Pliocene) of Jamaica
Contr. Tert. Quatern. Geol. 35(1-4) 147-160 1 fig., 2 tabs, 4 pis. Leiden, April 1998
Otolith-based fishes from
the Bowden shell bed (Pliocene) of Jamaica:
systematics and palaeoecology
Gary Layne+Stringer
NORTHEASTLOUISIANA UNIVERSITY
MONROE, LOUISIANA, U.S.A.
Stringer, Gary Layne. Otolith-based fishes from the Bowden shell bed (Pliocene) ofJamaica: systematics and palaeoecology. In: Donovan, S.K.
2 4 (ed.). The Pliocene Bowden shell bed, southeast Jamaica. — Contr. Tert. Quatern. Geol., 35(1-4): 147-160,1 fig., tabs, pis. Leiden, April
1998.
otolith studies have the to substantial Investigations of fossil teleosteanotoliths from the Caribbean are quite restricted, although potential provide
shell bed, Bowden are described herein information on the palaeontology of the region. The fish otoliths from the Bowden Formation, Jamaica,
in order to ascertain the ichthyological fauna and related palaeoecological conditions. Previous investigations of otoliths for the Caribbean are
addressed and support the importance ofthis study. Approximately 1,650otoliths representing at least 38 teleost families and 68 species were obtained from the shell bed at the type locality of the Pliocene Bowden shell bed. Comparison of the identifiedotoliths to the bathymetric
distributions of closely related Recent taxa revealed a diversifiedassociation with shallow-water marine forms (including euryhaline species),
and shelf forms with and elements. The also contained forms neritic species, middle to outer some upper slope pelagic otolith assemblage
commonly associated with reef environments. The otolith assemblage of the Bowden shell bed is quite similar to the Recent Caribbean
ichthyological fauna with some notable exceptions (three Pacific forms).
Key words — Bowden shell bed, otoliths, fossil fishes, otolith morphology, systematics, palaeoecology.
G.L. Stringer, Department ofGeosciences, Northeast Louisiana University, Monroe, Louisiana 71209, U.S.A.
Contents can contribute untold and misleading numbers of skeletal
elements to the thanatocoenosis. Since an individual fish can
Introduction and contribute two saccular otoliths to the fossil record, the previous investigations p. 147 only
Material and methods 148 of otoliths is a more precise indicator of abundance p. presence and identificationof otoliths 148 (Stringer, 1986). Morphology p. studies from the Caribbean that include Systematic palaeontology p. 149 Palaeontological
teleostean otoliths limitedand detailed Palaeoecology based on otoliths p. 151 are quite descriptions
Comparisons to otherotolith faunas of the of fossil otolith assemblages are almost nonexistent. This is
especially true for Jamaica, despite the of thick Caribbean p. 153 presence
of Cretaceous to Quaternary marinesedimentary Conclusions p. 154 sequences
154 rocks (Purdy et al., 1996). Some such as the molluscs Acknowledgements p. groups,
References and other invertebrates, are well documented from the Plio- p. 154
cene Bowden shell bed in Jamaica (Donovan et al., 1995).
of otolith studies in the However, a survey Tertiary Carib-
Introduction and previous investigations bean Faunal Province clearly illustrates the lack of extensive
research in this region. Fitch & Barker (1968) mentionedthe
Fossil teleosteanotoliths have been utilised in a very limited occurrence of a morid otolithfrom Jamaica, and Fitch (1969)
manner in the investigation of the palaeontology of the noted that the Bowden shell bed in Jamaica containedotoliths
Caribbean region, although fossil otoliths have the potential of lanternfish(family Myctophidae).
to contribute meaningful informationon the fish fauna and Clarke & Fitch (1979), in a study of Cenozoic teuthoid palaeoecology. The use of fossil teleosteanotoliths provides (cephalopod) statoliths, reported that a 225 kg sample from
a more detailed and accurate representation of the ichthy- the Bowden shell bed in Jamaica yielded 25,OCX) otoliths.
fauna ological than the exclusive use of skeletal components Fitch (1969) noted that the otoliths probably represented
such as teeth, scales, spines and vertebrae (Nolf, 1985; more than 110 kinds of fish belonging to 50-60 families.
Breard & Stringer, 1995). Otoliths can also provide a more Some of the forms mentioned by Fitch included bonefish accurate interpretation of abundance. Fishes, sharks and rays (Albula), croakers (iMicropogon, Larimus and Equetus), 148
soldierfish (Myripristis and Holocentrus), cardinalfish(Apo- processed by the Vokes in obtaining the otoliths. Otoliths Florida gon), jawfish(Opisthognathus), pearlfish (Carapus), her- were also provided by Roger Portell of the Museum The rings, anchovies, gobies, goatfish, gurnards, cuskeels, catfish, ofNatural History (University of Florida, Gainesville).
forms. Fitch otoliths obtained from codlets and a few deeper-water Unfortunately, were a sample weighing approxi-
10 A small of matrix 2 never formally describedthe otoliths from the Bowden shell mately kg. sample (approximately kg)
Dr Jon ofOkaloosa-Walton Commu- bed. Upon his death, the otoliths were placed in the John E. was supplied by Bryan The Bowden shell bed Fitch Collection of Research Fossil Otoliths housed at the nity College (Fort Walton, Florida).
Institut des Sciences Los Angeles County Museum of Natural History (Robert otolith collection from the royal
ofthe data naturelles de Belgique (IRScNB, Brussels) was examined on Lavenberg, pers. comm.). A check pre-catalogue
Dirk Nolf. addition to the material at the Los Angeles County Museum revealed that the vast a loan supplied by Dr In
bed otolith material identi- described examination of the Bowden majority of the Bowden shell was above, a preliminary
data indicated shellbed material of the late John Fitch was made in 1984. fied only to family level. The pre-catalogue a
the material of Dr Fitch could not be total of only 4,946 otoliths, not the 25,000reported by Clarke Unfortunately, re-
examined for this & Fitch (1979). In addition, the inventory indicated 89 taxa study.
kinds 110 It The need for bulk for otolith is or of otoliths, not the taxa previously reported. sampling investigations
illustrated when the number of otolithsand is is uncertain whether the collection is the same, but, if it is, clearly species
for the various The 10 yielded there is a discrepancy of over 20,000 otoliths. compared samples. kg sample
An assemblage consisting of 2,074 otoliths from the over 200 otoliths and 30 species, while the 2 kg sample
middle Miocene Gatun Formation of Panamawas described yielded about 30 otoliths and 14 species. The 70 kg sample
the in Gillette (1984). The assemblage consisted of 26 taxa of produced over 1,200otoliths and 55 taxa. By comparison,
marine fishes identifiedby John Fitch. Unfortunately, Fitch largest sample in the study of the Neogene otoliths from the
Dominican Nolf & supplied only a list of the otoliths and was not able to com- Republic by Stringer (1992) was ap- the otoliths and the plete the study before his death. Gillette presented the list proximately 120kg which yielded most
number of In both of these exactly as provided by Fitch, but did add some annotations, largest species (62 species).
studies well definite correlation exists and also noted that the specimens were unavailable for as as many others, a size and the number of renewed study. A few other studies have mentioned the between sample species recognised.
from The wet-screened a of sieve occurrence of otoliths, including Schubert (1909) samples were using variety mesh Standard will Panama, Leriche (1938) from Colombia, Weiler (1959) from sizes. Sieves from 30 to 40 (U.S. Sieves) retain almost all otoliths. Sieve sizes Mexico and Casier (1966) from Barbados. Stringer (1992b) species represented by
otoliths from the Bowden shell bed than 25 mesh lose fish with small oto- published an abstract on greater may species
in Jamaica and noted at least 55 taxa of teleosts. liths and should be avoided. Otoliths for this study were
examined a binocular stereozoom with There are only two comprehensive studies of fossil using microscope
otoliths from the entire Caribbean region (Nolf, 1976; Nolf 10x-20x eyepieces with variable power (lx-2.5x). Generally,
& Stringer, 1992). Nolf (1976) described 66 otolith-based the otoliths from the Bowden shell bed are well preserved,
and in size from less than 1 to 8 teleost taxa from the Neogene of Trinidad. Most of the taxa mostly complete, range mm
in came from various Miocene formations, but there were a few length.
