V O L . 1 2 0 N O . 5 / 6 M AY / J U N E 2 0 0 8 S UE • IN H S I S I • IN E U T SS T H IS IS S U China n r e t s We axes rel j a tarctic n A n i fire Ice on j umed h x e n sca Franci Mt. Diablo in E • IN T HIS I ISSN 0016-7606 Eocene climate record of a high southern latitude continental shelf: Seymour Island, Antarctica Linda C. Ivany† Department of Earth Sciences, Syracuse University, Syracuse, New York 13244, USA Kyger C. Lohmann Franciszek Hasiuk Department of Geological Sciences, University of Michigan, Ann Arbor, Michigan 48109, USA Daniel B. Blake Alexander Glass* Department of Geology, University of Illinois, Urbana, Illinois 61801, USA Richard B. Aronson Ryan M. Moody Dauphin Island Sea Lab, Dauphin Island, Alabama 36528, USA ABSTRACT A high-resolution record of Eocene paleotemperature variation is preserved within the high southern latitude, marine shelf succession of the La Meseta Formation on Seymour Island, off the NE side of the Antarctic Peninsula. 87Sr/86Sr ratios of bivalve shell carbonate indicate that the La Meseta Formation spans virtually the entire Eocene, and suggest the presence of an early middle Eocene unconformity. Paleoclimatic and paleoceanographic inferences are based on the stable oxygen and carbon isotope values of two genera of bivalves collected with a high degree of stratigraphic resolution within the formation, and with multiple replicate samples from each horizon. δ18O data indicate roughly 10 °C of cooling from the early Eocene climatic optimum (~15 °C) through the end of the Eocene (minimum ~5 °C), much of which took place in two comparatively short intervals at ca. 52 and ca. 41 Ma. Many features of the isotope curves generated from this Eocene shelf section are apparent in δ18O and δ13C data from the Southern and global oceans, including warm intervals that likely correspond to the early and middle Eocene climatic optima (EECO and MECO). † E-mail: [email protected] *Present address: Division of Earth and Ocean Sciences, Duke University, Durham, North Carolina 27708, USA A rapid middle Eocene shift to much more positive values is the most significant in the section and reflects a drop to universally cooler temperatures in the late middle and late Eocene that might also be associated with a short-lived glacial advance. However, even using a somewhat depleted value for δ18O of seawater in the Antarctic peninsular region, average Seymour Island shelf-water paleotemperatures did not reach freezing before the end of the Eocene. δ13C data similarly reflect the documented middle Eocene surface ocean enrichment followed by more negative values, but depletion is much more pronounced on Seymour Island and persists for the remainder of the Eocene, suggesting a combination of upwelling, metabolic effects, and/or atypical carbon cycling on the shelf in this region. Isotope data capture information about changes in the paleoenvironment that also had consequences for the biota, as published paleontological records document marked change in the nature of terrestrial and marine biota at this time. The fact that middle Eocene cooling and biotic turnover in the Peninsular region correspond well in time to the proposed initial opening of Drake Passage suggests that the formation of gateways, in addition to changes in pCO2, had significant consequences for the Earth’s climate system during the Paleogene. Keywords: stable isotope, Eocene, climate, bivalve, Antarctica, strontium. INTRODUCTION The Eocene Epoch encompasses the climate transition from the warmest interval of the Cenozoic through progressive cooling that culminated in the initiation of Antarctic glaciation in the earliest Oligocene. This fundamental shift led to the establishment of present-day oceanographic and atmospheric circulation patterns, and set the stage for evolutionary turnovers that shaped the modern marine biota. The global climate record of this interval is reasonably well constrained by studies of open ocean sequences (e.g., Zachos et al., 2001, and many others), but it remains unclear how representative these records are of conditions on continental margins, from which most of the macrofaunal marine fossil record is derived. This is particularly true at high latitudes, where exposed stratigraphic sections are rare, and where climate change was most severe. In this paper, we present a record of high-latitude shelf paleotemperatures from the Eocene La Meseta Formation on Seymour Island, Antarctica. The Seymour Island section is the only place known in all of Antarctica where Eocene shelf sediments are exposed in outcrop and in stratigraphic continuity. Because ongoing coring efforts on the shelf (Shallow Drilling on the Antarctic Continental Margin [SHALDRIL] and Antarctic Geological Drilling [ANDRILL]) have yet to successfully recover Eocene sediments due to the difficulties of drilling through ice and penetrating the overlying mantle of glacial debris, this section remains critical for GSA Bulletin; May/June 2008; v. 120; no. 5/6; p. 659–678; doi: 10.1130/B26269.1; 11 figures; 2 tables. For permission to copy, contact [email protected] © 2008 Geological Society of America 659 Ivany et al. low-water settings is needed. Carbonate macrofossils such as mollusks provide high-quality records of skeletal δ18O and the shelf environment in which they lived because sampling can be done within cross sections of shells, taking care to avoid any material that may be secondary or diagenetically altered—a luxury that is not possible with microfossils. Despite complications arising from variation in the δ18O of nearshore waters and depositional hiatuses imposed by relative changes in base level, these data can provide accurate estimates of shelf paleotemperatures that offer excellent complements to open ocean records and thereby help to complete the puzzle of climate history during this most interesting interval of Earth’s history. studies of high-latitude climate in the Paleogene greenhouse world. Our paleotemperature record is derived from the stable isotope values of bivalve carbonate collected throughout the section. Samples are well constrained stratigraphically and taxonomically, and include multiple replicate samples from each horizon, and thus provide for a significantly more detailed record than earlier work. 87Sr/86Sr ratios of mollusk carbonate allow samples to be positioned on the Eocene seawater Sr-evolution curve in order to augment existing biostratigraphically based age control. In addition, because the section is fossiliferous, these data allow for an integrated examination of the response of the Antarctic shelf biota to Eocene climate change, which we hope will facilitate more detailed studies of paleontological trends within the formation. A record of Eocene shelf paleotemperatures from the high southern latitudes will help to refine climate models and thereby further understanding of the mechanisms responsible for global climate cooling. High-latitude records are particularly necessary to constrain pole-to-equator thermal gradients and how they evolved over the course of the Eocene. In addition, the potential for δ18O values of planktonic foraminifera to underestimate surface-water paleotemperatures due to precipitation of secondary calcite in cold deep waters or pore waters (Zachos et al., 1994; Pearson et al., 2007) suggests that examining independent types of records from Eocene shal- km Seymour Island is located roughly 100 km east of the Antarctic Peninsula near its northern end (Fig. 1). Fossiliferous marine outcrops on the NE third of the island span much of the Eocene, and thus provide an ideal opportunity to study the effects of cooling on high-latitude, marine shelf environments and their associated faunas. The Eocene La Meseta Formation (Elliot and Trautman, 1982) is a shallow marine succession composed of sandstones, mudstones, and shell banks that accumulated in a variety of inner shelf environments (Sadler, 1988; Porębski, 1995, 2000; Marenssi et al., 1998, 2002). The 100 N 0 GEOLOGIC SETTING Seymour Island South Shetland Islands James Ross Island a nsul Peni rctic Anta Weddell Sea Figure 1. The location of Seymour Island in the James Ross Basin off the east side of the Antarctic Peninsula. 660 unit is confined within a fault-bounded NW-SE– trending basin on the east side of the Antarctic Peninsula. Underlying Cretaceous and Paleocene shelf sediments of the James Ross Basin were tilted gently to the SE by uplift associated with arc volcanism along the west side of the Peninsula, and dissected prior to the onset of La Meseta Formation deposition (Elliot, 1988; Macellari, 1988). The formation has been described as a compound, incised–valley-fill system (Porębski, 2000). Internal architecture is complex, with stacked and interfingering sediment lenses and occasional slumps of variable thickness. Consequently, and because the floor of the basin at the channel’s axis is not exposed, formation thickness has been difficult to measure, with estimates ranging from just over 300 m to an expanded composite section estimate of over 700 m (Elliot, 1988; Marenssi et al., 1998). Petrographic study of the La Meseta Formation (Marenssi et al., 2002) and diagenetic analysis of underlying units on the nearby island of Vega (Pirrie et al., 1994) demonstrate that the section has not been buried more than 1 km and has undergone minimal diagenetic alteration. Stratigraphy and Depositional Environment The original three-part lithologic division of the formation by Elliot and Trautman (1982) was expanded to seven distinct, mappable, lithologic units, or Telms (“Tertiary Eocene La Meseta”) by Sadler (1988). More recently, several authors have attempted to recognize genetically related, unconformity-bound, depositional sequences reflecting the interplay of fault-controlled subsidence and eustatic change in sea level. Porębski (1995, 2000) defined three large sequences, the boundaries of which he believes to be dominantly eustatically controlled. Smaller scale transgressive packages within, however, are more likely to be associated with tectonic processes, since the basin is several hundred meters deep yet facies are dominantly shallow marine throughout (Porębski, 1995). The six allomembers of Marenssi et al. (1998) are consistent with those of Porębski, but more finely subdivided. He sees eustasy as a more important control on deposition, and calls for fluvial erosion at the base of each sequence (Marenssi, 2006). A common feature to all, however, is the recognition of a significant depositional hiatus at the base of Telm 4, a surface associated with ravinement, perhaps following incision, and the generation of a glauconite- and phosphate-rich lag. Different authors have variously stressed estuarine, deltaic, and channel or valley fill interpretations for the depositional setting of different parts of the La Meseta Formation (e.g., Elliot and Trautman, 1982; Sadler, 1988; Geological Society of America Bulletin, May/June 2008 Eocene climate change on the Antarctic shelf Wiedman and Feldman, 1988; Stilwell and Zinsmeister, 1992; Porębski, 1995, 2000; Marenssi et al., 1998, 2002). The shallow-water, tidally influenced nature of component facies, the evidence for channelization, and the fault-bounded nature of the entire unit have led recent workers to favor the estuarine model (Porębski, 1995, 2000; Marenssi et al., 1998). While such interpretations imply (but do not necessarily require) an influx of fresh water from an associated river, the diverse nature of the invertebrate fauna preserved throughout the unit suggests accumulation under relatively normal marine conditions. An exploration of the variability in stable isotope values, both spatially and temporally, will help to resolve uncertainties