2014 Mestas NunŢez 2014
2014 Mestas NunŢez 2014
2014 Mestas NunŢez 2014
Paleoceanography
RESEARCH ARTICLE A mechanism for freshening the Caribbean Sea
10.1002/2013PA002515
in pre-Ice Age time
Key Points: Alberto M. Mestas-Nuñez1 and Peter Molnar2
• Caribbean to Pacific moisture transport
decreases during El Niño events 1
Department of Physical and Environmental Sciences, Texas A&M University-Corpus Christi, Corpus Christi, Texas, USA,
• El Niño-like SSTs could account for a less 2
saline Caribbean before Ice Age time Department of Geological Sciences and Cooperative Institute for Research in Environmental Sciences, University of
• These results are consistent with an Colorado Boulder, Boulder, Colorado, USA
Isthmus of Panama long before 4 Ma
Abstract Many believe that the Central American Seaway closed near 4 Ma and that that closure led
to increased salinity in the Caribbean Sea and stronger Meridional Overturning Circulation in the Atlantic,
Correspondence to:
A. M. Mestas-Nuñez,
which facilitated the waxing and waning of ice sheets in the Northern Hemisphere. We offer an alternative
alberto.mestas@tamucc.edu explanation for Caribbean salinification. The atmosphere transports approximately 0.23 Sv (1 Sv = 106 m3s 1)
of fresh water (moisture) from the Caribbean to the Pacific today, but that amount varies by >20% during
Citation:
El Niño–Southern Oscillation events. Regressions of moisture transport against the Niño-3 index, a measure
Mestas-Nuñez, A. M., and P. Molnar of the sea surface temperature in the eastern tropical Pacific, show less moisture transport from the Caribbean
(2014), A mechanism for freshening the during El Niño events than average. Abundant evidence indicates that at 3–4 Ma the eastern tropical Pacific
Caribbean Sea in pre-Ice Age time,
Paleoceanography, 29, 508–517,
was 3.5–4°C warmer than today, and if so, an extrapolation of such regressions suggests that smaller moisture
doi:10.1002/2013PA002515. transport across Central America might account for paleoceanographic inferences of a smaller salinity difference
between the Caribbean and Pacific at that time. Accordingly, that decreased salinity difference at ~3–4 Ma
Received 27 MAY 2013 would not require blockage of relatively fresh Pacific water at ~2–4 Ma by the closure of the Central American
Accepted 9 MAY 2014
Accepted article online 14 MAY 2014
Seaway, but rather would be consistent with a transition from El Niño to La Niña-like conditions in the eastern
Published online 11 JUN 2014 tropical Pacific around that time.
1. Introduction
Many discussions of both modern climate and paleoclimate assign a key role to the Atlantic meridional
overturning circulation (AMOC), and the high salinity of surface water exiting the Caribbean Sea and flowing
northward into the North Atlantic seems to be an essential ingredient of this circulation. Accordingly,
hypotheses for climate changes on virtually all timescales appeal to processes that affect that salinity. Many
have inferred that a closing of the Central American Seaway (1) blocked fresher water from the Pacific and
allowed saline water to accumulate in the Caribbean (Figure 1a) before moving northward, and then (2) the
change from fresher to more saline surface water in the Caribbean played a crucial role in the transition from
essentially no Northern Hemisphere ice sheets to recurring ice ages since ~ 3 Ma [e.g., Bartoli et al., 2005;
Berggren and Hollister, 1974; Haug and Tiedemann, 1998; Kaneps, 1979; Keigwin, 1982; Sarnthein et al., 2009;
Weyl, 1968]. In this paper we present an alternate mechanism for the salinification of the Caribbean ~3–4 Ma
that does not depend on a simultaneous closing of the Central American Seaway. We first review the
paleoceanographic evidence for the emergence of the Seaway. Then we discuss the evolution of the
temperature and salinity differences between the Pacific and the Atlantic Oceans. Finally, we review the
relationship between El Niño–Southern Oscillation (ENSO) and interocean moisture fluxes and use it to
account for the evolution of the interocean salinity contrast.
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1
during December–March and significant positive
correlations during July–August (Figure 3). Results
0.5
for the Niño-3.4 index are similar.