from the Pliocene. The fauna consisted mainly of neritic
forms, together with a few mesopelagic elements and demer-
MORPHOLOGY IDENTIFICATION OF OTOLITHS sal (bottom dwelling) fishes from the upper slope. Nolf & AND
Stringer (1992) reported a highly diversifiedteleost fauna of
84 species for the Neogene of the Cibao Valley area, north- To utilise otoliths to reconstruct fossil fish faunas and to
the ern Dominican Republic. The study included numerous determinepalaeoecological conditions, identificationof
collecting localities from four Miocene-Pliocene units (Bai- otolith is essential. Salientmorphological features are used to
toa, Cercado, Gurabo and Mao Formations). Palaeoenviron- identify the otoliths. Although most bony fishes contain three
and mental conditions based on the present-day bathymetric otoliths in each of the two labyrinths (the sagitta, lapillus
distribution of related teleost taxa were determinedfor each asteriscus), the sagitta is used for identification purposes in
the of the formations. the majority of cases. The sagitta tends to be the largest of
face. otoliths and has a characteristic pattern on the inner
This pattern is usually indicative of the family, and in many
MATERIAL AND METHODS examples the genus and species (Nolf, 1985).
The innerface of the sagitta has numerous salient mor-
features. The inner face is the side The Bowden otoliths for this study were supplied by several phological inward-facing
otoliths of the that is connected and individualsand institutions. The largest percentage of sagitta to nerve endings usually
has characteristic it known the sulcus was provided by Drs Harold and Emily Vokes of Tulane a pattern on as (Fig. 1). the University, New Orleans (Louisiana). Much of their material The sulcus is usually the most conspicuous feature on
from the innerface of the The sulcus be undividedin the was obtained during the collectionof molluscs type sagitta. may other locality. It is estimated that at least 70 kg of sample was otoliths of some fishes and divided into two parts in 149
species. If the sulcus is divided, the anterior portion is known additionto modernteleosteanotoliths, comparisons were also
the Domini- as the ostium, while the posterior portion is known as the made to fossil Miocene-Plioceneotoliths from cauda. can Republic and Trinidad for this study.
Fossil otoliths should be assigned to living taxa where
possible. This requires not only excellent Recent comparative
collections, but also well-preserved fossil material. The
identification of fossil otoliths can be limited by several
conditionof the conditions. Sometimes, the preservation or
the fossil otolith may limit identification. In some cases,
factor. variability within a species may be a restricting
also be restricted due the limited Otolithidentificationmay to
knowledge of Recent forms. For example, in this study, the
knowledge of Recent gobiid otoliths from the Caribbean is
too inadequate to identify the fossil gobiids at the generic or
species level. Therefore, the gobiids are identified as form
groups withinthe family Gobiidae. Another problem related
to fossil otolith identification has recently been identified by
otoliths of Schwarzhans (1994), who found that the some
dimor- species of fish (Recent and fossil) may exhibit sexual
phism. However, this sexual dimorphism of otoliths has only
in of the Fig. 1. General morphology ofthe saccular otolith (sagitta). been clearly recorded one family teleosts, Ophidii-
numbers of otoliths dae. Finally, in some species, large
Other features of the described needed for prominent sagitta are by representing all stages of growth are species referring to the margins. The dorsal, anterior, ventral and determination.As noted by Nolf & Stringer (1992), extensive posterior margins can be recognised on the sagitta, although series of large otoliths are often required to identify good
unclear. from the their exact boundaries may be Projections species-diagnostic criteria. This is illustrated by Diaphus
otolith named the periphery of the are according to margin on otoliths in which only adultotoliths larger than 2 mm show
dorsal dome is which they are located. Therefore, a a projec- characteristic features (Brzobohaty & Nolf, 1995). Some tion from the dorsal margin. The rostrum and antirostrum are species have otoliths with such generalised morphology that
anterior A rim located above prominent projections. ridge or even large specimens lack diagnostic features. the sulcus is the crista and below the sulcus taxonomic superior a ridge Extinct genera and species may also present is the crista inferior. A above the sulcus is referred otoliths extinct forms and do depression problems. Some represent not of the to as the area. Colliculum deposits (raised portions have living representatives. This problem has been ap-
Cauda referred sulcus floor) in the ostium and are to as proached in several different ways. One method is to desig- anteriorand posterior colliculum, respectively. nate a fossil genus which is commonly doneby palaeontolo-
the otolithis another The shape or outline of morphologi- gists working in other groups. However, the presumed
used for taxonomie Geometric terms otoliths cal feature purposes. 'extinct' form may be a Recent genus of which the such and often utilised of as circular, elliptical subtriangular are are unknown. Considering the imperfect knowledge
describe otolith In addition the of the ifworkers do to shapes. to shape Recent fish otoliths, this is a distinct possibility otolith, the of the also known as fossil otoliths to shape margin, sculpturing, not take great care to extensively compare
be be entire lobed, Another method, Nolf is may diagnostic. Margins may (smooth), Recent genera. proposed by (1985), Otolith sinuate, crenulate, denticulate, serrate or irregular. the use of plural genitive names. In this method, species margin sculpturing, as well as many other morphological which do not fit in known Recent generaare cited by plural
described in detail Smale features, were illustratedand by et genitive names, preceded by the word 'genus'. The method
various al. (1995). Morphological features of otoliths from proposed by Nolf is commonly utilised by many otolith families (congrids, myripristids and albulids) were illustrated palaeontologists, especially those in Europe (Steurbaut, 1984; in Stringer (1979). Brzobohaty, 1986; Nolf & Cappetta, 1989; Radwanska,
determined The taxonomie position of a fossil otolith is 1992). by comparison to otoliths of identified Recent fishes or by comparison with in situ otoliths in identified fossil fish
PALAEONTOLOGY skeletons. Since fossil fish skeletons with in situ otoliths are SYSTEMATIC extremely rare, the vast majority offossil otolith identifica-
otoliths at least 38 teleost tion is based on a comparison with otoliths of Recent forms. Approximately 1,650 representing familiesand obtained from the shell bed of As noted by Nolf (1985), the precision of fossil otolith 68 species were
the Bowden Formation. All of the identifications of this identification will depend largely upon the knowledge of
based the or the saccular otoliths Recent otoliths. Several museums have Recent comparative investigation are on sagittae
of the teleost for the where the collections with 2,(XX) to 6,(XX) species of fish otoliths. In except family Plotosidae, 150
lapillus or utricular otolith is utilised for identificationof the this study follows that of Nelson (1984, 1994). A list of all
Plotosus. the catfish In catfish order Siluriformes, the lapillus otolith-based teleosts identifiedfrom the type locality of the
is the largest and most diagnostic of the otoliths, and is Bowden shellbed in Jamaica is presented in Table 1.
utilisedfor taxonomic purposes. The classification scheme of
Family Taxon Plate and figure number
Albulidae Albula sp.sp. PI. 1, Fig. 1 [Nolf&[Nolf & Stringer, 1992,1992,pi.pi. 9, figsfigs 1, 2)2]
Heterenchelydae PythonichthysPythonichthys sp. PI. 1,Fig. 2 [Nolf & Stringer, 1992,1992,pi.pi. 9, figsfigs 8-10]
Congridae AriosomaAriosoma balearica Pi.PI. 1, Fig. 3 [Nolf & Stringer, 1992,1992,pi.pi. 9, fig.fig. 3]
RhyncocongerRhyncoconger flava PI. 1,1,Fig. 4 [Nolf & Stringer, 1992,1992, pi.pi. 9, figs 11-13]
Rhechias tysanochila Pi.PI. 1, Fig. 5 [Nolf & Stringer,Stringer, 1992,1992, pi.pi. 9, figsfigs 6, 7]7]
Gnathophis sp. [Nolf, 1976,pi. 2, figs 5-7]
PI. 7 Paraconger sp 1, Fig.