in the interpretation of depositional setting. Note though that, because geochemical data are derived from carbonate macrofossils, they say nothing about the conditions that prevailed during deposition of intervening more barren intervals. While we made a concerted effort to sample every fossiliferous horizon we encountered in vertical sections, it is possible that erosional truncation along channel bases removed portions of the section that might be preserved elsewhere along strike. More extensive discussions of sedimentology, stratigraphic relations, and inferred depositional environments of stratigraphic subunits are found in Elliot and Trautman (1982), Sadler (1988), Stilwell and Zinsmeister (1992), Porębski (1995, 2000), and Marenssi et al. (1998, 2002), as well as below in the context of isotope geochemistry. Age Age control within the La Meseta Formation has been based primarily on biostratigraphy and suggests that deposition spanned much of the Eocene, but uncertainty remains about the age of particular units within the formation. Wrenn and Hart (1988) used dinoflagellate assemblages to identify a late early Eocene and a late-middle to late Eocene interval separated by the discontinuity at the base of Telm 4. Askin et al. (1991), Cocozza and Clarke (1992), and Askin (1997) support these bounds on the base and top of the unit with terrestrial and marine palynomorphs, although they place most of the formation within the middle Eocene. Diatoms in the lower part of the formation also yield an early Eocene age estimate (Harwood, 1985, 1988). Mollusk faunas support an Eocene age, with a struthiolariid gastropod from Telms 4 and 5 being indicative of the late Eocene (Stilwell and Zinsmeister, 1992). Hall (1977) used dinoflagellate assemblages to suggest a late Eocene to early Oligocene age for the top of the formation, and Fordyce (1989) concurred based on the presence of a cetacean skeleton with mysticete affinities (Mitchell, 1989). More recently, workers have used strontium isotope stratigraphy to augment the existing biostratigraphy. Dutton et al. (2002) present strontium isotope ratios from a limited number of mollusk shells from Telms 2, 5, and 7 that suggest early, middle, and late Eocene ages, respectively. Marenssi (2006) concurs with data from the base of Telm 2 that yield an early Eocene age. Dingle and Lavelle (1998) and Ivany et al. (2006) report strontium isotope ratios from shells at the top of the formation that are consistent with ages at or just below the Eocene-Oligocene boundary. Here, we report additional strontium isotope ratios from mollusks collected in stratigraphic order throughout the section in the hopes that they will help resolve some of the inconsistencies in age estimates within the formation. Previous Paleoclimate Interpretations Paleontological studies of La Meseta floral and faunal assemblages suggest cool to warmtemperate conditions during deposition of the unit (Askin and Fleming, 1982, 1997; Case, 1988; Doktor et al., 1996; Francis and Poole, 2002; Stilwell and Zinsmeister, 1992; Wiedman et al., 1988). The presence of fossil driftwood through much of the section (Fig. 2) demonstrates that conditions were mild enough to support the growth of significant temperate rain forests on the peninsula (Case, 1988; Francis and Poole, 2002; Poole and Cantrill, 2006). Dingle et al. (1998) used clay mineral assemblages to recognize a shift from warm, wet conditions to cooler and drier at the top of the formation. Paleotemperatures have been estimated using δ18O values of early marine cements (from the high-Mg calcite linings of Teredolites borings; Pirrie et al., 1998) and an assortment of invertebrate macrofossils (Gaździcki et al., 1992; Ditchfield et al., 1994) collected within the formation. All yield temperature estimates ranging between 5 and 15 °C and generally show cooling where there is stratigraphic control. Recently, Dutton and colleagues (2002) constructed a temperature record based on δ18O values of shell carbonate from individuals of the bivalve genus Cucullaea, in which they infer an overall pattern of Eocene cooling from ~15 °C in Telms 2–5 to ~10.5 °C in Telms 6 and 7. Shells were pooled from localities resolved only to the level of Telm, however, and hence yielded only six paleotemperature estimates for the Telm 2–7 section. Data Figure 2. Fossil wood collected from low in Telm 5 (locality 01–93), La Meseta Formation. This section of a large trunk was washed out to sea and its outside surface colonized by shipworms (wood-boring bivalves) while still floating, producing the trace fossil Teredolites. The wood ultimately sank and was preserved with a marine benthic fauna in a nearshore environment. Such material is common in the formation up through Telm 5, indicating the presence of forests on land. Telm—Tertiary Eocene La Meseta. 1 cm Geological Society of America Bulletin, May/June 2008 661 Ivany et al. ’W 40 Cape Wiman ’S 13 Telm 4 o 56 o 64 o 15 64 o 14 64 Bahia Lopez de Bertodano ’S (Paleocene) ’S ey Cross Vall presented here reinforce this overall trend toward cooler temperatures but, because of much higher stratigraphic resolution, we are able to identify structure in the data and offer a more detailed record of climate change during this important period of Earth history. The reader is referred to Dutton et al. (2002) for additional information regarding the preservation and chemistry of individual shells from this unit. Telm 5 Telm 3 Telm 2 o 56 Lan ding MATERIALS AND METHODS Strip Base Marambio Field Collection and Specimen Preparation 662 W Larsen Cove (Paleocene) Macrofossils were collected in situ from shelly facies within the La Meseta Formation on Seymour Island and their locations recorded using global positioning system (GPS) coordinates and/or positioned on the U.S. Geological Survey (USGS) topographic map 64056-T5-TM-010 produced in 1995. Relative stratigraphic position and correlation with other samples were determined in as much detail as possible. Localities are mainly confined to the north and west of the meseta that gives the formation its name (Fig. 3). Wherever possible, multiple localities were collected within a single stratigraphic horizon in order to assess heterogeneity that may reflect differences in time averaging or spatial variability in environmental conditions. Sampled horizons for which there are replicates within Telms 4, 5, most of 6, and 7 could be physically traced along strike. Two vertical sections were sampled within Telm 3 (Fig. 3, one to the west and one to the east) between which precise correlation could not be ascertained; their stratigraphic placement in the Results section is based on the similarity of the isotope trajectories between them. One locality within Telm 6 was stratigraphically ambiguous but its relative position inferred based on equivalent elevation with another site between which the beds dipped slightly, requiring that one be younger than the other. The most abundant and/or consistently present macrofossil groups were collected for isotope analysis, including the arcid bivalve Cucullaea (C. raea and C. donaldi) and the venerid bivalve Eurhomalea (E. antarctica, E. newtoni, and E. florentinoi), both shallow infaunal suspension feeders, the naticid gastropod Polinices subtenuis, a semi-infaunal predator of other mollusks, and the terebratellid brachiopod Bouchardia antarctica, an epifaunal suspension feeder (Fig. 4). Species within genera are similar morphologically and occur in the same facies, and thus are presumed to have very similar ecologies and physiologies. Fossils were collected from 54 localities distributed within 27 stratigraphic horizons throughout the formation. ’ 36 Telm 7 (Shoreline deposits, alluvium) Telm 6 Penguin Bay Telm 4 Alluvium, slump Telm 3 Telm 7 Telm 2 Telm 6 Telm 1 Telm 5 isotope sample locality 0 1000m Figure 3. Geologic map of the NE portion of Seymour Island (based on Sadler, 1988), showing units of the Eocene La Meseta Formation and locations of sampling localities for this study. The meseta that gives rise to the formation’s name is capped by alluvium overlying Telm 7 and is occupied by Argentina’s Base Marambio. Telm—Tertiary Eocene La Meseta. Individuals were sectioned along the maximum growth axis (or as much as could be revealed in a flat plane through the body whorl for naticids) and polished to reveal growth banding and internal architecture. Sections were visually evaluated for quality of preservation as indicated by presence of original crystal structure and absence of cracks and diagenetic cement. Selected sections were scanned using cathodoluminescent microscopy, and X-ray diffraction was performed on mollusk subsamples to ensure primary aragonite mineralogy. Cracks, discolored regions, or other apparent shell defects that may reflect diagenetic alteration were avoided. Only the best preserved specimens or portions of specimens were used for isotope analysis. Sampling All samples for geochemical analyses were collected from polished cross sections, not external shell surfaces. Initially, we sampled three to seven individuals of each taxon from each of two different collecting localities in order to assess sampling methodology and differences in stable isotope values among taxa. The two bivalve genera (Cucullaea and Eurhomalea) were selected for the remainder of the study because they were most consistently present through the section and their internal growth structure was the most straightforward for microsampling. For all remaining localities for which there is good stratigraphic control, three to five individuals from each bivalve genus (where possible) were prepared and sampled for stable isotope geochemistry. Estimates of average annual δ18O and δ13C for each sampled shell were attained by drilling a path of sampled carbonate across a number of growth bands so as to obtain an average “bulk” composition representing several years of growth. Three to five bulk samples were collected from each individual in this manner. Consecutive growth-band–parallel samples were also collected across a number of growth increments from two individuals of each bivalve taxon from the same locality in order to reveal differences in seasonal (intraannual) growth patterns. Shells were sampled on the high-precision Merchantek (New Wave) MicroMill computer-controlled milling systems at Syracuse University and at the University of Michigan. Resulting powdered samples were analyzed directly or split for paired stable isotopic and Sr-isotopic analysis. Geological Society of America Bulletin, May/June 2008 Eocene climate change on the Antarctic shelf B A C D Figure 4. Taxa used in this study for geochemical analysis. (A) the arcid bivalve Cucullaea raea; (B) the terebratellid brachiopod Bouchardia antarctica; (C) the naticid gastropod Polinices subtenuis; (D) the venerid bivalve Eurhomalea newtoni. Scale bars are 1 cm; same scale for B, C, and D. Strontium Isotope Analysis In an effort to improve temporal control within the La Meseta Formation, 87Sr/86Sr ratios of skeletal aragonite were analyzed from 43 bivalves distributed throughout the section (Telms 2–7). Samples for Sr isotopic analysis were dissolved in HNO3 and eluted through a quartz cation exchange column using Sr-specific resin (Eichrom). Between 50 and 100 ng of Sr were dried down, re-dissolved in 1 μL HCl, and loaded onto tungsten single filaments with Ta2O5 powder and 1 μL 0.3 M H3PO4. Analyses were performed on the Finnigan MAT 262 thermal ionization mass spectrometer at the University of Michigan’s Radiogenic Isotope Geochemistry Laboratory. Sample splits were retained for stable isotope analysis as well. While the marine 87Sr/86Sr curve (McArthur et al., 2001) shows only limited (and