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Anodorhynchus Macaws as a followers of extinct megafauna

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... This species ingests whole palm fruits and during the rumination process, pulp is stripped off, with only the exocarp and mesocarp being consumed while the endocarp is regurgitated or defecated intact in latrines. Tapir latrines frequently present a high number of palm fruits with intact seeds inside the endocarps (Yamashita 1997;Quiroga-Castro & Roldán 2001), which are usually able to germinate after passing through digestive tract of tapirs (Capace et al. 2013). In Pantanal and Amazonia, latrines are usually at areas susceptible to seasonal floods, which may prevent seed germination (Quiroga-Castro & Roldán 2001). ...
... Palm seeds are an essential food source for macaws, especially for Anodorhynchus species that live preferentially in areas where this plant is abundant (Brandt & Machado 1990;Yamashita & Valle 1993;Yamashita 1987;. Although macaws are able to remove fruits directly from palm trees (Brandt & Machado 1990), they frequently obtain seeds from latrines and also cattle ruminations, which were suggested as evidence of their relationship with the extinct megafauna (Yamashita 1997;Pires et al. 2014). In fact, the habit of Anodorhynchus macaws of following cattle and also tapir herds is thought to be a behaviour derived from adaptations to follow the now extinct megaherbivores, which disappeared around 10,000 years ago (Yamashita 1997). ...
... Although macaws are able to remove fruits directly from palm trees (Brandt & Machado 1990), they frequently obtain seeds from latrines and also cattle ruminations, which were suggested as evidence of their relationship with the extinct megafauna (Yamashita 1997;Pires et al. 2014). In fact, the habit of Anodorhynchus macaws of following cattle and also tapir herds is thought to be a behaviour derived from adaptations to follow the now extinct megaherbivores, which disappeared around 10,000 years ago (Yamashita 1997). ...
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When foraging on a specific range of fruits or seeds, animals can represent a natural selection force, affecting plant population realized fecundity. Most studies have focused on characteristics of dispersed seeds, but the higher predation of seeds of specific size or shape is also an important force that influences the populations' realized fecundity and can help to understand patterns of plants genetic variability. In the Brazilian Pantanal, we investigated the predation of Acuri (Attalea phalerata) palm fruits by macaws, whose seeds go intact through tapirs digestive tract and then are deposited on latrines, where they are often consumed by seed predators. We found 251 fruits, containing between 1 and 5 seeds per fruit; 170 were intact and 81 had been partially damaged by macaws, mostly those containing 3 or 4 seeds per fruit and with thin endocarps. We argue that the higher consumption of fruits with these traits reflects a trade-off in the energy spent by macaws to open the fruits and the energy provided by them. Higher predation on 3-4 seeded fruits favors the selection of 1-2 seeded fruits, revealing a selection force opposed to the one exerted by bruchids and small rodents, which favors multi-seeded palm fruits. Frugivorous and seed predator animals play important roles in ecological processes involving plant populations, affecting individuals' reproductive success and selection for particular traits. Frugivorous and seed eating animals may choose which fruiting species to feed on, which plant individual to forage on and which fruits among those available in the tree they will consume. These choices may represent natural selection driving forces, when animals show preference for a specific size or shape of fruits or seeds, since they may influence plants' realized fecundity (Jordano 1995).
... Its patchy distribution in remote and inaccessible habitats, high daily and seasonal mobility, and longevity have impeded research aimed at understanding its conservation status and needs. Its diet ties it to a number of palm species; it depends heavily on Sterculia apetala trees for nest sites, and it shows a clear preference for open wooded, riparian and forest-edge habitats, often near pastures (Guedes, 2004;Pinho & Nogueira, 2003;Scherer-Neto et al., 2019;da Silva et al., 2019;Tella et al., 2020;Yamashita, 1997), but the precise factors governing its responses to land use change, and how these might differ in different parts of its huge range, are unknown (BirdLife International, 2019). Nevertheless, owing to population increases in the Pantanal (e.g. ...
... Aware that the macaw is specialized on the fruits of several palms, for example Acrocomia and Attalea spp. (summarized in Collar et al., 1992; see also Tella et al., 2020), we also included a combined palm/tree occurrence predictor layer, consisting of the summed habitat suitability values from eight separate species distribution models for seven palms (Acrocomia aculeata, Acrocomia totai, Attalea eichleri, Attalea maripa, Attalea phalerata, Attalea speciosa and Syagrus coronata) as well as one tree species (Sterculia apetala) known to be important for food and nesting, respectively (Collar et al., 1992;Johnson et al., 1997;Pinho & Nogueira, 2003;Pizo et al., 2008;Yamashita, 1997). ...