Engraulidae Engraulidaeindet.indet. [Nolf& Stringer, 1992,pi. 9, fig. 16]
Plotosidae Plotosus sp.sp. [Nolf& Stringer, 1992,pi. 10,10, fig. 3]3]
SternoptychidaeSternoplychidae Polyipnus sp. PI. 1, Fig. 6 [Nolf& Stringer,Stringer, 1992,1992, pi. 10, fig.fig. 3]
ValenciennellusValenciennellus not sp.sp. not figuredfigured
Synodontidae Saurida caribbaea PI. 2, Fig. 1 [Nolf& Stringer,Stringer, 1992,1992, pi. 10, fig. 3]
Myctophidae Diaphus brachycephalusbrachycephalus PI. 2, Fig. 2 [Nolf& Stringer, 1992,pi. 10, figs 4-9]4-9]
* 11 PI. 2, 3 & 10, Diaphus sp. 2, Fig. [Nolf Stringer,Stringer, 1992,pi. 10, figs 18-23]
Diaphus sp. notfiguredfigured
Symbolophorus aff. veranyi [Brzobohaty & Nolf, 1996, pi.pi. 8,8, figs 17-19]
Hygophum aff. benoilibenoiti [Nolf& Stringer, 1992, pi. 10,10, fig. 32]32]
Hygophumsp.sp. not figuredfigured
MyctophumMyctophum sp. [Nolf& Stringer, 1992, pi. 10,10, figs 14, 15]
Lobianchia PI. sp. PI. 2, Fig.Fig. 55
PI. 44 & Bregmacerotidae Bregmaceros sp. PI. 2, Fig.Fig. [Nolf Stringer,Stringer, 1992,pi. 11, figs 7-8]
GadidaeGadidac Gadiculus labiatuslabiatus [Nolf, 1980,1980, fig. 49H]
Ophidiidae LepophidiumLepophidiumlatesulcatum PI. 2, Fig.Fig. 6 [Nolf & Stringer, 1992,pi. 11, fig. 11]11 ]
LepophidiumLepophidiumcervinumcervinum [Nolf, 1980,1980,pi.pi. 3,3, figsfigs 1-8]
Otophidiumrobustumrobustum PI. 2,2, Fig.Fig. 7 [Nolf& Stringer, 1992,pi. 12, figs 4,4,5]5]
Otophidiumrobinsirobinsi PI. 3,3, Fig.Fig. 1 [Nolf& Stringer, 1992,pi. 11, figs 16-19]
BrotulaBrotula aff. clarkaeclarkae [Nolf& Stringer, 1992,1992,pi.pi. 11, fig.fig. 13]
LepophidiumLepophidiumsp.sp. not figuredfigured
PI. 2 1, CarapidaeCarapidac Carapus sp. 3,3, Fig.Fig. [Nolf, 1980,pi. 1, fig. 3]
EchiodonEchiodon sp. |Nolf,[Nolf, 1980,pi. 1, figs. 8,8,11]11]
Bythidtidae 'genus Dinematichthyinorum'Dincmatichthyinorum'smithvanizismilhvanizi [Nolf& Stringer, 1992,1992, pi.pi. 12, figsfigs 11,12]11, 12]
'genus Dinematichthyinorum' sp. notnot figured
Batrachoididae & 11, Porichthys sp. [Nolf(Nolf Stringer, 1992,1992, p. 11, fig. 3]3]
AtherinidaeAthcrinidac AtherinomorusAlherinomorus stipes [Nolf|Nolf& Stringer, 1992,1992,pi.pi. 12, figsfigs 13-16]13-16]
HemiramphidaeHcmiramphidae Hyporhamphus sp. [Nolf & Stringer, 1992,1992, pi.pi. 12, figsfigs 17,18]17, 18]
Anomalopidae PhthanophaneronPhthanophaneronsp. PI. 3,3, Fig.Fig. 33
Holocentridae Holocentrus & sp. [Nolf|Nolf Stringer, 1992,1992, pi.pi. 12, fig.fig. 25]
Scorpaenidae Scorpaenidaeindet.indet. |Nolf[Nolf& Stringer, 1992,1992, pi.pi. 12, fig.fig. 26]
Triglidae Prionotus cf. carolinus [Nolf|Nolf& Stringer, 1992,1992, pi.pi. 12, fig.fig. 27]
Acropomatidae ParascombropsParascombropscf.cf. spinosus PI. 3,3, Fig. 55
Serranidae & 1992, 13, Diplectrumsp.sp. [Nolf Stringer, 1992,pi.pi. 13, fig.fig. 7]
PriacanthidaePriacanthidac Pristigenys cf. altusalms PI. 3,3, Fig. 44
* * & Apogonidae Apogon sp.sp. 1 [Nolf& Stringer, 1992,1992, pi.pi. 13, fig.fig. 10]10]
2* & Apogon sp.sp. [Nolf Stringer, 1992,1992, pi. 13, fig.fig. 13]13]
3*3* & Apogon sp.sp. [Nolf Stringer, 1992,1992, pi. 13, figsfigs 15,16]15, 16]
Apogon sp.sp. 4*4* [Nolf& Stringer, 1992,1992, pi. 13, fig. 12]
PI. 6 & LutjanidaeLutjanidac PristipomoidesPrislipomoidessp.sp. 3,3,Fig.Fig. [Nolf|Nolf Stringer, 1992,pi. 14, figs 2-4]2-4]
Rhomboplites aurorubens PI. 4, Fig. 1
Gerreidae * EucinostomusEucinostomus sp. 11 [Nolf & Stringer, 1992, pi. 14, fig.fig. 7]
cf. Moharra sp.sp. not figured
Haemulidae Haemulon 2 sp. PI. 4,4, Fig.Fig. [Nolf& Stringer, 1992, pi. 14, figs 16-19]
Sciaenidae Protosciaena trewavasae PI. 4,4,Fig.Fig. 3 [Schwarzhans, 1993, figs 344, 345]
Umbrina & sp. [Nolf Stringer, 1992,1992, pi. 14, fig.fig. 18]18]
Bathyclupeidae Bathyclupea sp. [Nolf, 1976,1976, pi.pi. 9,9, fig.fig. 4]
& Mugilidae Mugil sp. [Nolf Stringer, 1992,1992, pi.pi. 15, fig.fig. 11]
Sphyraenidae & Sphyraena sp. [Nolf Stringer, 1992,1992, pi.pi. 17, fig.fig. 1]1]
Opisognathidae LonchopisthusLonchopislhus micrognathusmicrognathias PI. 4,4, Fig.Fig. 44 [Nolf& Stringer, 1992,pi. 15, figs 7-9]7-9] 151
Uranoscopidae Uranoscopidae indet. not figured
Gobiidae Gobiidae * sp. 1*1 PI. 4, Fig. 5 [Nolf& Stringer, 1992,pi.pi. 17, fig. 12]
Gobiidae 3* 6 sp. PI. 4, Fig. 6 [Nolf& Stringer, 1992,pi. 17, fig. 14]
Gobiidae indet. not figured
Bothidae Citharichthys sp. [Nolf, 1976,pi. 9, fig. 12]
Bothidae indet. [Nolf & Stringer,Stringer, 1992,pi. 17, figs 15,16]15, 16]
BothidaeBothidae indet. not figured
Bothidae indet. not figured
PleuronectidaePlcuronectidae Pleuronectidae indet. [Nolf & Stringer, 1992,pi. 17, fig. 20]
Cynoglossidae Cynoglossidae indet.indet. [Nolf(Nolf & Stringer, 1992,pi. 17, fig. 17]17]
Soleidae Soleidae indet. [Nolf& Stringer, 1992,pi. 17, figs 18,18,19]19]
* Species designatedwith numbersnumbers refer to specimens described and figuredin Nolf&Nolf& Stringer (1992). Plate andfigure numbers enclosed in brackets refer
topreviously figuredmaterial.
