Viviana Barreda

Patagonian Vegetation Turnovers during the Paleogene-Early Neogene: Origin of Arid-Adapted Floras

The structure of the living Patagonian flora, dominated by the steppe, is a direct consequence of past climatic and tectonic events. These arid-adapted communities were widespread during the Late Neogene, but their origin in Patagonia can be traced back to the Paleogene. Vegetational trends throughout Paleocene-Miocene time are based on available paleobotanical and palynological information. Four major supported stages in vegetation turnovers are recognized: (1) Paleocene and Early Eocene floras were rainforest-dominated, including many angiosperms with warm-temperate affinities (e.g., palms, Juglandaceae, Casuarinaceae). However, mainly in the Early Eocene, some geographic areas influenced by warm but drier conditions are suggested by the occurrence of certain taxa (e.g., Anacardiaceae). These areas containing arid-adapted floras would have arisen in Patagonian inland regions, in a generally wet continent. (2) The Middle Eocene-Early Oligocene interval was distinguished by the invasion ofNothofagus forests. Progressive replacements of megathermal communities by meso- and microthermal rainforest are documented.Nothofagus forest expansion suggests a marked cooling trend at this time, although some megathermal elements (AquifoliaceaeIlex, Tiliaceae-Bombacaceae, Sapindaceae) were still present at the beginning of this period. Arid-loving taxa have not been recorded in abundance. (3) Late Oligocene-Early Miocene floras were characterized by the occurrence of shrubby-herbaceous elements belonging to Asteraceae, Chenopodiaceae, Ephedraceae, Convolvulaceae, Fabaceae, and Poaceae. They began to give a modern appearance to plant communities. Xerophytic formations would have occupied coastal salt marshes and pockets in inland areas. Megathermal angiosperms of the Rubiaceae, Combretaceae, Sapindaceae, Chloranthaceae, and Arecaceae occurred mainly during the Late Oligocene. Forests of Nothofagaceae, Podocarpaceae, and Araucariaceae are still documented in extra-Andean Patagonia; however, a contrast between coastal and inland environments may have developed, particularly in the Miocene. (4) Middle-Late Miocene records show an increasing diversity and abundance of xerophytic-adapted taxa, including Asteraceae, Chenopodiaceae, and ConvolvulaceaeCressa/Wilsonia. Expansion of these xerophytic taxa, coupled with extinctions of megathermal/nonseasonal elements, would have been associated with both tectonic and climatic forcing factors, led to the development of aridity and extreme seasonality. These arid-adapted Late Miocene floras are closely related to modern communities, with steppe widespread across extra-Andean Patagonia and forest restricted to the western humid upland regions.ResumenPrincipales tendencias de la vegetación en Patagonia durante el Paleógeno-Neógeno temprano: origen de las floras adaptadas a condiciones de aridez. La estructura de la flora patagónica actual, dominada por la estepa, es consecuencia directa de los eventos tectónicos y climáticos a los que ha estado sometida. Estas comunidades, adaptadas a condiciones de extrema aridez, se expandieron durante el Neógeno tardío, aunque su origen en Pagatonia pudo haber ocurrido en el Paleógeno. En base a la información paleobotánica y palinolíogica disponible se sustentan las cuatro etapas principales de cambios en la vegetación a través del intervalo Paleoceno-Mioceno: 1-Paleoceno-Eoceno Temprano, con floras dominadas por selvas, incluyendo angiospermas con afinidades megatérmicas (ej. palmeras, Juglandaceae, Casuarinaceae). En el Eoceno Temprano, en algunas áreas geográficas habrían prevalecido condiciones cálidas pero áridas según surge de la presencia de taxones con estos requerimientos (ej. Anacardiaceae). Estos parches xerof íticos se habnían desarrollado en el interior de la Patagonia dentro de un entorno general húmedo. 2-Eoceno Medio-Oligoceno Temprano, caracterizado por la expansión de los bosques deNothofagus. Se documentó un progresivo reemplazo de comunidades megatérmicas por bosques meso y microtérmicos dominados porNothofagus y podocarpáceas, indicando un marcado enfriamiento. Al principio de este intervalo, sin embargo, todavía se reconocen algunos elementos megatérmicos (AquifoliaceaeIlex, Tiliaceae-Bombacaceae, Sapindaceae); los taxones xerofíticos, en cambio, son muy escasos. 3-Oligoceno Tardío-Mioceno Temprano, determinado por la presencia de elementos herbáceo-arbustivos de Asteraceae, Chenopodiaceae, Ephedraceae, Convolvulaceae, Fabaceae, Poaceae, que empezaron a dar una apariencia moderna a las comunidades vegetales. Las formaciones xerofíticas habrían ocupado ambientes costeros como marismas o parches abiertos en áreas internas. Angiospermas megatérmicas como Rubiaceae, Combretaceae, Sapindaceae, Chloranthaceae y Arecacea están bien representadas, en particular en el Oligoceno Tardío. Los bosques de Nothofagaceae, Podocarpaceae y Araucariaceae todavía estarían presentes en la Patagonia extra-andina, pero ya existiría un marcado contraste entre los ambientes continentales y costeros. 4-Mioceno Medio-Tardío, definido por un marcado incremento en la diversidad y abundancia de taxones xerofítícos incluyendo Asteraceae, Chenopodiaceae y ConvolvulaceaeCressa/Wilsonia. La expansión de estas formas y la extinción de elementos megatérmicos, no estacionales, habrían estado asociadas a factures tectónicos y climáticos que condujeron al desarrollo de aridez y extrema estacionalidad. Las floras áridas del Mioceno Tardío se encuentran estrechamente relacionadas con las comunidades modernas, con la estepa expandida en la Patagonia extra-andina y los bosques restringidos a la región húmeda, occidental, de los Andes.

