Costa Rica.
Stellifer depressifrons n. sp.
(Plate 17, Figs. 5)
Holotype (and unique specimen): Pl. 17, Fig. 5, NMB P1125, San
José de Cocodite, Paraguaná Peninsula, Venezuela, Cantaure
FM, late Burdigalian to Langhian.
Name: From depressus (Latin = depressed) and frons
(Latin = forehead) referring to the very compressed outline
of the otolith.
Diagnosis: OL : OH = 1.3. Anterior rim slightly inclined
below angle at mid-section, predorsal spine sharp, relatively
short. Ventral rim flat at mid-section. Ostium small, short.
Cauda curved downwards posteriorly at about 60°.
Description: Compact, moderately thick otolith of 5.2 mm
length. OH : OT = 1.7. Dorsal rim shallow, regularly curved,
highest above anterior part of cauda, with distinct step be-
hind predorsal spine where a dorsal furrow meets dorsal
rim. Ventral rim flat except for steeply upturned posterior
section. Anterior rim near vertical, ventrally slightly inclined
backwards, with sharp, short predorsal spine and protruding
angle instead of spine at middle section of anterior rim.
Predorsal spine slightly bent outwards, middle and ventral
portion of anterior rim strongly bent outwards. Posterior rim
with broad, short, massive, angular postdorsal projection.
Inner face strongly bent along the horizontal axis with
a curvature index of 17 %. Ostium shallow, very small and
short, positioned about
1
/
3
below ostial-caudal joint. Distinct
furrow present above ostial colliculum leading up to cauda.
Cauda very deep, not narrowing towards ostial-caudal
joint, posterior portion of cauda bent downward at about
60°. [The following sulcus proportions measured on views
from inner face, resulting in short values for ostium due
to its strongly outward bent position.] OCL : OCH about
0.65; CCL : OCL about 3.5; caudal curvature index = 0.5;
ostial-caudal interspace 43 %. No dorsal depression, but
distinct edge or terrace and furrow running close to dorsal
rim of otolith from step above ostium where it meets dorsal
rim to near posterior tip of otoliths where it fades, almost
straight and hence considerably departing inwards below
middorsal rim; no ventral furrow. Outer face flat in view
from ventral but somewhat bulged posteriorly in view from
posterior.
Discussion: Stellifer depressifrons is one of the most com-
pressed otoliths of Stellifer, indicating that such advanced
morphologies were already around in early to middle
Miocene times. Other compressed otoliths of Recent species
either have a longer predorsal spine reaching the level of
the projection at the middle of the anterior rim (S. rastrifer,
S. stellifer, S. zestocarus) or have a clearly developed spine
at the middle section of the anterior rim (S. chaoi). None of
these Recent specimens show a backward inclined ventral
section of the anterior rim.
Distribution: Late Burdigalian to Langhian of Venezuela.
Stellifer onzole n. sp.
(Plate 17, Fig. 6)
Holotype: Pl. 17, Fig. 6, NMB P1126, PPP 3470, Río Santiago,
Ecuador, Onzole FM, Messinian.
/
Plate 17
Figs. 1-2. Stellifer abbreviatus n. sp. 1, holotype, NMB P1119;
2, paratype, NMB P1120; Costa Rica, Rio Banano FM, late Miocene.
Figs. 3-4. Stellifer bicornutus n. sp. 3, holotype, NMB P1123;
4, paratype, NMB P1124; Costa Rica, Rio Banano FM, late Miocene.
Fig. 5. Stellifer depressifrons n. sp.; holotype, NMB P1125,
Venezuela, San José de Cocodite, Cantaure FM, early Miocene.
Fig. 6. Stellifer onzole n. sp.; holotype, NMB P1126, Ecuador,
Onzole FM, late Miocene.
Fig. 7. Stellifer illecebrosus GILBERT, 1898; coll. MNHN, off
Peru, Recent.
Fig. 8. Stellifer ephelis CHIRICHIGNO, 1974; coll. MNHN, off
Peru, Recent.
Fig. 9. Stellifer chaoi AGUILERA, SOLANO & VALDEZ, 1983; coll.
OA, off Venezuela, Recent.
Fig. 10. Stellifer ericymba (JORDAN & GILBERT, 1882); AFORO
database, fish ID 7346, Recent.
Fig. 11. Stellifer lanceolatus (HOLBROOK, 1855); AFORO data-
base, fish ID 6819, Recent.
Fig. 12. Stellifer stellifer (BLOCH, 1790); BMNH 1930.19.30.86-
87, Brazil, off Santos, Recent.
Fig. 13. Stellifer mancorensis CHIRICHIGNO, 1962; coll. MNHN,
off Peru, Recent.
Fig. 14. Stellifer oscitans (JORDAN & GILBERT, 1882); coll. OA,
Pacific off Panama, Recent.
Fig. 15. Stellifer fuerthii (STEINDACHNER, 1876); coll. Aguirre,
off Ecuador, Recent.
Fig. 16. Stellifer pizarroensis HILDEBRAND, 1946; coll. MNHN,
off Peru, Recent.
Fig. 17. Stellifer rastrifer (JORDAN, 1889); coll. OA, off Ven-
ezuela, Recent.
Fig. 18. Stellifer wintersteenorum CHAO, 2001; coll. WS, Pacific
off Mexico, Recent.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
71
Palaeo Ichthyologica 14
1a
1b
1c
1d
1e
1f
2
3a
3b
3c
3d
3e
4a
4b
4c
3f
5a
5b
5c
6a
6b
6c
7a
7b
7c
8a
8b
9 10
11
12a
12b
12c
13a
13b
13c
14a
14b
15
16
17
18
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
2 mm
Stellifer abbreviatus
Stellifer onzole
Stellifer depressifrons
Stellifer bicornutus
Stellifer illecebrosus
Stellifer ephelis
Stellifer chaoi
Stellifer ericymba
Stellifer lanceolatus
Stellifer
stellifer
Stellifer pizarroensis
Stellifer rastrifer
Stellifer
wintersteenorum
Stellifer fuerthii
Stellifer
mancorensis
Stellifer oscitans
72
Paratypes: 2 specimen; 1 specimen NMB P1127, PPP 3467, Río
Santiago, Ecuador, Onzole FM, Messinian; 1 specimen NMB
P361, location NMB 16845, Río Cana, Dominican Republic,
Cercado FM, late Tortonian-Messinian.