undulatory) variation throughout much of the Eocene, multiple skeletal values distributed throughout the section combined with stratigraphic constraints on their relative ages allow for the construction of a coherent chronostratigraphy into which to place stable isotope values and interpret climate change. Age assignments were made by fitting Sr-isotopic data to the 08/2004 version of LOWESS 3, the Phanerozoic Srisotope lookup table for global marine seawater (Howarth and McArthur, 1997; McArthur et al., 2001). Differences in inferred age for a given 87 Sr/86Sr ratio between this study and Dutton et al. (2002) result from the difference in timescale to which the global marine curve is calibrated; our data are plotted relative to the Gradstein et al. (2004) timescale. Best inferred ages are constrained by both 87Sr/86Sr ratios and relative stratigraphic position. When samples are plotted in stratigraphic order on the marine curve, best estimates of their ages are determined by adjusting age such that, wherever possible, values fall directly on the curve or their analytical errors intersect it. In cases where multiple ratios are derived from the same horizon, they are plotted such that they straddle the curve. Relative stratigraphic position is maintained in all cases. Error bars on age estimates derived solely from the LOWESS (locally weighted regression scatterplot smoother) fit, generally ~±2 m.y., are not plotted here, as stratigraphic position places more stringent constraints on age. Note though that poor knowledge of the marine curve during the early Eocene in particular introduces the greatest amount of error on these estimates. New data from Hodell et al. (2007) may help to resolve some of this uncertainty. Stable Isotope Analysis remove volatile contaminants and water. They were then reacted with anhydrous H3PO4 at 76 ± 2 °C in a Kiel automated carbonate preparation device coupled directly to the inlet of a Finnigan MAT 251 or 253 mass spectrometer at the University of Michigan’s Stable Isotope Laboratory. 17 O-corrected data are adjusted for acid fractionation and source mixing by calibration to a bestfit regression line defined by National Bureau of Standards (NBS)-18 and NBS-19 standards. Data are reported in ‰-variation relative to the Vienna PeeDee Belemnite (VPDB) standard. Measured precision is better than 0.1‰ for both δ13C and δ18O. The bulk-sample data set represents 907 analyses from 187 shells. An additional 288 samples were collected during high-resolution analysis of two specimens of Cucullaea and two of Eurhomalea. Paleotemperature Determinations Paleotemperatures are calculated using the empirically determined paleotemperature equation of Grossman and Ku (1986) for biogenic aragonite, adjusted to correct for the difference between “average marine water” and standard mean ocean water (SMOW) (e.g., Kobashi et al., 2003): Samples for stable isotopic analysis were roasted in a vacuum for one hour at 200 °C to Geological Society of America Bulletin, May/June 2008 T(°C) = 20.6 – 4.34 [δ18OARAGONITE – (δ18OSeawater −0.2)] 663 Ivany et al. Average global δ18OSW values are generally given as −1‰ for an ice-free Eocene world (Shackleton and Kennett, 1975; Zachos et al., 1994), and estimates of δ18OSW from paired δ18OCALCITE and Mg/Ca ratios of benthic foraminiferans also vary around −1‰ (Lear et al., 2004). The ocean is heterogeneous, however, and here we use a value of −2‰ for δ18OSW based on estimates from Huber et al. (2003) specific to the Eocene Antarctic Peninsula region derived from a coupled ocean-atmosphere climate model. Values may have been as low as −3‰ (Huber et al., 2003), and if we use this value, calculated temperatures are cooler by ~4.5 °C. If more positive δ18OSW values are employed to account for proposed accumulations of Eocene ice (e.g., Lear et al., 2000; Tripati et al., 2005), estimated temperatures are somewhat warmer. If local shelf water δ18O values were further depleted from mixing with fresh water, calculated temperatures will be too warm. It is unlikely that evaporative enrichment is a significant concern in this high-latitude setting. RESULTS Inter-Taxon Comparisons Stable isotope ratios of the four different taxa (two bivalves, one gastropod, and one brachiopod) from the same collections at two different localities were compared to evaluate potential 5 differences among taxa (Fig. 5). The naticid Polinices showed the greatest spread in values, with samples from the same location exhibiting at least 2‰ of variation in both carbon and oxygen isotope values. Their semi-infaunal habit may introduce more variation in δ13C due to differences in δ13C of dissolved inorganic carbon (DIC) between the water column and pore fluids, which might include more negative carbon derived from the decomposition of organic material. In addition, and perhaps more likely, their predatory trophic strategy and resulting difference in metabolic rate from suspensionfeeding taxa (Huebner and Edwards, 1981) may result in the incorporation of variable amounts of isotopically negative, respired CO2 into shell carbonate. Difference in δ13C of tissue for prey taxa of different trophic levels introduces yet another variable. The δ13C values of Polinices overlap the negative end of the range in variation exemplified by the other taxa, suggesting that one or more of these variables may play a role in its composition. It is unclear, however, why δ18O should show such a range, since the reservoir and its incorporation into shell material is not affected by biological processes. One possibility is growth rate—if Polinices were growing so fast that individual bulk samples did not in fact represent an average of multiple years, and/or if individuals stopped growing for variably long intervals of each year, and/ or if growth rates were at times fast enough Polinices Bouchardia Cucullaea Eurhomalea 4 to introduce kinetic disequilibrium effects, the δ18O values of our samples would show similar high variability. Rapid, seasonally interrupted, and prey-dependent growth is characteristic of modern Polinices from the coast of Massachusetts, USA (Edwards and Huebner, 1977), and may be true of Eocene taxa as well. This hypothesis requires additional high-resolution sampling to evaluate and, even if substantiated, it would rule out Polinices as a reliable and consistent recorder of annual average temperatures. Although they present an interesting paleoecological puzzle, for this reason they were avoided in construction of the long-term climate record despite their ubiquity in collections throughout the unit. The brachiopod Bouchardia shows a tighter clustering of δ18O data, but examination by cathodoluminescence revealed the common presence of secondary luminescent cement filling the punctae of shells. Hinge regions appear to be relatively free of such cement, suggesting that sampling difficulties could be overcome if focused on these regions, but even a small potential for contamination by isotopically depleted diagenetic cement makes them a less than ideal candidate taxon for long-term temperature reconstruction. In addition, the inconsistent relationship between data from brachiopods and the two bivalve taxa is difficult to explain, and so they were not investigated further for this particular study. 1 3 δ13C δ13C 0 2 -1 1 -2 0 B A -2 -1 0 1 2 -3 -2 -1 δ O 18 0 1 2 δ O 18 Figure 5. Differences in isotopic composition of bulk samples from different taxa at two localities within Telm 5. Polinices is a naticid gastropod, Bouchardia is an articulate brachiopod, Cucullaea is an arcid bivalve, and Eurhomalea is a venerid bivalve. Bouchardia has a calcite shell, the others are aragonite. Telm—Tertiary Eocene La Meseta. 664 Geological Society of America Bulletin, May/June 2008 Eocene climate change on the Antarctic shelf The bivalves Cucullea and Eurhomalea show roughly the same spread in δ13C values, and δ18O values are offset but to a similar degree in the two test localities. The straightforward nature of sampling with respect to the growth trajectory (a single cross section captures the whole life history, as opposed to naticids’ spiral growth), the lack of complicating diagenetic cement (as with the brachiopods), and the consistency of isotope results made these two genera the best taxa of choice for reconstruction of the long-term paleoclimate record. Cucullaea has a thick, easy to sample shell with clear growth banding, and has been used already for paleoclimate reconstruction (Dutton et al., 2002); therefore, these results will be directly comparable to this earlier work. Eurhomalea is a venerid bivalve, and venerids have been used often and with consistent success for stable isotope reconstruction of environmental conditions (e.g., Arthur et al., 1983; Jones and Quitmyer, 1996; Surge and Walker, 2006). The combination of the two taxa should therefore produce robust and reproducible results. An examination of bulk sample data from additional localities confirms that mean values from Cucullaea are ~0.4‰ more positive than those from Eurhomalea (Fig. 6), and hence will yield paleotemperatures cooler by ~2 °C. The origin of this taxon-specific offset was investigated through high-resolution microsampling within and across growth bands of two speci- mens of each bivalve from a single locality in Telm 5 (Fig. 7). This approach reveals that Cucullaea grows mainly during the Austral winter (Buick and Ivany, 2004), while Eurhomalea appears to capture more of the Austral summer. The average (bulk) composition of Eurhomalea shell carbonate is therefore more isotopically negative, yielding warmer average annual temperatures. Dark (translucent) growth bands, indicating slower growth due to physiological stress (Pannella and MacClintock, 1968), in Cucullaea correspond to more negative δ18O values and warm temperatures (Figs. 7A and 7B; Buick and Ivany, 2004), while in Eurhomalea they fall at δ18O maxima (cool temperatures, Fig. 7D). This is consistent with the truncation of the annual temperature cycle in the summer for Cucullaea but in the winter for Eurhomalea, giving rise to their different bulk isotopic compositions. The thin outermost layer of the shell in both taxa was often abraded or not preserved, and here the potential was greater for encountering diagenetic cements in surface cracks or altered shell in contact with matrix material. As a result, all samples for this study come from internal cross sections rather than the exposed shell surface. While most studies of shell geochemistry in Recent specimens have used material from the outer shell layer, Surge and Walker (2006) show that δ18O values from the middle shell layer are indistinguishable from those in the outer and hence provide a robust paleoclimate archive for ancient settings. In this study, bivalve carbonate was sampled from both the middle and inner shell layers. Compositions of microsampled carbonate from the middle and inner shell layers of the same specimen of Cucullaea (Figs. 7B and 7C) exhibit several differences. First, because the growth bands are much closer together in the inner shell layer, it is impossible to achieve here the temporal resolution attainable in the middle layer. Each individual sample is therefore more time-averaged, and hence the amplitude of the seasonal cycle is much reduced in the record from the inner shell. Secondly, the mean δ18O value for each region is identical, but δ13C is markedly more negative in the inner shell layer (Fig. 7C). This offset is likely due to the preferential incorporation of isotopically negative metabolic CO2 into the inner shell layer, where the mantle is in constant contact with the shell and where the seawater reservoir is more distant. While this poses no problem for the