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- Eurya stigmosa (Theaceae), a new and extinct record for the Calabrian stage of Madeira Island (Portugal): 40Ar/39Ar dating, palaeoecological and oceanic island palaeobiogeographical implicationsPublication . Góis-Marques, Carlos A.; Mitchell, Ria L.; Nascimento, Lea de; Fernández-Palacios, José María; Madeira, José; Sequeira, Miguel Menezes deThe general dynamic model of oceanic island biogeography (GDM) predicts the immigration, speciation and extinction of terrestrial biota through geological time on oceanic islands. Additionally, the glacial sensitive model of island biogeography (GSM) also predicts extinction due to eustatic and climate change within islands. However, well-documented and natural pre-Holocene plant extinctions are almost unknown for oceanic islands worldwide. To test these predictions, we have sampled the Early Pleistocene Porto da Cruz lacustrine and fluvial sediments for plant fossils that could confirm the GDM and GSM extinction predictions. Additionally, two new 40Ar/39Ar geochronological analyses were per formed, constraining the age of the sediments to 1.3 Ma (Calabrian). Among the fossils, Eurya stigmosa (R.Ludw.) Mai (Theaceae) seeds were recognised and studied by scanning electron microscopy (SEM). E. stigmosa is the first report of a natural (non-anthropogenic) extinct plant in the fossil record for Madeira Island, and for an oceanic island, confirming the GDM and GSM predictions. Eurya spp. palae obiogeography indicates wider distribution in Europe until the end of the Pliocene (2.58 Ma), becoming extirpated to small refugia and extinct thereafter. The Madeiran record expands the formerly unknown presence of E. stigmosa to the Macaronesian realm. The new dating of the deposit at 1.3 Ma (Calabrian) means that E. stigmosa in Madeira was already in a refugium. The extinction in Madeira is most probably a combination of island ontogeny and climate change due to Pleistocene glaciations. The palaeoecological role of this extinct shrub or tree is currently unknown, but it was a probably an element of the Madeiran laurel forest, as this community was already present in Madeira at least 1.8 My ago. This new information corroborates the predictive power of GDM and GSM and adds a new view on the importance of studying oceanic island palaeobotany, specially palaeocarpofloras.
- The Quaternary plant fossil record from the volcanic Azores Archipelago (Portugal, North Atlantic Ocean): a reviewPublication . Góis-Marques, Carlos A.; Nascimento, Lea de; Sequeira, Miguel Menezes de; Fernández-Palacios, José María; Madeira, JoséPlant fossils are known from the Azores Islands, yet poorly studied. We present a comprehensive bibliographical review for the archipelago. A first pre-scientific reference dates from late fifteenth century, while the first scientific description was reported in 1821, accounting for trunks in pyroclastic units and silicified plants within hydrothermal deposits. Throughout the second-half of the nineteenth century and the first-half of the twentieth century, prospection by naturalists and geological mapping work, led to the discovery and description of plant fossils in most islands. From the 1970s onwards, the taxonomic interest ceased, and plant fossils were used mainly for 14C dating. Recently, sediment cores from lakes and peatlands were used for palaeoecological reconstructions and to measure anthropogenic impacts. Generally, plant fossils are younger than 50 ka, although older fossils may exist. Azorean plant fossils include somatofossils of leaves, stems, logs and seeds preserved as impressions, compressions, adpressions, permineralizations, lava tree casts and mummifications. The taphonomy of macrofloral elements is usually related to explosive volcanic activity, while palynological record is associated with lake sediments and peat bogs. The persistence in palaeobotanical and palaeopalynological studies will decisively contribute to disentangle the paleodiversity, palaeoecology, and add crucial information on insular plant phylogeny and biogeography.
- Tracing insular woodiness in giant Daucus (s.l.) fruit fossils from the Early Pleistocene of Madeira Island (Portugal)Publication . Góis-Marques, Carlos A.; Nascimento, Lea de; Fernández‐Palacios, José María; Madeira, José; Sequeira, Miguel Menezes dePlants on oceanic islands can evolve insular syndromes such as secondary woodiness, a generalized trend found in island floras worldwide. This phenomenon occurs through evolution in situ. It is triggered by ecological and physiological stimuli that trans form herbaceous annuals into woody perennials. However, well-dated and informative fossils that could help track and frame the evo lution of this syndrome are lacking. Remarkably, in Madeira Island (Portugal), there are good examples of Apiaceae that evolved secondary woodiness, like the giant neoendemic Melanoselinum (≡ Daucus). Apiaceae has a very scarce fossil record, despite being a cosmopolitan family and an economically important crop. Here we describe the oldest Daucus s.l. fossil known to date and the first fossil evidence of a plant with insular woodiness. The fossils are preserved as mummified/compressed mericarps within 1.3-million year-old fluvio-lacustrine sediments of the Funchal unit, Upper Volcanic complex, near Porto da Cruz. We assign them to the extant neoendemic species Melanoselinum (≡ Daucus) decipiens. The mericarp morphology shows remarkable stasis since the Calabrian stage of the Early Pleistocene. Our results demonstrate that in the Madeiran Daucinae clade, insular woodiness developed at least 1.3 million years ago, indicating a coeval or earlier immigration to Madeira Island of a Daucus sp. Our results reinforce the role of palaeobotanical research in oceanic islands, supported by stratigraphy and geochronology studies, as a key element for the understanding of plant palaeobiogeography, ecology and evolution worldwide. We expect this contribution to shed light on the evolutionary origins of carrots, and related plant groups, an important element of human food, and to better comprehend the evolution of plant insular woodiness.