Name: Named after the type-formation, the Onzole FM.
The name is used as a noun in apposition.
Diagnosis: OL : OH = 1.55. Anterior rim projecting strong-
est at spine on mid-section above ostium; with less sharp,
short, more backward positioned predorsal spine. Dorsal
rim regularly curved, mildly convex. Ventral rim flat to
slightly concave at mid-section. Ostium moderately sized,
depressed, positioned entirely below ostial-caudal joint.
Cauda curved downwards posteriorly at about 70°.
Description: Moderately elongate, moderately thick otoliths
up to about 6.5 mm length. OH : OT = 1.7. Dorsal rim dis-
tinctly convex, regularly curved, with short concavity behind
short, stubby predorsal spine where a dorsal furrow meets
dorsal rim. Ventral rim flat to slightly concave except for
steeply upturned posterior section. Anterior rim blunt with
two spines, one short predorsal spine situated somewhat
backward and one stronger spine well below mid-section
and above ostium and marking maximum protrusion of
anterior tip; rounded ventral portion with ostium not pro-
truding. Predorsal spine slightly bent outwards, section with
spine at middle section of anterior rim and ventral portion
with ostium strongly bent outwards. Posterior rim with broad,
angular postdorsal projection.
Inner face strongly bent along the horizontal axis with a
curvature index of 20 %. Ostium shallow, small, moderately
short, positioned directly below ostial-caudal joint. Distinct
furrow present above ostial colliculum leading up to cauda.
Cauda very deep, not narrowing towards ostial-caudal
joint, posterior portion of cauda bent downward at about
70°. [The following sulcus proportions measured on views
from inner face, resulting in short values for ostium due to
its strongly outward bent position.] OCL : OCH about 0.75;
CCL : OCL about 2.3; caudal curvature index = 0.6; ostial-
caudal interspace 40 %. No dorsal depression, but distinct
edge or terrace and furrow running close to dorsal rim of
otolith from concavity above ostium where it meets dorsal
rim to near posterior tip of otoliths where it fades, running
at same curvature as dorsal rim and hence not departing
from it; no ventral furrow. Outer face slightly concave, with
small projection on dorsal field underpinning mediodorsal
angle.
Discussion: Stellifer onzole resembles otoliths of the Re-
cent S. illecebrosus, S. fuerthii, S. lanceolatus, S. oscitans
and S. wintersteenorum, but only S. lanceolatus shares the
ventral portion of the anterior rim projecting more than the
predorsal spine and the position of the second spine on the
anterior rim being positioned so much inferior. In these two
characters S. onzole also resembles the fossil S. bicornutus,
but differs from both in the more strongly bent dorsal rim
and the dorsal furrow not departing from its close course to
the dorsal rim. From S. lanceolatus it further differs in the
anterior-ventral margin being slightly inclined backwards
(vs. projecting as strong as the spine on the middle section
of the anterior rim).
Distribution: Messinian of Ecuador.
Lapilli of Stelliferini
Remarks: Lapilli of the Stelliferinae are almost as large
as their sagitta otoliths and show an amazing degree of
morphological diversification not usually observed in lapilli
otoliths. We therefore assume that their lapilli may also
be identifiable at the species level, given, however, that a
detailed investigation would support our assumption. We
have figured a few Recent specimens as example and for
comparison: Odontoscion xanthops (pl. 18, fig. 8), Ophi-
oscion scierus (pl. 18, fig. 7), Stellifer ephelis (pl. 18, fig. 4),
Stellifer fuerthii (pl. 18, fig. 5) and Stellifer microps (pl. 18,
fig. 6). As can be seen from these few examples, the lapilli
of the Stellifer species are more compressed with a deep
ventral rim when compared to those of Odontoscion and
Ophio scion. However, the lapillus of Stellifer microps shares
the morphology more with Ophioscion than the other Stel-
lifer species again supporting the hypothesis that it should
be separated from Stellifer.
Ophioscion sp. (lapillus)
(Plate 18, Fig. 3)
Material: 1 specimen PPP 1164, NMB P1128, Río
Chico, Darien, Panama, Tuira FM, early Tortonian.
Discussion: A slightly eroded specimen that resembles well
the Recent specimen of O. scierus figured for comparison.
Stellifer sp. (lapillus)
(Plate 18, Figs. 1-2)
Material: 2 specimens PPP 1726, NMB P1129-1130, Río Banano,
Costa Rica, Río Banano FM, late Pliocene.
Discussion: Two very distinctive and large lapilli character-
ized as Stellifer lapilli by their compressed shape, outline
and utricular sulcus. They differ from the known Stellifer
lapilli in the very slender predorsal process, the blunt pos-
terior rim and the rather intense marginal ornamentation
on the outer face.
Plate 18
Figs. 1-2. Stellifer sp. lapilli; Costa Rica, Rio Banano FM, late
Pliocene.
Fig. 3. Ophioscion sp. lapillus; Panama, Darien, Tuira FM, late
Miocene.
Fig. 4. Stellifer ephelis CHIRICHIGNO, 1974; lapillus; coll. MNHN,
off Peru, Recent.
Fig. 5. Stellifer fuerthii (STEINDACHNER, 1876); lapillus; coll.
Aguirre, off Ecuador, Recent.
Fig. 6. Stellifer microps (STEINDACHNER, 1864); lapillus; coll.
MNHN, Atlantic off Colombia, Recent.
Fig. 7. Ophioscion scierus (JORDAN & GILBERT, 1884); lapillus;
coll. WS, Pacific off Mexico, Recent.
Fig. 8. Odontoscion xanthops GILBERT, 1898; lapillus; coll. Agu-
irre, off Ecuador, Recent.
Figs. 9-12. Xenotolithus retrolobatus AGUILERA & SCHWAR-
ZHANS, 2014. 9, holotype, MPEG-1811-V; 10-12, paratypes,
MPEG-1812-V; Brazil, Atalaia beach, Pirabas FM, early Miocene.
Figs. 13-16. Xenotolithus sasakii SCHWARZHANS, 1993. 14, 15,
paratypes, NMB P 1202-1203; 13, 16, NMB P1197-1198; Trini-
dad, Manzanilla FM, late Miocene.