Cretaceous/Paleogene Floral Turnover in Patagonia: Drop in Diversity, Low Extinction, and a Classopollis Spike

Nearly all data regarding land-plant turnover across the Cretaceous/Paleogene boundary come from western North America, relatively close to the Chicxulub, Mexico impact site. Here, we present a palynological analysis of a section in Patagonia that shows a marked fall in diversity and abundance of nearly all plant groups across the K/Pg interval. Minimum diversity occurs during the earliest Danian, but only a few palynomorphs show true extinctions. The low extinction rate is similar to previous observations from New Zealand. The differing responses between the Southern and Northern hemispheres could be related to the attenuation of damage with increased distance from the impact site, to hemispheric differences in extinction severity, or to both effects. Legacy effects of the terminal Cretaceous event also provide a plausible, partial explanation for the fact that Paleocene and Eocene macrofloras from Patagonia are among the most diverse known globally. Also of great interest, earliest Danian assemblages are dominated by the gymnosperm palynomorphs Classopollis of the extinct Mesozoic conifer family Cheirolepidiaceae. The expansion of Classopollis after the boundary in Patagonia is another example of typically Mesozoic plant lineages surviving into the Cenozoic in southern Gondwanan areas, and this greatly supports previous hypotheses of high latitude southern regions as biodiversity refugia during the end-Cretaceous global crisis.

Eocene Patagonia Fossils of the Daisy Family

Fossil evidence suggests that daisies and sunflowers may have originated in South America more than 47 million years ago. Fossil capitula and pollen grains of Asteraceae from the Eocene of Patagonia, southern Argentina, exhibit morphological features recognized today in taxa, such as Mutisioideae and Carduoideae, that are phylogenetically close to the root of the asteracean tree. This fossil supports the hypothesis of a South American origin of Asteraceae and an Eocene age of divergence and suggests that an ancestral stock of Asteraceae may have formed part of a geoflora developed in southern Gondwana before the establishment of effective dispersal barriers within this landmass.

An extinct Eocene taxon of the daisy family (Asteraceae): evolutionary, ecological and biogeographical implications

