ABSTRACT
Resumen El orden Ciconiiformes incluye especies de aves zancudas de tamaño mediano a grande, con dinámicas de crecimiento corporal que pueden ser analizadas para esclarecer las tendencias microevolutivas del grupo. Las garzas (familia Ardeidae) provienen de un ancestro común, sin embargo, existen diferencias en la forma corporal de los adultos actuales y se ha sugerido un proceso de evolución por heterocronías. Sin embargo, los estudios de crecimiento previamente se han enfocado solo en las dimensiones lineales y no se han analizado los cambios alométricos. En el presente trabajo se describen los cambios de proporciones corporales durante el crecimiento en siete especies de ardéidos y se analizan bajo una hipótesis filogenética para identificar el patrón morfológico primitivo entre los géneros basales Butorides y Nycticorax. Para ellos se calcularon las proporciones pico / tarso en 353 pichones, medidos entre 1998 y 2006, y se evaluaron sus cambios en relación con la edad y el incremento del peso corporal. Las especies mostraron diferencias marcadas en la magnitud del cambio en las proporciones pico/tarso entre la eclosión y un momento análogo del crecimiento, excepto Bubulcus que tiende a crecer de forma casi isométrica. Los cambios alométricos al crecer conjuntamente con las proporciones al nacer, generan un crecimiento diferencial que produce las disímiles morfologías adultas que se expresan en el grupo de las garzas. La tendencia general de estos cambios es de un incremento ligero en el medio del crecimiento para luego descender a casi la misma proporción inicial. El crecimiento en las primeras etapas de vida tiende a ser más isométrico y las diferencias se acentúan en momentos más tardíos. La hipótesis de asumir a Nycticorax como morfotipo peramórfico es más parsimoniosa en las tendencias de cambio dentro del grupo, resultando en un alargamiento relativo con alometría positiva del pico y del tarso en todas las especies. Esta hipótesis sería consistente con una hipermorfosis gradual que alcanzaría su máxima expresión en Ardea.
Abstract The order Ciconiiformes include wading bird species of sizes from medium to high, with body growth dynamic that can be analyzed to enlighten micro evolutionary trends. Egrets and herons (family Ardeidae) evolved from a common ancestor, but there are differences in adult body shapes, and their evolution has been suggested to be based on heterochronic processes. However, previous researches on growth have focused only in lineal dimension, and alometric changes have not been studied. In the current paper I described changes in body proportions during growth in seven ardeid species, and analyzed body growth under a phylogenetic point of view, to identify the primitive morphology pattern among genus Butorides and Nycticorax. For this purpose, I calculated bill/ tarsus rate in 353 nestlings, measured between 1998 and 2006, and assessed their changes with age and body weight. All species showed marked differences in proportion changes extension between hatching and an analogous growth moment, except Bubulcus that grows almost isometrically. Alometric changes during growth and at hatch, generate a differential growth that produced the different adult morphologies expressed among egrets and herons. The general trends were toward a slight increase in the middle of the growth period up to a lowering to almost the same initial proportions. Growth in the first life stages tends to be more isometric and differences get higher latter in growth. The hypothesis of Nycticorax as peramorphic morfotype is more parsimonious with changes trends in the group, resulting in a relative extremities extension with positive alometry in bill and tarsus in all species. This hypothesis is consistent with a gradual hipermorphosis that reaches a maximum expression in Ardea. Rev. Biol. Trop. 65 (4): 1347-1357. Epub 2017 December 01.
ABSTRACT
Intraclutch egg size variation in birds depends on many ecological factors and on the evolutive history of each species. In wading birds, a trend to smaller eggs with laying order has been described, but comparative reports are scarce. In this study, egg size variation patterns were described for nine Egrets and Heron species nesting in Birama’ Swamp, Cuba. The patterns were described using external dimensions of 3 142 eggs from 1 875 nests of Butorides virescens, Bubulcus ibis, Ardea alba, Nycticorax nycticorax, Nyctanassa violacea and four Egretta species, taken in the field between 1998 and 2006. Results showed that eggs were 4.9-10% of adult weight and had volume variation coefficients between 6-9%. There were no general and consistent interspecies relationship between clutch size and egg sizes. Average volumes tend to get smaller with laying order, but it is not statistically detectable in Butorides and Bubulcus. Last egg was between 0.2% and 15% smaller than the first, showing an inverse relationship with it. Intraclutch asymmetry is light in E. thula and fluctuating around null in Bubulcus. Size only predicted laying or hatching order for the last egg, in nests with more than two eggs, with 72.4% of confidence.