... Our results indicate a complex, changing distributional pattern of the Hyacinth Macaw in response to changing land cover over its range, with marked differences across the three biomes it inhabits. (Tella et al., 2020;Yamashita, 1997). Certainly, the known importance of palm stands to the species should not be underestimated (Yamashita & Valle, 1993). ...
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Aim To explore spatiotemporal changes in Hyacinth Macaw Anodorhynchus hyacinthinus distribution and the impacts of land use change over 25 years, across its vast range in central/eastern South America. Location Brazil, Bolivia and Paraguay, South America, covering almost 3 million km². Methods We use a novel, multi‐temporal species distribution model, to combine both year‐specific occurrence records and land use/cover data in a single model that is subsequently projected over a land cover time series. We investigate changes in geographic range over 25 years and potential drivers of range extent at multiple spatial scales (10 and 30 km) and time‐lags (current and 10 years previously). We also assess protected area coverage and impacts from degazettement within the remaining range and highlight priority areas to search for undiscovered populations. Results The modelled range of the species increased by 75,000 km² to over 500,000 km² between 1995 and 2019, during which period agriculture and pasture increased by 600,000 km² within the species' extent of occurrence. Habitat suitability was influenced most strongly by distribution of palms, forest cover and changes in pasture over 10 years and usually by predictors measured at larger spatial scales. Just 8% of the macaw's modelled range falls within protected areas, and more than 20,500 km² of protected areas overlapping with the range have been degazetted in the last 40 years. We highlight key areas to search for undiscovered populations in under‐sampled and remote areas, especially led by community citizen science initiatives involving indigenous groups and protected area staff. Main conclusions Novel modelling methods, combining multi‐temporal occurrence records and land cover data, appropriate for small sample sizes per year, have revealed habitat dynamics and changes to the range of this threatened species over 25 years. The method may have wide applicability for a range of species, including elusive and poorly studied species.
... A pesar de que d'Orbigny (en Bourjot Saint-Hilaire 1837-38) indica que no bajaban al suelo, seguramente si lo hacían para alimentarse. Este comportamiento es frecuente entre sus parientes cercanos, y de hecho es una característica que los diferencia de otros guacamayos (Yamashita 1997). Mientras que la mayor parte del grupo desciende, uno o más individuos permanecen en perchas elevadas, vigilando los alrededores para detectar eventuales amenazas (Yamashita 1987, Yamashita y Valle 1993. ...
... El uso de estos centinelas constituye una eficaz estrategia anti-depredatoria. Los guacamayos azules 6 se distinguen además por tener una dieta altamente especializada que se basa en el consumo de dátiles o "coquitos" de palmeras (Yamashita y Valle 1993, Yamashita 1997. La mandíbula de estos loros recuerda un cincel de surco cuadrado y el filo cortante del "cincel" es utilizado a modo de cuña para partir los dátiles con cortes transversales. ...
... Estos sitios habrían sido buscados activamente por los guacamayos con el fin de aprovechar tal fuente de alimento. En la actualidad esta función facilitadora es realizada por el ganado doméstico (Yamashita 1997). Cabe mencionar que, con respecto a sus referidas asociaciones de hábitat y supuesta dieta especializada, algunas de estas ideas han sido criticadas por estar basadas en información extremadamente limitada (Teixeira y Papavero 2016). ...
... The hyacinth macaw Anodorhynchus hyacinthinus (hereafter HM) is the largest parrot species in the world, while the congeneric Lear's macaw A. leari (hereafter LM) is phenotypically similar but somewhat smaller [26]. They have the strongest beaks among parrots, allowing them even to crack the nuts of large-sized palm fruits after defleshing them and discarding the mesocarp [25,27]. Therefore, these macaws might not be contributing to legitimate seed dispersal despite having been observed transporting fruits in flight using their beaks or feet [17,23]. ...
... As previously mentioned, Anodorhynchus macaws have been observed transporting fruits picked from the mother plants to distant perches for handling and consumption [17,23], but it is unknown whether all seeds are then predated, or a fraction of them survive predation and thus contributes to effective primary seed dispersal. On the other hand, these macaws have been reported actively searching for and predating on large palm seeds excreted by livestock, which has been interpreted as an evolutionary adaptation to exploit seeds excreted by extinct megafauna [27,29]. Similarly, it is unknown whether this tertiary dispersal process, following secondary dispersal by livestock, translates into effective seed dispersal. ...