Table 1. List of all otolith-based teleosts (class Osteichthyes) identified from the shell bed ofthe Pliocene Bowden Formation in Jamaica.
The vast majority of the identified taxa represent forms PALAEOECOLOGY BASED ON OTOLITHS
that are presently known from the Atlantic Ocean including
the Gulf of Mexico and the Caribbean area. Three notable Studies have indicated that fossil otolith assemblages will exceptions are the catfish genus Plotosus, the ophidiid spe- reflect, with reasonable certainty, the ichthyological fauna cies Brotula clarkae and the lanterneye fish Phthanophan- inhabiting an area during a specific interval of geologic time
eron, all of which are known from the Pacific. The lapilli (Schwarzhans, 1984; Stringer, 1992a; Breard & Stringer, from Plotosus compare remarkably well with those of the 1995). As a general principle, the otoliths will represent a Recent species Plotosus anguillaris (see Nolf & Stringer, part of the biocoenosis. General palaeoecological parameters 1992, pi. 10, fig. 2). The otoliths from Brotula clarkae are can be determinedby utilising data on the preferred habitats also distinctiveand since the very diagnostic they represent exercised of comparable modem fishes. Greater care must be only Brotula species with a prominent rostrum (see Nolf, with older Palaeogene and Cretaceous otolith assemblages 1980, pi. 2, fig. 3). The otoliths fromPhthanophaneron seem because of fewer taxonomic affinities with Recent genera.
to be closely related toPhthanophaneron harveyi [Rosenblatt Also, fewer Cretaceous faunas have been studied (Hud- & 1976) which is foundin the Gulfof Califor- Montgomery, dleston & Savoie, 1983; Nolf & Dockery, 1990; Stringer, nia and the eastern tropical Pacific (Nelson, 1994). The 1991;Nolf& Stringer, 1996; Rana, 1996). Nolf(1985) noted of these three Pacific forms is relicts presence interpreted as that otolith associations will reflect the fishes present in a of the western Tethys fauna. No new species are introduced qualitative sense, but otolith faunas are not a quantitative
from the Bowden shell bed at this time. Several new species measure of the biocoenosis. However, Nolf & Cappetta
may be represented, but adequate growth series were not (1989) suggested a more objective method of interpreting available to make this determinationfor several taxa, such as indicated otoliths. Nolf& Brzobo- palaeobathymetry as by andLobianchia all of the Lepophidium sp. sp. Essentially, otoliths haty (1992) described a method utilising that was
taxa from the Bowden shell bed have been described and suited for the bathymetric interpretation of associations with illustrated Nolf & previously, mainly by Stringer (1992). deep neritic or deep-water fishes.
However, two taxonomic changes should be noted from Nolf The general bathymetry of the Bowden shell bed can be
& Stringer (1992). Hildebrandia was synonymised with deduced by the ratio of neritic fishes to myctophids and Rhynchoconger Smith (1989) in his revision of Recent by macrourids. Nolf & Brzobohaty (1992) noted that otolith western Atlantic eels, and the form reported as Cte- conger associations with both neritic species and myctophids, but nosciaena latecaudata Protosciaena probably represents few or no macrourids, are indicative of neriticenvironments
trewavasae described and illustrated in Schwarzhans as exposed to oceanic influence. The proportion of myctophid References illustrations of the (1993). to previous taxa as otolithsbecomes higher in the deeper part of the continental well as plates in the present study are presented in Table 1. shelf. The otolith assemblage from the Bowden shell bed
taxa examined in the are illustrated Significant present study contains neritic species and myctophids with no macrourids,
in Pis 1-4. and is interpreted as indicative of a neritic environment
In some cases, the present knowledge of Recent Carib- exposed to oceanic influence. beanotoliths is limited allow for identification too to beyond Except for a few taxa, all of the studied otoliths from the the level. There is also the of vari- family problem extreme Bowden shell bed exhibit close affinities to the Recent fauna
in the families Gobiidae, Bothidae and ability among species in the Caribbean, and seem to belong to the same climatic
Pleuronectidae. In the form Gobiidae, groups were desig- and environmentalrealm as their present-day relatives. Based nated while other such the bothids and groups as pleuronec- on the otolith association from the Bowden shell bed, palaeo-
tids were identified with ascertained to family. Species designated bathymetric parameters can be by comparisons
numbers, such as 1, refer to forms described and Diaphus sp. to related modern forms. General parameters followed by in Nolf& figured Stringer (1992). Nolf & Stringer (1992) in their study of the Neogene fish 152
also followed in this of that is utilised. References for fauna from the Dominican Republic are species genus bathymetric
the data for Recent forms include Bohlke & study. In the case of extant or closely-related species, Chaplin (1968), Hoese & Moore bathymetry indicated for the fossil species is that of the Darnell et al. (1983), Dawson (1966),
Recent species. (1977), Nelson (1984, 1994), Poll (1953, 1954,1959), Rob- and Smith & Kana- bins et al. (1986), Schwarzhans (1993)
distribution of zawa (1977). The present-day bathymetric
related Recent taxa is shown for the fossil otoliths in Table 2.
otolith-based taxa from the Bowden shell bed in m. 0 200 300 400 500 600 Forty-nine Taxa/Depth in m. 0 100 Albula Albula sp. in Table 2. were utilised in developing the palaeobathymetry
Pythonichthyspythonichthys sp. It shouldbe noted that the dotted lines in Table2 indicate the Ariosoma balearicabaleanca
and that 'E' RhyncocongerRhyncocongerflava diurnal migration of mesopelagic species an
Rhechias tysanochilatysarnKhila estuaries denotes fishes that may occur in during some stage Gnathophis sp. of their life. Nelson noted that the abundance of Plotosus E (1994) peak Plotosus sp. E~.— the PoiyipnusPolyipnus sp. most myctophid species is between 300-1,200 m during
ValenciennellusVatenciennellussp. 10-100 day, but this peak abundancechanges to m at night.
SauridaSaurida caribbaeacaribbaea Basically, the method described by Nolf & Brzobohaty Diaphus brachycephalusbrachycephalus (1992) is followedin analysing the palaeobathymetry. Thirty- Diaphus sp. 11 foundin the 0-100 seven of the 49 potential forms (76%) are Diaphus sp. which is the m (inner to middleshelf). This percentage, HygophumHygophum aff. H.//. range
benoiti highest for all depth levels, strongly indicates that the most Hygophum sp. 0-100 This is further probable water depth was m. supported MyctophumMyctophum sp. by the progressive decrease in percentages for depths greater Lobianchiabobiancma sp. the of than 200 m. However, presence mesopelagic forms, BregmacerosBregmaceros sp. the of GadiculusGadiculuslabiatusSab'uitus especially the myctophids, signifies proximity open
LepophidiumlêpOptdauËM oceanic conditions. The majority of the taxa present in waters latesulcatumlatesulcatum
LepophidiumLepophidium than 400 m are myctophids. cervinumcervinum deeper
robusium OtophidiumOtophidium robustum In addition to the graphical representation of the bathy- Otophidium robinsirobinsi indicated by otoliths in Table 2, it is interesting to Brotulaaff. B. metry claricaeclarkae fish There comparethe bathymetrical distributionof the taxa. Lepophidiumsp.sp.
