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FRONTIERS IN PALAEONTOLOGY THE ORIGINS OF MOLLUSCS by JAKOB VINTHER Schools of Earth Sciences and Biological Sciences, University of Bristol, Life Sciences Building, 24 Tyndall Avenue, BS8 1TQ, Bristol, UK; e-mail: jakob.vinther@bristol.ac.uk Typescript received 3 September 2014; accepted in revised form 20 October 2014 Abstract: The interrelationships and evolutionary history of molluscs have seen great advances in the last decade. Recent phylogenetic studies have allowed alternative mor- phology-based evolutionary scenarios to be tested and, most significantly, shown that the aplacophorans are sister group to polyplacophorans (chitons), corroborating palaeontologi- cal and embryological evolutionary scenarios in which apla- cophorans are secondarily simplified from a chiton-like ancestor. Aplacophoran morphology therefore does not represent the plesiomorphic condition for molluscs as a whole. The mollusc crown group radiated in the Early Cam- brian, and rapidly thereafter, stem lineages to the major mol- luscan classes emerged: cephalopods, gastropods, bivalves (= pelecypods), monoplacophorans, rostroconchs (inferred stem scaphopods) and aculiferans. This attests to the fast, adaptive radiation of the crown group during the Cambrian explosion. Kimberella from the latest Ediacaran exhibits several molluscan traits, which justifies its position as a molluscan stem-group member, rather than as a more basal Lophotrochozoan. The interrelationships among the conchiferan molluscs are still a matter of contention and require further palaeontological and molecular phylogenetic scrutiny. Key words: Mollusca, Aculifera, Cambrian explosion, phy- logenomics, molecular palaeobiology, small shelly fossils. THE overall framework for understanding fundamental molluscan interrelationships has finally reached a stage in which several robust molecular phylogenetic analyses con- verge on similar topologies confirming certain hypotheses of early molluscan evolution. Relaxed molecular clocks provide means to test the completeness of the fossil record and to evaluate evolutionary scenarios. Recent fossil dis- coveries have been adding to the picture of how the mol- luscan body plan has evolved. Particularly, illustrative cases are highlighted in groups, such as cephalopods (Kr€oger et al. 2011), which showcase a major transition from exter- nally shelled orthocones to the internally shelled coleoids through the Phanerozoic. Molecular and palaeontological studies have also presented convincing evidence that mol- luscs evolved planktotrophy convergently in two major clades, the bivalves and gastropods (Peterson 2005; N€utzel et al. 2006) linked to changes in suspension feeding inten- sity. Another recent advance has been the identification of the vermiform and morphologically simple aplacophorans as a derived clade (Sutton et al. 2012; Vinther et al. 2012a), rather than a primitive grade of molluscs (Salvini- Plawen and Steiner 1996; Haszprunar 2000), which forces a reassessment of the ancestral morphology of the phylum. Molluscs are ecologically important invertebrates, which occupy a broad range of niches in marine, freshwater and terrestrial environments. They can be sessile, agile, infaunal, nektonic and planktonic; they can be car- nivores, scavengers, herbivores, grazers, photosymbiotic and suspension feeders. Several extinct clades were important and abundant marine denizens such as the Palaeozoic/Mesozoic ammonoids or the Cretaceous ru- dists (Skelton 1978). The most diverse and disparate clades are the bivalves, gastropods and cephalopods, and the least diverse and disparate are the scaphopods, monopla- cophorans, chitons and aplacophorans. MOLLUSCAN PHYLOGENY AND EVOLUTION While molluscs are morphologically distinct and well studied, early attempts to obtain a stable molecular phy- logeny of the phylum (Winnepenninckx et al. 1996; Pass- amaneck et al. 2004; Giribet et al. 2006) failed to support established morphological hypotheses, or even molluscan monophyly. However, the advent of phylogenomic studies (Dunn et al. 2008) and better informed choice of molecu- lar loci for analyses (Vinther et al. 2012a) enabled the recovery of molluscan monophyly. Furthermore, three independent studies in 2011 (Fig. 1) recovered the clade © The Palaeontological Association doi: 10.1111/pala.12140 19 [Palaeontology, Vol. 58, Part 1, 2015, pp. 19–34] Aculifera (Polyplacophora + Aplacophora) (Kocot et al. 2011; Smith et al. 2011; Vinther et al. 2012a) and there- fore could reject the Testaria hypothesis (Salvini-Plawen and Steiner 1996; Haszprunar 2000; Salvini-Plawen 2006), in which aplacophorans are primitive and chitons and conchiferan molluscs are united in a monophyletic shell plate-bearing clade. Conchiferan monophyly, which is justified from a mor- phological viewpoint (e.g. Nielsen 2012), has been recov- ered in some recent analyses (Kocot et al. 2011; Smith et al. 2011) using extensive genetic sampling (300–1100 gene fragments), but with little coverage of each loci. Using only seven gene loci, but gene fragments with high phylogenetic informativity and more than 80% gene sequence coverage, conchiferans were recovered as para- phyletic with cephalopods as sister group to the Aculifera. This topology was robust to a series of sensitivity tests, such as long-branch taxon exclusion, dissection of fast- evolving sites from the data set and heterogenous taxon exclusion, which would be expected