... Parrots are able to exploit multiple foraging opportunities, encompassing virtually all plant parts in all maturation stages of a widesr variety of species [54]. However, Anodorhynchus macaws can be considered palm specialists [26][27][28][29], and, as we expected, they play a key role as dispersers of mostly large-fruited palms. Our results contributed to filling a large knowledge gap of palmfrugivore interactions, recently highlighted by Muñoz et al. [22], and supported their suggestion that parrots might play a role as palm seed dispersers. ...
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The extinction of ecological functions is increasingly considered a major component of biodiversity loss, given its pervasive effects on ecosystems, and it may precede the disappearance of the species engaged. Dispersal of many large-fruited (>4 cm diameter) plants is thought to have been handicapped after the extinction of megafauna in the Late Pleistocene and the recent defaunation of large mammals. We recorded the seed dispersal behavior of two macaws (Anodorhynchus hyacinthinus and Anodorhynchus leari) in three Neotropical biomes, totaling >1700 dispersal events from 18 plant species, 98% corresponding to six large-fruited palm species. Dispersal rates varied among palm species (5%–100%). Fruits were moved to perches at varying distances (means: 17–450 m, maximum 1620 m). Macaws also moved nuts after regurgitation by livestock, in an unusual case of tertiary dispersal, to distant perches. A high proportion (11%–75%) of dispersed nuts was found undamaged under perches, and palm recruitment was confirmed under 6%–73% of the perches. Our results showed that these macaws were legitimate, long-distance dispersers, and challenge the prevailing view that dispersal of large-fruited plants was compromised after megafauna extinction. The large range contraction of these threatened macaws, however, meant that these mutualistic interactions are functionally extinct over large areas at a continental scale.
... There is very scarce information about the destination microhabitat of spat seeds. Domestic species that forage at day often ruminate at night in pens or farmyards (e.g., Troup, 1921;Yamashita, 1997), where there is little or no probability of seed germination. Brodie et al. (2009a) conclude that sambar deer regurgitated higher numbers of seeds of Choerospondias axillaris, but muntjak was the only disperser moving them to open microhabitats, where germination was enhanced. ...
... In Brazil, >50% of seeds of the palm Acrocomia aculeata chewed by cattle were infested by bruchids, and this proportion increased to 99% after 27 days; regurgitated seeds had a significant higher rate of insect predation than seeds of non-chewed fruit (Scariot, 1998). Additionally, piles of seeds of the palms Acrocomia aculeata, Attalea phalerata, and Syagrus coronata regurgitated by cattle were regularly visited by macaws (Anodorhynchus leari and A. hyacinthinus) to feed on them, cracking the nuts (Yamashita, 1997). This last author hypothesizes that macaws could track the movements of the extinct Pleistocene megaherbivores in order to collect the large seeds they dispersed. ...
... (Sánchez-Valverde, 1785, 35.(Scariot, 1998), 36. (Slater anddu Toit, 2002), 37.(Sridhara et al., 2016), 38.(Tiver et al., 2001), 39.(Troup, 1921), 40.(Wilson and Clarke, 1962), 41.(Yamashita, 1997), 42.(Yasuda et al., 2005). ...
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Given their strong masticatory system and the powerful microbial digestion inside their complex guts, mammalian ruminants have been frequently considered seed predators rather than seed dispersers. A number of studies, however, have observed that ruminants are able to transport many viable seeds long distances, either attached to the hair or hooves (i.e., epizoochory) or inside their body after ingesting them (i.e., endozoochory). However, very few studies have investigated a modality of endozoochory exclusive to ruminants: the spitting of usually large-sized seeds while chewing the cud. A systematic review of the published information about this type of endozoochory shows a marked scarcity of studies. Nonetheless, at least 48 plant species belonging to 21 families are dispersed by ruminants in this manner. Most of these plants are shrubs and trees, have fleshy or dry fruits with large-sized seeds, and are seldom dispersed via defecation. Many cases have been observed in tropical areas, where more frugivorous ruminant species occur, but other records are from temperate and dry areas, covering thus all continents except Antarctica. Twenty-one species of ruminants from 18 genera have been reported as endozoochore spitters. They involve domestic and wild species belonging to the families Tragulidae, Cervidae, and Bovidae. This suggests that almost any ruminant species could potentially eat fruits and regurgitate large hard seeds during rumination. Likely, this seed dispersal mechanism has been neglected due to the difficulty of observing rumination behavior and locating spat seeds. Further research on the potential of wild and domestic ruminant species as long-distance seed dispersers through spitting seeds from the cud appears particularly important given their increasing pervasiveness and abundance worldwide.