Carapus taxa that be found in estuarineenvironments. Campus sp. are eight may
EchiodonEchiodonsp. indicative ofthe Twenty-eight of the taxa are restricted to or
EE" PorichthysPorichthys sp. shelf environment (generally 0 to 200 m). Several fish taxa AtherinomorusAtherinomorus stipes stipes - environment. Recent also suggest the presence of a reef HyporhamphusHyporhamphus sp.
Holocentrus cardinalfishes Holocentrus sp. squirrelfishes (Holocentridae) and (Apogoni-
cf. Parascombrops cf. characterised dae) as well as the Dinematichthyini are mainly spinosus
Diplectrum sp. as reef fishes in the Gulf of Mexico (Hoese & Moore, 1977). Pristigenys cf. u//m.taltus Nelson (1984, 1994) also stated that squirrelfishes and Apogon sp. 1 the shoreline and 100 Hoese Apogon sp.sp. 2 soldierfishes occur between m.
Apogon 3 sp.sp. & Moore (1977) stated that squirrelfishes are a tropical
Apogon sp.sp. 4 family represented by several species found, so far, only on Pristipomoides sp.sp.
Eucinostomus Er the offshore reefs in the Gulfof Mexico. Bohlke & Chaplin
sp. 1I
cfcf. MoharraMoharra EE" stated that and soldierfishes are reef sp.sp. (1968) squirrelfishes
Haemulonliacmulon sp. sp. fishes which occur in tropical waters around the world. They ProtosciaenaProiosciaena foundin crevices and in the trewavasae noted that these forms are caves
Umbrinai sp. rocks and reef. water taxa are mainly mesopelagic EE" Deeper Sphyraena sp. forms. LonchopisthusLonchopislhus ~T7E (Myctophidae) with no bathybenthic - micrognathusmicrognathia Gobiidae 11 EE" One form, Plotosus, was found in sp. seemingly enigmatic
Gobiidae 3 EE" sp. the otolith assemblage. Plotosus is an Indo-Pacific genusof
Gobiidaeindet. EE" and freshwater envi- catfish that occurs in neritic, estuarine
ronments Its is not surprising since taxa (Berra, 1981). presence Table 2. Present-day bathymetric range of teleostean it also found in the ofthe Dominican Republic represented in the Bowden shell bed, Jamaica. Dotted was Neogene
Nolf & This occurrence is probably lines indicate the nightly occurrence of mesopelagic by Stringer (1992). that related the Caribbean and the Central species near the surface. An E represents fishes to Tertiary province
in their life closure of the may occur estuaries at some stage of cycle. America seaway that existed prior to seaway
Brotula in the Pliocene (Keigwin, 1978). Plotosus, as well as
taxa taxa indeterminate to the For extinct or as species, aff. clarkae and Phthanophaneron sp., are interpreted as
global bathymetric distribution indicated for all Recent relicts of the ancient western Tethys fauna. 153
be from the which somewhat low Other palaeoecological indications may also gleaned recognised sample seems of otoliths. from the otolith-basedtaxa of the Bowden shell bed. Almost considering the size of the sample and number
in the Recent Caribbean and One is that with smallerotoliths lost all of the taxa are present fauna, explanation species were
forms. the due the sieve size This is further represent tropical and subtropical Therefore, to (25 mesh). supported by
is consideredto be similar to the modem the of sciaenid otoliths which tend to be climatic realm very large percentage
of 26-28°C in size. The fauna included distributed in Caribbean with surface sea water temperatures larger many taxa
both the Atlantic and Pacific. The sciaenids represented over (Dunn et al., 1981). The taxa associated with reef environ- 50% of the total number of otolithsand contained eight of the ments in the otolith assemblage support this observation.
twenty-six described taxa. Two Haemulon-type species were Based on the otoliths, the depositional environment was and constituted 15% of the characterised by normal marine salinity (approximately 35 the second most common types
total number of otoliths. No other percentages are given for %c), fairly low energy and mostly soft bottoms.
the other and of the taxa identifiedto The comparison of the fossil taxa from the Bowden shell taxa none are species.
these factors, it is difficult to bed to the bathymetric distributions of closely related Recent Due to quantitatively compare
forms the Gatun Formation with other faunas in the Caribbean. forms reveals an assemblage with shallow-water A diverse otolith fauna described from the (including several euryhaline species), reef inhabitants, and was Neogene of Trinidad Nolf collected at 30 middle to outer shelf forms with some upper slope and by (1976). Samples were
localities from nine Miocene and Pliocene formations in pelagic elements. The otolith assemblage, although predomi-
and otoliths. Sixty-six taxa were nantly shallow water (0-100 m), seems to represent a combi- Trinidad, produced 1,339
described from the otoliths with most of them from nationof several ecological environments. A driftedor mixed coming
Caldwell for the Bow- the Miocene units, mainly the Brasso, Nariva and Manzanilla assemblage was suggested by (1965) consisted of neritic fishes den shell bed, but Clarke & Fitch (1979) rejected this idea in Formations. The fauna primarily with few and demersal fishes from the favour of a 'mass mortality' deposit similar to those resulting a mesopelagic upper The and sciaenids from dinoflagellate blooms. Examining modern zoo- slope. ophidiids, pomadasyids represented
environment for the the most The most abundant taxa included indeter- geographical situations, a probable species.
bed reconstructed. Ifthe Bowden shell minate Gobiidae, Diaphus dumerili and Bregmaceros. Bowden shell may be
the otoliths from the of Trinidadresem- bed was deposited in the shallow, nearshore marine waters of Generally, Neogene
shelf to bled the present-day Caribbean fauna. an island with a very narrow region quickly dropping
situation and oceanic this would Some of the best faunas for would a slope open conditions, potential comparisons from the explain the combinationof several ecological environments. be those studied by Nolf & Stringer (1992) Neogene
in of the northern Dominican However, it should be The euryhaline, reef and shelf forms would be common Republic.
notedthat the materialfrom the Dominican the narrow shelf region, and the neritic and deeper-water Neogene Repub-
oceanic conditions lic represented otoliths from four different formations forms would be present due to the open rang-
shelf Pickerill al. in from Miocene for the lower adjacent to the narrow area. et (1996) ing age early-middle (NN4 Baitoa middle Pliocene for the concluded that the shallow-marineand relatively deep-water Formation) to (NN15 upper
Mao It must also be considered that 887 faunas ofthe Bowden shell bed were indicative of depostion Formation). samples localities in the northern from sediment gravity flows. Turbidite deposition would were collected from hundreds of
Dominican with 109 localities yielding otoliths account for the combinationof several ecological conditions Republic
et In Tulane University (New indicated by the teleostean otoliths. This interpretation would (Saunders al., 1986). addition, collected hundreds of localities in the be consistent with other palaeontological (invertebrate and Orleans, Louisiana) Dominican and otoliths vertebrate), sedimentary and tectonic studies of the Carib- same area of the Republic supplied bean and Central America (Briggs, 1974; Casey et al., 1975; for study. The otolith fauna from the Dominican Dunn etal., 1981; Duque-Caro, 1990; Gillette, 1984; Keig- largest Republic from the late Miocene NN11 Zone in the Cercado win, 1976, 1978; Smith, 1991; Whitmore & Stewart, 1965; was Formationand consisted of 62 The otoliths from the Woodring, 1966). species.
Cercado Formation indicateda shallow marine environment.
