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Geology; March 2009; v. 37; no. 3; p. 215-218; DOI: 10.1130/G25377A.1
© 2009 Geological Society of America
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Isotopic analysis of coexisting Late Jurassic fish otoliths and molluscs: Implications for upper-ocean water temperature estimates

G.D. Price1, D. Wilkinson1, M.B. Hart1, K.N. Page1 and S.T. Grimes1

1 School of Earth, Ocean, and Environmental Sciences, University of Plymouth, Drake Circus, Plymouth PL4 8AA, UK


   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 MATERIAL AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES CITED
 
The {delta}18O compositions of well-preserved Jurassic fish otoliths from Wootton Bassett, UK, provide upper-ocean paleotemperatures that are comparable with those derived from the isotopic analysis of fish tooth phosphates, providing independent scrutiny of such paleotemperatures. {delta}18O otolith temperatures in excess of 30 °C also rival temperatures associated with the middle Cretaceous thermal maximum. The negative carbon isotopes of the otoliths may point to a freshwater influence and potentially migratory nature of the fish. However, given the large departures from equilibrium fractionation toward more negative carbon values reported from modern marine fish, we consider our temperature interpretations to be robust and representative of the marine depositional environment. Depleted {delta}13C values, we believe, suggest that the otoliths examined in this study belong to fish with high metabolic rates.


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
MATERIAL AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES CITED
 
The oxygen isotope compositions of foraminifera (e.g., Huber et al., 1995; Wilson et al., 2002), brachiopod shells (e.g., Picard et al., 1998), and molluscs (e.g., Anderson et al., 1994; Price and Page, 2008) are frequently used to estimate temperatures of past oceans. The oxygen isotope compositions of fossil fish teeth have also been widely used as an upper-ocean temperature proxy (e.g., Kolodny and Raab, 1988; Anderson et al., 1994; Picard et al., 1998; Lécuyer et al., 2003; Pucéat et al., 2007), providing new insights on the thermal structure of Jurassic and Cretaceous oceans. Significantly, temperatures derived from {delta}18Ophosphate of fish teeth are typically higher by ~3–4 °C than temperatures inferred from planktic foraminifera {delta}18O (Pucéat et al., 2007) and ammonite {delta}18O (e.g., Anderson et al., 1994). The reasons behind this may relate to cold bottom-water diagenetic alteration of carbonates (e.g., Pearson et al., 2001), leading to underestimates of temperatures derived from chalk-hosted foraminifera, or that Mesozoic fish did not precipitate phosphate according to the phosphate-water fractionation of modern fish, leading to erroneous temperature interpretations (e.g., Anderson et al., 1994). Indeed, relatively little is known about species-specific fractionation effects for biogenic fish phosphate (Vennemann et al., 2001), and Hiatt and Budd (2001) suggest there is no universal agreement upon a phosphate-water paleotemperature equation. Alternatively, inferred temperatures derived from fish may not be representative of their final resting place due to the migratory nature of some fish. In order to examine the veracity of fish-derived paleotemperatures, exceptionally preserved Jurassic (Oxfordian-Kimmeridgian) fish otoliths, together with molluscs (oysters, belemnites, and ammonites) and benthic foraminifera, were isotopically analyzed from a mudspring in Wootton Bassett, UK, an unusual fossil lagerstätte.

Fish otoliths are relatively common in the fossil record and represent an underexploited resource with respect to isotopic studies, especially with respect to the Jurassic and Cretaceous periods. Research has suggested that otoliths contain an isotope record that can be interpreted in relation to the past life of an individual fish, including information on paleodiet, metabolism, and migratory behavior as well as information on the thermal history of a region (e.g., Kalish, 1991; Patterson, 1999; Sherwood and Rose, 2003; Zazzo et al., 2006). Interpretation of carbon and oxygen isotope records in fish otoliths and other biogenic carbonates requires an understanding of potential isotopic disequilibria between ambient waters and carbonates. Otoliths, the stato-acoustic organs of bony (teleost) fish, importantly are precipitated, with respect to oxygen, in equilibrium with environmental water (Kalish, 1991; Thorrold et al., 1997).


