Laurel S. Collins

Closure of the Isthmus of Panama: The near-shore marine record of Costa Rica and western Panama

The final closure of the Isthmus of Panama at ∼3.5 Ma divided the American tropical ocean into two separate and different oceanographic regions. Consequences for the marine biota were profound, but, hitherto, correlation of the Pacific and Caribbean coastal sections has not been precise enough to track biologic patterns. We present here a correlation of 31 sections from the Pacific and Caribbean coasts of Costa Rica and western Panama. Using calcareous nannofossils and planktonic foraminifera at both the tops and bottoms of each formation, we estimate that the Caribbean section ranges from 8.2 Ma to 1.7 Ma; and the Pacific sequence, from 3.6 Ma to <1.7 Ma. These intervals bracket postulated dates for final closure of the Isthmus and provide the first well-dated record of middle and late Pliocene faunas from the region. The Caribbean and Pacific sections include very different environments of deposition, yet there is sufficient overlap and diversity of habitats to permit meaningful biological comparisons. On the Caribbean side, formations tied together by the overlap of the upper Pliocene markers Sphenolithus abies and Pseudoemiliana lacunosa (3.5 Ma to 3.6 Ma) range from very shallow to shallow inner shelf (<200 m) and upper slope (200-800 m). The Pacific coast sections were mostly deposited in a trench slope environment, which is absent on the Caribbean side. These sections fortuitously include abundant thick intra-formational slumps containing shallow-water fauna more appropriate for biological comparison with the Caribbean biota. Similarly, the ∼1.9 Ma to 1.5 Ma interval, well constrained by various taxa, includes middle- to outer-shelf, and inner-shelf to upper-slope deposits on the Caribbean side, and marginal-marine to inner-shelf deposits on the Pacific coast. Using our new biostratigraphic framework to correlate previously poorly constrained mollusc collections, we show that evolutionary divergence of the Pacific and Caribbean near-shore marine faunas had occurred by 3.5 Ma. This strongly suggests that the Isthmus was effectively closed by this time.

The Geology of the Darien, Panama, and the late Miocene-Pliocene collision of the Panama arc with northwestern South America

The geology of the Darien province of eastern Panama is presented through a new geologic map and detailed biostratigraphic and paleobathymetric analysis of its Upper Cretaceous to upper Miocene sediments. The sequence of events inferred from the stratigraphic record includes the collision of the Panama arc (the southwestern margin of the Caribbean plate) and South American continent. Three tectonostratigraphic units underlie the Darien region: (1) Precollisional Upper Cretaceous–Eocene crystalline basement rocks of the San Blas Complex form a series of structurally complex topographic massifs along the northeastern and southwestern margins of the Darien province. These rocks formed part of a >20 m.y. submarine volcanic arc developed in a Pacific setting distant from the continental margin of northwestern South America. The northerly basement rocks are quartz diorites, granodiorites, and basaltic andesites, through dacites to rhyolites, indicating the presence of a magmatic arc. The southerly basement rocks are an accreted suite of diabase, pillow basalt, and radiolarian chert deposited at abyssal depths. Precollisional arc-related rocks, of Eocene to lower Miocene age, consist of 4000 m of pillow basalts and volcaniclastics, and biogenic calcareous and siliceous deep-water sediments. They consist of the Eocene-Oligocene Darien Formation, the Oligocene Porcona Formation and the lower-middle Miocene Clarita Formation. Postcollisional deposits are mostly coarse- to fine-grained siliciclastic sedimentary rocks and turbiditic sandstone of upper middle to latest Miocene age. This 3000 m thick sedimentary sequence is deformed as part of a complexly folded and faulted synclinorium that forms the central Chucunaque-Tuira Basin of the Darien. The sedimentary package reveals general shallowing of the basin from bathyal to inner neritic depths during the 12.8−7.1 Ma collision of the Panama arc with South America. The sediments are divided into the upper middle Miocene Tapaliza Formation, the lower upper Miocene Tuira and Membrillo Formations, the middle upper Miocene Yaviza Formation, and the middle to upper Miocene Chucunaque Formation. The precollisional open marine units of Late Cretaceous–middle Miocene age are separated from the overlying postcollisional sequence of middle to late Miocene age by a regional unconformity at 14.8−12.8 Ma. This unconformity marks the disappearance of radiolarians, the changeover of predominantly silica deposition from the Atlantic to the Pacific, the initiation of the uplift of the isthmus of Panama, and the onset of shallowing upward, coarser clastic deposition. This pattern is also recorded from the southern Limon Basin of Caribbean Costa Rica to the Atrato Basin of northwestern Colombia. By the middle late Miocene, neritic depths were widespread throughout the Darien region, and a regional unconformity suggests completion of the Central American arc collision with South America by 7.1 Ma. No Pliocene deposits are recorded from either the Darien or the Panama Canal Basin, and no sediments younger than 4.8 Ma have been identified in the Atrato Basin of Colombia, suggesting rapid uplift and extensive emergence of the Central American isthmus in the latest Miocene. Northward movement of the eastern segment of the Panama arc along a now quiescent Panama Canal Zone fault during Eocene-Oligocene time may have dislocated the pre-collision arc. Since collision, the portion west of this fault (Chorotega Block) has remained stable, without rotation; to the east, in the Darien region, compression has been accommodated through formation of a Panama microplate with convergent boundaries to the north (North Panama deformed belt) and south (South Panama deformed belt), and suturing with South America along the Atrato Valley. Deformation within the microplate has been accommodated in the Darien province by several major left-lateral strike-slip faults that were active until the early Pliocene, since when the plate has behaved rigidly.