Estimates of the linear regression coefficients
Correlation
between Atlantic-to-Pacific moisture flux
0
anomalies in g s 1 and Niño-3 SST anomalies in
°C using data for the months with significant
correlations (see Figure 3) are shown in Figure 4.
−0.5
The regression coefficient for December–March
is 4.59 ± 1.88 × 1010 g s 1 °C 1 (Figure 4a) and
−1 for July–August is 4.52 ± 1.68 × 1010 g s 1 °C 1
J F M A M J J A S O N D (Figure 4b). The overall regression coefficient
Figure 3. Correlation coefficients between Atlantic-to-Pacific
estimated by combining data from the
monthly moisture flux anomalies and Niño-3 monthly SST December–March and July–August periods is
anomalies for all calendar months. Correlations that are significant 2.45 ± 2.02 × 1010 g s 1 °C 1 (Figure 4c). A
with greater than 95% confidence are indicated with solid circles. similar value of the overall regression coefficient
( 2.12 × 1010 g s 1 °C 1) can be obtained by first
computing the regression coefficients for each of the individual months involved (December–March, July,
and August) and then taking their average. The value 0.0245 Sv °C 1 for Niño-3 when multiplied by ~3.5–4°C
is ~0.086–0.098 Sv, which is roughly half of the 0.2 Sv inferred by Schmittner et al. [2000] for moisture
transport from the entire Atlantic (20°S–20°N) and hence not exclusively across Central America, during
El Niño events. Prange et al. [2010] exploited
11 DJFM similar logic to explain changes in salinity
x 10
Moisture flux (g sec−1)
2
Atlantic. These latter three fluxes, however,
0 do not affect the salinity of the Atlantic, for
all of this moisture does not represent a
−2 net loss/gain of Atlantic’s water. Moreover,
b the high flux of surface water through
−4
the Caribbean, ~28 Sv [Johns et al., 2002],
−2 −1 0 1 2 3
suggests that surface water ~150 m deep
x 1011 DJFMJA and ~1000 km wide entering the Caribbean
Moisture flux (g sec−1)
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Paleoceanographic data from the Ceara Rise, ~4°N and north of the mouth of the Amazon, suggests that
near 4 Ma, that region became less saline [Billups et al., 1998; Ravelo and Wara, 2004; Steph et al., 2006a].
Consistently, using Xie and Arkin’s [1996] precipitation data set, Saravanan and Chang [2000] showed that
during El Niño events a band of enhanced precipitation lies north of an arid region over and east of the
Amazon Basin. One might imagine that the freshening north of the mouth of the Amazon near 4 Ma and
the northward flow of surface water might reduce salinity of the North Atlantic. Estimates of the stream
function for flow of surface water through the Caribbean, however, suggest that most of the water entering
the sea comes from the North Atlantic [Johns et al., 2002]. Thus, the freshening of equatorial water near the
Amazon Basin does not seem likely to have affected the salinity of the North Atlantic.
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In this study, the paleo-observation that a salinity gradient between the Pacific and Atlantic developed
around ~4.0 Ma is based on the assumption that the Caribbean salinity is a good representation of the
tropical Atlantic. As such, our results seem to be in contrast with those of Schmittner et al. [2000], who showed
enhanced rather than reduced freshwater flux out of the Atlantic during warm ENSO events. Schmittner et al.,
however, based their inference on surface freshwater flux (precipitation minus evaporation) estimates from
the National Centers for Environmental Prediction (NCEP)-National Center for Atmospheric Research (NCAR)
reanalysis [Kalnay et al., 1996], which are known to be of poor quality. Indeed, Mestas-Nuñez et al. [2005]
showed that although the freshwater fluxes from the NCEP-NCAR reanalysis were not useful for moisture
budget studies of the Caribbean and Gulf of Mexico, the vertically integrated moisture fluxes from the same
reanalysis were acceptable, and those are the ones used in the present study. In addition, Schmittner et al.
did not consider the seasonality of the ENSO relationships.
7. Conclusions
During El Niño events, atmospheric moisture transport from the Caribbean Sea to the Pacific Ocean decreases
in winter and increases in summer, but such that less moisture leaves the Caribbean during El Niño than
during most years. Regressions of such moisture transport against the Niño-3 SST anomaly index quantify
these relationships (Figure 4).