BACKGROUND AND AIMS Morphological, molecular and biogeographical information bearing on early evolution of the sunflower alliance of families suggests that the clade containing the extant daisy family (Asteraceae) differentiated in South America during the Eocene, although palaeontological studies on this continent failed to reveal conclusive support for this hypothesis. Here we describe in detail Raiguenrayun cura gen. & sp. nov., an exceptionally well preserved capitulescence of Asteraceae recovered from Eocene deposits of northwestern Patagonia, Argentina. METHODS The fossil was collected from the 47·5 million-year-old Huitrera Formation at the Estancia Don Hipólito locality, Río Negro Province, Argentina. KEY RESULTS The arrangement of the capitula in a cymose capitulescence, the many-flowered capitula with multiseriate-imbricate involucral bracts and the pappus-like structures indicate a close morphological relationship with Asteraceae. Raiguenrayun cura and the associated pollen Mutisiapollis telleriae do not match exactly any living member of the family, and clearly represent extinct taxa. They share a mosaic of morphological features today recognized in taxa phylogenetically close to the root of Asteraceae, such as Stifftieae, Wunderlichioideae and Gochnatieae (Mutisioideae sensu lato) and Dicomeae and Oldenburgieae (Carduoideae), today endemic to or mainly distributed in South America and Africa, respectively. CONCLUSIONS This is the first fossil genus of Asteraceae based on an outstandingly preserved capitulescence that might represent the ancestor of Mutisioideae-Carduoideae. It might have evolved in southern South America some time during the early Palaeogene and subsequently entered Africa, before the biogeographical isolation of these continents became much more pronounced. The new fossil represents the first reliable point for calibration, favouring an earlier date to the split between Barnadesioideae and the rest of Asteraceae than previously thought, which can be traced back at least 47·5 million years. This is the oldest well dated member of Asteraceae and perhaps the earliest indirect evidence for bird pollination in the family.

Paleo-Antarctic rainforest into the modern Old World tropics: The rich past and threatened future of the “southern wet forest survivors”

UNLABELLED • PREMISE OF STUDY Have Gondwanan rainforest floral associations survived? Where do they occur today? Have they survived continuously in particular locations? How significant is their living floristic signal? We revisit these classic questions in light of significant recent increases in relevant paleobotanical data.• METHODS We traced the extinction and persistence of lineages and associations through the past across four now separated regions-Australia, New Zealand, Patagonia, and Antarctica-using fossil occurrence data from 63 well-dated Gondwanan rainforest sites and 396 constituent taxa. Fossil sites were allocated to four age groups: Cretaceous, Paleocene-Eocene, Neogene plus Oligocene, and Pleistocene. We compared the modern and ancient distributions of lineages represented in the fossil record to see if dissimilarity increased with time. We quantified similarity-dissimilarity of composition and taxonomic structure among fossil assemblages, and between fossil and modern assemblages.• KEY RESULTS Strong similarities between ancient Patagonia and Australia confirmed shared Gondwanan rainforest history, but more of the lineages persisted in Australia. Samples of ancient Australia grouped with the extant floras of Australia, New Guinea, New Caledonia, Fiji, and Mt. Kinabalu. Decreasing similarity through time among the regional floras of Antarctica, Patagonia, New Zealand, and southern Australia reflects multiple extinction events.• CONCLUSIONS Gondwanan rainforest lineages contribute significantly to modern rainforest community assembly and often co-occur in widely separated assemblages far from their early fossil records. Understanding how and where lineages from ancient Gondwanan assemblages co-occur today has implications for the conservation of global rainforest vegetation, including in the Old World tropics.

Fossil pollen records reveal a late rise of open-habitat ecosystems in Patagonia

The timing of major turnovers in terrestrial ecosystems of the Cenozoic Era has been largely interpreted from the analysis of the assumed feeding preference of extinct mammals. For example, the expansion of open-habitat ecosystems (grasslands or savannas) is inferred to have occurred earlier in Patagonia than elsewhere because of the early advent of high-crowned teeth (hypsodont) mammals ∼26 Ma ago. However, the plant fossil record from Patagonia implies another evolutionary scenario. Here we show that the dominance of key open-habitat species--amaranths, Ephedra, asters and grasses--occurred during the last 10 Ma, about 15 Ma later than previously inferred using feeding/habitat ecology of extinct mammals. This late rise of open-landscapes in southern South America brings into question whether the expansion of open-habitat vegetation could have been the prime factor of high-crowned mammal diversification.