   MATERIAL AND METHODS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES CITED
 
The samples for this study were obtained from Wootton Bassett mudsprings, Wiltshire, UK (Fig. 1), located in the northern part of the Anglo-Paris Basin. During the Late Jurassic the area was part of an extensive epeiric sea joining the Tethys Ocean to the Boreal Sea to the north. The fossils, brought to the surface by the springs, include ammonites (which have retained their original iridescent aragonitic nacre), belemnites, bivalves, gastropods, echinoids, serpulids, and wood. From this suite of fossils, isotopic analyses have been performed on oysters (Deltoideum delta), belemnites (Cylindroteuthis sp.), ammonites (aulacostephanids), "glassy" calcitic benthic foraminifera (Lenticulina muensteri), "glassy" aragonitic benthic foraminifera (Epistomina tenuicostata), and fish otoliths (Otolithus [Leptolepididarum] cf. simplex). The mudsprings are sited along the axis of a syncline, and confined groundwater from the Coral Rag aquifer, passing up through overlying clays, is probably the force driving the mudsprings (Bristow et al., 2000). The erupted material contains biostratigraphically diagnostic fossils indicating sourcing from the Ampthill and Kimmeridge clays (of late Oxfordian to early Kimmeridgian age) immediately underlying the site at a depth of around 9 m (Harding et al., 2000). This stratigraphic interval spans just the Pseudocordata to Baylei Zones (Wright, 2003) and is thus comparable with other studies (e.g., Anderson et al., 1994). Preservation of fauna was evaluated by scanning electron microscope (SEM) and trace element geochemistry. Our trace elemental analyses revealed low Fe (<247 ppm) and Mn (<15 ppm) in all belemnite rostra and <4536 ppm Fe and <359 ppm Mn for the oysters, which is consistent with a good state of preservation of these fossils. SEM analyses demonstrate that otoliths and ammonites are composed of well-preserved pristine aragonite (Figs. 2A and 2B). Both the aragonitic and calcitic foraminifera, despite having an external glassy appearance, reveal test interiors filled with calcite spar and pyrite octahedra and framboids (see also Hart et al., 2006). Evidence of minor recrystallization and homogenization of test walls is also evident (Figs. 2C and 2D).


Figure 01
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  		Figure 1. Sketch map showing the occurrence of Oxfordian-Kimmeridgian rocks in the UK and the location of Wootton Bassett.

 


Figure 02
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  		Figure 2. Scanning electron microscope photomicrographs. A: Otolith microstructures revealing structures similar to other well-preserved otoliths (e.g., Dufour et al., 2000). B: Nacreous layers in ammonite sample (an aulacostephanid) showing typical hexagonal tablets and vertical arrangement. C: Microstructure of calcitic foraminifera Lenticulina muensteri showing partial recrystallization of wall texture and test infilling. D: Microstructure of aragonitic foraminifera Epistomina tenuicostata showing wall texture and pyrite octahedra.

 
Stable isotopes of oxygen and carbon were determined (at the University of Plymouth) on a VG Instruments Optima Isotope Ratio Mass Spectrometer with a Multiprep Automated Carbonate System using 200–300 µg of carbonate. Isotopic results were calibrated against NBS-19. Isotopic data are reported using the per mil notation relative to the Vienna Peedee belemnite (VPDB) standard. Reproducibility for both {delta}18O and {delta}13C was better than ±0.1{per thousand}, based upon multiple sample analyses. All isotope and geochemical data are given in Table DR1 in the GSA Data Repository1.


   RESULTS AND DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
 RESULTS AND DISCUSSION
CONCLUSIONS
 REFERENCES CITED
 
The {delta}18Ocarbonate and {delta}13 Ccarbonate values are illustrated in Figure 3. The {delta}18O scatter is most pronounced within the otolith and ammonite data. Because fish and ammonites are nektonic organisms and growth is likely to occur periodically over several months or years depending on species, part of this scatter potentially arises from vertical and horizontal migration in conjunction with seasonal temperature variations. However, the analytical approach adopted here, whereby whole otoliths are analyzed, as well as molluscan fragments representative of the entire well-preserved portion of the shell, results in a degree of time-averaging of thermal information. Potential sourcing from the Ampthill and Kimmeridge clays may also introduce scatter within the data.


Figure 03
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  		Figure 3. Crossplot of {delta}18O and {delta}13C data from Jurassic fauna from Wootton Bassett.