Paleoceanographic evidence suggests that at 3–4 Ma the eastern tropical Pacific Ocean was warmer by ~4°C
than it commonly is today [e.g., Dekens et al., 2007; Lawrence et al., 2006; Wara et al., 2005], but comparable to
what it is during large El Niño events, and that the difference in salinity between the Pacific and Caribbean
was smaller by ~1‰ [e.g., Steph et al., 2006a]. Extrapolations of the regressions to Niño-3 indices of 2°C and 4°C
suggest that moisture transports would have been 21–42% less than the present-day average. The effects of
such moisture transport on Caribbean salinity are similar in magnitude to those estimated from ocean GCM
runs with only shallow Central American Seaways. Reasonable extrapolations of the regressions, to 0.049 or
0.098 Sv, do not yield such a large value of atmospheric moisture transport, but they show them to be of the
order of magnitude that might yield a large difference in climate.
Recent geologic studies suggest that the Isthmus of Panama may have formed as long ago as 15–20 Ma [Montes
et al., 2012a, 2012b]. Lower moisture transport across that region because of a warmer eastern Tropical Pacific
at ~3–4 Ma might account for the smaller salinity difference between the Caribbean Sea and Pacific Ocean at
that time. A later transition to La Niña-like conditions and associated increase in moisture transport from the
Caribbean to the Pacific could account for the salinification of the Caribbean ~2–4 Ma without invoking a
simultaneous closing of the Central American seaway. Our proposed mechanism for Caribbean freshening in
pre-Ice Age times may bear on future climates. In a warming world, the zonal interocean exchange of moisture
at low latitudes may contribute to the slowdown of the AMOC.
Acknowledgments References
One of the authors (P.M.) was stimulated
Allmon, W. D., G. Rosenberg, R. W. Portell, and K. S. Schinder (1993), Diversity in Atlantic coastal plain mollusks since the Pliocene, Science,
by discussions with C. Montes and R.F.
260, 1626–1629.
Stallard. A.C. Ravelo provided advice on
Allmon, W. D., G. Rosenberg, R. W. Portell, and K. Schindler (1996), Diversity of Pliocene-recent mollusks in the western Atlantic: Extinction,
how to structure our arguments, and
origination, and environmental change, in Evolution and Environment in Tropical America, edited by J. B. C. Jackson, A. F. Budd, and
she, J.C.H. Chiang, and an anonymous
A. G. Coates, pp. 271–302, Univ. of Chicago Press, Chicago, Ill.
reviewer offer constructive criticism of
Barreiro, M., G. Philander, R. Pacanowski, and A. Fedorov (2006), Simulations of warm tropical conditions with application to middle Pliocene
the manuscript. The research was
atmospheres, Clim. Dyn., 26, 349–365.
supported in part by the National
Bartoli, G., M. Sarnthein, M. Weinelt, H. Erlenkeuser, D. Garbe-Schönberg, and D. W. Lea (2005), Final closure of Panama and the onset of
Science Foundation (NSF) under grant
northern hemisphere glaciation, Earth Planet. Sci. Lett., 237, 33–44.
AGS-1136466 and while the other author
Berggren, W. A., and C. D. Hollister (1974), Paleogeography, paleobiogeography and the history of circulation in the Atlantic Ocean, in Studies
(A.M.) was serving at NSF. Any opinion,
in Paleo-Oceanography, Spec. Publ. No. 20, edited by W. W. Hay, pp. 126–186, Soc. Econ. Paleontol. Mineral., Tulsa, Okla.
findings, and conclusions or recommen-
Billups, K., A. C. Ravelo, and J. C. Zachos (1998), Early Pliocene climate: A perspective from the western equatorial Atlantic warm pool,
dations expressed in this publication
Paleoceanography, 13, 459–470, doi:10.1029/98PA02262.
are those of the authors and do not
Brierley, C. M., and A. V. Fedorov (2010), Relative importance of meridional and zonal sea surface temperature gradients for the onset of the
necessarily reflect the views of NSF.
ice ages and Pliocene-Pleistocene climate evolution, Paleoceanography, 25, PA2214, doi:10.1029/2009PA001809.