to eliminate or mini- mize rooting or long-branch attraction artefacts (Vinther et al. 2012a). Here, the Conchifera are accepted as mono- phyletic due to the congruence with morphological hypotheses. Analyses generally recover a sister group relationship between bivalves and gastropods (Kocot et al. 2011; Vin- ther et al. 2012a) with scaphopods as a sister group to this clade. However, Smith et al. (2011) recovered scaph- opods as a sister group to gastropods, while in a corri- gendum (correcting faults in the pipeline that miscoded certain amino acids) instead recovered scaphopods as a sister group to bivalves (Smith et al. 2013). Cephalopods have been recovered as a sister to all other conchiferans (Kocot et al. 2011), together with the monoplacophorans (Smith et al. 2011) or, as mentioned before, as sister group to the Aculifera (Vinther et al. 2012a). The differ- ence between these topologies is mainly in terms of where the root has been inferred in the analysis (Fig. 1). A recent study using mitochondrial genomes also recov- ers Aculifera and Conchifera (Osca et al. 2014). A recent study (St€oger et al. 2013) revisiting the data set of Giribet et al. (2006) with more taxa, could not recover Aculifera or Conchifera, which can be ascribed to the little phylogenetic informativity of the gene selection and Ap lac op ho ra Po lyp lac op ho ra Mo no pla co ph or a Ce ph alo po da Sc ap ho po da Ga str op od a Biv alv ia Smith et al. 2013, corrigendum Smith et al. 2011 Kocot et al. 2011 Vinther et al. 2012 Aculifera Conchifera F IG . 1 . Phylogeny of molluscs, a review and consensus of three recent studies. Ambiguous taxon placements are indicated by stippled lines and a reference to the given study. 20 PALAEONTOLOGY , VOLUME 58 resulting relationships recovered mainly driven by long-branch attraction artefacts (i.e. clustering of the long-branched cephalopods and aplacophorans as well as scaphopods). THE ACULIFERA Aplacophorans are derived from footed, eight-plated ancestors Former studies, based on morphology, pointed to the sim- ple morphology ofaplacophorans as evidence for their primitive status in the molluscs, giving them importance for understanding early molluscan evolution (Salvini-Plawen and Steiner 1996; Haszprunar 2000; Salvini-Plawen 2006; Shigeno et al. 2007). In particular, some scenarios pre- sented aplacophorans as a paraphyletic basal grade, render- ing their morphology plesiomorphic for the molluscs. The chitons in such scenarios bridged the morphological gap between aplacophorans and the remaining conchiferan molluscs. However, the fossil record could not corroborate this scenario (Runnegar and Pojeta 1974), which otherwise shows the appearance of conchiferan groups in the earliest Cambrian, while chitons appear in the Late Cambrian and aplacophorans then lacked fossil representatives. Others have contemplated the possibility of aplacopho- rans being secondarily derived as a sister group to chitons (Scheltema 1993, 1996). Furthermore, evidence suggested that aplacophorans could have evolved from a chiton-like ancestor (Ivanov 1996). Specifically, embryological study of both neomeniomorphs (Solenogastres) and chae- todermomorphs (Caudofoveata) demonstrated a sevenfold dorsal iteration expressed as a series of naked regions in a post-larval individual (Scheltema and Ivanov 2002) or as transverse ridges containing calcium carbonate-secreting cells in the chaetodermomorph Chaetoderma nitidulum (Nielsen et al. 2007). Fossil discoveries made in the new millennium have been of tremendous importance. A fossil vermiform animal Acaenoplax hayae (Fig. 2D), from the Wenlock (Silurian) Herefordshire Lagerst€atte (Fig. 2D) with seven dorsal shell plates and a posterior ventral plate (Sutton et al. 2001, 2004), fitted the scenario in which aplacophorans evolved from a chiton-like ancestor through secondary reduction and loss of the foot and dorsal shell plates (Vinther et al. 2012a). Additional fossils were discovered and forms inter- preted as polyplacophorans re-examined. These were shown to have a rounded transverse profile with a mantle extending ventrally leaving only a shallow groove for a foot and no ventral mantle cavity (Sutton and Sigwart 2012). The fossils are from the Late Ordovician and pre- serve eight shell plates (Donovan et al. 2010, 2011). The fossils therefore provide a minimum timing for the diver- gence between aplacophorans and chitons with their derived aplacophoran traits. As chiton-like forms, which could subtend both chitons and aplacophorans, are known from deposits no older than the latest Cambrian (Runnegar et al. 1979; Vendrasco and Runnegar 2004; Pojeta et al. 2010; but see Vendrasco et al. 2009), these provide a putative maximum age for the divergence of Aculifera, the common ancestor to chitons and aplacoph- orans. Scheltema (1993) pondered that this clade would have diversified in the Early Ordovician. The molecular clock analyses of Vinther et al. (2012a) did indeed recover an Early Ordovician divergence of the Aculifera robustly, irrespective of clock and substitution models or the inclusion of cephalopods as a contentious sister group to Aculifera. Therefore, an emerging scenario holds that Aculifera is an Early Ordovician clade, which evolved from an ancestor, which has sclerites and eight dorsal shell plates. Subsequently, an additional seven-plated stem aplacophoran, Kulindroplax (Fig. 2E), has been unearthed in the Herefordshire Lagerst€atte (Sutton et al. 2012). This form