... Su principal dispersor es el ganado vacuno, que recorre largas distancias en campos de pastura o cuando éste es transportado como ganado vivo para diversas regiones, llevando frutos en su interior (Scariot, 1998). Otros animales que contribuyen a la distribución de semillas de A. aculeata son murciélagos, monos, roedores como carpinchos (capibara), antas, ratas; y aves como tucanes, guacamayos, guacamayo jacinto (Arara), que en muchos casos aprovechan los frutos previamente rumiados por los bovinos (Yamashita, 1997;Scariot, 1998). ...
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La presencia de palmeras de cocotero denominadas “Mbocayá” o “Mbokaja” – Acrocomia aculeata (Jacq.) Lodd. ex Mart.– sinonimia Acrocomia totai Mart, es muy común en el Paraguay y se encuentra distribuida en gran parte del territorio nacional. Generalmente todas las partes de la planta son útiles para varios usos, sin embargo, los frutos o cocos, son los más utilizados a nivel industrial, porque son materia prima para varios productos, la almendra para producir aceites destinados a la industria cosmética, el carozo para combustible o carbón activado y la pulpa para aceite y forraje, por citar los más relevantes. Tradicionalmente se da mayor importancia a la almendra, descuidando la calidad de la pulpa que se deteriora durante la recolección y el almacenamiento, lo cual conlleva a la obtención de aceite de elevada acidez, limitando su uso posterior. Considerando las propiedades físicas y químicas del aceite de pulpa y su importancia para el rubro de la producción de biodiesel, fue desarrollado el Proyecto 14-INV-093 “Sistema de colecta y almacenamiento de frutos de Acrocomia aculeata y aprovechamiento del aceite de la pulpa con miras a su utilización como materia prima para la obtención de biodiesel” por la Facultad de Ciencias Químicas de la UNA con la colaboración de EMBRAPA Agroenergía (Brasil), y cofinanciado por el FONACIDE a través de CONACYT de Paraguay. Cumpliendo con uno de los objetivos del proyecto, este manual presenta las principales alternativas para la recolección y almacenamiento de frutos del mbokaja, en base a los resultados obtenidos experimentalmente por el equipo de investigadores y las prácticas empleadas en otros países o publicadas por otros investigadores, y que son consideradas factibles de ser adaptadas en el Paraguay. En cuanto al procesamiento para la extracción de aceite, se describe la tecnología empleada actualmente en la industria paraguaya y como alternativa se resumen algunos procesos estudiados a escala de laboratorio y planta piloto, acompañados de las características químicas y físicas del aceite. Por último, se describen algunos procesos de obtención de biodiesel convencional y se presenta una propuesta para obtención de biodiesel empleando el método de esterificación y transesterificación en condiciones supercríticas, que permite procesar materias primas con alto contenido de ácidos grasos libres y agua, sin catalizadores. Agradecemos a CONACYT por el cofinanciamiento y orientación pertinente para la gestión del proyecto, al Laboratorio de Procesos Químicos de EMBRAPA Agroenergía con sede en Brasilia, por la destacada colaboración en cuanto a capacitación, uso de laboratorio y por compartir experiencias muy importantes. Agradecimiento especial a la Facultad de Ciencias Agrarias - UNA por permitir el acceso a las plantaciones de Mbokaja del Campo Experimental y por el asesoramiento correspondiente. A los directivos, docentes e investigadores, estudiantes de grado, postgrado y personal de mantenimiento de la Facultad de Ciencias Químicas - UNA por aceptar el desafío y apoyar la ejecución de un trabajo muy complejo y multidisciplinario de mucho interés para el país. A las empresas Oleaginosas Industrializadas S.A. (OISA), Base Industrial Salinas S.A. y Aceitera Industrial S.A., por permitir el acceso a sus industrias y proveer información técnica de mucho valor.
... Por exemplo, o condor-andino Vultur gryphus, que hoje depende da carniça de camelídeos neotropicais e mamíferos marinhos na Patagônia e Andes, ocorria não muito tempo atrás em Minas Gerais, onde deveria ser dependente das carcaças de grandes mamíferos (Alvarenga 1998). Araras-azuis Anodorhynchus provavelmente sofreram reduções na suas áreas de distribuição devido à ruptura dos sistemas de dispersão de sementes de palmeiras que coevoluíram com grandes mamíferos (Yamashita 1997). Na América do Norte, várias espécies de aves necrófagas hoje extintas, incluindo os incríveis Teratornithidae, eram dependentes dos grandes mamíferos e os acompanharam à extinção (Grayson 1977, Feduccia 1996 e o mesmo deve ter ocorrido na América do Sul. ...