Budd, A. F., K. G. Johnson, and T. A. Stemann (1996), Plio-Pleistocene turnover and extinctions in the Caribbean reef-coral fauna, in Evolution and
Environment in Tropical America, edited by J. B. C. Jackson, A. F. Budd, and A. G. Coates, pp. 168–204, Univ. of Chicago Press, Chicago, Ill.
Cane, M. A., and P. Molnar (2001), Closing of the Indonesian seaway as a precursor to east African aridification around 3–4 million years ago,
Nature, 411, 157–162.
Cannariato, K. G., and A. C. Ravelo (1997), Plio-Pleistocene evolution of eastern tropical Pacific rate of circulation and thermocline depth,
Paleoceanography, 12, 805–820, doi:10.1029/97PA02514.
MESTAS-NUÑEZ AND MOLNAR ©2014. American Geophysical Union. All Rights Reserved. 514
Paleoceanography 10.1002/2013PA002515
Chaisson, W. P., and A. C. Ravelo (2000), Pliocene development of the east–west hydrographic gradient in the tropical Pacific, Paleoceanography,
15, 497–505, doi:10.1029/1999PA000442.
Chiang, J. C. H., and B. R. Lintner (2005), Mechanisms of remote tropical surface warming during El Niño, J. Clim., 18, 4130–4149.
Coates, A. G., and J. A. Obando (1996), The geologic evolution of the Central American Isthmus, in Evolution and Environment in Tropical America,
edited by J. B. C. Jackson, A. F. Budd, and A. G. Coates, pp. 21–56, Univ. of Chicago Press, Chicago, Ill.
Coates, A. G., J. B. C. Jackson, L. S. Collins, T. M. Cronin, H. J. Dowsett, L. M. Bybell, P. Jung, and J. A. Obando (1992), Closure of the Isthmus of
Panama: The near-shore marine record of Costa Rica and western Panama, Geol. Soc. Am. Bull., 104, 814–828.
Collins, L. S. (1996), Environmental changes in Caribbean shallow waters relative to the closing tropical American seaway, in Evolution and
Environment in Tropical America, edited by J. B. C. Jackson, A. F. Budd, and A. G. Coates, pp. 130–167, Univ. of Chicago Press, Chicago, Ill.
Collins, L. S., A. G. Coates, W. A. Berggren, M.-P. Aubry, and J.-J. Zhang (1996a), The late Miocene Panama isthmian strait, Geology, 24, 687–690.
Collins, L. S., A. F. Budd, and A. G. Coates (1996b), Earliest evolution associated with closure of the Tropical American seaway, Proc. Natl. Acad.
Sci. U.S.A., 93, 6069–6072.
Collins, T. (1996), Molecular comparisons of transisthmian species pairs: Rates and patterns of evolution, in Evolution and Environment in
Tropical America, edited by J. B. C. Jackson, A. F. Budd, and A. G. Coates, pp. 303–334, Univ. of Chicago Press, Chicago, Ill.
Conkright, M. E., et al. (1998), World Ocean Database 1998 Documentation and Quality Control, National Oceanographic Data Center,
Silver Spring, Md.
Cook, K. H., and E. K. Vizy (2010), Hydrodynamics of the Caribbean Low-Level Jet and its relationship to precipitation, J. Clim., 23, 1477–1494.
Dayem, K. E., D. C. Noone, and P. Molnar (2007), Tropical western Pacific warm pool and maritime continent precipitation rates and their
contrasting relationships with the Walker Circulation, J. Geophys. Res., 112, D06101, doi:10.1029/2006JD007870.
Dekens, P. S., A. C. Ravelo, and M. D. McCarthy (2007), Warm upwelling regions in the Pliocene warm period, Paleoceanography, 22, PA3211,
doi:10.1029/2006PA001394.
Durán-Quesada, A. M., L. Gimeno, J. A. Amador, and R. Nieto (2010), Moisture sources for Central America: Identification of moisture sources
using a Lagrangian analysis technique, J. Geophys. Res., 115, D05103, doi:10.1029/2009JD012455.