Fig. 17. Xenotolithus semiostialis n. sp.; holotype, NMB P1131,
Trinidad, Brasso FM, early
-
middle Miocene.
Figs. 18-19. Xenotolithus sineostialis n. sp.; 18, holotype, NMB
P1132, Trinidad, Manzanilla FM, late Miocene; 19 (r), paratype,
NMB P666, Trinidad, Gros Morne FM, early Pliocene.
/
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
73
Palaeo Ichthyologica 14
1 mm
1 mm
1 mm
1 mm
Xenotolithus semiostialis
Xenotolithus sineostialis
Xenotolithus sasakii
Xenotolithus retrolobatus
1a
1b
1c
2
3a
3b
4a
4b
5a
5b
6a
2 mm
Stellifer sp.
Ophioscion sp.
Stellifer ephelis
Stellifer fuerthii
Stellifer microps
Ophioscion scierus
19a
2 mm
6b
7a
7b
8a
8b
9a
9b
9c
10
11
12
15a
15b
18c
18d
16a
16b
17a
17b
17c
19b
18a
18b
9d
13a
13b
14
2 mm
Odontoscion
xanthops
74
Tribe indet. of Stelliferinae
Genus
†
Xenotolithus
SCHWARZHANS, 1993
Remarks: When Schwarzhans (1993) established the
fossil otolith-based genus
†
Xenotolithus he felt not abso-
lutely certain that it should represent indeed a sciaenid,
because its sulcus morphology is so aberrant. The type
species, Xenotolithus sasakii was the only one then known
and it is remarkable for its very wide and extremely short
ostium without postostial lobe, the anterior portion of the
cauda being bent upwards and the posterior portion of the
cauda being directed downward at a relatively low angle
and being straight, though somewhat widened. In 2014 a
second species was described by A
guilera & Schwar-
zhans as Xenotolithus retrolobatus showing a more typical
sciaenid ostium which is a little wider and shows a distinct
postostial lobe. Therefore, it is now certain that we are
dealing with an extinct sciaenid lineage probably within or
related to the Stelliferinae and characterized by a highly
autapomorphic sulcus pattern. Now, two more species are
being described in the following – Xenotolithus semiostialis
n. sp. and Xenotolithus sineostialis n. sp. – which have an
even more reduced ostium than X. sasakii.
†
Xenotolithus retrolobatus
AGUILERA & SCHWARZHANS, 2014
(Plate 18, Figs. 9-12)
2014
†
Xenotolithus retrolobatus – AGUILERA & SCHWARZ-
HANS: figs. 11.8-11.15
Material: 180 specimens (including holotype and paratypes)
Atalaia beach, Brazil, Aquitanian to early Burdigalian, MPEG-
1811-1813-V.
Description: Moderately elongate, rather thin otoliths up
to about 10 mm length. OL : OH = 1.5-1.7; OH : OT about
2.0. Dorsal rim shallow anteriorly and gently curved, poste-
riorly considerably bending downwards; ventral rim deeper,
regularly curved anteriorly, posteriorly almost straight as-
cending, smooth; anterior rim broadly rounded; posterior
tip sharply pointed, shifted towards ventral.
Inner face moderately convex with distinctly supramedian
sulcus. Ostium short and compressed, wide, particularly
ventrally, with well developed distinctly backward extended
postostial lobe. Cauda narrow and deepened, anterior
portion concave, bending upward, then bedding downward
in an obtuse angle of about 120-130° into a long, more
deepened, straight, fusiform shaped, inclined longer stretch
of cauda running parallel to posterior part of dorsal rim.
OCL : OCH = 0.55-0.75; CCL : OCL = 3.0- 3.5; caudal
curvature index about 0.65-0.75; ostial-caudal interspace
measure to mid-section of downturned cauda as fixed point
is about 50 %. Dorsal depression small, only above anterior,
concave part of cauda; no ventral furrow. Outer face flat
to slightly convex, smooth.
Discussion: The well developed postostial lobe characteriz-
es this species as the most plesiomorphic within Xenotolithus.
Distribution: Aquitanian to early Burdigalian of Brazil.
†
Xenotolithus sasakii SCHWARZHANS, 1993
(Plate 18, Figs. 13-16)
1993 Xenotolithus sasakii – SCHWARZHANS: figs. 403-406
Material: 11 specimens (and previously investigated holotype and
paratypes); PPP 2667, 2670 (figured specimens NMB P1197-
1198), PPP 2674, Cb. 1633 (paratypes, NMB P1202-1203)
San José River, SM 1399 (paratype, NMB P663) Pointe-à-Pierre,
Trinidad, Manzanilla FM, San José Mb, late Tortonian.
Description: Moderately elongate, rather thin otoliths up
to about 13 mm length. OL : OH = 1.5-1.7 (down to 1.35
in juveniles); OH : OT = 2.5-3.0. Dorsal rim shallow ante-
riorly and gently curved, at about mid-section considerably
bending downwards and thereafter straight; ventral rim
slightly deeper, regularly curved anteriorly, posteriorly almost
straight ascending, smooth; anterior rim broadly rounded to
blunt; posterior tip sharply pointed, shifted towards ventral.
Inner face moderately convex with distinctly supramed-
ian sulcus. Ostium extremely short and compressed, wide,
both dorsally and ventrally, without postostial lobe. Cauda
narrow and deepened, anterior portion concave, bending
upward, then bending downward in a sharp angle of about
90° into a long, more deepened, straight, fusiform shaped,
inclined longer stretch of cauda running parallel to posterior
part of dorsal rim. OCL : OCH = 0.2-0.3; CCL : OCL = 6.0-
7.5; caudal curvature index about 0.55-0.65; ostial-caudal
interspace measure to mid-section of downturned cauda as
fixed point is about 50 %. No dorsal depression visible, but
sharp edge along middorsal rim; no ventral furrow. Outer
face flat, rather smooth.
Ontogeny: Two very small otoliths of slightly more than
2 mm length allow recognition of certain ontogenetic
changes. The small otoliths are considerably more com-
pressed than the large ones from 5 mm length onwards
(OL : OH = 1.35 vs. 1.5-1.7), the downturned part of the
cauda is relatively longer and the ostium not as wide.
