Jessica H. Arbour

TESTING FOR ANCIENT ADAPTIVE RADIATIONS IN NEOTROPICAL CICHLID FISHES

Most contemporary studies of adaptive radiation focus on relatively recent and geographically restricted clades. It is less clear whether diversification of ancient clades spanning entire continents is consistent with adaptive radiation. We used novel fossil calibrations to generate a chronogram of Neotropical cichlid fishes and to test whether patterns of lineage and morphological diversification are congruent with hypothesized adaptive radiations in South and Central America. We found that diversification in the Neotropical cichlid clade and the highly diverse tribe Geophagini was consistent with diversity‐dependent, early bursts of divergence followed by decreased rates of lineage accumulation. South American Geophagini underwent early rapid differentiation in body shape, expanding into novel morphological space characterized by elongate‐bodied predators. Divergence in head shape attributes associated with trophic specialization evolved under strong adaptive constraints in all Neotropical cichlid clades. The South American Cichlasomatini followed patterns consistent with constant rates of morphological divergence. Although morphological diversification in South American Heroini was limited, Eocene invasion of Central American habitats was followed by convergent diversification mirroring variation observed in Geophagini. Diversification in Neotropical cichlids was influenced by the early adaptive radiation of Geophagini, which potentially limited differentiation in other cichlid clades.

Ecological variation in South American geophagine cichlids arose during an early burst of adaptive morphological and functional evolution

Diversity and disparity are unequally distributed both phylogenetically and geographically. This uneven distribution may be owing to differences in diversification rates between clades resulting from processes such as adaptive radiation. We examined the rate and distribution of evolution in feeding biomechanics in the extremely diverse and continentally distributed South American geophagine cichlids. Evolutionary patterns in multivariate functional morphospace were examined using a phylomorphospace approach, disparity-through-time analyses and by comparing Brownian motion (BM) and adaptive peak evolutionary models using maximum likelihood. The most species-rich and functionally disparate clade (CAS) expanded more efficiently in morphospace and evolved more rapidly compared with both BM expectations and its sister clade (GGD). Members of the CAS clade also exhibited an early burst in functional evolution that corresponds to the development of modern ecological roles and may have been related to the colonization of a novel adaptive peak characterized by fast oral jaw mechanics. Furthermore, reduced ecological opportunity following this early burst may have restricted functional evolution in the GGD clade, which is less species-rich and more ecologically specialized. Patterns of evolution in ecologically important functional traits are consistent with a pattern of adaptive radiation within the most diverse clade of Geophagini.

Testing of the effect of missing data estimation and distribution in morphometric multivariate data analyses.

Missing data are an unavoidable problem in biological data sets and the performance of missing data deletion and estimation techniques in morphometric data sets is poorly understood. Here, a novel method is used to measure the introduced error of multiple techniques on a representative sample. A large sample of extant crocodilian skulls was measured and analyzed with principal component analysis (PCA). Twenty-three different proportions of missing data were introduced into the data set, estimated, analyzed, and compared with the original result using Procrustes superimposition. Previous work investigating the effects of missing data input missing values randomly, a non-biological phenomenon. Here, missing data were introduced into the data set using three methodologies: purely at random, as a function of the Euclidean distance between respective measurements (simulating anatomical regions), and as a function of the portion of the sample occupied by each taxon (simulating unequal missing data in rare taxa). Gowers distance was found to be the best performing non-estimation method, and Bayesian PCA the best performing estimation method. Specimens of the taxa with small sample sizes and those most morphologically disparate had the highest estimation error. Distribution of missing data had a significant effect on the estimation error for almost all methods and proportions. Taxonomically biased missing data tended to show similar trends to random, but with higher error rates. Anatomically biased missing data showed a much greater deviation from random than the taxonomic bias, and with magnitudes dependent on the estimation method.

Morphometric and genetic analyses of two sympatric morphs of Arctic char (Salvelinus alpinus) in the Canadian High Arctic

Multivariate morphometric analyses were used to examine variation in head, body, and fin shape between two sympatric morphotypes of Arctic char (Salvelinus alpinus (L., 1758)) from Lake Hazen, Ellesmere Island, Nunavut, Canada. Population structure of the Lake Hazen Arctic char was examined using five microsatellite loci. The “small” morph was found to have a larger (primarily deeper) head, larger and more elongate fins, and a deeper lateral profile than the “large” morph. The morphs also differed in allometric growth patterns. The large and small morphs do not appear to represent genetically distinct populations. The head morphology of the Lake Hazen small and large morphs exhibited similarities to benthic and pelagic morphs (respectively) from other lakes. We hypothesize that the large morph may be adapted to high-efficiency swimming and that the small morph may be adapted to low-efficiency, high-acceleration swimming. Such functional trade-offs are not uncommon among fish specializing in dispersed or m...