Enfield, D. B., and D. A. Mayer (1997), Tropical Atlantic SST variability and its relation to El Niño–Southern Oscillation, J. Geophys. Res., 102,
929–945, doi:10.1029/96JC03296.
Farris, D. W., C. Jaramillo, G. Bayona, S. A. Restrepo-Moreno, C. Montes, A. Cardona, A. Mora, R. J. Speakman, M. D. Glascock, and V. Valencia (2011),
Fracturing of the Panamanian Isthmus during initial collision with South America, Geology, 39, 1007–1010.
Fedorov, A. V., P. S. Dekens, M. McCarthy, A. C. Ravelo, P. B. deMenocal, M. Barreiro, R. C. Pacanowski, and S. G. Philander (2006), The Pliocene
Paradox (Mechanisms for a Permanent El Niño), Science, 312, 1485–1489.
Fedorov, A. V., C. M. Brierley, and K. Emanuel (2010), Tropical cyclones and permanent El Niño in the early Pliocene epoch, Nature, 463, 1066–1070.
Fedorov, A. V., C. M. Brierley, K. T. Lawrence, Z. Liu, P. S. Dekens, and A. C. Ravelo (2013), Patterns and mechanisms of early Pliocene warmth,
Nature, 496, 43–49.
Giannini, A., Y. Kushnir, and M. A. Cane (2000), Interannual variability of Caribbean rainfall, ENSO, the Atlantic Ocean, J. Clim., 13, 297–311.
Godfrey, J. S. (1989), A Sverdrup model of the depth-integrated flow for the ocean allowing for island circulations, Geophys. Astrophys. Fluid
Dyn., 45, 89–112.
Groeneveld, J., S. Steph, R. Tiedemann, D. Garbe-Schönberg, D. Nürnberg, and A. Sturm (2006), Pliocene mixed-layer oceanography for Site 1241,
18
using combined Mg/Ca and δ O analyses of Globigerinoides sacculifer, in Proc. ODP, Sci. Results, vol. 202, edited by R. Tiedemann et al.,
pp. 1–27, Ocean Drilling Program, College Station, Tex.
Groeneveld, J., D. Nürnberg, R. Tiedemann, G.-J. Reichart, S. Steph, L. Reuning, D. Crudeli, and P. Mason (2008), Foraminiferal Mg/Ca increase in
the Caribbean during the Pliocene: Western Atlantic Warm Pool formation, salinity influence, or diagenetic overprint?, Geochem. Geophys.
Geosyst., 9, Q01P23, doi:10.1029/2006GC001564.
Haug, G. H., and R. Tiedemann (1998), Effect of the formation of the Isthmus of Panama on Atlantic thermohaline circulation, Nature,
393, 673–676.
Haug, G. H., R. Tiedemann, R. Zahn, and A. C. Ravelo (2001), Role of Panama uplift on oceanic freshwater balance, Geology, 29, 207–210.
Huybers, P., and P. Molnar (2007), Tropical cooling and the onset of North American glaciation, Clim. Past, 3, 549–557.
Jackson, J. B. C., P. Jung, A. G. Coates, and L. S. Collins (1993), Diversity and extinction of the tropical American mollusks and emergence of the
Isthmus of Panama, Science, 260, 1624–1626.
Jackson, J. B. C., P. Jung, and H. Fortunato (1996), Paciphilia revisited: Transisthmian evolution of the Strombina group (Gastropoda:
Columbellidae), in Evolution and Environment in Tropical America, edited by J. B. C. Jackson, A. F. Budd, and A. G. Coates, pp. 234–270,
Univ. of Chicago Press, Chicago, Ill.
Johns, W. E., T. L. Townsend, D. M. Fratantoni, and W. D. Wilson (2002), On the Atlantic inflow to the Caribbean Sea, Deep Sea Res., Part I,
49, 211–243.
Kalnay, E., et al. (1996), The NCEP/NCAR 40-Year Reanalysis Project, Bull. Am. Meteorol. Soc., 77, 437–471.
Kaneps, A. G. (1979), Gulf Stream: Velocity fluctuations during the late Cenozoic, Science, 204, 297–301.