Discussion: Xenotolithus sasakii is more advanced than
X. retrolobatus, which is primarily visible in the extremely
short and high ostium without postostial lobe.
Distribution: Late Tortonian of Trinidad.
†
Xenotolithus semiostialis n. sp.
(Plate 18, Fig. 17)
Holotype: Pl. 18, Fig. 17, NMB P1131, PPP 2661, beyond bridge
over Gorge River, Trinidad, Brasso FM, late Burdigalian to
early Langhian.
Name: Referring to the ostium, which appears to be only
half of the usual shape through the lack of a ventral lobe.
Diagnosis: Ostium very short, extending only upwards from
ostial-caudal joint. Anterior part of cauda distinctly concave.
Ventral rim of otolith moderately convex.
Description: Elongate, moderately thick otolith of 6.8 mm
length (posterior tip missing). OH : OT = 1.9. Dorsal rim
shallow and gently curved throughout, posteriorly more
downward oriented and nearly straight; ventral rim mod-
erately curved, particularly anteriorly, smooth; anterior rim
broadly rounded, dorsally pronounced; posterior tip not
preserved in unique holotype.
Inner face markedly convex with distinctly supramedian
sulcus. Ostium short and compressed, extending solely
dorsal of ostial-caudal joint. Cauda narrow and deepened,
anterior portion deeply concave, bending upward, then
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
75
Palaeo Ichthyologica 14
bedding downward in an obtuse angle of about 110° into
a long, more deepened, straight, fusiform shaped, inclined
longer stretch of cauda running parallel to posterior part
of dorsal rim. OCL : OCH = 0.4; CCL : OCL = 6.3; caudal
curvature index about 0.6; ostial-caudal interspace can not
be measured. No dorsal depression visible, but sharp edge
along middorsal rim; no ventral furrow. Outer face convex,
slightly less than inner face, smooth.
Discussion: Xenotolithus semiostialis differs from the
younger X. sasakii in the shape of the ostium, which only
extends upwards of the ostial-caudal joint (vs. extending
upwards and downwards in about equal amounts) and in
being thicker and (probably) more elongate. In the aspect
of the reduction of the ostium X. semiostialis is more ad-
vanced than X. sasakii indicating that at least two lineages
of this peculiar extinct otolith-based sciaenid genus existed
in parallel.
Distribution: Late Burdigalian to Langhian of Trinidad.
†
Xenotolithus sineostialis n. sp.
(Plate 18, Figs. 18-19)
Holotype: Pl. 18, Fig. 18, NMB P1132, PPP 2673, Point Paloma
beach, Trinidad, Manzanilla FM, Montserrat Mb., Messinian.
Paratypes: 3 specimens NMB P666, Trinidad, K 12059, Mogue
River, Gros Morne FM, early Pliocene.
Name: Referring to the ostium, which appears to be almost
missing at first sight.
Diagnosis: OL : OH = 1.75-1.9. Ostium very short and
small, extending only upwards from ostial-caudal joint.
Anterior part of cauda gooseneck-lobate. Ventral rim of
otolith shallow, nearly flat.
Description: Elongate, rather thin otoliths up to about
5.6 mm length. OH : OT = 1.9-2.0. Dorsal rim shallow
and gently curved throughout, posteriorly more downward
oriented, somewhat undulating; ventral rim nearly flat,
only anteriorly curved but not expanded, smooth; anterior
rim broadly and regularly rounded; posterior tip sharply
pointed, shifted towards ventral.
Inner face moderately convex with distinctly supramedian
sulcus. Ostium very short, small, compressed, sometimes
barely visible, extending solely dorsal of ostial-caudal joint.
Cauda narrow and deepened, anterior portion lobate,
gooseneck-shaped, bending upward, then bedding down-
ward in an obtuse angle of about 110-120° into a long,
more deepened, straight, fusiform shaped, inclined longer
stretch of cauda running subparallel to posterior part of
dorsal rim. OCL : OCH = 0.2-0.3; CCL : OCL = 9 to about
15; caudal curvature index about 0.7-0.8; ostial-caudal
interspace can not be measured. No dorsal depression
visible, but sharp edge along middorsal rim; no ventral
furrow. Outer face flat to slightly convex, smooth.
Discussion: Xenotolithus sineostialis clearly shows the
most advanced otolith morphology in this already strongly
aberrant genus characterized by the ventrally completely
reduced, extremely narrow, sometimes barely visible os-
tium, and an anterior portion of the cauda, which shows a
peculiar gooseneck-type curvature and almost touches the
anterior rim of the otolith. It is also more elongate than the
other species, may be except for the incomplete holotype
of X. semiostialis, from which it has derived. Xenotolithus
sineostialis represent the last known species of this enigmatic
and peculiar extinct sciaenid genus.
Distribution: Messinian and early Pliocene of Trinidad.
8. Faunal reconstruction
8.1 Sciaenid otoliths from the Neogene of tropical America –
expressions from a center of diversity
The Sciaenidae are an important fish group inhabiting
temperate to tropical shallow coastal waters and estuaries
throughout the world (SASAKI, 1989). In tropical America,
their otoliths are the most abundant faunal component in
almost any shallow water Neogene sediment and show
an amazing degree of diversification (Text-fig. 6), currently
being represented by 85 species, including 20 Recent spe-
cies recorded as fossil. The highly diverse assemblage from
tropical America covers all major Recent groups of the
region plus one extinct genus group (Xenotolithus group).
Sciaenid otoliths are also common in similar Neogene sedi-
ments of North America (MÜLLER, 1999; HUDDLESTON
& TAKEUCHI, 2006, 2007; TAKEUCHI & HUDDLESTON,
2008a,b) with 32 species, thereof 11 Recent species re-
corded as fossil. The total known record of Neogene sciaenid
otolith-based species from the Americas thus amounts to
117 species as compared to 169 Recent species (and 36
genera) (ESCHMEYER & FONG, 2015) in the same region.
This shows that America is the richest region of the world
in sciaenid biodiversity and that it has been a very rich and
diverse sciaenid evolutionary center since at least Neogene
times. Furthermore, most fossil sciaenid species from the
American Neogene are represented by indigenous Ameri-
can genera, indicating that tropical America has acted as
a major center of sciaenid diversity largely separated from
other potential contemporaneous sciaenid hotspots such as
in the Tethyan, tropical East Atlantic and/or Indo-West Pacific
realms for the last 20 million years at least.