δ18O-derived paleoclimate record, it may introduce an added source of variation for the long-term δ13C record. Intra-Horizon Comparisons While some variation exists within and between shells from a given locality that reflects a combination of seasonal and interannual temperature variation during each time-averaged interval of deposition, the range of values exhibited by shells at different localities from Cucullaea Eurhomalea 2 * 1 δ18O * 0 * * * * * -1 -2 00 -1 * * 00 Figure 6. Comparison of δ18O values from co-occurring Cucullaea and Eurhomalea bivalves. With very few exceptions, Cucullaea is more positive than Eurhomalea. On average, the difference is 0.42 ± 0.1‰. Localities are not arranged in stratigraphic order. -11 0 -17 1- 04 1-87 1-12 1-29 1-30 1- 45 1- 46 1-53 1-54 - 004 - 005 - 600 0 0 0 0 0 0 0 0 0 0 03 03 03 Locality Geological Society of America Bulletin, May/June 2008 665 Ivany et al. Isotope Value (VPDB) 3.0 A Cucullaea 3 2.5 2.0 1.5 1.0 0.5 0.0 -0.5 0 2 4 6 8 10 12 14 Distance (mm) B 2.5 18 δ O Cucullaea 1 2.0 Isotope Value (VPDB) Isotope Value (VPDB) 13 δ C 3.0 1.5 1.0 0.5 0.0 -0.5 0 1 2 3 4 5 6 7 C Cucullaea 1 inner shell layer 1.5 1.0 0.5 0.0 -0.5 1 8 5 3.0 3.0 D Eurhomalea 4 2.0 1.5 1.0 0.5 0.0 40 25 50 60 Eurhomalea 5 0.5 0.0 -1.0 30 21 1.0 -1.0 20 17 1.5 -0.5 10 13 2.0 -0.5 0 E 2.5 Isotope Value (VPDB) 2.5 9 Sample Number Distance (mm) Isotope Value (VPDB) 16 0 5 Sample Number 10 15 20 25 30 35 Sample Number Figure 7. Microsampled isotope data from two specimens of Cucullaea (A–C) and two of Eurhomalea (D and E) from the same locality in Telm 5. Dashed horizontal lines at 1.5‰, 0.5‰, and −0.5‰ are merely to aid in visual comparison; note that the vertical axes for Eurhomalea (D and E) extend to more negative values than those for Cucullaea. Vertical gray bars in A, B, and D denote dark growth bands. Isotope values from both Cucullaea specimens are very similar (A and B), as are values from both Eurhomalea specimens (D and E). However, δ18O values of Eurhomalea extend to more negative (warmer) values than Cucullaea and do not record the more positive (cooler) values. Growth slow-downs or cessations marked by growth bands occur in the summer for Cucullaea but the winter in Eurhomalea. Inner shell-layer carbonate from Cucullaea 1 (C) shows a smaller range of variation than the middle shell layer (B) due to closer spacing of growth increments and lower temporal resolution of samples. Mean value of δ18O is similar between transects, but δ13C is markedly more negative in the inner shell layer, suggesting vital effects associated with incorporation of respired CO2. Telm—Tertiary Eocene La Meseta. 666 Geological Society of America Bulletin, May/June 2008 Eocene climate change on the Antarctic shelf the same stratigraphic horizon is nearly always statistically indistinguishable (Table 1). This is true even for shells collected from transgressive lag deposits that may represent significant intervals of time. Telm 4, equivalent to the base of the Cucullaea I Allomember of Marenssi et al. (1998), is one such horizon (“4.5” in Table 1). Another is a laterally extensive horizon marked by the abundance of small to moderate-sized Cucullaea bivalves found near the top of Telm 5 (“5.9” in Table 1). Unlike most other horizons, these two are easily recognized, and can be traced throughout much of the outcrop area. Their sedimentology makes them suboptimal for reconstructing climate information due to their potentially time-averaged nature, and therefore they were not heavily sampled for geochemical analysis. Nevertheless, the fact that shells from different localities yield similar overall ranges of values suggests that the amount of time represented by accumulated shell is not significant and/or does not encompass much secular variation in temperature. These results suggest that conditions across the depositional surface at a given time were reasonably consistent from place to place, and therefore that any temporal change in the average δ18O-derived paleotemperature is likely to represent a meaningful shift in Eocene peninsular climate rather than simply the incomplete sampling of a heterogeneous environment at each stratigraphic level. Stratigraphic and Strontium Isotopic Constraints on Age Before presenting stable isotope data in a stratigraphic context, we first explore the potential of strontium isotope ratios to provide additional age control within the formation. Secular variation in the 87Sr/86Sr ratio of marine waters is reflected in the chemistry of carbonate precipitates through time. In portions of the record where that ratio is well constrained and changing rapidly, the 87Sr/86Sr ratio of carbonate precipitated in fully marine seawater is an extremely useful tool for age control (e.g., Howarth and McArthur, 1997; McArthur et al., 2001). The Eocene Epoch, however, presents difficulties in that the marine ratio undulates around a value of ~0.70775; thus there is not a unique age solution for any single measured ratio. In addition, data constraining the marine curve for the early and early middle Eocene are very sparse (McArthur et al., 2001; but see new data from Hodell et al., 2007), and therefore assigned ages in this part of the section must be considered hypotheses to be tested with additional approaches. While unequivocally resolving Eocene ages using strontium isotope stratigraphy is a daunting task under the best of circumstances, the TABLE 1. ANALYSIS OF VARIANCE RESULTS FOR COMPARISONS OF MEAN δ18O VALUES BETWEEN CORRELATIVE LOCALITIES WITHIN A STRATIGRAPHIC HORIZON Stratigraphic Taxon Number of Number of F-statistic p-value horizon localities analyses 7.7 Eurhomalea 2 40 1.06 0.310 7.5 Cucullaea 2 39 0.79 0.380 7.5 Eurhomalea 2 45 7.03 0.011 6.3 Eurhomalea 3 29 2.66 0.089 6.1 Eurhomalea 5 56 1.03 0.399 5.9 Cucullaea 2 45 0.97 0.331 5.7 Cucullaea 3 36 12.83 0.000 5.7 Eurhomalea 3 46 3.00 0.060 5.3 Cucullaea 2 25 0.06 0.814 4.5 Cucullaea 3 14 0.78 0.481 3.5 Cucullaea 2 21 0.05 0.824 Note: Horizons are coded as Telm (“Tertiary Eocene La Meseta”) number with relative position indicated by the decimal. Only those horizons for which there were multiple localities where the same taxon was analyzed are included. Tests indicating significant difference between localities within horizons are marked by an asterisk (*). TABLE 2. 87Sr/86Sr RATIOS AND ANALYTICAL ERROR FROM LA MESETA FORMATION BIVALVES 87 Telm Stratigraphic Locality Sample Sr/86Sr Error Best inferred age horizon for horizon (m.y.) 7 7.7 01-82 C1 0.707780 0.000009 34.0 7.6 01-55 E2 0.707726 0.000014 37.3 7.4 01-99 C1 0.707741 0.000011 37.5 7.2 01-50 E2 0.707731 0.000011 39.1 6.4 01-10 E1 0.707748 0.000011 6 6.4 01-10 E3 0.707738 0.000020 41.0 6.2 01-63 E2 0.707799 0.000010 6.2 01-63 E3 0.707852 0.000010 6.1 01-68 E4 0.708158 0.000010 6.1 01-8 E2 0.707840 0.000013 5.9 01-73 C5 0.708243 0.000010 48.8 5 5.9 01-70 C1 0.707704 0.000016 5.8 01-35 E1 0.707732 0.000010 49.0 5.8 01-35 C3 0.707715 0.000010 5.7 01-29 C1 0.707781 0.000013 5.7 01-29 E4 0.708187 0.000013 5.7 01-30 C1 0.707874 0.000010 5.5 01-46 C1 0.707712 0.000010 49.4 5.5 01-46 E4 0.707733 0.000013 5.4 01-45 C2 0.707710 0.000012 50.8 5.4 01-45 E1 0.707708 0.000011 5.3 01-43b C1 0.707827 0.000011 51.0 4 4.8 01-42 C20 0.707769 0.000011 4.5 01-27 C20 0.707723 0.000011 52.5 3.8 00-17 C2 0.707771 0.000011 52.8 3 3.8 00-17 E1 0.707740 0.000012 3.5 01-12a E3 0.707717 0.000010 53.0 3.5 01-12 C1 0.707848 0.000012 3.5 01-12 E3 0.707762 0.000012 3.2 01-22 E1 0.707712 0.000010 53.2 3.1 01-23 E3 0.707698 0.000009 53.5 3.1 01-23 E1 0.707736 0.000011 3.1 01-23 E4 0.707791 0.000013 2 2.5 01-04 C3 0.707744 0.000011 54.0 2.5 01-04 C2 0.707738 0.000012 Note: Samples are arranged in stratigraphic order by locality. Telm—“Tertiary Eocene La Meseta”; C—Cucullaea; E—Eurhomalea. Best inferred age is determined by both 87Sr/86Sr and relative stratigraphic position. Error is ±2σ. See text for discussion. nearshore facies that characterize the La Meseta Formation pose additional concerns. Mixing of marine waters with freshwater carrying an 87 Sr/86Sr signature that differs from the marine value may be difficult to recognize given the undulatory nature of the curve during the Eocene. Without estimates of the 87Sr/86Sr ratio of Peninsula basement rocks and the concentration of Sr in Eocene runoff, it is impossible to determine how salinity variation would affect the 87Sr/86Sr ratio of nearshore carbonates (e.g., Bryant et al., 1995). Continental runoff is generally more radiogenic while Eocene marine 87 Sr/86Sr values are particularly nonradiogenic; therefore, the influence of freshwater influx could be significant. Nevertheless, while there is some variation, most La Meseta Formation samples produce isotope ratios that fall within the range for typical Eocene seawater (Table 2, Fig. 8A), suggesting that any salinity variation Geological Society of America Bulletin, May/June 2008 667 Ivany et al. Telm 7 Telm 6 Telms 4, 3, 2 Telm 5 0.70785 0.70780 87 Sr 86 Sr (unconformity) 0.70775 Eurhomalea Cucullaea 0.70770 Ivany et al. 2006 Dutton et al. 2002 A 35 40 45 Age (my) 50 55 7 0.70785 Sr 4 86 y = -0.17x + 13.95 r 2 = 0.78 5 y = -1.24x + 69.39 r 2 = 0.94 3 2 Sr 6 87 Relative Strat. Position 0.70790 y = -0.23x + 17.09 r 2 = 0.85 B 0.70780 0.70775 0.70770 C 0.70765 35 40 45 50 Age (my) is relatively minor. In addition, freshwater influx would be expected to affect not only the 87Sr/86Sr ratio, but also the stable isotope values of carbonate. There is, however, no consistent relationship between the 87Sr/86Sr ratio and the mean δ18O value of shell carbonate (Fig. 8C). Furthermore, the regular and low-amplitude variation in δ18O exhibited in microsampled bivalves is more consistent with seasonal temperature variation than with fluctuating salinity. Lastly, Dutton et al. (2002) demonstrate that multiple 87Sr/86Sr measurements on the same Cucullaea shell are statistically indistinguishable, suggesting little if any variation in salinity occurred over timescales of an individual animal’s lifetime. As this taxon can live for over a century (Buick and Ivany, 2004), the within-shell consistency of Srisotope values suggests sediment accumulation 668 55 -1.5 r 2 = 0.0042 -0.5 0.5 18 δ OVPDB in a marine environment of consistently normal salinity. These geochemical data therefore support interpretations based on faunal composition (Wiedman and Feldmann, 1988; McKinney et al., 1988; Blake and Zinsmeister, 1988; Stilwell and Zinsmeister, 1992; Filkhorn, 1994; Baumiller and Gaździcki, 1996; Blake and Aronson, 1998) and published geochemical data (Dutton et al., 2002) that, while shallow and nearshore, the La Meseta Formation accumulated under basically normal marine salinities. 