The late Miocene Panama isthmian strait

Miocene sediments of the Caribbean Gatun and Chagres formations, Panama Canal basin, were deposited within an archipelagic strait that connected Caribbean and Pacific waters. Shallow-water (∼ 25 m) benthic foraminifera of the Gatun Formation have a strong Caribbean affinity, indicating that a significant interoceanic, biogeographic barrier had formed at ∼ 8 Ma. However, benthic foraminifera of the overlying Chagres Formation are bathyal and markedly Pacific in affinity, indicating that at ∼ 6 Ma, waters of the Panama isthmian strait deepened to ∼ 200–500 m and Pacific bathyal waters flowed into the Caribbean. The Chagres Formation crops out at the Caribbean entrance to the Panama Canal in a large wedge of cross-laminated sandstone and coquina. The cross-laminations and coarse grain size indicate high-energy currents atypical of bathyal settings. We infer that a jet of the Pacific North Equatorial Countercurrent–Equatorial Undercurrent passed through the Panama isthmian strait to deposit these sediments on the Caribbean side. This later entry of Pacific taxa into the Caribbean had no apparent effect on the subsequent composition of Caribbean faunas.

Diversity and Extinction of Tropical American Mollusks and Emergence of the Isthmus of Panama

The gradual closure of the Panamanian seaway and the resulting environmental change stimulated an increase in Caribbean molluscan diversity rather than the mass extinction hypothesized previously on the basis of inadequate data. Upheaval of molluscan faunas did occur suddenly throughout tropical America at the end of the Pliocene as a result of more subtle, unknown causes. There is no necessary correlation between the magnitude of regional shifts in abiotic conditions and the subsequent biological response.

Formation of the Isthmus of Panama

Independent evidence from rocks, fossils, and genes converge on a cohesive narrative of isthmus formation in the Pliocene. The formation of the Isthmus of Panama stands as one of the greatest natural events of the Cenozoic, driving profound biotic transformations on land and in the oceans. Some recent studies suggest that the Isthmus formed many millions of years earlier than the widely recognized age of approximately 3 million years ago (Ma), a result that if true would revolutionize our understanding of environmental, ecological, and evolutionary change across the Americas. To bring clarity to the question of when the Isthmus of Panama formed, we provide an exhaustive review and reanalysis of geological, paleontological, and molecular records. These independent lines of evidence converge upon a cohesive narrative of gradually emerging land and constricting seaways, with formation of the Isthmus of Panama sensu stricto around 2.8 Ma. The evidence used to support an older isthmus is inconclusive, and we caution against the uncritical acceptance of an isthmus before the Pliocene.