Keigwin, L. D., Jr. (1978), Pliocene closing of the Isthmus of Panama, based on biostratigraphic evidence from nearby Pacific Ocean and
Caribbean Sea cores, Geology, 6, 630–634.
Keigwin, L. D., Jr. (1982), Isotope paleoceanography of the Caribbean and east Pacific: Role of Panama uplift in late Neogene time, Science,
217, 350–353.
Keller, G., C. E. Zenker, and S. M. Stone (1989), Late Neogene history of the Pacific-Caribbean gateway, J. South Am. Earth Sci., 2, 73–108.
Kirby, M. X., and J. B. C. Jackson (2004), Extinction of a fast-growing oyster and changing ocean circulation in Pliocene tropical America,
Geology, 32, 1025–1028.
Knowlton, N., L. A. Weigt, L. A. Solórzano, D. K. Mills, and E. Bermingham (1993), Divergence in proteins, mitochondrial DNA, and reproductive
compatibility across the Isthmus of Panama, Science, 260, 1629–1632.
Lawrence, K. T., Z.-H. Liu, and T. D. Herbert (2006), Evolution of the eastern tropical Pacific through Plio-Pleistocene glaciation, Science,
312, 79–83.
Lunt, D. J., P. J. Valdes, A. Haywood, and I. C. Rutt (2008), Closure of the Panama Seaway during the Pliocene: Implications for climate and
Northern Hemisphere glaciation, Clim. Dyn., 30, 1–18.
MacFadden, B. J. (2006), North American Miocene land mammals from Panama, J. Vert. Paleontol., 26, 720–734.
MacFadden, B. J., M. X. Kirby, A. Rincon, C. Montes, S. Moron, N. Strong, and C. Jaramillo (2010), Extinct Peccary “Cynorca” Occidentale
(Tayassuidae, Tayassuinae) from the Miocene of Panama and correlations to North America, J. Paleontol., 84, 288–298.
Martin, E., and C. Schumacher (2011), The Caribbean Low-Level Jet and its relationship with precipitation in IPCC AR4 Models, J. Clim.,
24, 5935–5950.
MESTAS-NUÑEZ AND MOLNAR ©2014. American Geophysical Union. All Rights Reserved. 515
Paleoceanography 10.1002/2013PA002515
McDougall, K. (1996), Benthic foraminiferal response to the emergence of the Isthmus of Panama and coincident paleoceanographic changes,
Mar. Micropaleontol., 28, 133–169.
Mestas-Nuñez, A. M., and D. B. Enfield (2001), Eastern equatorial Pacific SST variability: ENSO and non-ENSO components and their climatic
associations, J. Clim., 14, 391–402.
Mestas-Nuñez, A. M., C. Zhang, and D. B. Enfield (2005), Uncertainties in estimating moisture fluxes in the Intra-Americas Sea, J. Hydrometeorol.,
6, 696–709.
Mestas-Nuñez, A. M., D. B. Enfield, and C. Zhang (2007), Water vapor fluxes over the Intra-Americas Sea: Seasonal and interannual variability
and associations with rainfall, J. Clim., 20, 1910–1922.
Molnar, P. (2008), Closing of the Central American Seaway and the Ice Age: A critical review, Paleoceanography, 23, PA2201, doi:10.1029/
2007PA001574.
Molnar, P., and M. A. Cane (2002), El Niño’s tropical climate and teleconnections as a blueprint for pre-Ice Age climates, Paleoceanography,
17(2), 1021, doi:10.1029/2001PA000663.
Molnar, P., and M. A. Cane (2007), Early Pliocene (Pre-Ice Age) El Niño-like Global Climate: Which El Niño?, Geosphere, 3(5), 337–365.
Montes, C., et al. (2012a), Evidence for middle Eocene and younger emergence in Central Panama: Implications for Isthmus closure,
Geol. Soc. Am. Bull., 124, 780–799.
Montes, C., G. Bayona, A. Cardona, D. M. Buchs, C. A. Silva, S. Morón, N. Hoyos, D. A. Ramírez, C. A. Jaramillo, and V. Valencia (2012b),
Arc-continent collision and orocline formation: Closing of the Central American seaway, J. Geophys. Res., 117, B04105, doi:10.1029/
2011JB008959.