Incomplete specimens in geometric morphometric analyses

Summary The analysis of morphological diversity frequently relies on the use of multivariate methods for characterizing biological shape. However, many of these methods are intolerant of missing data, which can limit the use of rare taxa and hinder the study of broad patterns of ecological diversity and morphological evolution. This study applied a mutli-data set approach to compare variation in missing data estimation and its effect on geometric morphometric analyses across taxonomically variable groups, landmark position and sample sizes. Missing morphometric landmark data were simulated from five real, complete data sets, including modern fish, primates and extinct theropod dinosaurs. Missing landmarks were then estimated using several standard approaches and a geometric-morphometric-specific method. The accuracy of missing data estimation was determined for each estimation method, landmark position and morphological data set. Procrustes superimposition was used to compare the eigenvectors and principal component scores of a geometric morphometric analysis of the original landmark data, to data sets with A) missing values estimated, or B) simulated incomplete specimens excluded, for varying levels of specimens incompleteness and sample sizes. Standard estimation techniques were more reliable estimators and had lower impacts on morphometric analysis compared with a geometric-morphometric-specific estimator. For most data sets and estimation techniques, estimating missing data produced a better fit to the structure of the original data than exclusion of incomplete specimens, and this was maintained even at considerably reduced sample sizes. The impact of missing data on geometric morphometric analysis was disproportionately affected by the most fragmentary specimens. Missing data estimation was influenced by variability of specific anatomical features and may be improved by a better understanding of shape variation present in a data set. Our results suggest that the inclusion of incomplete specimens through the use of effective missing data estimators better reflects the patterns of shape variation within a data set than using only complete specimens; however, the effectiveness of missing data estimation can be maximized by excluding only the most incomplete specimens. It is advised that missing data estimators be evaluated for each data set and landmark independently, as the effectiveness of estimators can vary strongly and unpredictably between different taxa and structures.

Morphology and efficiency of a specialized foraging behavior, sediment sifting, in neotropical cichlid fishes.

Understanding of relationships between morphology and ecological performance can help to reveal how natural selection drives biological diversification. We investigate relationships between feeding behavior, foraging performance and morphology within a diverse group of teleost fishes, and examine the extent to which associations can be explained by evolutionary relatedness. Morphological adaptation associated with sediment sifting was examined using a phylogenetic linear discriminant analysis on a set of ecomorphological traits from 27 species of Neotropical cichlids. For most sifting taxa, feeding behavior could be effectively predicted by a linear discriminant function of ecomorphology across multiple clades of sediment sifters, and this pattern could not be explained by shared evolutionary history alone. Additionally, we tested foraging efficiency in seven Neotropical cichlid species, five of which are specialized benthic feeders with differing head morphology. Efficiency was evaluated based on the degree to which invertebrate prey could be retrieved at different depths of sediment. Feeding performance was compared both with respect to feeding mode and species using a phylogenetic ANCOVA, with substrate depth as a covariate. Benthic foraging performance was constant across sediment depths in non-sifters but declined with depth in sifters. The non-sifting Hypsophrys used sweeping motions of the body and fins to excavate large pits to uncover prey; this tactic was more efficient for consuming deeply buried invertebrates than observed among sediment sifters. Findings indicate that similar feeding performance among sediment-sifting cichlids extracting invertebrate prey from shallow sediment layers reflects constraints associated with functional morphology and, to a lesser extent, phylogeny.