Early Neogene history of the Central American arc from Bocas del Toro, western Panama

A newly discovered sequence of lower to middle Miocene rocks from the eastern Bocas del Toro archipelago, western Panama, reveals the timing and environment of the earliest stages in the rise of the Isthmus of Panama in this region. Two new formations, the Punta Alegre Formation (lower Miocene, Aquitanian to Burdigalian) and the Valiente Formation (middle Miocene, Langhian to Serravallian), are here named and formally described. The Punta Alegre Formation contains a diagnostic microfauna of benthic and planktic foraminifera and calcareous nannofossils that indicate deposition in a 2000-m-deep pre-isthmian neotropical ocean from as old as 21.5-18.3 Ma. Its lithology varies from silty mudstone to muddy foraminiferal ooze with rare thin microturbidite layers near the top. The Valiente Formation, which ranges in age from 16.4 to ca. 12.0 Ma, lies with slight angular unconformity on the Punta Alegre Formation and consists of five lithofacies: (1) columnar basalt and flow breccia, (2) pyroclastic deposits, (3) coarse-grained volcaniclastic deposits, (4) coral-reef limestone with diverse large coral colonies, and (5) marine debris-flow deposits and microturbidites. These lithofacies are interpreted to indicate that after ca. 16 Ma a volcanic arc developed in the region of Bocas del Toro and that by ca. 12 Ma an extensively emergent archipelago of volcanic islands had formed. 3 9 Ar/ 4 0 Ar dating of basalt flows associated with the fossiliferous sedimentary rocks in the upper part of the Valiente Formation strongly confirms the ages derived from planktic foraminifera and nannofossils. Paleobathymetric analysis of the two new formations in the Valiente Peninsula and Popa Island, in the Bocas del Toro archipelago, shows a general shallowing from lower- through upper-bathyal to upper-neritic and emergent laharic and fluviatile deposits from ca. 19 to 12 Ma. The overlying nonconformable Bocas del Toro Group contains a lower transgressive sequence ranging from basal nearshore sandstone to upper-bathyal mudstone (ca. 8.1-5.3 Ma) and an upper regressive sequence (5.3-3.5 Ma). A similar paleobathymetric pattern is observed from the Gatun to Chagres Formations (12-6 Ma) in the Panama Canal Basin area and in the Uscari, Rio Banana, Quebrada Chocolate, and Moin Formations (8-1.7 Ma) in the southern LimOn Basin of Costa Rica.

Latitudinal difference in biodiversity caused by higher tropical rate of increase

Tropical diversity has generally exceeded temperate diversity in the present and at points in the past, but whether measured differences have remained relatively constant through time has been unknown. Here we examine tropical vs. temperate diversities from the Neogene to Recent using the within-habitat diversity measure Fishers alpha of Cenozoic benthic foraminifera from the temperate Central Atlantic Coastal Plain and the tropical Central American Isthmus. During the Neogene, the mean value of alpha at temperate latitudes increased 1.4 times or 40%, whereas in the tropics it increased 2.1 times or 106%. Thus, while both areas exhibit an increase of diversity with time, past differences in the rate of increase have generated a more pronounced gradient today (164%) than existed in the Miocene (80%). These data disagree with the suggestion that the world reached an equilibrium number of species during the Paleozoic and demonstrate the need to consider both temperate and tropical components in global diversity assessments.

Carbon, nitrogen and phosphorus storage in subtropical seagrass meadows: Examples from Florida Bay and Shark Bay