 
Paleotemperatures were calculated using the equation of Anderson and Arthur (1983) for molluscan calcite and water, Grossman and Ku (1986) for the oxygen isotope fractionation between molluscan aragonite and water, and the isotope fractionation equation of Thorrold et al. (1997) for fish otoliths. Calculation of paleotemperatures from {delta}18O values of biogenic carbonate relies upon an estimation of the {delta}18Oseawater value. In environmental settings where continental ice volume is at a minimum and evaporation or freshwater inputs are minor factors, a sea water value of –1.0{per thousand} (VSMOW) is typically used (e.g., Anderson et al., 1994; Picard et al., 1998; Price and Mutterlose, 2004). The mean isotope values and paleotemperatures, shown in Table 1, for the oysters (15.6 °C) and belemnites (14.4 °C) are identical within statistical uncertainty. The mean paleotemperatures for ammonites (24.5 °C) and otoliths (28.9 °C) are on the other hand substantially higher and distinct from the oysters and the belemnites and also from each other. If a degree of homogeneity is assumed with respect to {delta}18Oseawater at this location, then these data define a depth-related temperature profile whereby the oysters and belemnites provide the coolest temperatures, indicative of bottom waters, and the ammonites and otoliths the warmest values, suggestive of near-surface waters. Such a predicted range of paleotemperatures is not inconsistent with other studies of the Jurassic (e.g., Anderson et al., 1994; Price and Page, 2008). The belemnites (Cylindroteuthis sp.) in terms of temperature would appear to be nektobenthic organisms, living alongside the oysters, rather than surface dwellers. Given the tight cluster of belemnite-derived isotope data (Fig. 3), a nonmigratory, territorial mode of life may also be inferred.


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  		TABLE 1. SUMMARY OF ISOTOPIC DATA

 
The fish otolith mean paleotemperature being higher than the ammonite temperature is consistent with previous studies where temperatures derived from fish (via {delta}18Ophosphate) gave higher temperatures than those inferred from carbonate {delta}18O data (e.g., Anderson et al., 1994; Pucéat et al., 2007). The otolith-derived temperatures are also consistent with other temperature interpretations based upon the oxygen isotope composition of phosphate from vertebrate tooth enamel from the Anglo-Paris Basin (e.g., Lécuyer et al., 2003). However, as a note of caution, applying an alternative temperature equation to the fish otolith data (e.g., Patterson et al., 1993) lowers the apparent temperatures by ~4 °C, thus bringing them into line with the mean ammonite paleotemperature. However, as the Thorrold et al. (1997) equation is based upon a marine species of fish raised over a range of temperatures, it would hence appear more appropriate to use in this study, as opposed to the equation of Patterson et al. (1993), which was based on empirical data derived from freshwater fish raised in aquaria or captured from well-characterized environments. Ultimately, the choice of equation governs the final temperature result and interpretation. Bearing this in mind, if our otolith temperature interpretations, based upon the Thorrold et al. (1997) equation, are robust (i.e., ~4 °C higher than ammonite-derived paleotemperatures), this suggests that paleotemperatures derived from the {delta}18Ophosphate of fish teeth (e.g., Kolodny and Raab, 1988) are also likely to be good estimates of ambient temperatures and not overestimates as a result of nonequilibrium fractionation as Anderson et al. (1994) suggest. Furthermore, our otolith-derived sea surface paleotemperature results add to the view that the Late Jurassic was a very warm interval in Earth history. Indeed, temperatures in excess of 30 °C at a paleolatitude of ~30°N rival temperatures associated with the middle Cretaceous thermal maximum. Such high temperatures are also consistent with data from general circulation models of Late Jurassic climate (e.g., Sellwood and Valdes, 2008). A number of studies (e.g., Hudson and Martill, 1991; Anderson et al., 1994; Price and Page, 2008) have, however, suggested, on the basis of stable isotope data, that an enhanced freshwater component of Jurassic surface waters may be potentially responsible for anomalously high surface seawater temperatures at this time.