resembles well-known fossils previously described as chitons, such as Chelodes (Cherns 1998a, b), but also pre- serves no visible foot, except for a diminutive ventral ridge. The loss of plesiomorphic characters in aplacophorans is chimaeric. While chaetodermomorphs have retained a pair of gills in their posterior mantle cavity, the neo- menimorphs have reduced gills to a series of papillae. Meanwhile, neomenimorphs retain a narrow ciliated foot, which is lost in chaetodermomorphs. The observations that Kulindroplax and Acaenoplax seem to completely lack a foot (Sutton et al. 2001; Sutton et al. 2004; Sutton et al. 2012) and have posterior structures in a mantle cavity sug- gest that they are stem chaetodermomorphs. Furthermore, their possession of shell plates implies convergent loss of these in the two modern aplacophoran groups. These results would indicate that aplacophorans diverged before the deposition of the Herefordshire Lagerst€atte (Wenlock, c. 426 Ma). The molecular clock analyses of Vinther et al. (2012a) estimate a divergence of the Aplacophora of around 450 Ma, which is congruent with this hypothesis. Both stem-group aplacophorans and stem-group chitons have members with highly conical shells, which harbour one or two internal lacunae. These forms were classified as an extinct group of chitons, Palaeoloricata, by Bergenhayn (1955). It is apparent that this group is paraphyletic, sub- tending aplacophorans, chitons and the aculiferan stem. Chiton evolution Chitons retained the plesiomorphic complement of eight shell plates in a row. A putative early stem chiton is the remarkable Ordovician Echinochiton (Pojeta et al. 2003; Pojeta and DuFoe 2008; Fig. 2C). Fossil evidence VINTHER : MOLLUSC ORIG INS 21 suggests that the crown group radiated in the Carbonif- erous (Sigwart 2009) about 350 Ma. Molecular clock analyses are congruent with the fossil record and esti- mate divergences at c. 340 Ma (Vinther et al. 2012a, b). A subgroup of chitons, the Chitonida, are represented by stem-group forms from the Early Permian around 270–275 Ma, and molecular clock analysis similarly estimates the Chitonida to have diverged by about 268 Ma (Vinther et al. 2012a, b). A major transition happened in the skeleton of chi- tons during the lead up to the crown group. Anterior projections of the medial shell layer form so-called sutural laminae, which serve for firmer embedding of the shell plates within the tissues. The evolution of this feature in the Devonian or Carboniferous coincides with an increased predation landscape and the Devonian Nekton Revolution (Klug et al. 2010). Chitons evolved from vermiform molluscs (still possessing a foot and A B C D HG JI M N OL K FE 22 PALAEONTOLOGY , VOLUME 58 elaborate mantle cavity) with conical, projecting shell plates into more dorsoventrally flattened forms with more closely appositioned shell plates, which are more firmly embedded with sutural laminae. This evolutionary trend continued in the Chitonida, which, by the Permian, had evolved lateral projections of the lower shell layer (articulamentum) in addition to the anterior sutural laminae for even firmer shell attachment. Preda- tion pressure increased in shallower waters (Vermeij 1987) and presumably drove the evolution of this clade. In modern oceans, there is a distinct correlation with Chitonida generally inhabiting shallower and more exposed environments compared to the more plesiomor- phic lepidopleurids, which now live in deeper and more cryptic environments. As modern chitons radiated to inhabit shallow energetic environments with less fossili- sation potential, they in effect evolved themselves out of the fossil record; Cenozoic and Mesozoic occurrences of chitons are heavily biased towards deeper water lepi- dopleurids (Sirenko 2006). While the skeleton is extremely conserved in crown chitons, one particular group of stem chitons took the exploration of a multiplated scleritome to an extreme. The multiplacophorans (Hoare and Mapes 1995) have seven transverse shell fields, with the anterior and poster- ior shell plates representing single valves and the five intermediate regions divided intothree distinct sections (Fig. 2F; Vendrasco et al. 2004). Multiplacophorans had appeared by the Silurian (Aurivillius 1892) and disap- peared in the Permian (Hanger et al. 2000). Some confu- sion has existed with respect to their affinity after the first recognition of their unequivocal affiliation with the Polyplacophora (Vendrasco et al. 2004). This stems from their amalgam of characters, which are shared with cer- tain crown members of chitons. While Vendrasco et al. (2004) assigned multiplacophorans to stem-group chitons, arguing that these particular features were likely to have evolved prior to the diversification of the crown, Puchal- ski et al. (2009) chose to assign them to crown group chi- tons based on the possession of the same features. However, this would push the origin of the crown group of chitons back from the Carboniferous to the Silurian, for which there is no fossil evidence despite relatively good sampling (Cherns 2004). The congruence between the Carboniferous appearance of fossil crown group chitons and molecular clock estimates rejects an older, hidden history of crown chitons and argues for multipla- cophorans as stem-group chitons that evolved certain crown group characteristics convergently (Vinther et al. 2012b). Stem aculiferans: Halkieria and the sachitids The ancestral aculiferan arguably looked like a chiton with eight shells in a row (Vinther et al. 2012a). Fossils F IG . 