87 Sr/86Sr ratios in the lower part of the section tend to be more ambiguous than those above. Telm 2 yielded values from four different specimens (two from this study and two from Dutton et al., 2002) that are virtually identical and pin the age of this part of the section at ~54 m.y. 87Sr/86Sr ratios from Telms 3–5, 1.5 Figure 8. (A) 87Sr/86Sr ratios from the La Meseta Formation relative to the global marine seawater curve (gray band encompasses 95% confidence limits around curve, LOWESS Version 4: 08/03; McArthur et al., 2001) and consistent with their relative stratigraphic positions. Error bars on the y-axis reflect analytical error. Associated error on age estimates during the Eocene generally approximates ±2 m.y.; those error bars are not plotted here for clarity, and due to additional controls on age by stratigraphic position that are not considered in the LOWESS (locally weighted regression scatterplot smoother) fit. (B) Age model for the La Meseta Formation based on the best estimates of age derived from A (Table 2). The section was divided into three intervals and best-fit lines were determined for each. Y axis values reflect relative stratigraphic positions (Telm numbers), so differences in slope do not directly relate to change in sedimentation rate. (C) Relationship between 87 Sr/86Sr ratio and mean δ18Ο value for sampled shells. Telm— Tertiary Eocene La Meseta. however, are much more variable. While the majority of values can be placed such that their analytical error bars intersect the marine curve, some horizons (e.g., the base of Telm 3) show a large range of variation in presumably correlative samples. Several samples are more radiogenic than expected, suggesting the potential for limited freshwater mixing. Acknowledging this uncertainty, 87Sr/86Sr ratios from Telms 2 through 5 are nearly all low enough to ensure that they represent the early Eocene minimum in marine 87Sr/86Sr values, rather than the latemiddle Eocene saddle in the marine 87Sr/86Sr curve. This suggests that deposition of the lower part of the formation took place in the early Eocene, an interpretation consistent with estimates derived from dinoflagellate (Wrenn and Hart, 1988; Cocozza and Clarke, 1992) and Geological Society of America Bulletin, May/June 2008 Eocene climate change on the Antarctic shelf diatom (Harwood, 1985, 1988) biostratigraphy. Telm 5, however, is apparently older than expected based on molluscan and microfossil biostratigraphy, only just crossing into the base of the middle Eocene. While Sadler (1988) and Stilwell and Zinsmeister (1992) had speculated that the basal unconformity of the La Meseta Formation may have been generated by a major sea-level lowstand at 49.5 Ma (Vail et al., 1980; Haq et al., 1987), these strontium isotope data confirm that deposition of La Meseta Formation sediments began several millions of years earlier. Miller et al. (2005) report an early Eocene eustatic rise following a relative Paleocene stillstand that may correspond to the initiation of La Meseta Formation marine deposition, but the potential competing influence of local tectonism makes this correlation tentative at best. A potential explanation for 87Sr/86Sr variability in the lower part of the section is the incorporation of small amounts of siliciclastic material as contaminants in samples of shell carbonate. Mollusks can entrain minute amounts of sediment in their shells as they grow, and siliciclastic sediments tend to be quite radiogenic. If a small amount of this material is leached during sample preparation, its influence on the 87Sr/86Sr of the sample could be significant. Mud is more likely to be incorporated than sand, and because the lower part of the section is muddier than above, this may explain the more variable ratios there. It does not appear that differing degrees of diagenesis within the section can explain the higher strontium variability in Telms 3–5. Preservation of shell material is equally good here as in the upper part of the formation, and shells tend to be larger and thicker, facilitating greater ease of microstructural evaluation and selective sampling. There has been some discussion, both informally and in the literature, about the stratigraphic and age relationships between Telms 2 and 3 (e.g., Sadler, 1988; Stilwell and Zinsmeister, 1992; Marenssi et al., 1998; Porębski, 1995, 2000). The muds and coarser grained lenses of Telm 2 may represent a deeper-water facies that everywhere predates and underlies Telm 3, and the lack of clear shallow-water sedimentary structures in Telm 2 supports this interpretation. Alternatively, the Telm 2 muds may instead be a lower-energy facies that accumulated along the margins of the trough that constrains the La Meseta Formation, while Telm 3 sands are at least in part laterally correlative to the Telm 2 muds and represent the higher-energy facies in the axis of the trough. The base of Telm 3 is not exposed in the axis, thus it is unknown whether there are Telm 2 muds below, but field relations suggest that the two units are at least in part correlative (Stilwell and Zinsmeister, 1992; Porębski, 1995). While the strontium isotope ratios from Telm 2 are consistent and unambiguous, the variation in ratios from Telm 3 makes age estimates of the two units overlap and, therefore, they are consistent with either scenario. Stable isotope values from Telm 2 material, however, are distinct from those in the lower part of Telm 3 (see next section), suggesting that the two units are not time equivalent, at least in portions of the island sampled for this study. Telm 4 clearly represents a ravinement surface and associated transgressive lag characterized by abundant phosphate pebbles and glauconite, and is conformably overlain by sediments of the ensuing Telm 5 transgression. The fact that fossils from Telm 4 include a significant number of reworked shells could explain the similarity of their ratios to values from Telm 3, although an effort was made to sample in situ (articulated) specimens. Because of the potential for reworking, this unit was not a focus of attention for geochemistry, and therefore data are limited, but immediately overlying shells from Telm 5 produce ratios that suggest an age for the base of the unit of ca. 51 Ma. Porębski (2000) and Marenssi (2006) propose that the Telm 4 disconformity may correspond to the previously mentioned 49.5 Ma sea-level lowstand. This is a plausible scenario, given the uncertainty associated with 87Sr/86Sr values in this setting. However, shells from multiple localities at the base of Telm 5 consistently yield values that correspond to the ca. 51 Ma minimum in marine strontium isotope ratios, and the Gradstein et al. (2004) timescale to which the marine strontium isotope curve is calibrated now places the (formerly 49.5 Ma) lowstand closer to 48.5 Ma. If this is correct, then there is no obvious manifestation of that eustatic lowstand in the La Meseta section. Pekar et al. (2005) note the presence of a sequence-bounding unconformity at 50.9 Ma on the Tasman Rise that likely correlates with a hiatus in the New Jersey, USA, shelf record (Miller et al., 1998), suggesting eustasy as a driver. This may correlate with the Telm 4 hiatus as constrained here, and if so, implies that this aspect of the Seymour Island section reflects eustatic sea-level change rather than regional patterns of tectonism and base-level change (see also Porębski, 1995). While the sedimentology of Telm 4 suggests a hiatus of some duration, strontium isotope ratios suggest no more than ~2 million years of section could be missing. Age control based on strontium isotopes is much better in the upper part of the formation. Specimens from Telm 7 and most of Telm 6 yield consistent and unambiguous ratios that establish an age ranging from late middle Eocene (ca. 41 Ma) up through virtu- ally the Eocene-Oligocene boundary (Fig. 8A; Ivany et al., 2006). That values fall consistently on the marine curve corroborates a normal marine inference based on sedimentology and stratigraphic relations (Porębski, 1995, 2000). Samples from the lowermost part of Telm 6 are radiogenic, suggesting the potential for freshwater mixing, but sediments appear to be in stratigraphic and temporal continuity with the overlying sediments, and we observed no lithologic differences sufficient to suggest any significant difference in depositional setting. This part of the section will be discussed in more detail below. A late-middle to late Eocene age for the upper part of the La Meseta Formation is consistent with existing biostratigraphy. Stratigraphic constraints on samples and comparison of their 87Sr/86Sr ratios with the global marine seawater curve suggest the presence of a middle Eocene (Lutetian) unconformity between Telms 5 and 6, spanning from roughly 48 to 43 Ma. Sedimentologic evidence for such a hiatus, however, is not immediately apparent. Scoured surfaces and concentrated shelly horizons are present throughout most of the section (Sadler, 1988; Porębski, 1995, 2000) but, aside from the base of Telm 4, none are associated with features that clearly record significant missing time. Shells in these layers are generally in good condition and often articulated, indicating much less time averaging than in Telm 4. A more laterally persistent shell bed with a scoured base is present near the top of Telm 5 (likely the base of Marenssi’s Cucullaea II Allomember; Dingle et al., 1998), suggesting a break in sedimentation, but lithofacies and biofacies as well as geochemistry clearly place it within Telm 5 rather than the overlying Telm 6 sands. The 87 Sr/86Sr ratios from shells in this bed (horizon “5.9” in Table 2) suggest that in fact the base of this layer may represent the aforementioned sea-level lowstand, now positioned at roughly 48.5 Ma. The short-lived nature of this particular lowstand (Haq et al., 1987) may explain the lack of a significant temporal hiatus evident in 87 Sr/86Sr ratios from below and above. The facies change from Telm 5 to Telm 6, marked by an influx of rust-weathering, sandbearing shell beds, low-angle trough crossbedding, and Ophiomorpha burrow networks, appears to correspond to the interval of missing time suggested by our strontium isotopic data. This is the most significant lithologic and facies shift in the section. Porębski (1995, 2000) notes the presence of an angular unconformity associated with the erosive base of a channel between Telm 6 strata and the underlying Telm 5, consistent with an interval of time not represented by sediment, but this is the first explicit suggestion of an unconformity of some Geological Society of America Bulletin, May/June 2008 669 Ivany et al. duration at this level. Miller et al. (2005) infer a substantial eustatic sea-level fall ca. 42–43 Ma based on backstripping data from the New Jersey coastal plain, and this may be responsible for the unconformity noted here. Field observations, however, suggest that Telm 5 and 6 lithologies may be interbedded in some areas, implying correlative ages not supported by 87Sr/86Sr data or the hypothesized unconformity. Porębski (1995) too notes that the unconformable contact between Telms 5 and 6 may in fact be conformable in the south-southwest part of the outcrop belt, and correlate to a horizon within what has been mapped as Telm 5. The transition, however, is poorly exposed due to slumping and hence it is difficult to rule out any single interpretation at this time. It is possible that the apparent gap in time recorded by strontium isotope values is at least in part a function of incomplete sampling, because shell beds are not consistently present throughout the unit and exposure was intermittent. Telm 5 sediments younger than those we recovered may be preserved in a different part of the section where they were not as severely truncated by the base of Telm 6. While Telm 7 is a bit finer grained and the bedding more even and laterally continuous than in Telm 6, both units appear to be shoreface deposits. The somewhat more variable and nearshore nature of Telm 6 may be associated with deposition in a bay-mouth bar (Porębski, 1995), while Telm 7 deposits accumulated just seaward. 87Sr/86Sr ratios give no indication of a significant time gap at the base of Telm 7, as suggested by Porębski (2000) from patterns of onlap and facies change. In the following section, isotope data are plotted stratigraphically using a combination of best age estimates from Table 2 and linear interpolation for intervening horizons from the age model in Figure 8B. Long-Term Stable Isotope Record and Eocene Paleotemperatures Within the temporal context provided by strontium isotope ratios and stratigraphic relationships, values of δ18O of shell aragonite indicate warm early Eocene temperatures with a maximum of ~15 °C near the base of Telm 3 (Fig. 9A). Telm 2 samples below (ca. 54 Ma) suggest slightly cooler temperatures, particularly if data from Eurhomalea are considered, although comparatively few samples constrain this inference (four shells total, only one of which is from Eurhomalea). Estimated temperatures drop within Telm 3 to ~10–11 °C, and remain essentially stable at this value through the end of Telm 5 (earliest middle Eocene). 