Las variaciones intranidada de la talla de los huevos de las aves responden a numerosos factores ecológicos y de la historia evolutiva de las especies. En las zancudas aunque se ha descrito una disminución de la talla con el orden de puesta, los trabajos comparativos entre especies son escasos. En este trabajo se describen los patrones de variación del tamaño de los huevos de nueve especies de garzas (Aves: Ardeidae) nidificantes en la ciénaga de Birama, Cuba. Los patrones se describen a partir de las dimensiones de 3 142 huevos, pertenecientes a 1 875 nidos de Butorides virescens, Bubulcus ibis, Ardea alba, Nycticorax nycticorax, Nyctanassa violacea y cuatro especies de Egretta, tomados en la ciénaga de Birama, Cuba, entre 1998 y 2006. Los huevos representaron entre el 4.9-10% del peso adulto y tuvieron un coeficiente de variación del volumen entre el 6-9%. No existió una relación general consistente entre especies entre el tamaño de puesta y las dimensiones de los huevos. El volumen promedio del huevo tiende a disminuir con el orden de puesta, pero no es estadísticamente detectable en Butorides y Bubulcus. El último huevo fue entre un 0.2 y un 15% menor que el primero, disminuyendo su volumen relativo con la talla del primer huevo. La asimetría intranidada es menos marcada en E. thula y prácticamente fluctuante alrededor de cero en Bubulcus. En general, la talla del huevo solo puede predecir el orden de puesta para el último huevo en nidadas mayores de dos con un 72.4% de confianza.
Subject(s)
Animals , Female , Male , Beak/anatomy & histology , Birds/classification , Eggs , Tarsus, Animal/anatomy & histology , Cuba , Nesting BehaviorABSTRACT
Egg shape is difficult to quantify due to the lack of an exact formula to describe its geometry. Here I described a simple algorithm to characterize and compare egg shapes using Fourier functions. These functions can delineate any closed contour and had been previously applied to describe several biological objects. I described, step by step, the process of data acquisition, processing and the use of the SHAPE software to extract function coefficients in a study case. I compared egg shapes in three birds' species representing different reproductive strategies: Cuban Parakeet (Aratinga euops), Royal Tern (Thalasseus maximus) and Cuban Blackbird (Dives atroviolaceus). Using 73 digital pictures of eggs kept in Cuban scientific collections, I calculated Fourier descriptors with 4, 6, 8, 16 and 20 harmonics. Descriptors were reduced by a Principal Component Analysis and the scores of the eigen-values that account for 90% of variance were used in a Lineal Discriminant Function to analyze the possibility to differentiate eggs according to its shapes. Using four harmonics, the first five component accounted for 97% of shape variances; more harmonics diluted the variance increasing to eight the number of components needed to explain most of the variation. Convex polygons in the discriminant space showed a clear separation between species, allowing trustful discrimination (classification errors between 7-15%). Misclassifications were related to specific egg shape variability between species. In the study case, A. euops eggs were perfectly classified, but for the other species, errors ranged from 5 to 29% of misclassifications, in relation to the numbers or harmonics and components used. The proposed algorithm, despite its apparent mathematical complexity, showed many advantages to describe eggs shape allowing a deeper understanding of factors related to this variable.
Subject(s)
Birds , Fourier Analysis , Ovum , Algorithms , Animals , Principal Component AnalysisABSTRACT
Avian postnatal growth has received considerable attention and its ecological implications have been deeply analyzed. In this current paper, I describe the patterns of culmen and tarsus growth, as well as of weight gain patterns in eight species of herons and egrets (Aves: Ardeidae) found in the Birama Swamp in Eastern Cuba. Between 1998 and 2006,714 nestlings of the following species were measured every two days: Butorides virescens, Bubulcus ibis, Egretta thula, E. tricolor, E. caerulea, E. rufescens, Ardea alba and Nycticorax nycticorax. Logistic and Gompertz equations were adjusted to data using non-lineal regression models with adult values as the asymptote. For each species, the following were determined and recorded: growth rate, age at inflexion, instantaneous growth rates at each age interval, and time taken to reach 90% of adult size. Reported hatchling sizes were similar in other localities, with a variation coefficient ranging between 10-19%. At hatch, each species exhibited differing sizes relative to adult values. In all cases, Gompertz equations were best fitted to explain more variance and lesser residuals. Rates of weight change and tarsus growth were alometrically related to the log of adult weight. Two main growth processes were identified: a physical extension in dimensions of each measurement reflecting inter-specific morphometric differences, and a lineal increase of the growth period from Green Heron to Great Egret. The Black-crowned Night Heron, Cattle Egret and Reddish Egret exhibited some unique measurement characteristics in comparison to the remaining members of the family. All results support the hypothesis that hypermorphosis, as the main evolutionary process in the microevolution of Ardeidae, is caused by a delayed final moment of growth.