Sciaenid otoliths are also common in the late Eocene
and Oligocene rocks of the US Gulf Coast. MÜLLER (1999),
NOLF (2003) and NOLF & STRINGER (2003) listed a total
of 10 sciaenid otolith-based species. These are all very
basal, plesiomorphic forms more or less resembling spe-
cies of Umbrina and Pachyurus and mostly representing
related extinct genera. This may serve as an indication that
the family Sciaenidae as such is geologically rather young
when compared to other percoids. It must have undergone
a rapid radiation probably during the time span from Eo-
cene to early Miocene. LO et al. (2015) arrived at similar
conclusions and estimated the sciaenid crown group to
76
Text-figure 6. Chronostratigraphic range chart of sciaenid species identified by otoliths.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
77
Palaeo Ichthyologica 14
78
have originated about 27.3 Ma (early late Oligocene) in
tropical America. In order to accommodate for the above
mentioned paleontological records from the area, which
were not fully incorporated in LO et al.’s study, we would
however argue for a somewhat older origin during late
Eocene (37-40 Ma).
The tropical American Sciaenidae have been the
subject of several recent molecular genetic studies that
aimed at assessment of phylogenetic interrelationships and
reconstruction of divergence times (e. g., VINSON et al.
2004, VERGARA-CHEN et al. 2008, SANTOS et al. 2013,
BARBOSA 2014). Evidence from mitochondrial genome
analyses in XU et al. (2014) suggests that the New World
may be the region where sciaenid fishes originated and
from where they dispersed, which is in agreement with
the hypothesis first expressed by SASAKI (1989) and as
recently supported by LO et al. (2015) and, as may be
deduced from above discussion, is also congruent with
our observations on the fossil record of sciaenid otoliths.
XU et al. (2014) also discussed the origin of Sciaenidae,
which following their Bayesian relaxed clock model would
indicate the origination of sciaenids during the late Juras-
sic to early Cretaceous period and postulated the origin of
sciaenids to have occurred 208 Ma, which however would
be in late Triassic. In our view such early origination time
of Sciaenidae is highly unlikely and not supported by fossil
evidence. The suborder Percoidei, the order Perciformes
and the superorder Acanthopterygii all are not being
recorded before Cenomanian, i. e. 100 Ma (PATTERSON,
1993). In fact the time interval of late Triassic and Jurassic
is populated only by stem Teleostei (ARRATIA, 1996) which
are not attributable to any crown teleost groups such as
the Sciaenidae.
8.2 Miocene sciaenid otoliths in South American
as indicators of the paleoenvironment
Estuaries and coastal lagoons have played an important role
for the marine faunal evolution, and coastal physiography
and oceanography are very important forcing elements for
paleobiogeography of fishes adapted to such environments,
like the Sciaenidae. Sciaenidae are particularly adapted to
soft and sandy bottomed near-shore marine and brackish
environments, where they sometimes occur in large shoals,
but are missing from most oceanic islands. Only a few
species and genera are known from deeper shelf environ-
ments. In addition, there are four genera recognized from
the freshwaters of the Americas.
During the early and middle Miocene (23-11 Ma), a
large back-arc basin subsided to the east of the northern
Andean orogeny and developed into the Amazonian mega-
wetland Pebas system (HOORN et al., 2010; VONHOF &
KAANDORP, 2010; WESSELINGH, 2006; WESSELINGH &
SALO, 2006) that opened to the north onto Venezuelan
terrain. The early stages at the opening of the proto-Pebas
during early Miocene are represented by the Cantaure FM,
while the late Miocene Urumaco FM represents the marine
facies at the opening during the Pebas development. The
proto-Pebas and the Pebas systems were the scenes of an
active evolution of wetlands biota that were rich in land,
swamp and freshwater animals and have been the subject
of a great variety of recent research works (see special
editions of Scripta Geologica, no. 133, 2006, ed. WES-
SELINGH; Urumaco & Venezuelan Paleontology, 2010, eds.
SANCHEZ-VILLAGRA, AGUILERA & CARLINI; Amazonia:
Landscape and species evolution, 2010, eds. HOORN &
WESSELINGH). The Pebas system mouth obviously rep-
resented a favorable environment for estuarine adopted
sciaenids, witnessed by the diverse sciaenid assemblage
found in the early Miocene Cantaure and Castilletes FMS
and the late Miocene Urumaco FM (Text-fig. 6). The Castil-
letes FM of Colombia is particularly remarkable for being
composed almost entirely of sciaenid and ariid otoliths,
while the Cantaure FM of northwestern Venezuela largely
contains the same sciaenid species but in a much more
diverse and richer marine teleost fauna. The late Miocene
Urumaco FM is similarly rich and diverse in sciaenids many
of which represent the same lineages already evident in
the Cantaure FM, with the notable exception of the genus
Cynoscion, which occurs in the Urumaco FM for the first
time.
Changes and rearrangements of the hydrographic set-
ting of the Amazon river basin (POTTER, 1997; FIGUEIREDO
et al., 2009; HOORN et al., 1995, 2010; SHEPARD et al
2010) and the Orinoco river mouth (DIAZ DE GAMERO,
1996; ROD, 1981) determined the reconfiguration of the
South American continental hydrographic drainage sys-
tems and the establishment of the giant transcontinental
western Amazon River flux to the Atlantic Ocean during
the late Miocene (FIGUEIREDO et al., 2009). The Amazon
Basin complex in Northeastern South America includes the
Amazon, Orinoco and Esequivo river drainage systems into
the Atlantic Ocean. The Amazon and the Orinoco rivers
developed in response to the initiation of the uplift of the
Central and Northern Andes (HOORN et al., 1995, 2010;
DÍAZ DE GAMERO 1996; POTTER 1997; GARZIONE et
al. 2008; FIGUEIREDO et al., 2009; POTTER & SZATMARI
2009; BERMÚDEZ et al., 2011). Based on lithological
evidence, a major change in the hydrographic basin oc-
curred at about 10 Ma with the onset of the Amazon fan
building out into the Atlantic Ocean (FIGUEIREDO et al.,
2009; HOORN et al. 2010) and subsequently with the
onset of the Orinoco fan at about 5 Ma. Trinidad, located
at the northeastern margin of South America, is character-
ized during the Miocene and Pliocene by a sequence of
progressive siliciclastic deposits related to the freshwater
outflow from the paleo-Orinoco into the paleo-Gulf of Paria
after the calcareous silts of the late Miocene Manzanilla FM,
which were deposited in an inner sublittoral environment.