2 . Fossil early crown group molluscs. A, Halkieria evangelista Conway Morris and Peel, 1990 (MGUH 30887), a stem-group acu- liferan from the Early Cambrian Sirius Passet Lagerst€atte, Buen Formation, North Greenland (newly collected specimen from a 2011 expedition). B, Maikhanella multa (left, SMNH X2232) and Siphogonuchites (right, GPIN 106788) from the Early Cambrian Bayan-Gol Formation; note that this stem aculiferan has a shell plate consisting of merged sclerites. C, Echinochiton dufoei Pojeta, Eernisse et al., 2003 (USNM 517482), from the Ordovician Forreston Mb, Grand Detour Formation, cast. D, the total group aplacophoran Acaenoplax hayae Sutton, Briggs et al., 2001 (virtual reconstruction) from the Silurian (Wenlock) of Herefordshire, a putative stem chae- todermomorph (reprinted with permission from MacMillan Publishers: Nature). E, the total group aplacophoran Kulindroplax perissoko- mos Sutton, Briggs et al., 2012 (virtual reconstruction) from the Silurian (Wenlock) of Herefordshire (reprinted with permission from MacMillan Publishers Ltd: Nature). F, the multiplacophoran (stem polyplacophoran) Protobalanus spinicoronatus Vinther, Jell et al., 2012b from the Devonian (Early Givetian) Silica Formation of Lucas County, Ohio; the image is a re-articulated reconstruction based on a segmented lCT scan (Vinther et al. 2012b). G, Watsonella crosbyi, Grabau 1900, from the Early Cambrian of Siberia (SEM image) a univalved stem bivalve. H, Pojetaia runnegari Jell, 1980, from the Early Cambrian of South Australia (SEM image), a hinged and bival- ved stem bivalve. I, Mellopegma schizocheras Vendrasco, Kouchinsky et al., 2011b (CPC 40416) from the Middle Cambrian Gowers Formation of Australia. J, Stenotheca drepanoida SMNH Mo167627 from the Early Cambrian Ajax limestone. These forms (I, J) have been interpreted as early putative rostroconchs and thus stem scaphopods. K, the stem cephalopod Plectronoceras cambria (Walcott, 1905) from the Upper Cambrian of China (USNM 57819, holotype left and paratype right), one mile west of Tsi-Nan, Shantung (Yo- chelson, Flower et al., 1973). L, the chambered helcionellacean Tannuella elinorae Brock and Paterson, 2004 (SAMP 340146, holotype), from the Early Cambrian Mernmerna Formation of the Flinders Ranges, South Australia; this univalved mollusc possesses distinct ter- minal chambers (but lacks a siphon), which suggest that it might be a stem cephalopod. M–O, the stem gastropod Pelagiella atlantoides (Matthew, 1894) USNM 298724 from the Early Cambrian Hanford Brook, New Brunswick; the specimen preserves distinct dorsolateral pedal muscle scars (black arrows) surrounded by a grooved line, suggesting the extent of the mantle cavity (white arrows); muscle scars suggest that this form was an untorted stem gastropod, but exhibiting a c. 10 degree rotation (Runnegar 1981); M, view from the left; N, dorsal view; O, view from right. Images courtesy of Stefan Bengtson (B), Mark D. Sutton (D, E), Michael J. Vendrasco (G–J), John Paterson (L). Abbreviations: MGUH, Geological Museum of Copenhagen, Denmark; SMNH, Swedish Museum of Natural History; USNM, United States National Museum of Natural History; SAMP, South Australian Museum; CPC, Commonwealth Palaeontological Collections, Geoscience Australia, Canberra; GPIN, Nanjing Institute of Geology and Palaeontology, Academica Sinica. V INTHER : MOLLUSC ORIG INS 23 fitting this description can be traced back to the Late Cambrian (Runnegar et al. 1979; Vendrasco and Runne- gar 2004; Pojeta et al. 2010). However, the aculiferan fos- sil record should extend back to at least the Early Cambrian when their sister group, the conchiferans, radi- ated. A plethora of lepidote molluscs (sachitids) are known from the Cambrian and include forms such as the halkieriids (Fig. 2A) and siphogonuchitids (Fig. 2B) (Bengtson and Missarzhevsky 1981). Initially, the exact nature of these forms was difficult to decipher due to the lack of articulated specimens in the small shelly deposits that they are typically extracted from (Bengtson and Con- way Morris 1984). However, the discovery of articulated halkieriids in the Early Cambrian Sirius Passet (Conway Morris and Peel 1990, 1995) demonstrating a body plan with several morphologically distinct sclerites in zones and a large shell growing by marginal accretion at the front and back put an end to this palaeontological mys- tery. Other forms have been discovered since then with a different number of shell plates (Conway Morris and Ca- ron 2007). Some forms have shell plates with sclerites embedded into the shell plates (Bengtson 1992; Fig. 2B). Several characteristics in these forms are shared with acu- liferans, such as their sclerite growth and replacement, and the presence of a complex internal canal system in either the sclerites or the shell plates, which are derived from mantle papillae (Vinther and Nielsen 2005; Vinther 2009). A number of shell plates described in isolation likely belong to the sachitids and indicate a highly vari- able scleritome (Conway Morris and Caron 2007; Pater- son et al. 2009; Vendrasco et al. 2009). The sachitids extend the aculiferan fossil record back to the earliest Cambrian (Terreneuvian) (Maloof et al. 2010) and thus fill the gap between the earliest conchiferan fossils and the appearance of aculiferans. Two other fossils worth mentioning are the wiwaxiids and Odontogriphus. Wiwaxia is now known as body fos- sils