670 Immediately above the middle Eocene unconformity, δ18O values decrease to −1.0‰ in the lower part of Telm 6, recording a shortlived return to temperatures more typical of the base of Telm 3 (~15 °C). Immediately above this (ca. 41 Ma), δ18O increases rapidly, reflecting a drop in temperature to ~7 °C by the end of Telm 6. Temperatures continue to cool into Telm 7 and reach an Eocene minimum value of ~5 °C by ca. 37 Ma. Upper Telm 7 (late Eocene) temperatures exhibit a modest warming back to ~7–8 °C. For comparison, water temperature around the Antarctic Peninsula today is ~−2 °C. With the possible exception of Telm 2, the overall trend in stable oxygen isotope values is consistent between the two bivalve genera examined (Fig. 9A), providing assurance that the observed secular pattern is robust. Interestingly, the characteristic offset between Cucullaea and Eurhomalea is less apparent in the upper part of the section. This could indicate that the taxonomic difference in preferred season of growth decreases upsection, or that the seasonal range of temperature variation had decreased to a point where summer and winter temperatures were not as distinct as in Telm 5, and therefore any seasonal bias in growth had only a minimal effect on the average δ18O value of shells. These hypotheses need to be evaluated with high-resolution microsampling of shells from Telm 7. While it may be tempting to interpret the range of isotope values from shells within any stratigraphic horizon as an indication of the seasonal range of temperature variation at that time, it should be remembered that this range is also influenced by the degree of time averaging within the horizon, the amount of interannual variation in temperature, and the range of temperatures over which the taxon produces shell material. Because all of these variables might change through the section, with or without any accompanying change in mean temperature or seasonal range in temperature, caution is advised in making inferences about changes in seasonality through time. Seasonality can only be confidently addressed with records from multiple microsampled shells distributed throughout the section, reinforced with complementary independent records based on paleobotanical (e.g., Francis and Poole, 2002; Gandolfo et al., 1998a, 1998b, in Case, 2007) or clay mineralogical (e.g., Dingle et al., 1998) data. Initial results of microsampling on shells in Telms 3, 5, and 7 suggest a significant decrease in seasonality in the coldest parts of Telm 7, associated especially with a decrease in summer temperatures (Ivany, 2007; Miklus, 2008). Stable carbon isotope values fall between −2.0‰ and +2.0‰ from Telm 2 up through the lower part of Telm 6 (Fig. 9B). A short- lived excursion to values near −6‰ takes place within Telm 6 above the negative values of δ18O and coincident with the shift back to more positive δ18O and cooler temperatures (ca. 41 Ma). Above this, δ13C values from Eurhomalea remain on the low side (~−3 to −4‰), while those from Cucullaea return to near zero by the top of the section. Variation in δ13C within a horizon is considerable, yet the seasonal range of δ13C in microsampled shells from Telm 5 is generally less than 2‰ (Fig. 7). The large spread of values within horizons could reflect a combination of real variation in the composition of DIC at the scale of time averaging and variation introduced by sampling from both the inner and middle shell layers of bivalves. Despite the variability within the section, the trend toward more negative values in Telms 6–7 is highly robust (p <.0001). PALEOENVIRONMENTAL IMPLICATIONS Data on stable isotopic values and strontium isotope ratios provide constraints on the depositional environment of the La Meseta Formation. Most importantly, the dominantly marine values of shell carbonate offer little evidence in support of a traditional deltaic or river-mouth estuary setting. The consistent range of δ18O variation and complete lack of any pronounced negative oxygen isotope values argues strongly against such an interpretation. In addition, no covariation exists between δ18O and δ13C values (see also Dutton et al., 2002), as would be expected with seawater-freshwater mixing (e.g., Ingram et al., 1996). Even at the base of Telm 6, where the potential for freshwater influx seems highest based on the negative δ18O values and radiogenic strontium ratios, mean δ18O values (constrained by 114 analyses from 29 clams at 10 different localities) are no more variable than elsewhere in the section. This is unlike what would be expected in a brackish-water setting, particularly at high latitude where precipitation can be expected to be more distilled and hence isotopically negative. Instead, these data in combination with the coast-normal, fault-bounded nature of the basin suggest accumulation in a relatively small, tectonically generated graben with minimal freshwater flow reflecting a limited catchment area. The general compositional maturity of sediments (Marenssi et al., 2002; field observations) suggests to us that sediments were likely delivered not by an associated river but instead primarily by longshore transport along the coast combined with mass wasting and localized runoff from valley margins. Sediments accumulated here due to subsidence associated with downdropping of the graben rather than Geological Society of America Bulletin, May/June 2008 Eocene climate change on the Antarctic shelf Ice-Free Temperature (°C) 5 10 15 Figure 9. Stable oxygen (A) and carbon (B) isotope values for all individual analyses from every bivalve shell from all horizons, plotted by taxon. Intensity of color reflects stacking of multiple data points. Age is plotted according to strontium isotope stratigraphy presented in Figure 8 and Table 2. proximity to the source of sediment delivery. The predominantly landward paleoflow directions in at least parts of the formation combined with the “conspicuous absence” of the expected fluvial deposits associated with lowstands (Porębski, 1995) and the strong tidal influence throughout the unit support this interpretation. The great challenge of working in shelf settings is determining which of the observed changes seen in the section can be attributed to local facies shifts and which reflect more significant trends driven by climatic and/or oceanographic change (even assuming that changes due to variable diagenetic alteration can be ruled out). Both can be manifest as changes in lithofacies, isotope values, and faunal composition, but the first might simply represent a shift in depositional setting along the shelf while the second indicates a wholesale change in conditions that affects all (or most) shelf environments simultaneously. Without widely distributed, correlative exposures both along strike and updip and downdip where appropriate, it can be difficult to distinguish the effects of these two possibilities. This is a consideration for the Seymour Island section, since the area of outcrop is limited and there is limited potential for finding correlative sections exposed elsewhere in the region (but see Askin et al., 1991). Nonetheless, we have sampled the same horizons along strike wherever possible so as to better understand the variation within the environment at the time of deposition. By comparing geochemical data with lithologic and faunal properties, we can begin to resolve differences between faciesimposed and climate-imposed trends. The shift in δ18O values from more negative toward more positive in Telm 3 likely reflects real climate change, as there is effectively no change in lithofacies associated with it. Both taxa exhibit a consistent change of a similar magnitude through time, the trend is maintained Geological Society of America Bulletin, May/June 2008 671 Ivany et al. through four different sampling horizons, and there is no difference in variability among horizons. This is more consistent with a trend from warm to cooler temperatures. Likewise, the secular variation in δ18O seen in the upper part of Telm 6 and all of Telm 7 is likely a real climate signal, because these samples all yield strontium isotope values that consistently fall on the marine curve and, for the most part, samples are from similar lithofacies. As discussed below, it is possible that some of this δ18O variation could be due to changes in ice volume, but it is not likely to be associated simply with facies shift on the shelf. The unusual isotopic variation in the lower part of Telm 6, however, is more difficult to resolve. There is a marked facies change to cross-bedded sand associated with the Telm 5–6 contact. The depleted oxygen isotope values and radiogenic 87Sr/86Sr ratios at the base of Telm 6 in association with this facies change suggest that all these phenomena could be due to a change in depositional setting to a locus more proximal to a freshwater source. However, the lack of evidence for riverine discharge elsewhere in the unit (see above) makes it difficult to argue for here. The Telm 6 δ18O depletion more likely reflects a real climate signal than a facies change. The fact that Cucullaea is virtually absent at sample sites during this interval might also be an indication of warmer (rather than fresher) water, since this taxon grows preferentially during the winter months and therefore might prefer cooler conditions. In general then, we maintain that most of the oxygen isotope variation exhibited in these data can be attributed to climate change rather than significant environmental and facies shifts along the shelf. Because there is little convincing evidence for significant freshwater influx to the depositional setting, relative changes in water depth or distance from open water are unlikely to result in changes in δ18O that could be misinterpreted as temperature change. PALEOCLIMATIC AND PALEOCEANOGRAPHIC IMPLICATIONS Antarctic Shelf Paleotemperatures These data provide the first taxonomically consistent record of paleotemperature variation with this level of stratigraphic resolution throughout the Eocene in an Antarctic marine shelf section. The overall trend in the La Meseta Formation is broadly consistent with the pattern of increasing δ18O and cooling temperature reported by Gaździcki et al. (1992) and Dutton et al. (2002), but these data reveal a 672 much greater level of complexity in δ18O and temperature through the Eocene section than anticipated from the earlier, lower resolution work. [Note that our inferred temperatures are cooler than Dutton et al. (2002) for equivalent carbonate isotope values due to a difference in assumed δ18O of seawater (−2‰ versus −1‰); see Fig. 11 for the effect on temperature of different presumed δ18OSW values.] Absolute paleotemperature estimates from these La Meseta Formation bivalves are in good agreement with data from high southern latitude surface waters summarized by Zachos et al. (1994). Several features of the global (Zachos et al., 2001) and Southern Ocean (Stott et al., 1990; Bohaty and Zachos, 2003) marine records of δ18O-based paleotemperatures are evident in this high-resolution, shelf data set. First, given the uncertainty in strontium isotope age estimates, it is likely that maximum temperatures near the base of Telm 3 reflect the early Eocene climatic optimum (EECO). Strontium