More precisely, 37 of the 44 taxa usable for palaeobathyme-
COMPARISONS TO OTHER OTOLITH FAUNAS OF THE try (84%) occur in waters from 0 to 100 m. Many of the
CARIBBEAN neritic taxa do not live at depths exceeding 50 m and at least
11 species are regular inhabitantsof euryhaline environments
estuaries. of the Although the number of described otolith assemblages from such as lagoons or None deeper-water taxa
several of them and all of them occur in neritic the Caribbeanis quite limited, are appropri- are exclusively bathyal may Labrisomusand the ate for comparison. An otolithassemblage from the Miocene areas. Several taxa, such as Holocentrus,
described Gillette the of reef environ- Gatun Formation in Panamawas in (1984). Dinematichthyini, suggest proximity
this identifications but the of the forms confined to shallow As noted earlier in paper, preliminary ments, majority are
the fauna neritic environments with rather soft bottoms & were done by the late John Fitch, but was never (Nolf
formally described. Approximately 900 kg of sample was Stringer, 1992). The otolith from the Bowden shell bed wet-sieved and yielded 2,074 otoliths. Twenty-six taxa were assemblage 154
compares most closely with the otolith fauna from the late most closely with the otolith fauna from the late Miocene
Miocene NN11 Zone in the Cercado Formation in the Do- NN11 Zone in the Cercado Formation in the Dominican
minican Republic. Although the Bowden otolith assemblage Republic described by Nolf & Stringer (1992). The most
is younger in age (Pliocene, probably around NN13-NN14), notable difference in the two assemblages is the larger
there are many similarities between the two assemblages. number of sciaenid species in the Cercado assemblage (six
Someof the similaritiesinclude the number of total species versus only two in the Bowden shell bed fauna).
for both faunas, the percentage of potential forms found in
the 0-100 m range is almost the same (approximately 80% ACKNOWLEDGEMENTS for both faunas), the number of euryhaline inhabitants is
about the same, the number ofshallow water taxa occurring A special thanks to Dr Dirk Nolf(Institut royal des Sciences on the shelf is similar, several taxa that suggest a reef envi- naturellesde Belgique, Brussels), who provided invaluable ronment are present in both faunas and deeper water forms assistance in numerous ways including taxonomic identifica- are mainly mesopelagic with no bathybenthic forms. Both tions, loaned material and suggestions for the study. Thanks assemblages containPlotosus, an Indo-Pacific catfish genus to all ofthe people who graciously provided otoliths from the which is interpreted as a relict element of the ancient western Bowden shellbed for research; Dr Harold Vokes, Dr Emily Tethys fauna. A significant difference in the Bowden shell Vokes, Mr Roger Portell and Dr Jon Bryan. Thanks to Dr bed and the NN11 Zone in the Cercado Formation is the Frank Pezold of Northeast Louisiana University (Monroe, number of sciaenids. The Bowden shell bed has only two Louisiana) for information on the bathymetry of Recent species of sciaenids while the Cercado has at least six sciae- fishes in the Gulf of Mexico. Thanks to the reviewers for nid species represented. their constructive critique of the manuscript. Special appre-
ciation is extended to Dr Stephen K. Donovan(University of
the West Indies, Mona) for his invitationto participate in this CONCLUSIONS of this project and for his advice regarding numerous aspects
study. His detailed editing of the draft manuscript was most A detailed study of approximately 1,650 fossil teleostean appreciated. otoliths from the type locality ofthe Pliocene Bowden shell
bed, Bowden Formation (Jamaica) revealed at least 38 teleost
families and 68 species. A definite correlation was found to REFERENCES exist between the sample size and the number of species
identified.All of the identificationswere based on the sagit- Berra, T., 1981. An atlas of distribution of the freshwater fish the saccular otoliths for the tae or except family Plotosidae, familiesof the world. Lincoln (University of Nebraska Press), where the lapillus or utricular otolith was utilised. All of the 197 pp. identified taxa from the Bowden shell bed represent forms Bohlke, J. & C. Chaplin, 1968. Fishes of the Bahamas and adjacent that are presently known from the Atlantic Ocean (including tropical waters. Wynnewood(Livingston Publishing Co.), 771 the Gulf of Mexico and Caribbean) except for three species. pp.
These three notable are the catfish Plotosus, exceptions genus Breard, S. & G. Stringer, 1995. Paleoenvironment of a diverse
the ophidiid species Brotula clarkae and the lanterneye fish marine vertebrate fauna from the Yazoo Clay (late Eocene) at
Caldwell Louisiana. — Transactions of the Phthanophaneron, all of which are known from the Pacific. Copenhagen, Parish,
relicts the ancient Gulf Coast Association of Geological Societies, 45: 77-85. These forms are interpreted as of western Breder, C.M. Jr., 1927. Scientific results of the first ocenographic Tethys fauna.
expedition of the ”Pawnee”, 1925. Fishes. — Bulletin of the Forty-nine of the otolith-based taxa from the Bowden Bingham Oceanographic Collection, 1(1): 1-90. shell bed were utilised to determinedetailed palaeobathyme- Briggs, J., 1974. Marine zoogeography. New York (McGraw-Hill), of the forms found try. Thirty-seven 49 potential (76%) were 475 pp. in middle of the the 0-100 m range (inner to shelf). Eight taxa Brzobohaty, R., 1986. Einige neue Arten von Knochenfischen may be found in estuarine environments, and 28 are restricted (Teleostei, Otolithen) aus dem Westkarpatischen Tertiar. — to indicativeof the shelf environment 0 to 200 or (generally Acta Musei Moraviae, 71: 55-71.
Several fish, such the holocentridsand also m). as apogonids, Brzobohaty, R. & D. Nolf, 1995. Diaphus otoliths from the Euro-
the of a reef environment. the — Bulletin de suggest presence Flowever, pean Oligocene (Myctophidae, Teleostei).
of the l'Institut des Sciences naturelles de Sciences presence mesopelagic forms, especially myctophids, royal Belgique, de
the of oceanic conditions. The la Terre, 65: 257-268. signifies proximity open
R. & D. 1996. Otolithes de otolith-based fishes suggest deposition in shallow, nearshore Brzobohaty, Nolf, myctophides (pois-
des terrains tertiaires revision des marine of island with shelf sons teleosteens) d'Europe: waters an a very narrow region
genres Benthosema, Hygophum, et to situation and oceanic Lampadena, Notoscopelus quickly dropping a slope open
Symbolophorus. — Bulletin de l'Institut royal des Sciences conditions. This palaeoenvironment, coupled with sediment naturelles de Belgique, Sciences de la Terre, 66: 151-176. gravity flows, would explain the combination of several Caldwell, D., 1965. A Miocene needlefishfrom Bowden, Jamaica. ecological environments indicated in the Bowden shellbed. — Quarterly Journal of the Florida Academy of Science, 28: The otolith from the Bowden shell bed assemblage compares 339-344. 155
Casey, R., K. McMillen & M. Bauer, 1975. Evidence for and — Proceedings of the Biological Society of Washington, 96:
paleooceanographic significance of relict radiolarian popula- 658-663.
tions in the Gulf of Mexico and Caribbean. — Geological Soci- Keigwin, L., 1976. Late Cenozoic planktonic foraminiferal
ety of America, Abstracts with Programs, 7:1022, 1023. biostratigraphy and paleoceanographyof the Panama Basin. —
Casier, E., 1966. Sur la faune ichthyologique de la formation de Micropaleontology, 22: 419-442.
et et Bissex Hill de la serie oceanique de Pile de la Barbade, sur Keigwin, L., 1978. Pliocene closing of the Isthmus of Panama, formations.— Pacific 1'age de ces Eclogae geologicae Helvetiae, 53: based on biostratigraphic evidence from nearby Ocean
493-515. — 630-634. and Caribbean Sea cores. Geology, 6:
Chao, L.N. & R.V. Miller, 1975. Two new species of sciaenid fishes Leriche, M., 1938. Contribution a l'etude des poissons fossiles des
from the Caribbean Sea and waters. Americaine (tribe: Sciaenini) adjacent pays riverains de la Mediterranee (Venezuela,
— Bulletin of marine Science, 25: 259-271. Trinite, Antilles, Mexique). — Memoires de la Societe paleon-
Clarke, M. & J.E. Fitch, 1979. Statoliths of Cenozoic teuthoid tologique Suisse, 61: 1-42.
cephalopods from North America. — Palaeontology, 22: 479- Nelson, J.S., 1984, Fishes of the world (2nd edition). New York
511. (John Wiley & Sons), 523 pp.