Early evolution of the angiosperm clade Asteraceae in the Cretaceous of Antarctica

Significance The flowering plant family Asteraceae (e.g. sunflowers, daisies, chrysanthemums), with about 23,000 species, is found almost everywhere in the world except in Antarctica. Asteraceae (or Compositae) are regarded as one of the most influential families in the diversification and evolution of a large number of animals that heavily depends on their inflorescences to survive (e.g. bees, hummingbirds, wasps). Here we report the discovery of pollen grains unambiguously assigned to Asteraceae that remained buried in Antarctic deposits for more than 65 million years along with other extinct groups (e.g. Dinosaurs, Ammonites). Our discovery drastically pushes back the assumed origin of Asteraceae, because these pollen grains are the oldest fossils ever found for the family. The Asteraceae (sunflowers and daisies) are the most diverse family of flowering plants. Despite their prominent role in extant terrestrial ecosystems, the early evolutionary history of this family remains poorly understood. Here we report the discovery of a number of fossil pollen grains preserved in dinosaur-bearing deposits from the Late Cretaceous of Antarctica that drastically pushes back the timing of assumed origin of the family. Reliably dated to ∼76–66 Mya, these specimens are about 20 million years older than previously known records for the Asteraceae. Using a phylogenetic approach, we interpreted these fossil specimens as members of an extinct early diverging clade of the family, associated with subfamily Barnadesioideae. Based on a molecular phylogenetic tree calibrated using fossils, including the ones reported here, we estimated that the most recent common ancestor of the family lived at least 80 Mya in Gondwana, well before the thermal and biogeographical isolation of Antarctica. Most of the early diverging lineages of the family originated in a narrow time interval after the K/P boundary, 60–50 Mya, coinciding with a pronounced climatic warming during the Late Paleocene and Early Eocene, and the scene of a dramatic rise in flowering plant diversity. Our age estimates reduce earlier discrepancies between the age of the fossil record and previous molecular estimates for the origin of the family, bearing important implications in the evolution of flowering plants in general.

Fossil pollen records indicate that Patagonian desertification was not solely a consequence of Andean uplift

The Patagonian steppe-a massive rain-shadow on the lee side of the southern Andes-is assumed to have evolved ~15-12 Myr as a consequence of the southern Andean uplift. However, fossil evidence supporting this assumption is limited. Here we quantitatively estimate climatic conditions and plant richness for the interval ~10-6 Myr based on the study and bioclimatic analysis of terrestrially derived spore-pollen assemblages preserved in well-constrained Patagonian marine deposits. Our analyses indicate a mesothermal climate, with mean temperatures of the coldest quarter between 11.4 °C and 16.9 °C (presently ~3.5 °C) and annual precipitation rarely below 661 mm (presently ~200 mm). Rarefied richness reveals a significantly more diverse flora during the late Miocene than today at the same latitude but comparable with that approximately 2,000 km further northeast at mid-latitudes on the Brazilian coast. We infer that the Patagonian desertification was not solely a consequence of the Andean uplift as previously insinuated.

Early history of Asteraceae in Patagonia: Evidence from fossil pollen grains

Abstract The Asteraceae are classified into three subfamilies: Barnadesioideae, Cichorioideae, and Asteroideae. It has been suggested that the southern South American subfamily Barnadesioideae is the basal branch of the phylogenetic tree of the family, and Patagonia is the ancestral area of Asteraceae. Here we explore the chronological records of some members of the family, with findings of Mutisiinae (tribe Mutisieae, subfamily Cichorioideae) from the Late oligocene (28–23 Ma) and of Barnadesioideae and Nassauviinae (Mutisieae) from the early Miocene (23–20 Ma), all recovered from marine deposits of Patagonia. even though the succession of fossil appearances (first Mutisieae) differs from that provided by molecular data (first Barnadesioideae), this new scheme offers additional evidence towards the understanding of the early history of ancestral Asteraceae.

Late Miocene Continental and Marine Palynological Assemblages from Patagonia

Publisher Summary This chapter focuses on the main trends of the Late Miocene Patagonian vegetation. Several forcing factors may have influenced Miocene vegetation trends to their current establishment. The development of the Antarctic Circumpolar Current was of particular importance, which prevented equatorial currents from penetrating into the southern polar regions. The result was the thermal insulation of the Antarctic continent and the development of the major ice sheet on western Antarctica by the Late Miocene. Palynological assemblages from the Barranca Final, Rio Negro and Puerto Madryn formations are integrated by marine—dinoflagellates, prasinophycean algae, and foraminiferal linings—and continental—pollen, spores, and freshwater algae—palynomorphs. Marine elements show a decreasing trend toward upper levels, while continental palynomorphs prevail. In continental palynomorphs, spore pollen assemblages are angiosperm dominated. The most abundant families are Chenopodiaceae, Convolvulaceae Cressa, and Asteraceae almost reaching 50% of the continental spectrum.