Seagrass meadows in Florida Bay and Shark Bay contain substantial stores of both organic carbon and nutrients. Soils from both systems are predominantly calcium carbonate, with an average of 82.1% CaCO3 in Florida Bay compared with 71.3% in Shark Bay. Soils from Shark Bay had, on average, 21% higher organic carbon content and 35% higher phosphorus content than Florida Bay. Further, soils from Shark Bay had lower mean dry bulk density (0.78 ± 0.01 g mL–1) than those from Florida Bay (0.84 ± 0.02 mg mL–1). The most hypersaline regions of both bays had higher organic carbon content in surficial soils. Profiles of organic carbon and phosphorus from Florida Bay indicate that this system has experienced an increase in P delivery and primary productivity over the last century; in contrast, decreasing organic carbon and phosphorus with depth in the soil profiles in Shark Bay point to a decrease in phosphorus delivery and primary productivity over the last 1000 y. The total ecosystem stocks of stored organic C in Florida Bay averages 163.5 MgCorg ha–1, lower than the average of 243.0 MgCorg ha–1 for Shark Bay; but these values place Shark and Florida Bays among the global hotspots for organic C storage in coastal ecosystems.

Controls on facies and sequence stratigraphy of an upper Miocene carbonate ramp and platform, Melilla basin, NE Morocco

Abstract Upwelling of cool seawater, paleoceanographic circulation, paleoclimate, local tectonics and relative sea-level change controlled the lithofacies and sequence stratigraphy of a carbonate ramp and overlying platform that are part of a temporally well constrained carbonate complex in the Melilla basin, northeastern Morocco. At Melilla, from oldest to youngest, a third-order depositional sequence within the carbonate complex contains (1) a retrogradational, transgressive, warm temperate-type rhodalgal ramp; (2) an early highstand, progradational, bioclastic platform composed mainly of a temperate-type, bivalve-rich molechfor facies; and (3) late highstand, progradational to downstepping, subtropical/tropical-type chlorozoan fringing Porites reefs. The change from rhodalgal ramp to molechfor platform occurred at 7.0±0.14 Ma near the Tortonian/Messinian boundary. During a late stage in the development of the bioclastic platform a transition from temperate-type molechfor facies to subtropical/tropical-type chlorozoan facies occurred and is bracketed by chron 3An.2n (∼6.3–6.6 Ma). Comparison to a well-dated carbonate complex in southeastern Spain at Cabo de Gata suggests that upwelling of cool seawater influenced production of temperate-type limestone within the ramp and platform at Melilla during postulated late Tortonian–early Messinian subtropical/tropical paleoclimatic conditions in the western Paleo-Mediterranean region. The upwelling of cool seawater across the bioclastic platform at Melilla could be related to the beginning of ‘siphoning’ of deep, cold Atlantic waters into the Paleo-Mediterranean Sea at 7.17 Ma. The facies change within the bioclastic platform from molechfor to chlorozoan facies may be coincident with a reduction of the siphoning of Atlantic waters and the end of upwelling at Melilla during chron 3An.2n. The ramp contains one retrogradational parasequence and the bioclastic platform three progradational parasequences. Minor erosional surfaces that bound the upper surface of the ramp and upper surface of the oldest platform parasequence are related to relative falls in sea level induced by local volcanism and associated tectonic uplift. These local relative falls had little influence on a broader-scale rise to stillstand in relative sea level that controlled development of the transgressive and early highstand systems tracts represented in the ramp and platform, respectively.

Neogene paleoenvironments of the Bocas del Toro Basin, Panama

Outcrops of upper Miocene to upper Pliocene sediments in the Bocas del Toro Basin, Caribbean coast of western Panama, contain a record of the emergence of the southern Central American isthmus. Paleodepths and characteristics of past sedimentary regimes within the basin were determined from benthic foraminiferal species assemblages. The present Valiente Peninsula shallowed from upper bathyal depths of about 300–500 m to outer neritic depths of 150–200 m between latest Miocene (5.6–6.5 Ma) and middle Pliocene (3.5–3.6 Ma) times. Outer neritic assemblages from about 100–150 m paleodepth on the island of Escudo de Veraguas indicate negligible emergence through the late Pliocene, and their present sea-level altitude suggests rapid Quaternary shallowing. Middle neritic foraminiferal assemblages of Cayo Agua island shallowed from depths of about 40–80 m to 20–40 m between early (4.5–5.0 Ma) and middle (3.5–3.6 Ma) Pliocene times. Cayo Agua assemblages indicate establishment of a carbonate regime in the southwestern Caribbean region prior to complete closure of the Caribbean–East Pacific seaway around 3.5 Ma.