Muñoz, E., A. J. Busalacchi, S. Nigam, and A. Ruiz-Barradas (2008), Winter and summer structure of the Caribbean Low-Level Jet, J. Clim.,
21, 1260–1276.
Nisancioglu, K. H., M. E. Raymo, and P. H. Stone (2003), Reorganization of Miocene deep water circulation in response to the shoaling of the
Central American Seaway, Paleoceanography, 18(1), 1006, doi:10.1029/2002PA000767.
Nof, D., and S. Van Gorder (2003), Did an open Panama Isthmus correspond to an invasion of Pacific water into the Atlantic?, J. Phys. Oceanogr.,
33, 1324–1336.
O’Dea, A., and J. B. C. Jackson (2002), Bryozoan growth mirrors contrasting seasonal regimes across the Isthmus of Panama, Palaeogeogr.
Palaeoclimatol. Palaeoecol., 185, 77–94.
O’Dea, A., and B. Okamura (2000), Intracolony variation in zooid size in cheilostome bryozoans as a new technique for investigating
palaeoseasonality, Palaeogeogr. Palaeoclimatol. Palaeoecol., 162, 319–332.
O’Dea, A., J. B. C. Jackson, H. Fortunato, J. T. Smith, L. D’Croz, K. G. Johnson, and J. A. Todd (2007), Environmental change preceded Caribbean
extinction by 2 million years, Proc. Natl. Acad. Sci. U. S. A., 104, 5501–5506.
O’Dea, A., N. Hoyos, F. Rodríguez, B. Degracia, and C. De Gracia (2012), History of upwelling in the tropical eastern Pacific and the paleogeography
of the Isthmus of Panama, Palaeogeogr. Palaeoclimatol. Palaeoecol., 348–349, 59–66.
Okamura, B., A. O’Dea, and T. Knowles (2011), Bryozoan modular growth and the retrospective analysis of environments, Mar. Ecol. Prog. Ser.,
430, 133–146.
Philander, S. G., and A. V. Fedorov (2003), Role of tropics in changing the response to Milankovich forcing some three million years ago,
Paleoceanography, 18(2), 1045, doi:10.1029/2002PA000837.
Prange, M., and M. Schulz (2004), A coastal upwelling seesaw in the Atlantic Ocean as a result of the closure of the Central American Seaway,
Geophys. Res. Lett., 31, L17207, doi:10.1029/2004GL020073.
Prange, M., S. Steph, M. Schulz, and L. D. Keigwin (2010), Inferring moisture transport across Central America: Can modern analogs of climate
variability help reconcile paleosalinity records?, Quat. Sci. Rev., 29, 1317–1321.
Ravelo, A. C., and M. W. Wara (2004), The role of the tropical oceans on global climate during a warm period and a major climate transition,
Oceanography, 17(3), 32–41.
Ravelo, A. C., D. H. Andreasen, M. Lyle, A. Olivarez Lyle, and M. W. Wara (2004), Regional climate shifts caused by gradual global cooling in the
Pliocene Epoch, Nature, 429, 263–267.
Ravelo, A. C., P. S. Dekens, and M. McCarthy (2006), Evidence for El Niño–like conditions during the Pliocene, GSA Today, 16(3), 4–11.
Saito, T. (1976), Geological significance of coiling direction in the planktonic foraminifera Pulleniatina, Geology, 4, 305–309.
Saravanan, R., and P. Chang (2000), Interaction between tropical Atlantic variability and El Niño–Southern Oscillation, J. Clim., 13, 2177–2194.
Sarnthein, M., G. Bartoli, M. Prange, A. Schmittner, B. Schneider, M. Weinelt, N. Andersen, and D. Garbe-Schönberg (2009), Mid-Pliocene shifts
in ocean overturning circulation and the onset of Quaternary-style climates, Clim. Past, 5, 269–283.
Schmittner, A., C. Appenzeller, and T. F. Stocker (2000), Enhanced Atlantic freshwater export during El Niño, Geophys. Res. Lett., 27, 1163–1166,
doi:10.1029/1999GL011048.
Schneider, B., and A. Schmittner (2006), Simulating the impact of the Panamanian seaway closure on ocean circulation, marine productivity
and nutrient cycling, Earth Planet. Sci. Lett., 246, 367–380.
