title: Phenotypic plasticity and morphological integration in a marine modular invertebrate.
abstract: BACKGROUND
  Colonial invertebrates such as corals exhibit nested levels of modularity, imposing a challenge to the depiction of their morphological evolution. Comparisons among diverse Caribbean gorgonian corals suggest decoupling of evolution at the polyp vs. branch/internode levels. Thus, evolutionary change in polyp form or size (the colonial module sensu stricto) does not imply a change in colony form (constructed of modular branches and other emergent features). This study examined the patterns of morphological integration at the intraspecific level. Pseudopterogorgia bipinnata (Verrill) (Octocorallia: Gorgoniidae) is a Caribbean shallow water gorgonian that can colonize most reef habitats (shallow/exposed vs. deep/protected; 1-45 m) and shows great morphological variation.
  
  
  RESULTS
  To characterize the genotype/environment relationship and phenotypic plasticity in P. bipinnata, two microsatellite loci, mitochondrial (MSH1) and nuclear (ITS) DNA sequences, and (ITS2) DGGE banding patterns were initially compared among the populations present in the coral reefs of Belize (Carrie Bow Cay), Panama (Bocas del Toro), Colombia (Cartagena) and the Bahamas (San Salvador). Despite the large and discrete differentiation of morphotypes, there was no concordant genetic variation (DGGE banding patterns) in the ITS2 genotypes from Belize, Panama and Colombia. ITS1-5.8S-ITS2 phylogenetic analysis afforded evidence for considering the species P. kallos (Bielschowsky) as the shallow-most morphotype of P. bipinnata from exposed environments. The population from Carrie Bow Cay, Belize (1-45 m) was examined to determine the phenotypic integration of modular features such as branch thickness, polyp aperture, inter-polyp distance, internode length and branch length. Third-order partial correlation coefficients suggested significant integration between polypar and colonial traits. Some features did not change at all despite 10-fold differences in other integrated features. More importantly, some colonial features showed dependence on modular features.
  
  
  CONCLUSIONS
  Consequently, module integration in gorgonian corals can be shifted, switched or canalized along lineages. Modular marine organisms such as corals are variations on a single theme: their modules can couple or decouple, allowing them to adapt to all marine benthic environments.
authors.affiliation: Laboratorio de Biología Molecular Marina-BIOMMAR, Departamento de Ciencias Biológicas-Facultad de Ciencias, Universidad de los Andes, Bogotá, Colombia. juansanc@uniandes.edu.co
authors.forename: Juan A
authors.initials: JA
authors.lastname: Sánchez
authors.forename: Catalina
authors.initials: C
authors.lastname: Aguilar
authors.forename: Daniel
authors.initials: D
authors.lastname: Dorado
authors.forename: Nelson
authors.initials: N
authors.lastname: Manrique
mesh_terms: D000818:Animals; D037421:Anthozoa / Q000033:anatomy & histology* / Q000235:genetics / Q000502:physiology*; D005075:Biological Evolution; D017753:Ecosystem; D005819:Genetic Markers / Q000235:genetics; D005838:Genotype; D008386:Marine Biology; D009792:Oceans and Seas; D010641:Phenotype
chemical_list: D005819:Genetic Markers