The sediments of the overlying Pliocene Springvale FM are
composed of silts and sands deposited during a shallow
water marine incursion and coastal swamp environments.
The subsequent Talparo FM consist of sands, sandy clays
and clays deposit in a marginal marine to brackish and
freshwater paleoenvironment (DONOVAN, 1994; NHM,
1996; KUGLER, 2001). Most of the late Miocene otolith
assemblages originate from rocks northwards and east-
wards of the paleo-Amazon delta, i. e. from the Gatun FM
of Panama westwards until the Manzanilla FM of Trinidad.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
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Palaeo Ichthyologica 14
These locations are all rich and diverse in sciaenids, but do
not differ greatly from the association in the Urumaco FM at
the Pebas opening. Therefore, we conclude that there was
a rather uniform sciaenid fauna along the northern shores
of South America during the late Miocene from Panama to
Trinidad.
Along the eastern Pacific shores of southern America,
chiefly in Ecuador, the hydrographic system was dominated
during the Neogene, like today, by the narrow coastal
plain with an intricate web of small rivers, most of them
intermittent, which come from the Andes cordillera. The
main flows are the northwestern Esmeraldas River and
the southwestern Guayas River. Otolith associations in the
late Miocene of Ecuador (Angostura and Onzole FMS) are
similar to the ones recorded from Caribbean shores from
Panama to Trinidad, but already show some indications of
vicariant evolution (see chapter 8.3. for further discussion).
We conclude that the northern shores of South America,
with the associated interior Pebas mega-wetlands and the
large and shifting river drainage systems chiefly of the
paleo-Amazon and paleo-Orinoco, offered a favorable
setting for the evolution of the Sciaenidae. These fishes
preferred estuarine environments, as evidenced by the
rich and diverse otolith-based sciaenid associations found
in the studied shallow marine formations of early to late
Miocene age in the region.
8.3 Tropical American sciaenid lineages that became extinct
during Neogene
The high degree of speciation and diversification of sciaenid
fishes in tropical America during the Neogene is contrasted
by a moderate number of extinctions of individual lineages
in sciaenid evolution that become evident only with the
currently available rich and dense fossil sciaenid otolith
record. Text-figure 7 summarizes our knowledge of sciaenid
otolith-based lineages from tropical America, which have
become extinct. Six extinct otolith-based sciaenid genera
are known so far from the Neogene of tropical America:
†
Amazonasciaena,
†
Frizzellithus,
†
Equetulus,
†
Protonebris,
†
Protolarimus and
†
Xenotolithus, the latter representing an
extinct group of insufficiently resolved relationship. This in-
dicates an early specialization in certain sciaenid tribes with
a wider and more diverse distribution in the late Paleogene
and early Miocene than today. Of these six extinct genera,
three (
†
Amazonasciaena,
†
Frizzellithus and
†
Protonebris)
are last recorded in the late Burdigalian to early Langhian
time interval. The extinction of
†
Amazonasciaena could be
linked to the change from the carbonate environment of
the Pirabas FM in Brazil to the clastic environments of the
Amazon river after its opening eastwards into the Atlantic
was established (AGUILERA et al., 2014). The other extinct
genera are last recorded from the late Miocene (
†
Equetulus
and
†
Protolarimus) and the early Pliocene (
†
Xenotolithus)
respectively. As is depicted in text-fig. 7, the late Miocene
time interval does not show up as one of particular extinction
stress for sciaenids, but rather as an interval with acceler-
ated evolution of these fishes (Text-fig. 6) probably caused
by the extension of favorable environments (Pebas opening,
Amazon delta development). Still, it could have proven to
be less favorable for certain sciaenid fishes adopted to the
less fluvial discharge dominated environments of the early
Miocene.
Apart from the extinct otolith-based genera mentioned
above, we have also observed certain inter-generic lineages
going extinct during latest Miocene and Pliocene. They are
related to the disappearance of lineages related to Callaus
cubaguanus, Polycirrhus trinidadensis, Larimus angosturae,
Isopisthus acer, Plagioscion ultimus and
†
Xenotolithus sine-
ostialis. Two of them went extinct in the Caribbean, but con-
tinued in the East Pacific (Callaus deliciosa vs. C. cubaguanus
and Polycirrhus dumerilii vs. P. trinidadensis). There are two
more lineages that disappeared earlier in the Neogene of the
Caribbean but persisted until nowadays in the East Pacific,
namely Paranebris bauchotae known as a fossil from the
early Tortonian of Darien, Panama and the Messinian of
Ecuador, and Ophioscion transitivus from the Burdigalian
to Langhian of Trinidad and the Dominican Republic and
related to the extant eastern Pacific O. vermicularis. Three
of the four lineages recognized to have become extinct in
the Caribbean but persisting in the East Pacific are thought
to represent plesiomorphic, primitive members within their
respective tribes (Polycirrhus dumerilii, Paranebris bauchotae
and Ophioscion vermicularis). Also three of the genera to
which they belong are nowadays restricted to the Eastern
Pacific (Callaus, Polycirrhus and Paranebris) and the fourth
has occasionally been reported as representing a mono-
specific genus of its own (Sigmurus GILBERT 1898) likewise
restricted to the East Pacific (SCHWARZHANS, 1993).
Another interesting aspect that emerges when studying
extinct sciaenid lineages at the species level as explained
above, is that many of them become extinct in the Carib-
bean during a narrow time interval of the late Pliocene.
These are Callaus cubaguanus, Larimus angosturae, Iso-
pisthus acer and Plagioscion ultimus, while only one line-
age is last recorded in the early Pliocene –
†
Xenotolithus
sineostialis. One of these lineages, Larimus angosturae, was
apparently amphi-American during the late Miocene, but
became extinct in the East Pacific prior to its extinction in
the Caribbean, while all the others were strictly Caribbean
in occurrence or have an extant eastern Pacific vicariant
species counterpart (Callaus cubaguanus vs. C. deliciosa).