Cuvier, G., 1816. Le Regne Animal distribue d'apres son organisa- Nelson, J.S., 1994. Fishes of the world (3rd edition). New York
& 600 tion, pour servir de base a l'histoire naturelle des Animaux et (John Wiley Sons), pp.
d'introduction a l'Anatomie Comparee. Les reptiles, les pois- Nolf, D., 1976. Les otolithes de teleosteens Neogenes de Trinidad
les les 532 — 69: 703-742. sons, mollusques et annelides, 1(2). Paris, pp. Eclogae geologicae Helvetiae,
Darnell, R.M., R.E. Defenbaugh& D. Moore, 1983.Northwestern Nolf, D., 1980. Etude monographique des otolithes des Ophidiifor-
Gulf Shelf bio-atlas: of the distribution actuels revision des fossiles a study of demersal mes et especes (Pisces, Teleostei).
fishes and penaeid shrimp of the soft bottoms ofthe continental — Mededelingen van de Werkgroep voor Tertiaire en Kwar-
shelf from the Rio Grande to the Mississippi River Delta. — taire Geologie, 17: 71-195.
Minerals Management Service, United StatesDepartment of the Nolf, D., 1985. Otolithi Piscium. In: H. Schultze (ed.). Handbook of
Interior File 82-04: 438 York Fischer Open Report, pp. Paleoichthyology. Stuttgart/New (Gustav Verlag),
Dawson, C., 1966. Observations on the anacanthine fish Bregmac- 1-145.
eros atlanticus in the north-central Gulf of Mexico. — Copeia, Nolf, D. & R. Brzobohaty, 1992. Fish otoliths as paleobathymetric
1966: 604, 605. indicators. — Paleontologia e Evolucio, 24/25: 255-264.
Delaroche, F., 1809. Observations sur les Poissons recueillis dans Nolf,D. & H. Cappetta, 1989. Otolithes de poissons du Pliocene du
lies Baleares Annates du sud-est de la France. — Bulletin de l'lnstitut des Sciences un voyage aus et Pythiuses.— royal
Museum national d'Histoire naturelle Paris, 13: 1-75. naturelles de Belgique, Sciences de la Terre, 58: 209-271.
Donovan, S.K., T.A. Jackson, H.L. Dixon & E.N. Doyle, 1995. Nolf, D. & D. Dockery, 1990. Fish otoliths from the Coffee Sand
Eastern and central Jamaica. — Geologists' Association Guide, (Campanian) of northeastern Mississippi. — Mississippi Geol-
1-62. 1-14. 53: ogy, 10(3):
Dunn, D., T. Moore & L. Keigwin, 1981. Atlantic-type carbonate Nolf, D. & G.L. Stringer, 1992. Neogene paleontology in the
stratigraphy in the late Miocene Pacific. — Nature, 291: 225- northern Dominican Republic: otolithsof teleostean fishes. —
227. Bulletins of American Paleontology, 102: 45-81.
Duque-Caro, H., 1990. Neogene stratigraphy, paleoceanography and Nolf, D. & G.L. Stringer, 1996. Cretaceous fish otoliths: a synthesis
paleobiogeography in northwest South America and the evolu- of the North American record. In: G. Arratia & G. Viohl (eds).
tion of the Panama Seaway. — Palaeogeography,Palaeoclima- Mesozoic Fishes: Systematics and Paleoecology. Proceedings
tology, Palaeoecology, 77: 203-234. of the First International Meeting on Mesozoic Fishes, Eichstatt,
Fitch, J.E., 1969. Fossil lanternfishotoliths of California, with notes Germany. Miinchen (Verlag Friedrich Pfeil): 433-459.
on fossil Myctophidae of North America. — Los Angeles Mu- Pickerill, R.K., D.G. Keighley & S.K. Donovan, 1996. Ichnology of
seum Contributions in Science, 173: 1-20. the Pliocene Bowden Formation of southeastern Jamaica. —
Fitch, J.E. & L. Barker, 1968. The fish Family Moridae in the Caribbean Journal of Science, 32: 221-232.
eastern North Pacific with noteson morid otliths, caudal skele- Poey, F., 1858-1861. Memorias sobra la historia natural de la Isla de
and the fossil — 70: 565-584. of sumarios Latinos Frances. tons, record. Fishery Bulletin, Cuba, acompanadas y extractos en
Gill, T., 1861. Catalogue of the fishes of the eastern coast of North — La Habana, 2: 337-442.
America, from Greenland to Georgia. — Philadelphia Academy Poll, M., 1953. Poissons III, Teleosteens malacopterygiens. —
of Natural Sciences, Proceedings for 1860: 1-63. Resultats scientifiques de l'Expeditionoceanographiquebeige
Gillette,D.D., 1984. A marine ichthyofauna from the Miocene of dans les Eaux Coheres Afficaines de l'Atlantique Sud, 4(2): 1-
Panama, and the Tertiary Caribbean Faunal Province. — Jour- 250.
nal of vertebrate Paleontology, 4: 177-186. Poll, M., 1954. Poissons IV, Teleosteens acanthopterygiens (pre-
— Resultats Goode, G.B. & T.H. Bean, 1896. Oceanic ichthyology, a treatise on miere partie). scientifiques de 1'Expedition oce-
dans les Eaux Cotieres Africaines de the deep-seaand pelagic fishes of the world, based chiefly upon anographique beige
the collections made by the steamers Blake, Albatross and Fish I'Atlantique Sud, 4(3a): 1-300.
Hawk in the northwestern Atlantic. — United States National Poll, M., 1959. Poissons V, Teleosteens acanthopterygiens
Museum, Special Bulletin 2: 1-553. (deuxieme partie). — Resultats scientifiques de I'Expedition
Hoese, H. & R. Moore, 1977. Fishes ofthe Gulf of Mexico, Texas, oceanographique beige dans les Eaux Cotieres Africaines de
and Louisiana, and adjacent waters. College Station, Texas I'Atlantique Sud, 4(3b): 1-417.
A & M 327 R.W., S.K. Donovan, R.K. Pickerill & H.L. Dixon, 1996. (Texas University Press), pp. Purdy,
Huddleston, R. & K. Savoie, 1983. Teleostean otoliths from the late Fish teeth from the Pleistocene of Jamaica. — Journal of verte-
Cretaceous (Maestrichtian age) Severn Formation of Maryland. brate Paleontology, 16: 165-167. 156
1992. Fish otoliths in the Middle Miocene of eels the Norlth Atlantic with Radwanska, U., (Bade- genus congrid from western re-
nian) deposits of southern Poland. — Acta geologica polonica, descriptions of Ariosoma analis, Hildebrandia guppyi and
42: 141-328. Rhechias vicinalis. — Bulletin of marine Science, 27: 530-543.
Rana, R.S., 1996. Additional fish otoliths from the Deccan Trap Smith, J., 1991. Cenozoic giant pectinids from California and the
associated beds near Caribbean ”Macrochlamis sedimentary exposed Rangapur, Ranga- Tertiary province: Lyropecten, ”,
reddi District, Andhra Pradesh. — Contributions to the XV In- Vertipecten, and Nodipecten species. — United States Geologi-
dian Colloquium on Micropaleontological Stratigraphy, 15: cal Survey Professional Paper, 1391: 1-155.