Shukla, S. P., M. A. Chandler, J. Jonas, L. E. Sohl, K. Mankoff, and H. Dowsett (2009), Impact of a permanent El Niño (El Padre) and Indian Ocean
Dipole in warm Pliocene climates, Paleoceanography, 24, PA2221, doi:10.1029/2008PA001682.
Shukla, S. P., M. A. Chandler, D. Rind, L. E. Sohl, J. Jonas, and J. Lerner (2011), Teleconnections in a warmer climate: The pliocene perspective,
Clim. Dyn., 37, 1869–1887.
Steph, S., R. Tiedemann, M. Prange, J. Groeneveld, D. Nürnberg, L. Reuning, M. Schulz, and G. H. Haug (2006a), Changes in Caribbean
surface hydrography during the Pliocene shoaling of the Central American Seaway, Paleoceanography, 21, PA4221, doi:10.1029/
2004PA001092.
Steph, S., R. Tiedemann, J. Groeneveld, A. Sturm, and D. Nürnberg (2006b), Pliocene changes in tropical east Pacific upper ocean stratification:
Response to tropical gateways?, in Proc. ODP, Sci. Results, vol. 202, edited by R. Tiedemann, et al., pp. 1–51, Ocean Drilling Program,
College Station, Tex.
Steph, S., R. Tiedemann, M. Prange, J. Groeneveld, M. Schulz, A. Timmermann, D. Nürnberg, C. Rühlemann, C. Saukel, and G. H. Haug (2010),
Early Pliocene increase in thermohaline overturning: A precondition for the development of the modern equatorial Pacific cold tongue,
Paleoceanography, 25, PA2202, doi:10.1029/2008PA001645.
Teranes, J. L., D. H. Geary, and B. E. Bemis (1996), The oxygen isotope record of seasonality in Neogene bivalves from the Central American
Isthmus, in Evolution and Environment in Tropical America, edited by J. B. C. Jackson, A. F. Budd, and A. G. Coates, pp. 105–129,
Univ. of Chicago Press, Chicago, Ill.
Vizcaíno, M., S. Rupper, and J. C. H. Chiang (2010), Permanent El Niño and the onset of Northern Hemisphere glaciations: Mechanism and
comparison with other hypotheses, Paleoceanography, 25, PA2205, doi:10.1029/2009PA001733.
Wang, C. (2007), Variability of the Caribbean Low-Level Jet and its relations to climate, Clim. Dyn., 29, 411–422.
MESTAS-NUÑEZ AND MOLNAR ©2014. American Geophysical Union. All Rights Reserved. 516
Paleoceanography 10.1002/2013PA002515
Wara, M. W., A. C. Ravelo, and M. L. Delaney (2005), Permanent El Niño-like conditions during the Pliocene warm period, Science, 309, 758–761.
Weyl, P. K. (1968), The role of the oceans in climatic change: A theory of the ice ages, Meteorol. Monogr., 8, 37–62.
Whyte, F. S., M. A. Taylor, T. S. Stephenson, and J. D. Campbell (2008), Features of the Caribbean low level jet, Int. J. Climatol., 28, 119–128.
Xie, P.-P., and P. A. Arkin (1996), Analyses of global monthly precipitation using gauge observations, satellite estimates, and numerical
model predictions, J. Clim., 9, 840–858.
Xie, S.-P., Y. Okumura, T. Miyama, and A. Timmermann (2008), Influences of Atlantic climate change on the tropical Pacific via the Central
American Isthmus, J. Clim., 21, 3914–3928.
Zaucker, F., T. F. Stocker, and W. S. Broecker (1994), Atmospheric freshwater fluxes and their effect on the global thermohaline circulation,
J. Geophys. Res., 99, 12,443–12,457, doi:10.1029/94JC00526.
Zhang, X., et al. (2012), Changes in equatorial Pacific thermocline depth in response to Panamanian seaway closure: Insights from a multi-model
study, Earth Planet. Sci. Lett., 317–318, 76–84.
MESTAS-NUÑEZ AND MOLNAR ©2014. American Geophysical Union. All Rights Reserved. 517