Molluscan Taxonomic Classes Mollusca: General Characteristics Bivalvia 7,650 sp Polyplacophora Cephalopoda Scaphopoda Other 5 Classes ~1100 Gastropoda and Aplacophora Monoplacophora Gastropoda 40,000 sp Bivalvia Very different, but with some important shared characters Cretaceous Triassic -Shell - Mantle Devonian - Ctenidium - Muscular foot Polyplacophora Rostriconchians Gastropoda Monoplacophora Bivalvia Cephalopoda Scaphopoda Fossil History of the Mollusca - Radular organ 1 Class Gastropoda SubClasses Prosobranchia Opisthobranchia Pulmonata Figure 1. Consensus phylogeny of mollusc classes as supported by the phylogenomic anlayses of Kocot et al.[1] and Smith et al.[2].The Mollusca are divided into two major clades: i) the Aculifera, which possess spicules and show repetition of structures along their antero-posterior axis; and ii) the Conchifera, which (at least primitively) possess a shell. Within Conchifera, there remains some uncertainty over the most likely position of the Scaphopoda (tusk shells). Monoplacophora was placed by the analysis of Smith et al.[2] as sister group of the Cephalopoda. The Cambrian fossil genus Latouchella is indicated at the base of the Conchifera. The Cambrian fossil genera Wiwaxia and Halkieria (reconstruction) possess dorsal sclerites and a radula-like structure and are supported as stem group molluscs by some palaeontologists. Modern Prosobranch Gastropod Evolution of Gastropoda: • Possibly from now extinct monoplacophoran (A class of molluscs represented today by one deep water genus) • Involves coiling and re-alignment of the shell • Also, in an independent evolutionary event, a twisting of the viscera relative to the A-P axis of the body as defined by the head and the foot: torsion 2 Coiling is a way to maximize volume while minimizing height A coiled (planospiral) shell is top-heavy so in most larger species the coil is angled relative to the A-P axis. Ancestral Pre-Torsional Torsion occurs during the development of the veliger Torsion Gastropod Modern Post-torsional Front view Early trochophore Side view Later trochophore Veliger Before torsion after torsion 3 Are there adaptive advantages to Torsion? Three hypotheses proposed 1. To bring adult coiled shell into better balance along a-p axis Mantle cavity is Beneath this area Are there adaptive advantages to Torsion? Two hypotheses have been proposed 1. To bring adult coiled shell into better balance along a-p axis 2. Allows larva to pull its head and velum into the mantle cavity first 3. New perspectives (Louise Page Univ. of Victoria) “asymmetry hypothesis” anterior re-arrangement/enlargement of the lateral mantle cavity in a monoplacophoran Design of experiments to test adaptive value of torsion (Pennington and Chia 1985) --Fed a choice of pre-torted and newly torted veliger larvae of the abalone Haliotis to seven potential predators: crab larvae, copepod, fish, ctenophore (2), hydromedusa (2) Results of Pennington and Chia 1985 Experiments Data are the mean number of larvae eaten. Torted larvae in white. Crab larvae copepod fish 10 20 30 hydromedusa spp A B ctenophore spp A B -- 5 replicates, 25 larvae of each type, allowed experiments to run for 15 hr Conclusions ??? 4 Problems Caused by Torsion Other Molluscan Groups: Opisthobranchia A problem of plumbing Evolutionary Tendencies: Archeogastropod Meso- and Neogastropods • Shell internal, reduced or lost • Limited torsion in development C. Bilateral D. Bipectinate E. monopectinate • Loss or reduction of mantle cavity • Loss of operculum Solutions to sanitary problems that resulted after torsion http://week.divebums.com/2008/Mar10-2008/index.html • Ctenidia replaced by gills F. Opisthobranchs Other Molluscan Sub Classes Opisthobranchia G. Pulmonates Varying degrees Of detorsion In Bubble Shell Opisthobranchs 5 What features seem to be adaptations to terrestrial life? Other Molluscan Sub Classes: Pulmonata land and f.w. snails and slugs very few marine Adaptations to life In terrestrial Environments? Simplified phylogeny of Gastropoda and Torsion 18s r- DNA phylogenies support the monophyly of Opisthobranchia and Pulmonata detorsion Shell loss Land slugs Pulmonata lungs Is Prosobranchia a valid taxon? Land snails Pulmonata coiled shell torsion Opisthobranchia Shell loss Gills detorsion Prosobranchia Monoplacophora ancestor In 28s and 18s rDNA phylogenies “pulmonates” are nested within the Opisthobranchia!! Bye- Bye Opisthobranchia!! Schroedl et al. 2010 Thalassas vol. 27:101-112 6