477-491. Steurbaut,E., 1984. Les otolithes de teleosteens de l'Oligo-Miocene
Reid, E., 1934. Two new congrid eels and a new flatfish.— Smith- d'Aquitaine(Sud-Ouest de la France). — Palaeontographica,
sonian MiscellaneousCollections, 91:1-11. A186:1-162.
Robbins, C.R., G.C. Ray & J. Douglass, 1986. A field guide to the Stringer, G., 1979. A study of the Upper Eocene otoliths of the
Atlantic coast fishes of North America. Boston (Houghton Mif- Yazoo Clay in Caldwell Parish, Louisiana. — Tulane Studies
flin 354 Co.), pp. in Geology and Paleontology, 15: 95-104.
Rosenblatt,KryptophaneronR.H. & W.L. Montgomery, 1976. Stringer, G., 1986. Teleostean otoliths and their paleoecological
from Schie- harveyi, a new anomalopid fish the eastern tropical Pa- implications at the Montgomery Landinglocality. In: J.
cific, and the evolution of the Anomalopidae.— Copeia, 1976: bout & W.A. van den Bold (eds). MontgomeryLanding site,
510-515. marine Eocene (Jackson) of central Louisiana. Proceedings of
Saunders, J.B., P. Jung & B. Biju-Duval, 1986. Neogene paleontol- a Symposium of the Gulf Coast Association ofGeological So-
Dominican 1. Baton ogy in the northern Republic. Field surveys, cieties. Rouge (GCAGS): 209-221.
and — Bulletins of Pa- Cretaceous lithology, environment, age. American Stringer, G., 1991. Upper (Maastrichtian) teleostean
leontology, 89: 1-79. otoliths from the Ripley Formation, Union County, Mississippi.
Schubert, R., 1909. Fischreste. In: A. Toula (ed.). Eine jungtertiare — Mississippi Geology, 11: 9-20.
Fauna von Gatun am Panama-Kanal. —Jahrbuch der kaiser- Stringer, G., 1992a. Late Pleistocene-early Holocene teleostean
lich-koniglichen geologischenReichsanstalt, 58: 687-691. otoliths from a Mississippi River mudlump. — Journal of verte-
and three of brate 12: Schultz, L.P., 1940. Two new genera new species Paleontology, 33-41.
with the other the fam- G. 1992b. Fossil teleostean otoliths Bowden cheilodipterid fishes, noteson genera of Stringer, from the
ily. — Proceedings of the United States National Museum, Formation of Jamaica. — Journal of the Mississippi Academy
88(3085): 403-423. of Sciences, 37: 41.
Schwarzhans, W., 1984. Fish otoliths from the New Zealand Taning, A.V., 1928. Synopsis of the scopelids in the north Atlantic.
Tertiary. — Report of the New Zealand Geological Survey, — Videnskabelige Meddelelser fra Dansk naturhistorisk
113: 1-269. Forening i Kpbenhavn, 86: 49-69.
Schwarzhans, W., 1993. A comparative morphological treatise of Weiler, W., 1959. Miozane Fisch-Otolithen aus der Bohrung S.
recent and fossil otolithsof the family Sciaenidae (Perciformes). Pablo 2 im Becken von Veracruz in Mexico. — Neues Jahr-
In: F. Pfeil (ed.). Piscium Catalogus (Part Otolithi Piscium, vol. buch fur Geologie und Paliiontologie, Abhandlungen, 109:147-
Friedrich 245 172. 1). Miinchen (Verlag Pfeil), pp.
Schwarzhans, W., 1994. Sexual and ontogenetic dimorphism in Whitmore, F. & R. Stewart, 1965. Miocene mammals and Central
otoliths the — 71-98. American — 148:180-185. of Family Ophidiidae. Cybium, 18: seaways. Science,
Smale, M.J., G. Watson & T. Hecht, 1995. Otolith atlas of southern Woodring, W., 1966. The Panama land bridge as a seabarrier. —
African marine fishes. — Ichthyological Mongographsof the American Philosophical Society Proceedings, 110: 425-433.
J.L.B. Smith Institute ofIchthyology, 1:1-253.
Smith, D.G., 1989. In: Fishes of the North Family Congridae. Manuscript received 31 January 1997, revised version accepted 6
Atlantic. — Foundationfor ma- Western Memoir of the Sears August 1997.
rine Research, l(part 9): 460-567.
Smith, D.G. & R. Kanazawa, 1977. Eight new species and a new
PLATE 1
All in from the Pliocene shell bed. in each followed the specimens figured Pls 1-4 are Bowden Figure numbers plate are by taxonomic identifica-
view and collection number. Material from the Institute des tion, location ofthe otolith in the fish (right or left side), (inner, outer, ventral) royal
collection is in order and do not have individual Sciences naturelles de Belgique are designated as IRScNB; this arranged systematic specimens
collection numbers. Figured specimens from the Florida Museum of Natural History (University of Florida, Gainesville) have the prefix UF, Division of that The scale bar beside each otolith and are reposited in the Type and Figured Collectionof the Invertebrate Paleontology museum.
figured is equal to 1 mm.
1. face Fig. Albulasp., right sagitta, inner (IRScNB).
2. left Fig. Pythonichthys sp., sagitta, inner face (UF 78951).
Fig. 3. Ariosoma balearica (Delaroche, 1809), right sagitta, inner face (UF 78952).
Fig. 4. Rhyncoconger flava (Goode& Bean, 1896), right sagitta, inner face (UF 78953).
Fig. 5. Rhechias tysanochila (Reid, 1934), left sagitta, inner face (UF 78954).
6. left inner Fig. Polyipnus sp., sagitta, face (UF 78955).
inner face Fig. 7. Paraconger sp., left sagitta, (UF 78956). 157\1
PLATE 1 158\2
PLATE 2
PLATE 2
Fig. 1. Saurida caribbaeaBreder, 1927, left sagitta, inner face (UF 78957).
Fig. 2. Diaphus brachycephalus Taning, 1928, right sagitta, inner face (UF 78958).
Fig. 3. face Diaphus sp. 1, right sagitta, inner (IRScNB).
4. Fig. Bregmaceros sp., left sagitta, inner face (UF 78959).
5. left face Fig. Lobianchia sp., sagitta, inner (IRScNB).
Fig. 6. Lepophidium latesulcatum Nolf & Stringer, 1992, left sagitta, inner face (UF 78960).
Fig. 7. Otophidium robustum Nolf & Stringer, 1992, left sagitta, inner face (UF 78961). 159\3
PLATE3
PLATE 3
Fig. 1. Otophidium robinsi Nolf & Stringer, 1992, right sagitta, inner face (UF 78962).
2. Fig. Carapus sp., right sagitta, inner face (UF 78963).
inner face Fig. 3. Phthanophaneron sp., left sagitta, (UF 78964).
Fig. 4. Parascombrops cf. spinosus Schultz, 1940, left sagitta, inner face (UF 78965).
Fig. 5. Pristigenys cf altus (Gill, 1861), right sagitta, inner face (IRScNB).
6. Fig. Pristipomoides sp., left sagitta, inner face (IRScNB). 160\4
PLATE 4
PLATE 4
Fig. 1.Rhomboplites aurorubens (Cuvier, 1816), right sagitta, inner face (IRScNB).
face Fig. 2. Haemulon sp., right sagitta, inner (UF 78966).
Fig. 3. Protosciaena trewavasae (Chao & Miller, 1975), left sagitta, inner face (UF 78967).
Fig. 4. Lonchopisthus micrognathus Poey, 1861, right sagitta, inner face (UF 78968).
Fig. 5. Gobiidae sp. 1, right sagitta, inner face (UF 78969).
6. Gobiidae inner face Fig. sp. 3, right sagitta, (UF 78970).