The {delta}13Ccarbonate values illustrated in Figure 3 show that the ranges of most groups, with the exception of the otoliths, overlap, with values between –0.45{per thousand} and +4.32{per thousand}, typical of marine environments. Both types of foraminifera also fall within this range, despite being diagenetically altered. The preservation of "original" foraminifera {delta}13Ccarbonate values during diagenesis is quite common and is likely to be due to the buffering effect of carbonate carbon on the diagenetic system, as this is the largest carbon reservoir. The otoliths, on the other hand, sit outside of this cluster and range from –4.20{per thousand} to –1.96{per thousand}. Although there is no simple relationship between carbon isotopes and the environment, {delta}13C of dissolved inorganic carbon (DIC) tends to decrease from marine to estuarine or freshwater settings (e.g., Patterson and Walter, 1994). The otolith results from this study plot squarely within the field defined by Patterson (1999) as estuarine conditions. As we can be sure they were deposited within a marine environment, this isotopic pattern may well result from a mixing of marine signals with those derived from freshwater environments. This may point to the otolith carbon isotope data (and oxygen isotopes) being not fully representative of their final resting place (i.e., the otoliths belonged to fish that migrated between fresh and marine waters). This potentially has implications with respect to the reliability of using fish material for isotopic reconstructions, unless the migratory or nonmigratory nature of the organisms can be established. However, it should also be noted that the {delta}13C values of otoliths (and molluscan shell material) represent a combination of {delta}13C from DIC combined with varying amounts of highly negative {delta}13C from metabolically derived CO2. Hence, biogenic carbonates have {delta}13C values that typically range from near equilibrium to significantly less than that of the {delta}13C of DIC (Anderson and Arthur, 1983). For example, small departures from equilibrium fractionation toward more negative values have been reported in Nautilus macromphalus (Auclair et al., 2004) and bivalves (Klein et al., 1996). Furthermore, Thorrold et al. (1997) report much larger departures (depletions of between 3{per thousand} and 7{per thousand} relative to equilibrium with ambient DIC) in the {delta}13C values of fish otoliths, potentially making precise inferences regarding the {delta}13C of DIC values difficult (cf. Patterson, 1999). This may reflect the fact that the growth of an otolith occurs in a microenvironment surrounded by thick tissue and is thus farther away from ambient seawater than, for example, the shell margin of Nautilus or a bivalve. If such large departures seen in modern fish otoliths can be applied to the Jurassic, they can easily account for the very negative isotope values observed in this study. The study by Kalish (1991) suggested that fishes with low metabolic rates or those living at low temperatures had {delta}13C values deposited near equilibrium, whereas fishes living at higher temperatures showed extreme depletion in otolith {delta}13C. Trends were also evident in {delta}13C values whereby fishes with presumably higher metabolic rates were significantly more depleted in {delta}13C (Kalish, 1991). Hence, the interpretation of warm temperatures coupled with depleted {delta}13C values suggests that the Jurassic otoliths examined in this study belonged to fish with high metabolic rates. Based upon modern fish (e.g., Sherwood and Rose, 2003), Jurassic fish with high metabolic rates may be associated with generally active organisms with a much greater food or calorie requirement, while low rates would be associated with comparatively less active, benthic feeders.


   CONCLUSIONS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
REFERENCES CITED
 
This is the first detailed study to isotopically analyze Jurassic marine fish otoliths and to provide an independent scrutiny of fish-derived paleotemperatures. The {delta}18O compositions of Jurassic fish otoliths provide upper-ocean paleo-temperatures that are comparable with those derived from the isotopic analysis of fish tooth phosphates (e.g., Lécuyer et al., 2003). Otolith paleotemperatures in excess of 30 °C also compare well with temperatures associated with the middle Cretaceous thermal maximum, and are warmer than coexisting carbonates, also consistent with previous studies (e.g., Anderson et al., 1994; Pucéat et al., 2007). Although the carbon isotope data of the otoliths point to a freshwater influence and potentially migratory fish (e.g., Patterson, 1999), given the large departures from equilibrium fractionation toward more negative values reported from modern marine fish (e.g., Thorrold et al., 1997), we consider the temperature interpretations based upon otoliths to be robust and representative of the marine depositional environment. Depleted {delta}13C values, we believe, suggest that the otoliths examined in this study belong to fish with high metabolic rates.


   ACKNOWLEDGMENTS
 
We are indebted to Wiltshire County Council and Natural England for permission to access and sample the site. This paper benefited from reviews of an anonymous reviewer, Christophe Lécuyer, and Debbie Thomas.


   FOOTNOTES
 
GSA Data Repository item 2009058, Table DR1, is available online at www.geosociety.org/pubs/ft2009.htm, or on request from editing{at}geosociety.org or Documents Secretary, GSA, P.O. Box 9140, Boulder, CO 80301, USA. Back


   REFERENCES CITED
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 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES CITED
 

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Received for publication 25 July 2008

Revised manuscript received 23 October 2008

Manuscript accepted 24 October 2008




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