... Their observations and collections resulted in relevant publications on the species found in the region (Travassos, 1940(Travassos, , 1941Travassos & Freitas, 1942;Pinto, 1948;Ruschi, 1955;Travassos et al., 1957;Sick, 1961). In 1970, specimen collections were made by the Museum of Fauna (formerly Brazilian Institute of Forestry Development) from Fazenda Descalvados, Mato Grosso (Tubelis & Tomas, 2003b), and a multitude of contemporary authors such as Antas (1983), Munn et al. (1989), Yamashita & Valle (1990), Guedes (1993), Antas (1994), Willis (1995), Guedes & Harper (1995), Antas & Nascimento (1996), Oliveira (1997), Yamashita (1997), Pinho (1998), and Tubelis & Tomas (1999) focused on the biology of species and communities through capture, direct observation, and identification of vocalizations (Tubelis & Tomas, 2003b). Additional inventories and records were collated by Weinberg (1984), Antas et al. (1986), Cintra & Yamashita (1990), Willis & Oniki (1990), Lago-Paiva & Willis (1994), Pacheco & Bauer (1994), and Strüssmann (1998). ...
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The Pantanal floodplains of Brazil are a region of rich biodiversity. To date, the true richness of the Pantanal avifauna has not been explored satisfactorily caused by a lack of studies in the region and, especially, by the divergence of opinion among the works published by various authors on the many species found in the region. This is due to the lack of criteria in examining records, both with regard to the reliability of the identifications and in the precise geographical allocation. Therefore, in the study, we collage findings from various studies and records created by us in the last few decades from 199 distinct locations to produce a list of birds in the Pantanal floodplains. We grouped the results into three lists: primary, secondary, and tertiary. We found that the avifauna of the Pantanal floodplain is composed of a total of 617 species, of which 571 (92%) have supporting records of occurrence (primary list) and 46 still lack documentation (secondary list). The number of species listed here for the Pantanal floodplain represents 32% of all avifauna known to the Brazilian territory. This reflects the importance of the biome, as part of the national territory, for the maintenance of a meaningful avifaunistic richness. Migratory birds (n = 183), notably northern ones (n = 43), are among the main players involved in ecological processes of nutrient cycling and dispersion of ARTICLE important pathogens between the two continents. With regard to conservation, 25 species are included in some category of threat in the lists of threatened species with global extinction. We hope that our list will help future researchers a more definitive approach when researching the avian fauna in this bountiful region.
... In the case of motacú, it is feasible that cattle could play a role in facilitating seedling regeneration. Cattle readily consume and disperse the fruits of Attalea palm species, depositing them in regurgitated heaps or dung piles [34,60,61]. The dung provides a mechanical barrier to bruchid beetle oviposition [62,63] leading to reduced rates of infestation [34,62] that may otherwise limit seedling growth in areas where fruits are densely deposited, e.g. ...
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Globally, forest-savanna mosaic landscapes are of significant conservation importance but have been widely impacted by human land-use. We studied how restoration, through cessation of long-term cattle grazing impacts (i) forest regeneration; (ii) forest understory structure and composition; and (iii) populations and diversity of large mammals and nocturnal birds, within naturally patchy gallery forests in the Beni Savannas of Bolivia. Comparing grazed and restored sites, we assessed the abundance and composition of tree functional types at different life stages (seedlings, saplings and adults), with focus on the region’s key palm species Attalea princeps (motacú). Additionally, we surveyed habitat structure and composition in the shrub and ground-layer, and monitored occurrence and encounter rates of large mammals and nocturnal birds along dusk and evening transects. We found evidence of lower recruitment of motacú palms on the grazed site and lower potential for natural motacú regeneration. Principal Components Analysis revealed forests on grazed sites had simpler, more open shrub-layers and altered ground-layer structure and composition including increased bare ground. Mammal species richness was greater on the restored site, and there were more declining, globally threatened and site-unique species. Species richness was similar for nocturnal birds within forests on both the grazed and restored site, but nearly all species tended to be encountered more frequently on the restored site. Our results suggest cattle negatively impact forest regeneration and alter the structure and composition of the shrub and ground layer with potential consequences for the diversity and abundance of wildlife. Our study represents one of only a handful completed in the Beni region of Bolivia to date. The Beni is currently under pressure from widespread, largely unregulated cattle ranching. Our results thus provide vital evidence to support development of restoration and conservation policy, and its integration with rangeland management in this threatened and critically understudied region.
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