isotope ratios from Telm 2 consistently yield age estimates of ca. 54 Ma, indicating that the EECO (roughly 51–53 Ma; Zachos et al., 2001) should be encompassed within the La Meseta Formation section. Maximum inferred temperatures in Telm 3 are ~15 °C, at least 4–5 °C warmer than earlier or later samples. The degree of warmth slightly exceeds that seen in the global deepocean record (Zachos et al., 2001). Inferred paleotemperatures at Seymour Island during the EECO are comparable to those inferred in the Arctic Ocean at the time based on TEX86 (~18 °C, Sluijs et al., 2006), but less than those suggested at the Paleocene-Eocene thermal maximum (PETM). Cooling on the Antarctic continent following the EECO at ca. 52 Ma is also inferred based on changes in the clay mineral assemblage of cores from the Southern Ocean (Robert and Kennett, 1992). Significantly, the age of Telm 2 samples, just postdating the PETM, suggests as well that this interesting event might also be recorded in Seymour Island sediments, perhaps manifest within Telm 1 and its unusual assemblage of calcitic fossils and deeply ironstained sediments [see Sadler (1988), Stilwell and Zinsmeister (1992), and Hara (2001) for descriptions]. The underlying Cross Valley Formation is upper Paleocene in age (Askin, 1988; Harwood, 1988). Inferred temperatures for the remainder of Telm 3, Telm 4, and Telm 5 are reasonably consistent, hovering around 10 °C for the rest of the early Eocene. The extreme but short-lived excursion toward negative δ18O values at the base of Telm 6 might also have a parallel in the open ocean, making the alternative explanation of a facies change to brackish water even less likely. Taken at face value, the δ18O shift indicates warming of shelf waters to temperatures comparable to those reached during the EECO, followed rapidly by ~7 °C of cooling. 87Sr/86Sr ratios from bivalves immediately above the negative excursion yield ages of 40.4 to 41.3 Ma, indicating that this warming could coincide with the middle Eocene climatic optimum (MECO, 41–42 Ma) observed in sediments deposited in the Southern Ocean on the Maud Rise and the Kerguelen Plateau (Bohaty and Zachos, 2003) (Figs. 10A and 10B). The amount of warming during this event on the Seymour Island shelf is ~5 °C, slightly more than that noted in Southern Ocean surface waters (Bohaty and Zachos, 2003). While the duration of warmth on Seymour Island is poorly constrained, samples that encompass it span a limited stratigraphic thickness (on the order of a few meters) and show no obvious evidence for disconformity within. The MECO as recognized in the Southern Ocean persists for less than a million years, but it is not inconceivable that such an event could be recognized in a shelf section. The pattern of variation in δ13C around the excursion is also consistent with, albeit more extreme than, that seen in the Southern Ocean (Figs. 10C and 10D) (Bohaty and Zachos, 2003), where a drop in δ13C occurs after peak MECO warmth is reached in the surface waters and coincident with the shift to cooler temperatures (see below). Whether this negative oxygen isotope excursion reflects warming temperatures, a flush of fresh water, or a combination of both, 87Sr/86Sr age estimates from immediately upsection indicate that the circumstances associated with the base of Telm 6 likely correlate with the MECO. Warmer temperatures might reasonably be expected to lead to an increase in precipitation and hence runoff, and so the negative δ18O values and shift to sandier sediments could represent a combination of both influences. Refinements in age estimates either in the La Meseta Formation or in the deep-sea record may call for an adjustment of this suggested correlation. However, there is no other clear interval of warm temperatures during the middle Eocene trend toward cooler temperatures represented in data thus far available from the Southern Ocean; therefore, it is likely that the excursion noted on Seymour Island correlates with the MECO, if indeed the negative Telm 6 δ18O values primarily reflect temperature. The ensuing shift toward substantially more positive values beginning ca. 41 Ma, hereafter the Middle Eocene Cooling, is the most significant interval of cooling in the Seymour Island section. Interestingly, Birkenmajer et al. (2005) report evidence for localized mountain glaciers in the South Shetland Islands that, while difficult to constrain, yield ages consistent with cooling at this time. In addition, Dallai et al. Geological Society of America Bulletin, May/June 2008 Geological Society of America Bulletin, May/June 2008 55 50 40 35 1.0 δ18O VPDB 0.0 -1.0 Eurhomalea Cucullaea A 2.0 1.5 0.0 -0.5 VPDB 0.5 δ18O 1.0 -2.0 0.0 VPDB δ13C 2.0 0.5 2.0 VPDB 1.0 1.5 δ13C Data from Bohaty and Zachos (2003) 2.5 (Holes 738B, 748B) Benthic (Holes 689B, 690B, 738B/C, 748B) Data from Bohaty and Zachos (2003) Fine fraction D (Holes 738B, 748B) Benthic (Holes 689B, 690B, 738B/C, 748B) -6.0 -4.0 C Fine fraction B Ice-Free Temperature (°C) 4 6 8 10 12 MECO 2 Figure 10. Stable oxygen and carbon isotope values of La Meseta Formation bivalves (plotted by genus) compared to data from the Southern Ocean (from Bohaty and Zachos, 2003). Pale symbols are mean values for each shell; bold symbols are average values and standard deviations for each horizon by taxon. (Α) δ18O values and calculated paleotemperatures using δ18OSW = −2‰; gray line is fitted through data from Eurhomalea, because values from that taxon more closely approximate mean annual temperature; (B) δ18O data from the Southern Ocean, showing middle Eocene climatic optimum (MECO); temperatures calculated using δ18OSW = −1‰; C); δ13C data from the La Meseta Formation, showing potential for divergence between taxa in Telm 7; (D) δ13C data from the Southern Ocean. Note the difference in the X axes between shelf and oceanic data, and that δ18O and δ13C axes increase in opposite directions. Telm—Tertiary Eocene La Meseta. Age (my) Late EOCENE Middle Early Ice-Free Temperature (°C) 5 10 15 Eocene climate change on the Antarctic shelf 673 Ivany et al. 35 -3 -2 δ18OSW -1 Late -4 50 (freezing) EOCENE Middle 45 (missing section) Early Age (my) 40 -4 -3 0 5 -2 Carbon Cycling on the Shelf -1 10 15 Temperature (°C) 20 Figure 11. Calculated paleotemperatures for a range of seawater compositions (−1‰ to −4‰). Mean temperatures do not reach freezing unless δ18OSW value exceeds −3‰. Winter temperatures presumably would be colder, however, and may support the existence of seasonal sea ice in the late middle and late Eocene. (2001) use δ18O and δD values of granitoid minerals altered by exchange with meteoric fluids in hydrothermal systems to suggest an interval of climate cooling in the Ross Sea of West Antarctica between 42 Ma and 38 Ma. Robert and Chamley (1991) and Robert and Kennett (1992) document an increase in illite at the expense of kaolinite and palygorskite in Southern Ocean cores, consistent with cooling during the middle Eocene (ca. 41 Ma). Thomas (2004) also suggests a link with high-latitude climate cooling to explain εNd data suggesting the reinstatement of Southern Ocean deepwater formation around 40 Ma. It is possible that the Middle Eocene Cooling inferred from Telm 6 oxygen isotope enrichment reflects not only falling temperatures but also limited growth of ice on the Antarctic continent at this time. Mg/Ca ratios from the Southern Ocean suggest the potential for ice growth on Antarctica starting ca. 40 Ma (Billups and Schrag, 2003). Tripati et al. (2005) have suggested a short-lived middle Eocene glaciation ca. 41 Ma based on benthic δ18O data from the equatorial Pacific Ocean. Edgar et al. (2007) note a correlative pattern in the equatorial Atlantic, but demonstrate that enrichment (≤0.6‰) is not enough to warrant the hypothesis (Tripati et al., 2005) of an ice sheet at both poles. Billups and Schrag (2003) similarly estimate that the ice volume contribution to the global δ18O signal at 674 is equivocal. The slight trend back to more negative δ18O values in bivalves preserved in the upper part of Telm 7 may reflect the interval of late Eocene warmth, or at least the interruption of cooling (Poag et al., 2003), recognized in both the Southern Ocean (Bohaty and Zachos, 2003) and the global benthic record (Zachos et al., 2001) just prior to the onset of glaciation in the early Oligocene. this time is ~0.4‰. A short-lived deepening of the carbonate compensation depth (CCD) noted in carbonate accumulation rates in the equatorial Pacific (Lyle et al., 2005) as well suggests that at least some ice growth occurred in concert with the middle Eocene positive δ18O shift, as glacioeustatic sea-level fall led to a shift in the locus of carbonate deposition from the shelves to the deep sea. However, even given that roughly 0.5‰ of the observed shift in δ18O on Seymour Island might be due to the growth of glacier ice in the interior of Antarctica, late middle and late Eocene mean temperatures are still universally cooler than in the earlier part of the record. As in the Southern Ocean record from Bohaty and Zachos (2003), minimum temperatures (most positive δ18O values) are attained ca. 37 Ma. This is roughly coincident with the earliest hypothesized evidence for glaciation in Greenland based on the presence of ice-rafted debris (Eldrett et al., 2007). Mean temperatures at this time approach 5 °C assuming a seawater δ18O value of −2‰, but would nearly reach freezing if seawater were 1‰ more depleted (Fig. 11). Since Huber et al. (2003) predict values as low as −3‰ east of the Peninsula from their early Eocene climate model, this is not out of the question. If so, conditions would likely be cool enough to support the existence of sea ice at least in the winter. The evidence for this in the sedimentologic record, however, The pattern of variation in δ13C through time on the shelf is less straightforward. As in the Southern Ocean (Bohaty and Zachos, 2003) and recently documented in the Italian Apennines (Jovane et al., 2007), δ13C values in the La Meseta Formation are generally positive during what we suggest to be the MECO (Figs. 10C and 10D), perhaps associated with increased burial of organic carbon during warm conditions. And like both of these records, values become more negative in concert with the subsequent cooling. However, the magnitude of this negative shift in δ13C on the Antarctic shelf is much larger and persists for much longer, making it clear that perturbations to the carbon cycle imposed by cooling were more pronounced or magnified in this shallow-water setting. Gaździcki et al. (1992) also recognize this significant shift toward more negative δ13C in the upper part of the La Meseta Formation, reporting values as low as −5‰ from the shells of both Cucullaea and Eurhomalea. Factors responsible for the large, late-middle Eocene negative shift in δ13C in the Seymour Island section are unclear, and more likely reflect aspects of the carbon cycle restricted to the shelf in this region. The long-term change to more negative values in the Southern Ocean following the MECO takes place synchronously in both surface and deep waters, indicating that it is likely due to a decrease in the proportion of organic carbon stored in the global sedimentary reservoir. This shift in the δ13C of exogenic carbon contributes to the pattern on the shelf, but only accounts for ~0.8‰ of the at least 6‰ shift in the Seymour Island section. The timing of the most precipitous drop in δ13C in both settings appears to correlate with a shift toward more radiogenic εNd values in the Atlantic Ocean, interpreted by Scher and Martin (2006) to reflect the initial opening of the Drake Passage and mixing of Pacific and Atlantic Ocean waters. That change in circulation