from a range of localities other than the Burgess shale (Conway Morris 1985; Smith 2014) including China (Zhao et al. 1994; Sun et al. 2014; Yang et al. 2014), Sibe- ria (Ivantsov et al. 2005) and Europe (Fatka et al. 2011). Wiwaxiid sclerites are also well known as ‘small organic bits’ (Butterfield 1990, 1994; Butterfield and Harvey 2012). Wiwaxia was early on recognized as a likely rela- tive of Halkieria (Bengtson and Conway Morris 1984). The radula in Wiwaxia was originally thought to consist of two bars (Conway Morris 1985), similar to a disti- chous aplacophoran radula (Scheltema et al. 2003). How- ever, recent studies using back-scattered electron imaging have provided a more detailed understanding of the radu- lar morphology, showing that both Wiwaxia and the unarmoured Odontogriphus had isolated denticles with a median tooth flankedby shoehorn-shaped lateral denti- cles (Smith 2012). CONCHIFERAN EVOLUTION Conchiferans have generally been assumed to be mono- phyletic from a morphological point of view. However, only recently have molecular phylogenies recovered con- chiferan monophyly (Kocot et al. 2011; Smith et al. 2011), although the relationships among its members remain in flux. Most conchiferan groups recognized as classes in the Linnean system can be traced back to the Cambrian small shelly fossil assemblages (Runnegar and Pojeta 1974). These in turn can be traced back to unival- ved forms, resembling monoplacophoran limpets, which arguably is the plesiomorphic condition for the group. These fossils are usually found as secondarily phospha- tized small shelly fossils (SSFs; Kouchinsky et al. 2012). This taphonomic window records a polyphyletic assem- blage of skeletal organisms ranging from submillimetric to a couple of millimetres in size. Based on these fossils, it has been speculated that early molluscs were generally microscopic (Haszprunar 1992), with implications for the evolution of the molluscan body plan. However, discover- ies of large, centimetre-sized halkieriids (Conway Morris and Peel 1990) and helcionellids (Mart�ı Mus et al. 2008) as crack-out specimens in shales demonstrate that early molluscs could be large and that the SSF taphonomic window discriminates towards smaller fossils (Creveling et al. 2014). There is still evidence that many early mol- luscs were relatively minute (B. N. Runnegar pers. comm. and pers. obs.) compared to Late Cambrian and Ordovi- cian forms. The intricate replacement of internal moulds (steinkerns) and partial replacements of the shell material make it possible to characterize their mineralogical ultra- structure, which provides additional information of potential phylogenetic relevance (Runnegar 1985; Kou- chinsky 1999, 2000; Feng and Sun 2003; Vendrasco et al. 2010, 2011a). An interesting study was performed using phosphatized steinkerns to characterize the larval shell (protoconch) through the Cambrian and Ordovician (N€utzel et al. 2006; N€utzel 2014). As the size of the pro- toconch reflects the size of the embryo, which is tightly correlated with either a prolonged planktotrophic devel- opment (small embryonic shell) or a short-lived, yolk feeding (lecithotrophic) stage (large embryonic shell), N€utzel et al. (2006) demonstrated that early molluscan groups were all yolk feeding during early ontogeny and only evolved planktotrophy in parallel at the onset of the Ordovician. This contrasts with ideas in which the ances- tor of bilaterians and animals should have primitively been planktotrophic (J€agersten 1972; Peterson et al. 1997; Nielsen 2013), which seems firmly refuted now (Haszpr- unar et al. 1995; Rouse 2000; Peterson 2005; Degnan and Degnan 2006). The parallel onset of planktotrophy in the Early Ordovician correlates with the Great Ordovician Biodiversification Event (GOBE; Harper 2006), which saw 24 PALAEONTOLOGY , VOLUME 58 the radiation and wide geographical dispersal of several molluscan classes. The onset of planktotrophy and the GOBE is correlated with a great radiation of sessile sus- pension feeders, which would have predated on larvae, and the evolution of planktotrophy thus could be viewed as driven by predator–prey feedback. Monoplacophoran roots Monoplacophorans were originally thought to be an extinct group of molluscs, but are now frequently recorded in deep-sea samples from across the globe (Lemche 1957; Lemche and Wingstrand 1959; War�en and Gofas 1996; Schr€odl et al. 2006). Monoplacophorans are unique among conchiferans in having a serialized body plan (Lemche 1957; Wingstrand 1985) with particular similarities to chitons in their muscle anatomy and serial- ization of the gills, while also possessing serialized kid- neys. Unequivocal monoplacophorans, identified by serial muscle scars in the shell, are known from the Late Cambrian (Stinchcomb 1986) and are present in the Bur- gess shale mollusc Scenella (Rasetti 1954; Runnegar and Pojeta 1974). Several monoplacophoran-like forms are known from older deposits. However, as all conchiferan groups can be traced back to univalved ancestors resem- bling monoplacophorans, many of these can be referred to stem lineages of specific classes or merely to the con- chiferan total group. There has been much discussion concerning the interpretation of these forms, as the fossils provide few characters, which provide for inferences about muscle scars, position of mantle cavity, direction of coiling relative to the body axis and whether a fossil form is torted or not (Runnegar 1981; Peel 1991; Parkhaev 