We present a comprehensive revision and synthesis of the higher-level classification of the barnacles (Crustacea: Thecostraca) to the genus level and including both extant and fossils forms. We provide estimates of the number of species in each group. Our classification scheme has been updated based on insights from recent phylogenetic studies and attempts to adjust the higher-level classifications to represent evolutionary lineages better, while documenting the evolutionary diversity of the barnacles. Except where specifically noted, recognized taxa down to family are argued to be monophyletic from molecular analysis and/or morphological data. Our resulting classification divides the Thecostraca into the subclasses Facetotecta, Ascothoracida and Cirripedia. The whole class now contains 14 orders, 65 families and 367 genera. We estimate that barnacles consist of 2116 species. The taxonomy is accompanied by a discussion of major morphological events in barnacle evolution and justifications for the various rearrangements we propose.

Scanning and transmission electron microscopy (SEM and TEM) were used to study lattice organs in facetotectan y‐cyprids from the White Sea and from Norwegian and Bahamian waters. The larvae represent at least four and possibly five different species of Facetotecta. Y‐cyprids have five pairs of lattice organs in the head shield (carapace) organized into two anterior pairs and three posterior pairs. Both groups of lattice organs are arranged around a large central pore. The facetotectan lattice organs are elongate areas with a longitudinal keel, just as in the Ascothoracida and some Cirripedia Acrothoracica. The terminal pore of the organs is situated posteriorly in all five pairs. TEM confirms that the organs have the same general morphology as in the Cirripedia and Ascothoracida, namely, a cuticular chamber into which project ciliary segments from the chemosensory cells. Unlike Cirripedia the cuticular roof of the chamber lacks any pores. We conclude that five pairs of lattice organs represent an autapomorphy for the Thecostraca, which supports the monophyly of this taxon. In the ground pattern the terminal pore is posterior in all five pairs. The anterior position of the pore in lattice organ pair 2 is apomorphic for the Cirripedia, while within this taxon an anterior position also in pair 1 is apomorphic for a monophylum comprising the Thoracica and the Rhizocephala. Minute pores in the roof of the organs is another apomorphy of the Cirripedia, but its elaboration into pores visible with SEM may have been subject to some homoplasy. Since lattice organs are omnipresent in the settling instar of the Thecostraca they probably serve a critical role for the function of these cypris or cypris‐like larvae.

Symbiotic relations and range of host usage are prominent in coral reefs and crucial to the stability of such systems. In order to explain how symbiotic relations are established and evolve, we used sponge-associated barnacles to ask three questions. (1) Does larval settlement on sponge hosts require novel adaptations facilitating symbiosis? (2) How do larvae settle and start life on their hosts? (3) How has this remarkable symbiotic lifestyle involving many barnacle species evolved? We found that the larvae (cyprids) of sponge-associated barnacles show a remarkably high level of interspecific variation compared with other barnacles. We document that variation in larval attachment devices are specifically related to properties of the surface on which they attach and metamorphose. Mapping of the larval and sponge surface features onto a molecular-based phylogeny showed that sponge symbiosis evolved separately at least three times within barnacles, with the same adaptive features being found in all larvae irrespective of phylogenetic relatedness. Furthermore, the metamorphosis of two species proceeded very differently, with one species remaining superficially on the host and developing a set of white calcareous structures, the other embedding itself into the live host tissue almost immediately after settlement. We argue that such a high degree of evolutionary flexibility of barnacle larvae played an important role in the successful evolution of complex symbiotic relationships in both coral reefs and other marine systems.

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