Selection towards different adaptive optima drove the early diversification of locomotor phenotypes in the radiation of Neotropical geophagine cichlids

BackgroundSimpson envisaged a conceptual model of adaptive radiation in which lineages diversify into “adaptive zones” within a macroevolutionary adaptive landscape. However, only a handful of studies have empirically investigated this adaptive landscape and its consequences for our interpretation of the underlying mechanisms of phenotypic evolution. In fish radiations the evolution of locomotor phenotypes may represent an important dimension of ecomorphological diversification given the implications of locomotion for feeding and habitat use. Neotropical geophagine cichlids represent a newly identified adaptive radiation and provide a useful system for studying patterns of locomotor diversification and the implications of selective constraints on phenotypic divergence in general.ResultsWe use multivariate ordination, models of phenotypic evolution and posterior predictive approaches to investigate the macroevolutionary adaptive landscape and test for evidence of early divergence of locomotor phenotypes in Geophagini. The evolution of locomotor phenotypes was characterized by selection towards at least two distinct adaptive peaks and the early divergence of modern morphological disparity. One adaptive peak included the benthic and epibenthic invertivores and was characterized by fishes with deep, laterally compressed bodies that optimize precise, slow-swimming manoeuvres. The second adaptive peak resulted from a shift in adaptive optima in the species-rich ram-feeding/rheophilic Crenicichla-Teleocichla clade and was characterized by species with streamlined bodies that optimize fast starts and rapid manoeuvres. Evolutionary models and posterior predictive approaches favoured an early shift to a new adaptive peak over decreasing rates of evolution as the underlying process driving the early divergence of locomotor phenotypes.ConclusionsThe influence of multiple adaptive peaks on the divergence of locomotor phenotypes in Geophagini is compatible with the expectations of an ecologically driven adaptive radiation. This study confirms that the diversification of locomotor phenotypes represents an important dimension of phenotypic evolution in the geophagine adaptive radiation. It also suggests that the commonly observed early burst of phenotypic evolution during adaptive radiations may be better explained by the concentration of shifts to new adaptive peaks deep in the phylogeny rather than overall decreasing rates of evolution.

Guianacara dacrya, a new species from the rio Branco and Essequibo River drainages of the Guiana Shield (Perciformes: Cichlidae)

A new species of Guianacara is described from tributaries of the Essequibo River and the rio Branco in Guyana and northern Brazil. Guianacara dacrya, new species, can be diagnosed from all congeners by the possession of a unique infraorbital stripe and by the shape of the lateral margin of the lower pharyngeal jaw tooth plate. Guianacara dacrya can be further distinguished from G. geayi, G. owroewefi, G. sphenozona and G. stergiosi by the possession of a thin midlateral bar, from G. cuyunii by the possession of dusky branchiostegal membranes and from G. oelemariensis by the possession of two supraneurals. This species differs from most congeners by the presence of white spots on the spiny portion of the dorsal fin, the placement of the midlateral spot, the presence of filaments on the dorsal, anal and in rare cases the caudal-fin and from at least the Venezuelan species by several morphometric variables. Guianacara dacrya is known from the Essequibo, Takutu and Ireng River basins of Guyana and possibly from the rio Uraricoera in the rio Branco basin in Brazil. A key to the species is provided.

A New Species of Bujurquina (Teleostei: Cichlidae) from the Rio Danta, Ecuador, with a Key to the Species in the Genus

Bujurquina pardus is described from 12 specimens collected in 1989 from the Río Danta in the upper reaches of the Río Tigre drainage in the Pastaza province of Ecuador. It is distinguished from all other Bujurquina by a pattern of small dark spots arranged in rows along its flanks. Bujurquina pardus can be further distinguished from all other Bujurquina by the combination of whitish hyaline dorsal-fin lappet margins, rather than black lappets margins, and a lateral band which reaches the dorsal edge of the caudal peduncle, rather than only reaching the soft dorsal-fin base. It can be further distinguished from many of its congeners by the presence of spotting on the spinous dorsal fin and in having short pectoral fins (<32% of SL). A key to all the currently described species of Bujurquina is provided. Bujurquina pardus se describe de 12 individuos colectados en 1989 en el Río Danta en la cuenca superior del Río Tigre, Provincia de Pastaza, Ecuador. Se distingue de todas las otras especies de Bujurquina por el patrón único de pequeños puntos oscuros dispuestos en filas a lo largo de los flancos. Bujurquina pardus se distingue además de todas las otras especies de Bujurquina por la presencia de extensiones blancas, en lugar de negras, en la aleta dorsal y una banda lateral que se extiende hasta el borde dorsal del pedúnculo caudal, en lugar de alcanzar solamente la base de la aleta dorsal blanda. La especie puede distinguirse de muchos de sus congéneres por la presencia de puntos oscuros en la porción espinosa de la aleta dorsal y por tener aletas pectorales cortas (<32% de la longitude estándar). Se provee una clave para todas las especies descritas de Bujurquina.