2008). Bivalve evolution Conchiferan molluscs with a bivalved scleritome con- nected by a dorsal hinge are represented in the Early Cambrian by forms such as Fordilla troyensis (Pojeta et al. 1973; Pojeta and Runnegar 1974) and the slightly older Pojetaia runnegari (Fig. 2H; Jell 1980; Runnegar and Bentley 1983). These forms were likely to be shallow bur- rowers (Runnegar and Bentley 1983), but the crown group radiated in the Early Ordovician and quickly evolved practically all the main modes of life exhibited by modern bivalves (Pojeta et al. 1978; Cope 2000). While molecular studies still are in some disagreement (Plazzi et al. 2011; Sharma et al. 2012), protobranchs are gener- ally recovered as a sister to all other bivalves (autolamelli- branchiata/autobranchiata), which is corroborated by the fossil record. Bivalves were generally less diverse and abundant compared to the ecologically similar brachio- pods in the Palaeozoic, with their relative abundances through the Phanerozoic being roughly inversely corre- lated (Clapham et al. 2006). Comparing bivalves and bra- chiopods from the perspective of their metabolic activity demonstrates bivalve dominance throughout most of the Phanerozoic (Payne et al. 2014). Bivalves can be traced back to laterally compressed univalved conchiferan forms, such as Watsonella (Fig. 2G; Pojeta and Runnegar 1974), and similarly, there are mor- phological and shell ultrastructural evidence to link Watsonella with the potentially more plesiomorphic An- abarella (Kouchinsky 1999). Further studies of shell microstructure in Fordilla, Pojetaia, Anabarella and Watsonella demonstrate that these early bivalves did not possess nacre in their shells, but rather foliated aragonite (Vendrasco et al. 2011a), nacreous mother of pearl appears to have evolved multiple times convergently in molluscs in the Ordovician period (Vendrasco et al. 2011a, 2013) as a response to the increased predation pressure of the GOBE. Scaphopods and the rostroconchs Scaphopods, or tusk shells, are a small group of conchif- erans with limited diversity and disparity. They possess a distinct, elongate conical shell and live infaunally as selec- tive detritus feeders. The fossil record of unequivocal scaphopods extends back to the Devonian or Carbonifer- ous (Peel 2004, 2006). Scaphopods probably evolved from the extinct clade of rostroconchs, consisting of two groups, the ribeirioids and the conocardioids. It was orig- inally proposed that scaphopods evolved from Ordovician ribeirioids (Runnegar and Pojeta 1974; Pojeta and Runne- gar 1976, 1979), but Peel (2004, 2006) demonstrated by close examination of the protoconch morphology and growth direction (exogastric vs. endogastric) of these forms that a derivation from younger conocardioid ro- stroconchs is more justifiable. Rostroconchs can be traced back to the Late Cambrian and potentially even back to the Early Cambrian with forms such as Mellopegma(Fig. 2L; Pojeta and Runnegar 1976; Runnegar and Jell 1976; Vendrasco et al. 2011b). However, much confusion exists with respect to the affinity of several laterally com- pressed Early Cambrian conchiferans and their potential affinity to either bivalves or rostroconchs (and thus scaphopods; see discussion in Vendrasco 2012). Some of this confusion might be resolved if scaphopods and bival- ves were to prove to be sister groups (Diasoma) as seen in the corrected analyses by Smith et al. (2013) as well as in some analyses of Osca et al. (2014), with all these later- ally compressed forms representing total group diaso- mans. V INTHER : MOLLUSC ORIG INS 25 Cephalopod evolution Cephalopods are nektonic predators, which are important constituents of the pelagic food chain. Modern cephalo- pod diversity is dominated by the coleoids (i.e. squids, cuttlefish and octopods), which have internalized or even secondarily lost their shell. The plesiomorphic, chambered external shell is retained in nautilids. The oldest unequiv- ocal cephalopod with a chambered phragmocone con- nected by a siphon is the Late Cambrian Plectronoceras (Yochelson et al. 1973; Chen and Teichert 1983; Fig. 2K). Several older, chambered shells have been proposed as candidates to subtend the cephalopod stem, such as Tan- nuella (Fig. 2L; reviewed in Kr€oger et al. 2011), but firm evidence remains elusive. A long-standing scenario holds that the two main extant lineages of cephalopods, the coleoids and nautilids, can be traced back to different Early Palaeozoic lineages, the orthocerids and oncocerids, respectively, which in turn diversified from each other in the Early Ordovician. However, molecular clock studies (Kr€oger et al. 2011; Warnke et al. 2011) consistently recover a divergence of the cephalopod crown at the Silurian–Devonian boundary. This is in fact remarkably congruent with the cephalopod fossil record, which sees the appearance of stem coleoids (bactritids and ammo- noids) and bona fide nautilids in the Devonian (Dzik and Korn 1992; Kr€oger and Mapes 2007) as well as the first fossil cephalopod beaks, a crown cephalopod apomorphy (Kr€oger et al. 2011). A general trend in cephalopod evo- lution is towards the loss of the shell and more efficient swimming by mantle pumping rather than shell pumping (Wells and O’Dor 1991). The timing of these transitions is closely matched by the evolutionary innovations and diversification of osteichthyan fishes (Packard 1972), which can be explained by evolutionary arms races among the two groups of pelagic