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Mixed bird flocks: patterns of activity and species composition in a region of the Central Andes of Colombia. Bandadas mixtas de aves: patrones de actividad y composición de especies en una región de la Cordillera Central de los Andes de Colombia. Rodríguez-Correa 1,3 and Manuela Restrepo-Chica 1. Programa de Licenciatura en Biología y Educación Ambiental. Facultad de Educación. Universidad del Quindío. Cra 15, Cll 12 N, Armenia, Quindío. Departamento de Biología Evolutiva. Facultad de Ciencias.
Universidad Nacional Autónoma de México. Apartado postal, México D. Recibido: 15 abril Aceptado: 09 septiembre Mixed what do birds compete for flocks are groups of individuals from different species that travel and forage together. Such groups are common in several bird communities around the world. We present species composition and activity patterns of mixed bird flocks in a region of the Central Andes of Colombia. We compared the number of species per flock, as well as the number of flocks among 3 different habitats.
We tested hypotheses concerning the flocks daily activity and the co-occurrences of species within them. We recorded 75 species, and the species number per flock varied from 4 to Our data suggest that habitat affects the number of flocks but not their species number, and that the activity of flocks is similar throughout the day. In addition, the association of birds in flocks is affected by interspecific facilitation, with some species co-occurrences found more times than expected by chance.
We hypothesize that some tanager species could have a role in flock cohesion. We witnessed 2 predator attacks upon flocks, a number of agonistic interactions among flock members, and squirrels following bird flocks. Our results meet some general patterns described for mixed what do birds compete for flocks. Las bandadas mixtas de aves son grupos de individuos what do birds compete for diferentes especies que viajan y forrajean what do birds compete for, y son comunes en varias comunidades de aves alrededor del mundo.
Presentamos la composición de especies y los patrones de actividad de las bandadas mixtas de aves en una región de la Cordillera Central de los Andes Colombianos. Hipotetizamos que algunas tangaras pueden tener un papel en la cohesión de las bandadas. Observamos 2 ataques de depredadores, así como varias interacciones hostiles entre miembros de las bandadas y la participación de ardillas. Nuestros resultados se ajustan a ciertos patrones descritos para las bandadas mixtas.
Heterospecific groups, defined as wild groups of animals made up of individuals of different species traveling and feeding together, are a common phenomenon what do birds compete for different habitats around the world. These groups have been what do birds compete for for birds, mammals, and fishes Morse, ; Terborgh, ; Haugaasen and Peres, ; Goodale et al. In general, heterospecific groups of birds have been studied in more detail Morse, Mixed bird flocks, also known as avian mixed-species flocks or mixed bird parties, are ubiquitous phenomena, and represent a conspicuous and prevalent characteristic of many habitats e.
Mixed bird flocks referred hereafter as MBFs result from attractions among their participants and not from an external concentration of food resources Morse, ; Powell, ; Sridhar et al. MBFs are different from bird feeding aggregations that form around clumped resources such as water, fruiting trees, or army ant swarms Diamond does eating meat cause prostate cancer Terborgh, ; Morse, ; Powell, ; Sridhar et al.
MBFs show wide variation in size, temporary cohesion, and strength of association Terborgh, ; Sridhar et al. The species that join MBFs have been divided into groups according to their role in the cohesion of the flock e. Two major non-mutually exclusive hypotheses have been proposed to explain why and how birds might benefit from flocking behavior e. The first suggests that birds can get feeding benefits from joining flocks by obtaining food more efficiently than when solitary Morse, ; Munn and Terborgh, ; Powell, ; Jullien and Clobert, ; Sridhar et al.
Alternatively, the second hypothesis considers the flock as a strategy for predator avoidance through several behavioral mechanisms Morse, ; Powell, ; Jullien and Clobert, ; Sridhar et al. Further, MBFs may simultaneously confer more than one benefit upon their members, and it is possible that each species obtains different benefits while staying in MBFs Morse, ; Moynihan, Nevertheless, the MBFs may also generate costs for their participants such as an increase in competition and aggressiveness, changes in foraging patterns, and increases in predation risk because these groups are more noisy and conspicuous than the solitary foragers Short, ; Jones, ; Hutto, ; Jullien and Clobert, In fact, previous studies have tested these hypotheses, but no consensus has arisen on the primary cause involved in the formation of MBFs e.
Altogether, MBFs in different habitats and at various latitudinal and altitudinal sites appear to have some features in common, but others are highly distinct Poulsen, Studies of MBFs have illustrated several patterns. First, the number of species in MBFs in 1 locality reflects the local bird richness Hutto, what do birds compete for Latta and Wunderle, ; Bohórquez, ; Brandt et al. Third, species typically acting solitary in 1 type of habitat e. Fourth, MBFs exhibit variation in their activity throughout the day Eguchi et al.