might have led to episodes of high productivity associated with upwelling of nutrient-rich bottom water in some regions (Anderson and Delaney, 2005; DiesterHaass and Zahn, 1996). Although negative δ13C values are consistent with increased upwelling, Geological Society of America Bulletin, May/June 2008 Eocene climate change on the Antarctic shelf a phenomenon also not unexpected in association with cooling, values in the La Meseta Formation become significantly more negative than those recorded from benthic foraminifera in the deep ocean, indicating that upwelling of isotopically depleted deep water onto the shelf cannot account for them alone. An influx of isotopically negative dissolved inorganic carbon (DIC) from the terrestrial realm is a possible explanation, but one would expect such a pulse to correlate with more negative δ18O values associated with freshwater runoff, and this is not the case. Another potential explanation involves the formation of sea ice that limited air-sea exchange, allowing depleted carbon generated by decomposition to accumulate in the water column. However, if there was enough water movement on the shelf to transport and deposit the sand that characterizes much of Telm 6, then it is also likely that circulation was vigorous enough to mix and ventilate the waters. Another alternative relies on the ecology of the bivalves themselves. It is possible that, as temperatures cooled rapidly, physiological stress led to incorporation of a fraction of isotopically depleted metabolic CO2 in the shell. This would likely be a short-lived response to rapidly changing conditions, and natural selection should have produced adaptations to the cooler conditions relatively quickly, making it difficult to explain why values remain comparatively negative upsection. The fact that δ13C values for the two genera diverge further upsection in Telm 7 (Fig. 10C) does suggest that they did not incorporate carbon into their shells in the same way, potentially supporting this, but the hypothesis has yet to be evaluated. Implications for Meridional Temperature Gradients In addition to providing high-latitude shelf paleotemperatures through the Eocene for climate reconstruction, these data can be compared to those from tropical settings to quantify the evolution of the pole-to-equator thermal gradient on shelves during the course of global cooling. This gradient is important because it reflects the processes associated with heat distribution on the surface of the planet, and therefore has significant implications for our understanding of the climate system (Zachos et al., 1994). Pearson et al. (2001, 2007) report tropical Eocene shelf temperatures that hover around 30–33 °C for the duration of the epoch, warmer and surprisingly stable in comparison to those based on deep-sea cores (e.g., Zachos et al., 1994). Based on this, paleotemperatures derived from La Meseta Formation bivalves suggest that the meridional temperature gradient at the EECO approximates 15 °C, and increases to near 25 °C at the coolest part of the late Eocene. These values are consistent with those reported by Pearson et al. (2001), and somewhat exceed those of Zachos et al. (1994) due to underestimates of tropical sea surface temperatures. The steepening of this gradient has consequences for the biogeographic distribution of marine shelf organisms (e.g., Crame, 1997). These data combined with those of Pearson et al. (2007) predict that thermally induced biogeographic differentiation peaked in the early Eocene shortly after the EECO, and again in association with cooling in the middle Eocene. This pattern might not entirely hold true for the northern hemisphere, however, because similar data from the length of the Atlantic Ocean suggest that development of cooler conditions in the middle Eocene was a phenomenon restricted to high southern latitudes (Robert and Chamley, 1991). IMPLICATIONS FOR THE BIOTA One of our underlying objectives in researching the climate record of the La Meseta Formation is to better understand the conditions experienced by Antarctic marine macrofauna during the course of Eocene cooling. Presumably, temperature is an important variable driving evolutionary and ecological change at high latitudes, but relationships during this time interval have not been clearly demonstrated. Here, we show that the most pronounced temperature change in the section is the middle Eocene cooling, occurring low in Telm 6 following a brief excursion to warmer conditions. A review of the published paleontological literature suggests that faunal and floral change within the La Meseta Formation is also highest at the Telm 5–6 transition. In the marine realm, molluscan data from Stilwell and Zinsmeister (1992) indicate a drop in taxonomic diversity as high as 50% at this time. Feldmann et al. (2003) similarly see a marked shift in the composition of the decapod fauna. Among marine vertebrates, Long (1992) notes the virtual elimination of sharks above Telm 5 (see also Case, 1992), and Case (2007) reports a significant increase in the abundance of penguin and cetacean material in the upper part of the formation. The transition appears to have had ecological as well as taxonomic consequences. Aronson et al. (1997, 2007) and Aronson and Blake (2001) report a change in the nature of predator-prey relationships at roughly the Telm 5–6 contact due to the appearance of dense populations of predation-sensitive, epifaunal suspension feeders. Werner et al. (2004) also relate morphological data from mollusks suggesting a drop in the importance of shell-crushing predators and a significant decline in the frequency of predatory drilling by naticid gastropods. Change evidently extended to the terrestrial flora and fauna as well, based on specimens washed in and preserved in marine sediments, confirming that changes in the marine assemblage are not simply related to facies change on the shelf but reflect a wholesale change in environmental conditions in the Peninsular region. Case (2007) documents a substantial drop in vertebrate diversity going into Telms 6 and 7, with changes particularly noted among the mammal and bird faunas. We also note based on field observations that fossil wood is not seen anywhere in the section above Telm 5, suggesting that climate change could mark the elimination of significant forest cover from the Peninsula. Although we recognize an unconformity at the Telm 5–6 contact based on 87Sr/86Sr values, and there is a coincident facies change to sandier sediments that might also have affected taphonomic conditions (e.g., Feldmann et al., 2003), the composition and character of the biota is sufficiently distinct on either side of this transition that we do not believe the difference can be explained by the simple accumulation of evolutionary change during the time missing from the stratigraphic section or by a shift in the local depositional and preservational environment. We hypothesize that the middle Eocene cooling of roughly 10 °C imposed metabolic constraints upon organisms with which they had not been faced for several tens of millions years. Whether temperature itself was the proximal cause, or associated changes in productivity on the shelf (Aronson et al., 2007) were more important (e.g., Dayton and Oliver, 1977), cooling initiated substantial ecosystem reorganization associated with the new physiological challenges of dealing with cold temperatures. The excellent fossil record of the La Meseta Formation allows for a detailed examination of the specifics of this relationship. We anticipate that future work will test aspects of this hypothesis and clarify the mechanisms through which climate change imposed such consequences for the Antarctic biota. CONCLUSIONS The agreement of our δ18O data from nearshore peninsular Antarctic habitats with those from the Southern Ocean, in both general pattern as well as detail, affirms the use of shallow shelf carbonate fossils as useful archives for paleoclimate reconstruction. In addition, the La Meseta Formation record demonstrates that climate perturbations recorded as far away as the Kerguelan Plateau (Bohaty and Zachos, 2003) were ubiquitous, high southern latitude phenomena, experienced on the shelf as well as in the open ocean. Likewise, aspects of the global Geological Society of America Bulletin, May/June 2008 675 Ivany et al. Eocene climate record are also evident in this high-latitude shelf section. Acknowledging the limitations of strontium isotopic age control, the most prominent events in the Seymour Island climate record include the EECO and subsequent cooling at ca. 52 Ma, the middle Eocene warming that we tentatively correlate with the MECO, and most prominently, Middle Eocene Cooling ca. 41 Ma. The cause(s) of the middle Eocene shift to warmer conditions and the subsequent reversal into Middle Eocene Cooling remains a matter for discussion. Bohaty and Zachos (2003) rule out methane release as the proximal cause for MECO warming because their transient δ13C excursion does not happen until after warming has already begun. They instead suggest a tectonic trigger, calling for CO2 release and greenhouse warming associated with ridge or arc volcanism. Scher and Martin (2006) suggest that cooling immediately following the MECO is coincident with the opening of the nascent Drake Passage, and that associated changes in circulation and productivity led to a drawdown of CO2 that may have accelerated cooling and brought about the proposed transient middle Eocene glaciation. It is possible that tectonism around Antarctica led to both these phenomena, first releasing CO2 to cause warming, and then, once a threshold is breached to allow for circum-Antarctic circulation, facilitating CO2 drawdown and cooling by subsequent upwelling and phytoplankton production. While decreasing CO2 is now favored as the trigger for rapid ice growth near the Eocene-Oligocene boundary (DeConto and Pollard, 2003), the correlation between the Middle Eocene Cooling recognized here in the Peninsular region and the opening of a shallow-water connection through the Drake Passage suggests that gateways nevertheless might have played an important role in global Paleogene cooling. ACKNOWLEDGMENTS We thank John Evans and Kurt Burmeister for help in the field, and Devin Buick, Nicole Miklus, Allison Cattani, and Cristina Story for microsampling bivalves in the lab. John Schue and Devin Buick evaluated growth banding in Eurhomalea and Cucullaea, respectively. Lora Wingate conducted all stable isotope analyses. Devin Buick and Jim Brower helped with specimen photography. Jim Zachos and Steve Bohaty offered thoughtful comments early on and made their data available. Bruce Wilkinson and Ellen De Man read drafts of the manuscript. Steve Bohaty and Greg Ludvigson reviewed the paper and made many helpful suggestions, and Hope Jahren handled the paper for Geological Society of America Bulletin. This research was supported by grants from the National Science Foundation (NSF)’s Office of Polar Programs to Ivany and Lohmann (OPP-0125409 and OPP-0125589), and a Research Experience for Undergraduate (REU) Program supplement to Ivany. 676 Fieldwork in the winters of 2000–2001, 2001–2002, and 2003–2004 was supported by NSF grants to Aronson and Blake (OPP-9908828, OPP-9908856, and ANT-0245563). Data from this study are available from the Antarctic Master Directory, under Paleoclimate–Land Records–Isotopes. REFERENCES CITED Anderson, L.D., and Delaney, M.L., 2005, Middle Eocene to early Oligocene paleoceanography from Agulhas Ridge, Southern Ocean (Ocean Drilling Program Leg 177, Site 1090): Paleoceanography, v. 20, p. PA 1013, doi:10.1029/2004PA001043. 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