predators. Note that the recent description of the Cambrian Nectocaris as a cephalopod (Smith and Caron 2010; Smith 2013) has been firmly refuted (Kr€oger et al. 2011; Mazurek and Zaton 2011; Runnegar 2011). Gastropod evolution Gastropods are characterized by the 180 degree torsion of the visceral mass and shell field relative to the head–foot. This means that the anus is above the head, whose little practicality has not gone unnoticed (Peel 1987, p. 306). The earliest forms, ascribable to the gastropod stem, are the Early to Mid-Cambrian pelagiellids and aldanellids (but see Dzik and Mazurek 2013). Muscle scars and pal- lial line imprints in a large specimen of Pelagiella (Fig. 2M) demonstrate an asymmetric arrangement of dorsolaterally inserted muscles, which suggests about 10 degrees of torsion (Runnegar 1981). Crown gastropods seem to have radiated in the latest Cambrian. Current molecular studies suggest that the patellogastropods and the vetigastropods form a basal clade (Smith et al. 2011; Vinther et al. 2012a; Smith et al. 2013), which is sister to the remaining gastropods. Within this latter clade, the following topology is seen (Neritimorpha (Caenogastro- poda + Heterobranchia)) (Zapata et al. 2014). EDIACARAN ROOTS AND KIMBERELLA Molluscs all have biomineralized skeletons, but some argue against homologizing the shell plates of conchifer- ans and aculiferans (Haas 1981; Scheltema 1988). It has been speculated that the shell plates of aculiferans could have evolved from amalgamation of sclerites as is evidenced in Maikhanella, which consists of sclerites of the genus Siphogonuchites (Bengtson 1992). This observa- tion also highlights the likely origin of the sensory aes- thete canal system in chitons. These structures are derived from papillae, which also secrete sclerites in the mantle (Bengtson 1992; Vinther 2009). Siphogonuchites and the other Cambrian sachitids exhibit hollow sclerites, often with a branching canal system within, and it has been argued that this canal system is similar in dimensions to modern chiton aesthetes (Vinther 2009). A case can still be made, however, for homology of conchiferan and aculiferan shell plates: both exhibit crossed lamellar ultrastructural fabric, which is a unique molluscan characteristic, suggesting that the latest common ancestor to crown molluscs had a biomineralized shell. While the molluscan crown group had diversified by the Terreneuvian (Stage 1) in the earliest Cambrian, there is little evidence to suggest that crown group molluscs existed in the Precambrian, as no skeletal fossils have been found even in the Nama group, which yields the first abundant material of presumed skeletized metazoans (Grotzinger et al. 2000; Wood et al. 2002). This provides a tight bracket for molluscan diversification of less than 15 million years (but see Runnegar 1987, for an alterna- tive explanation). A noteworthy fossil from the Ediacaran is Kimberella (Fig. 3A), which has been interpreted to be molluscan (Fedonkin and Waggoner 1997). While this was initially met with scepticism (Nielsen 2001), further studies have strengthened this hypothesis (Fedonkin et al. 2007; Ivant- sov 2009, 2010; Gehling et al. 2014). Kimberella is a cru- cial Ediacaran organism in that it exhibits a series of taphomorphs, which reveal a complex organism with a differentiated ventral and dorsal surface and internal anat- omy; often, these structures are superimposed. From these fossils, it is clear that Kimberella exhibits an anterior and posterior polarity and bilateral symmetry. The ventral 26 PALAEONTOLOGY , VOLUME 58 surface shows that there is a large ventral surface, which is smooth or transversely wrinkled, surrounded by con- centric body units, which can be interpreted as a foot and a mantle separated by a groove. These impressions resem- ble a molluscan body plan in having a distinct creeping sole (foot) and a surrounding mantle separated by a D BA C E F F IG . 3 . Kimberella and Kimberichnus. A, Kimberella quadrata from Erginskaya Formation, Zimnie Gory, Local Cluster Z1, ‘Kimberella lenses’ (PIN 3993/5136); specimen preserves the presumed ventral surface, compared to experiments in Seilacher (1999, fig. 7). B, Kim- berichnus teruzzi Ivantsov, 2013, holotype PIN 3993/5619; specimen preserves radial, paired scrape marks; notice the relief beyond the paired markings, demonstrating that the feeding trace extends beyond the activity of the paired tooth marks; rounded pellets on bed- ding planes have been interpreted as sand pellets from their feeding activity (Gehling et al. 2014). C, Kimberella quadrata (bf) associ- ated with Kimberichnus (tf) at the presumed anterior end (PIN 3493/5137), also figured in Fedonkin (2003). D, Kimberella quadrata, associated with Kimberichnus at the presumed anterior end (PIN 4853/333). E, Kimberella quadrata preserving internal anatomy; notice the preserved pharynx with paired pouches, arrowed (PIN 4853/326). F, Kimberella quadrata associated with Kimberichnus at the pre- sumed anterior end (PIN 4853/379). Abbreviations: PIN, PalaeontologicalInstitute, Moscow; bf, body fossil; tf, trace fossil. V INTHER : MOLLUSC ORIG INS 27 mantle cavity (Seilacher 1999; Seilacher et al. 2003; Fig. 3A). The dorsal surface exhibits a cuticular shield with tubercular nodes, which are particularly resistant to decay. Specimens are found in great abundance on sur- faces that also preserve distinct scraping traces named Kimberichnus Ivantsov, 2013 (Fig. 3B), which is also well known from Australia (Gehling et al. 2014). These traces suggest a mode of feeding on the microbial mats involv- ing two larger teeth (forming paired grooves), and a ser- ies of smaller denticles (as evidenced by the presence of relief outside the tooth marks indicating feeding over a larger area by a more elaborate feeding apparatus than 550 500 450 400 350 300 Cambrian Ordovician Silurian Devonian Carboniferous 250 PermianEdiacaran Ribeirioida Conocardioida Scaphopoda Cephalopoda Bivalvia Rostroconcha Gastropoda Conchifera Aculifera Aplacophora Polyplacophora Lepidopleurida Chitonida Chaetodermomorpha Neomenimorpha Monoplacophora Mollusca F IG . 