Our observations of Caribbean sciaenid otoliths confirm
the observations made by O’DEA et al. (2007: ‘Environ-
mental change preceded Caribbean extinction by 2 million
years’), according to which the main Caribbean extinction
event occurred 1.5 to 2 million years after the final uplift
of the Isthmus of Panama that has triggered the ‘Great
American Biotic Interchange’ (GABI) of land animals
(MARSHALL, 1988; WOODBURNE, 2010). The uplift of
the Central American land bridge (COATES & STALLAERD,
2013) interrupted the Pacific-Atlantic seaway and has driven
large-scale ocean circulation changes (STEPH et al., 2006;
SCHNEIDER & SCHMITTER, 2006; BUTZIN et al., 2011)
and produced environmental conditions that distinguishes
todays Pacific habitats and are characterized by productive
surface waters caused by coastal upwelling and abundant,
fast-growing suspension and detritus feeders on the sea
80
Event log
Text-figure 7. Stratigraphic range and evolution of selected sciaenid phylogenetic lineages as deduced from otoliths with particular
emphasis on extinction times. Blue = East Pacific records, orange = Caribbean records. Events depicted in the event log are based
on ROSETTI et al., 2013; FIGUEIREDO et al., 2009; VONHOF, & KAANDORP, 2010; COATES et al., 1992; WOODBURNE, 2010;
O’DEA et al., 2007.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
81
Palaeo Ichthyologica 14
Event log
82
† Equetulus
Polycirrhus
trinidadensis
Text-figure 8. Distribution of selected amphi-American otolith-based sciaenid species in tropi-
cal America during. Upper row = middle Miocene, light blue = Pebas paralic basin, medium
blue = Caloosahatchian bioprovince, dark blue = Garunian / Caribbean bioprovince. Lower
row = late Miocene, medium blue = Caloosahatchian bioprovince, dark blue = Garunian / Carib-
bean bioprovince, blue-green = central Eastern Pacific bioprovince. Paleogeaographic reconstruc-
tions are primarily based on COATES et al., 2003; COATES et al., 2004; ITTURALDE-VINENT &
MACPHEE, 1999; LANDAU et al., 2009; PINDELL & KENNAN, 2009; PINDELL & TARBUTT, 1995
and geological maps of the region.
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
83
Palaeo Ichthyologica 14
bottom. In contrast, the Caribbean habitats have become
progressively richer in carbonate, more nutrient-poor, and
with large coral reefs in shallow water, intercepted by ma-
jor, large river mouths with high freshwater discharge and
expanded estuarine environments (O’DEA et al., 2007;
LEIGH et al., 2013). However, the reason for the observed
delay in the perceived reaction of the marine biota to the
interception of the Pacific-Atlantic seaway and the subse-
quent environmental changes which were forced particularly
in the Caribbean remains unresolved and is the subject of
ongoing discussions (O’DEA et al., 2007).
A further aspect of sciaenid evolution that has emerged
from this and previous related studies (NOLF & STRINGER,
1992; AGUILERA & RODRIGUES DE AGUILERA, 2003) is the
presence in the marine Neogene of several species belong-
ing to genera that nowadays are restricted to freshwater,
i. e. Aplodinotus and Plagioscion. Aplodinotus nowadays is a
monospecific genus in the freshwaters of northern America,
while in the early Miocene it contained species in the near
shore marine environments of Venezuela and Brazil. Plagi-
oscion nowadays is a diverse genus in the tropical freshwater
schemes of South America with five valid extant species
(CASATTI, 2005). The fossil otolith-based record shows a rich
representation of the genus in marginal marine to estuarine
environments of the Miocene and Pliocene, representing
possibly three different individual lineages: P. travassosi
from the early Miocene of Brazil, P. urumacoensis, a giant
otolith-based species from the late Miocene Urumaco FM
of Venezuela at the Pebas opening and very similar to the
extant P. squamosissimus of the Amazon river scheme, and
the P. marinus – P. ultimus lineage reaching from early Mio-
cene to late Pliocene. Plagioscion ultimus represents the last
truly marine species of the genus and is known from near
the paleo-Orinoco river mouth in Trinidad, and, interest-
ingly, went extinct at about the same time during the late
Pliocene extinction event as discussed above. Plagioscion
ultimus appears to have been a dwarfed species, possible
indicating evolutionary stress prior to extinction.
We conclude that the highest extinction rate of line-
ages occurred in the Caribbean and appear to either
have been related to shifts and expansions of the large
river discharge systems during late Miocene or to a yet
insufficiently understood extinction event at the end of
the late Pliocene, about 1.5 to 2 million years after the
closure of the Isthmus of Panama (and the GABI). Cer-
tain sciaenid genera nowadays restricted to American
freshwater systems, like Aplodinotus and Plagioscion,
have representatives in marine Neogene strata of tropical
America indicating their marine origination.
8.4 Amphi-American sciaenid species and geminate sciaenid species pairs
The current consensus for the timing of the final closure of
the Pacific-Atlantic seaway by the uplift of the Isthmus of
Panama is between 3.5 and 3.2 Ma (COATES et al., 1992;
WOODBURNE, 2010). The period of time since then is
rather short at the evolutionary scale and many species
did not diverge much. For this reason many sciaenids were
considered as being the same on both sides of the isthmus,
until fine anatomical studies or, more recently, genetic
studies revealed that this was not the case. According to
current understanding there are no extant sciaenid species
distributed on either side of the Isthmus of Panama (FROESE
& PAULY, 2015).