- The first Ichneumonid fossil from the Early Pleistocene of Madeira Island (Portugal)Publication . Góis-Marques, Carlos A.; Jesus, José; Sequeira, Miguel Menezes; Madeira, JoséIn oceanic islands, well age-constrained deposits containing arthropod somatofossils (body fossils) are rare. However, when available, these are important for providing empirical and independent minimum ages for molecular phy logenetic dating and complementary data on taxonomy, evolution and palaeobiogeography information of the biological groups found as fossils. This is especially important for taxa that speciated within oceanic islands, many becoming single island endemics (SIE). Recently, associated with a 1.3 Ma (Calabrian) fluvial and lacustrine sedimentary deposit from Porto da Cruz in Madeira Island (Fig. 1), a wing, putatively identified as Hymenoptera, was found. Here we describe this wing fossil as belonging to Ichneumonidae, a group with ca. 30% of SIE in Madeira Island. Moreover, this is the first somatofossil of ichneumonid parasitic wasps found in Madeira Island and in Macaronesian islands (i.e. Azores, Madeira, Canaries and Cabo Verde). Since the 19th century, oceanic island attracted several naturalists due to the high probability of finding taxonomical novelties (e.g. Vieira, 2005). Darwin (1859) amplified this interest, as oceanic islands biota presented a central role to explain evolution. Today oceanic islands became the ideal locations to study evolution, biogeography and ecology (e.g. Whittaker et al., 2017). Madeira Island (Central Atlantic Ocean; Fig. 1), geologically a shield volcano of 7 Ma (Ramalho et al., 2015 and references therein), is considered an insect diversity hotspot where 3019 species and subspecies are known, of which 665 are SIE (Borges et al., 2008). This diversity is most probably explained by stepping-stone through palaeo-Macaronesian islands and isolation (Triantis et al., 2010; Fernández-Palacios et al., 2011). Palaeoentomological records are rare in Madeira. The only known record is from the Mio-Pleistocene deposit of São Jorge (see Góis-Marques et al., 2018), where Heer (1857) described an extinct coleopteran, Laparocerus wollastoni, based on fossilized elytra. Machado (2006) in a taxonomic review of Laparocerus considers this taxon as nomen dubium, due to the missing holotype and the impossibility of reapraising its taxonomy. On other Macaronesian archipelagos, espe cially in the Canaries Islands, several deposits with insect ichnofossils have been described (e.g. Edwards & Meco, 2000; Meco et al., 2011; La Roche et al., 2014). In Azores only xylophagous ichnoentomological traces in charcoal wood are known (Góis-Marques et al., 2019b). The fossil wing was found within laminated lacustrine fine sandstone, associated with plant fossils. The sediments are constrained by two 40Ar-39Ar dates to 1.3 Ma, Calabrian stage (Góis-Marques et al., 2019a). Fossils are kept in the palaeobotanical collection at the Madeira University herbarium (UMad-P) with the numbers UMad-P500a (part) and UMad-P500b (counter-part). The wing fossil was studied under a stereo microscope, and its identification was performed through several sources (e.g. Goulet & Huber, 1993) and specific guidebooks (Prehn & Raper, 2016). Wing description follows the Comstock-Needham system as described by Quicke (2015)
- Inventory and review of the Mio–Pleistocene São Jorge flora (Madeira Island, Portugal): palaeoecological and biogeographical implicationsPublication . Góis-Marques, Carlos A.; Madeira, José; Sequeira, Miguel Menezes deThe occurrence of plant fossils on Madeira Island has been known since the mid-nineteenth century. Charles Lyell and George Hartung discovered a leaf bed rich in Lauraceae and fern fossils at S~ao Jorge in 1854. The determinations were controversial but a full review was never performed. Here we propose possible geological settings for the fossiliferous outcrop, and present an inventory and a systematic review of the surviving specimens of the S~ao Jorge macroflora. The S~ao Jorge leaf bed no longer outcrops due to a landslide in 1865. It was possible to establish the two alternative volcano stratigraphical settings in the sedimentary intercalations from the Middle Volcanic Complex, ranging in age from 7 to 1.8 Ma. The descriptions of Heer (1857), Bunbury (1859) and Hartung & Mayer (1864) are reviewed based on 82 surviving specimens. From the initial 37 taxa, we recognize only 20: Osmunda sp., Pteridium aquilinum, Asplenium cf. onopteris, aff. Asplenium, cf. Polystichum, cf. Davallia, Woodwardia radicans, Filicopsida gen. et sp. indet. 1 and 2, Ocotea foetens, Salix sp., Erica arborea, cf. Vaccinium, Rubus sp, cf. Myrtus, Magnoliopsida gen. et sp. indet. 1 to 3, Liliopsida gen. et sp. indet. 1. Magnoliopsida gen. et sp. indet. 4 is based on one previously undescribed flower or fruit. The floristic composition of the S~ao Jorge fossils resembles the current floristic association of temperate stink laurel (Ocotea foetens) forest, suggesting a warm and humid palaeoclimate and indicating that laurel forests were present in Macaronesia at least since the Gelasian, a time when the palaeotropical geofloral elements were almost extinct in Europe.
