Molecular and Morphological Analysis of Enteromorpha Intestinalis and E. Compressa (Chlorophyta) in the British Isles

The very common green seaweeds Enteromorpha intestinalis (L.) Nees and E. compressa (L.) Nees are important fouling organisms and have commonly been used as indicators of eutrophication, but their taxonomic status is problematic. The genus presents extreme difficulties because there is wide intraspecific variation in morphology, but morphological differences between species are small and difficult to detect. In this study, molecular data were used in parallel with morphological characters to resolve the taxonomic problems. Phylogenetic analysis of sequences of the internal transcribed spacers ITS1 and ITS2 and the 5.8S gene distinguished two groups of samples, which were identified by morphological characters as E. compressa (branched) and E. intestinalis (normally unbranched). There was a low level of sequence divergence within each group of samples, but divergence between groups was as great as that between either of the two species and the outgroup E. prolifera. Clades representing E. compressa and E. intestinalis were also found in analyses of an independent molecular data set, chloroplast DNA restriction fragment length polymorphisms (RFLPs). Enteromorpha intestinalis and E. compressa represent two distinct, genetically divergent species. Reinterpretation of published studies shows that these species are reproductively isolated. However, E. compressa and E. intestinalis are sometimes very difficult to distinguish from each other and could be regarded as cryptic species. The presence or absence of branching was the most useful character distinguishing these two species, but there was an element of ambiguity because low salinity or salinity shock can induce branching in E. intestinalis. If environmental factors such as salinity are taken into account, branching can be used to identify the great majority of thalli correctly. This study therefore provides a basis for identifying the two most important marine fouling macroalgae and for their use in environmental monitoring and experimentation. Typification of these two Linnaean species showed that current usage of the names accords with the lectotype and protologue of both species. Samples that resembled E. usneoides did not form a clade in any of the trees, and constraining the data to support the monophyly of this group incurred a penalty. Enteromorpha usneoides appears to be an ecotype of E. compressa .