Obviously, the distribution of amphi-American spe-
cies in the past would offer some means for narrowing of
time constraints of the faunal separation. However, such
investigations are hampered by two effects: the lesser
knowledge of fossil marine faunas from the Pacific shores
of South America as compared to the Caribbean shores
and the similarity of the otoliths of the species involved
requiring very good preservation to facilitate observations
at that level of resolution, which is only the case in a few
instances summarized in text-figure 8. One of the most
common and widespread genera is
†
Equetulus known from
three species in the early and late Miocene. Two species,
E. fitchi and E. amazonensis are only known to the east of
the Pebas opening in Venezuela, while E. davidandrewi oc-
curs in Ecuador, Panama and western Venezuela. Another
species of wide regional distribution during the early late
Miocene (Tortonian) is Polycirrhus trinidadensis recorded all
the way from Ecuador, Panama, Venezuela and Trinidad,
but is nowhere very common. Larimus gatunensis is known
from the early late Miocene (Tortonian) of Ecuador, Panama
and Trinidad and a second Larimus species, L. pandus,
from the same time interval of Ecuador and Panama. In
younger, latest Miocene strata (Messinian) only two species
are known with an amphi-American distribution pattern,
Ophioscion amphiamericanus and Stellifer onzole, both from
Ecuador and the Dominican Republic. They represent the
youngest amphi-American records in sciaenid otoliths, but
this statement remains inconclusive for the time being due
to the lack of otoliths studied from Pliocene formations of
the Pacific realms of South America. Faunas studied from
the early Pleistocene of Ecuador and Pacific Panama do not
show any common species with those from the Caribbean
(COATES et al., 1992, JACKSON & O’DEA, 2013).
Jordan was long ago conscious of the fact that fishes
from the Pacific and the Atlantic shores of America were
different (JORDAN, 1908) and coined the term »geminate
Geminate sciaenid species of tropical America
East Pacific West Atlantic / Caribbean
Ctenosciaena peruviana Ctenosciaena gracilicirrhus
Umbrina xanti Umbrina broussonnetii
Pareques lanfeari Pareques acuminatus
Menticirrhus elongatus Menticirrhus littoralis
Larimus argenteus Larimus breviceps
Larimus pacificus Larimus fasciatus
Nebris occidentalis Nebris microps
Cynoscion albus Cynoscion acoupa
Cynoscion phoxocephalus Cynoscion leiarchus
Cynoscion reticulatus Cynoscion nothus
Isopisthus remifer Isopisthus parvipinnis
Macrodon mordax Macrodon ancylodon
Bairdiella armata Bairdiella ronchus
Corvula macrops Corvula batabana
Odontoscion xanthops Odontoscion dentex
Text-figure 9. List of geminate sciaenid species of tropical America.
Shaded are species with fossil otolith records.
84
species« to characterize those species that were apparently
very similar on both sides. Sciaenids are one of the most spe-
ciose taxa in the area, making the situation more confusing,
however several members of the family can be considered
as geminate species. It is difficult to ensure that the species
divergence observed is due to the isthmus barrier erection,
however we here consider, on the basis of morphometric
data, otolith shape and phylogenetic studies when available
(e. g. VERGARA-CHEN et al., 2009), that a shortlist of pairs
depicted in text-figure 9 are geminate species. We assume
that there are many more geminate sciaenid species within
the more speciose genera such as Stellifer or Ophioscion,
but these will require more detailed assessment, which would
be beyond the scope of this study. In text-figure 10 we have
Text-figure 10. Timing and evolution of certain geminate sciaenid species as deduced from otoliths. Blue = East Pacific records, or-
ange = Caribbean records. Timing of final closure of Isthmus of Panama following COATES et al., 1992 and WOODBURNE, 2010.
peruviana
gracilicirrhus
mexicana
deliciosa
cubaguanus
acer
††
remifer
parvipinnis
Ctenosciaena
Callaus
Isopisthus
occidentalis
microps
pacificus
fasciatus
pandus
dioneae
†
argenteus
breviceps
Nebris
Larimus
final closure of Isthmus of Panama
angosturae
gatunensis
angosturae
final closure of Isthmus of Panama
Pleistocene
Pleistocene
AguilerA, SCHWARZHANS & BÉAREZ: Otoliths of the Sciaenidae from the Neogene of tropical America
85
Palaeo Ichthyologica 14
visualized the origination of certain geminate pairs that are
well constrained in the Recent and that have a reasonable
fossil record. These refer to the three genera Ctenosciaena,
Isopisthus and Nebris, which each yield two extant species,
one on the Pacific the other on the Atlantic side of the
Americas, plus the genus Callaus, nowadays monospecific
on the Pacific side, but with an extinct geminate species in
the Caribbean, and two distinct lineages within the genus
Larimus, again each with an Atlantic and a Pacific species.
Notwithstanding the lack of evidence from the Pliocene of
the Pacific side it is readily evident that the stratigraphic
range of the geminate Atlantic species (Ctenosciaena gra-
cilicirrhus, Larimus fasciatus and Larimus breviceps) predate
the assumed final date of the closure of the Isthmus of
Panama by 2 to 5 Ma. The most recent common ancestors
(MRCA) relevant for four dichotomies of geminate species
(Ctenosciaena mexicana, Nebris dioneae, Larimus pandus
and Larimus gatunensis) all do not occur after the Torto-
nian, i. e. at least 4 Ma prior to the uplift of the Isthmus
of Panama. In one case, the MRCA and the Caribbean
geminate species existed for a short time span in parallel
on the Atlantic side (Larimus gatunensis and L. breviceps);
two other instances show extinct related species in addition
to the extant geminate pairs (Isopisthus acer and Larimus
angosturae), both becoming extinct after the late Pliocene
and one of them previously during Tortonian also having
been distributed on the Pacific side (Larimus angosturae).
Recently, the distribution of certain heavy mineral com-
positions from borehole and surface samples in northern
Colombia have been interpreted as indicating that the
closure of the Central American Seaway had occurred 10 to
13 million years earlier than current consensus, i.e. during
middle Miocene 13 to 15 Ma (MONTES et al. 2015). The
presence of amphi-American sciaenid species and common
ancestors of extant geminate species up to at least about
6 Ma however demonstrates that an active exchange of
marine biota must have existed during late Miocene.
We conclude from the stratigraphic distribution of the
youngest amphi-American species as well as the earli-
est occurrence of geminate species that the separation
of geminate eastern Pacific and Caribbean lineages in
sciaenids predates the final closure of the Pacific-Atlantic
seaway and the uplift of the Isthmus of Panama by
2 to 5 Ma, and indicates that the marine biota were then
already separated at a sufficient, possibly environmen-
tally induced level that triggered a geminate speciation.
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In der Reihe
Palaeo
Ichthyologica
erscheinen
zoologische und
paläontologische
Arbeiten zur
Systematik
Morphologie
Palökologie
Paläogeographie
Stratigraphie
der
Fische.
ISSN 0724-6331
ISBN 978-3-89937-207-6