4 . A time tree of molluscs summarizing discoveries from recent state-of-the-art molecular phylogenetic studies, clocks and the fossil record. 28 PALAEONTOLOGY , VOLUME 58 the visible paired teeth marks; Fig. 3B). Many specimens (Fedonkin et al. 2007; Ivantsov 2009, 2010, 2013; unpub. obs.) show the body fossil Kimberella in direct association with the Kimberichnus scrape marks (Fig. 3C, D, F). Sur- ficial locomotion trails are almost never observed in this mat ground environment, and observed traces are all feeding traces from organisms that are actively removing the mat (Sperling and Vinther 2010; Ivantsov 2013). However, some creeping trails are known connecting a feeding trace with a nearby body fossil (Fedonkin et al. 2007; Ivantsov 2009, 2010). Other features of Kimberella are molluscan, such as the presence of a structure resem- bling a digestive tract and a pharynx with a set of paired pouches that resembles oesophageal pouches (Fig. 3E; Vinther et al. 2012a); similar pouches are unknown in any lophotrochozoan other than molluscs. Unfortunately, Kimberella does not offer any characters for establishing character polarity in molluscan evolution as it does not possess a biomineralized skeleton and thus cannot establish whether the ancestor had multiple scle- rites and shell plates, or a single shell field, as in conchif- erans (Vinther et al. 2012a). Chitinous elements secreted by microvilli are present in a number of lophotrochozoans (annelids, brachiopods and some molluscs). Wiwaxia is a mollusc (Conway Mor- ris 1985), which possesses apparently unmineralized, likely chitinous, sclerites with a microvillar microstructure (But- terfield 1990) as well as a radula (Smith 2012). The lack of a mineralized skeleton and the presence of chitinous elements suggest that this form is a stem-group mollusc, which would polarize the aculiferan trait of having scle- rites in multiple distinct longitudinal/concentric zones, a plesiomorphy of the phylum. However, nonmineralic sclerites are observed in both chitons and aplacophorans and a microvillar microstructure is also observed in the organic pellicle covering aculiferan sclerites (Fischer et al. 1980, fig. 9). Furthermore, Wiwaxia is only known from Burgess Shale-type Lagerst€atten (Sun et al. 2014; Yang et al. 2014) and from small organic bits (Butterfield and Harvey 2012), which generally do not preserve any miner- alized components, complicating detailed inference of its skeletal nature as biomineralized or not. Odontogriphus is another ‘naked’ mollusc, whose morphology might per- tain to questions about the ancestral mollusc (Caron et al. 2006) or the evolution along the molluscan stem. In par- ticular, the radula is of interest and is comparable to the radula preserved in Wiwaxia (Smith 2012). CONCLUSIONS A general consensus has finally been reached concerning the fundamental relationships among molluscs. Aplacoph- orans were thought to be important extant members for understanding early molluscan evolution, but are highly derived and secondarily reduced (Vinther et al. 2012a). The fossil record in conjunction with molecular clock analyses shows the molluscs radiated rapidly during the Cambrian explosion (Fig. 4), while bivalves and gastro- pod crown groups radiated in the Ordovician Biodiversi- fication Event. Aplacophorans, chitons, scaphopods and cephalopod crown groups radiated later in the Palaeozoic. Some stem-group lineages were still remarkably disparate and diverse in the Early Palaeozoic, for example, cephalo- pods (Kr€oger et al. 2011). Monoplacophorans appear to have experienced a Late Cretaceous bottleneck (Kano et al. 2012), which can be ascribed to their colonization of the deep sea as a refugium, and the series of Cretaceous ocean anoxic events (Jenkyns 1980), which would have limited their distribution episodically. Several aspects of molluscan evolution still need scru- tiny: in particular, the sequence of character evolution, the nature of the transitions between major molluscan classes, especially within the Conchifera, and the evolution within them would benefit from more palaeon- tological discoveries and novel integrative studies in the light of the new phylogeny of molluscs. Acknowledgements. I am thankful to the editor Andrew Smith for inviting me to write this review and for his patient editorial sug- gestions. The reviewers provided invaluable feedback and made helpful corrections and suggestions, but any omission or mistake would be the responsibility of the author. I appreciate the hospi- tality of Andrei Yu Ivantsov and John Pojeta and for letting me study their collections. I thank Mike Vendrasco, John Pojeta, Bruce Runnegar, Stefan Bengtson, Christiane Todt, Chris Schand- er, Amelie Scheltema, Derek E. G. Briggs, Andreas Wanninger, Claus Nielsen, Kevin J. 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