Fifth, species with cohesive roles seem to occur in mono-specific flocks and exhibit some special plumage color pattern or particular behavioral traits Moynihan, ; Powell, ; Tubelis, ; Goodale and Beauchamp, To test the generality of these patterns, comparisons can be drawn with information gathered about MBFs in complex and highly diverse biological systems such as the What do birds compete for avifauna of northern South America, for which the documentation of this phenomenon is scarce Moynihan, ; Poulsen, ; Montero, ; Bohórquez, Considering the latter, we focused our efforts in this study to document some of the mentioned patterns of MBFs in a region of the Central Andes of Colombia by 1 describing species composition of MBFs; 2 comparing the number of species per MBF among different habitats; 3 checking for variation in the number of MBFs among different habitats and through time; 4 evaluating if there is evidence of cohesion in these groups by testing the hypothesis that species join MBFs randomly; and 5 placing our data in the framework of other studies conducted around the world.
Study area. This natural reserve is located in the Salento Municipality of Quindío Department on the western slope of the Central Andes of Colombia. The reserve encompasses ha, and is located within a zone of very humid sub-montane forest with steep topography. The landscape is composed of different habitats such as mature old growth forest, secondary old growth forest, early secondary forest, and commercial forestry plantations of Pinus sp.
In fact, some patches of such pastures still remain in the zone. Shrubs and is simple linear regression the same as correlation in the study area belong to more than 59 species from 30 families, of which Asteraceae, Cunoniaceae, Melastomataceae, and Rubiaceae are the most common. We conducted MBF observations along 3 sections ranging from 2 to 3 km in length.
Each section was located in a different type of habitat and was separated from other sections by 0. Section A was located in year-old second growth forest. This section is bordered by a road, and has a 10 to 15 m height canopy with some emerging trees that reach up what do birds compete for 21 m, and a dense what do birds compete for of shrubs up to 4.
Section B was located in year-old scrubby second growth. This section is crossed by a path, and harbors principally shrubs between 3. It also contains some scattered groves with trees up to 7. Section C was located in a mature old growth forest with low human intervention on a steep slope near a river Río Boquía. This section is crossed by a narrow path.
It is necessary to note that all these sections are intermixed in different degrees and do not represent a linear gradient. We chose these sections because they are representative of the habitats at La Patasola and reflect the condition of other montane forests along the Colombian Andes. We took precautions to ensure that the time of surveys in each section was the same. Despite the fact that the 3 sections were not separated by long distances what do birds compete for by some kind of barrier, we consider our observation scheme as useful for our aims related to describing patterns in MBFs among sections representing different habitats and through time, not centered on documenting differences related to spatial separation.
Data collection. Data collection was carried out by conducting 4 field trips between September of and February of Each field trip lasted 3 to 4 days, including observations for a total of hours 36 hours in each section. We defined a MBF as 1 group of 2 or more individuals of what do birds compete for species moving around actively, showing evidence of cohesion e. A similar definition for MBF was used in other studies e. The differentiation of the MBFs from another kind of birds groups, like frugivorous birds foraging in trees, is straightforward due to the conspicuousness of MBFs being enhanced by the fact that meaning of green manure in english tend to remain continually on the move Morse, MBFs in the Andes and other areas of South America seem to be loosely structured, presenting turnover of individuals and lacking territoriality Poulsen, ; Aleixo, ; Herzog et al.
It is suggestive that the same MBF is not maintained for a long time. However, to avoid taking data from the same MBF repeatedly we used 3 different strategies. First, we checked the direction of movement of each MBF and compared that with the next MBF detected, assuring that it was coming from a different direction. Second, we conducted observations in each section for only 3 hours daily, avoiding prolonged continuous observations in the same place.
Third, there were at least 10 days between each pair of field trips. We carried out observations along the main trail present in each section see above. We detected MBFs by observations through what do birds compete for, since there was extreme multi species bird activity. Once a MBF was located, it was followed and observed as long as possible, noting the time of detection and the species composition.
Other information on agonistic interactions among constituent members, predator attacks upon them, and the number of co-specific individuals was observed when the opportunity arose. For some MBFs we did not document the species composition; these cases were excluded from the analyses dealing with the number of species.
For the species observed, we followed the taxonomic treatment of Remsen et al. Data analyses. To verify if the set of species recorded joining MBFs were well represented in what do birds compete for observations, we used EstimateS 7. It was followed by a Mann Whitney pairwise comparison. Finally, we used 2 different approaches to test if species joined MBFs on a random basis. On one hand, we tested the effect of facilitation among species in the cohesion of MBFs, as a whole using CoOccur version 1.
For this analysis we grouped the species by family following Remsen et al. These comparisons were carried out only for species present in 4 or more MBFs. This last approximation and some variants have been used previously to test association among species in MBFs using sound effects in writing. Jones, ; Hutto, ; Latta and Wunderle, We detected a total of MBFs.
Of these, 71 were observed under conditions that what do birds compete for us to document their species composition, and were included in the analyses that consider the number of species. We recorded a total of 75 species joining MBFs Appendix.