- First evidence of ants (Hymenoptera, Formicidae) in the early Pleistocene of Madeira Island (Portugal)Publication . Marques, Carlos A. Góis; Correia, Pedro; Nel, Andre; Madeira, José; Sequeira, Miguel Menezes deTo be successfully established on oceanic islands, native ants (Hymenoptera: Formicidae) have to migrate from the mainland or from nearby islands, crossing the ocean barriers, to find a suitable habitat. Despite the general interest on oceanic islands biotas, nothing is known about the deep-time migration and settling of native ants in these insular ecosystems. Palaeoentomological studies on oceanic islands that could provide palaeobiological information on Formicidae are scarce. Here, we describe and illustrate the first fossil of an ant from the Macaronesian archipelagos (Atlantic Ocean), based on a partial forewing found within 1.3 Ma (Calabrian, Pleistocene) lacustrine sediments from Madeira Island, Portugal. Although unidentifiable beyond the family level, this fossil record provides a minimum age for the presence of ants in the Madeira archipelago. Palaeoecologically, this record indicates the presence of suitable habitats for ants during the early Pleistocene.
- The bicentenary of Georg Hartung, a German pioneer geologist, explorer and illustratorPublication . Marques, Carlos A.Góis; Sequeira, Miguel Menezes de; Madeira, JoséWe present a tribute to Georg Friedrich Karl Hartung (1821–1891), a less-known, non-academic German geologist, on his 200th birthday anniversary. Influenced by eminent 19th century scientific personalities, such as Oswald Heer, Charles Lyell, and Alexander von Humboldt, he performed pioneer geological observations and sampling in the Azores, Madeira, and Canary Islands volcanic archipelagos. Later in his life, he travelled to the USA and explored the Scandinavian countries. His scientific endeavours were published in several books and papers, many of them co-authored by academic German geologists and palaeontologists. His works on Macaronesia are deemed as classics, and many have been enriched by his detailed geological illustrations.
- Oceanic Island forests buried by Holocene (Meghalayan) explosive eruptions: palaeobiodiversity in pre-anthropic volcanic charcoal from Faial Island (Azores, Portugal) and its palaeoecological implicationsPublication . Góis-Marques, C. A.; Rubiales, J. M.; Nascimento, L.; Sequeira, Miguel Menezes de; Fernández-Palacios, J. M.; Madeira, J.In Faial Island (Azores Archipelago, North Atlantic Ocean), charcoalified and mummified wood fossils have been reported within late Holocene (Meghalayan) pyroclastic deposits from the Caldeira Formation. Due to their re cent age, a detailed study conveys a snapshot into Azorean palaeophytodiversity and palaeovegetation, ca. 7–5 centuries before the arrival of Portuguese settlers to the Azores Islands. Here we provide the first detailed ana tomical and taxonomical study of these wood fossils. In total, 41 samples were collected from seven localities, mainly from a ~1200 yr BP ignimbrite. Field work revealed autochthonous and paraautochthonous assemblages, with tree trunks in upright position. The anatomical study of the fossil woods resulted in the identification of Juniperus brevifolia, Laurus azorica, Myrsine retusa, Morella faya, Picconia azorica, Prunus lusitanica subsp. azorica, and Vaccinium cylindraceum. Two fossil assemblages are comparable to the proposed potential natural vegetation (PNV) for the Azores. Surprisingly, P. lusitanica subsp. azorica was the second most abundant fossil wood suggest ing that this tree was more abundant in a recent past in Faial Island and probably in the archipelago. This is cor roborated by historical accounts, and its modern scarcity was certainly anthropically driven. Identifying Holocene plant macrofossils is essential to properly reconstruct oceanic islands terrestrial palaeoecosystems, especially where forests with high percentage of entomophilous taxa are underrepresented in palaeopalynological limnic record. Further work is necessary to reconstruct Faial Island and Azores archipelago palaeovegetation which is essential to provide an ecosystem base-line for restoration and management.