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2. Phylogenetic hypothesis on phylogeny of the Paleozoic gastropods inferred from teleoconch morphology based on analysis of Wagner (1999b) (modifi ed from Figure 37, Wagner 1999b).
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This chapter describes the key feature of Paleozoic Gastropoda. It reviews ideas about the early evolution of gastropods and discusses inferences drawn from teleoconch and protoconch morphologies as well as possible relationships of extinct Paleozoic gastropod groups with living gastropods. It summarizes macroevolutionary trends of their teleoconch...
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... This character also brings the entire group closer to caenogastropods. On the other hand, Frýda and Manda (1997) and Frýda et al. (2008) documented well-preserved Murchisonia-like gastropods that have a protoconch typical of Vetigastropoda (i.e., consisting of about one whorl) suggesting placement in Vetigastropoda. The presence of nacre in the Ordovician "Murchisonia" (Mutvei 1983) further corroborates such a placement at least of early members assigned to this group. ...
High-spired Murchisonia-like slit-band gastropods are an important component of late Paleozoic gastropod faunas. Twenty-seven genera of such gastropods have been reported from the Permian, most of which representing the caenogastropod family Goniasmatidae. Only four genera, Trypanocochlea, Wannerispira, Laschmaspira, and Altadema
crossed the Permian/Triassic boundary. Based on the study of newly collected specimens and material from natural history collections, we studied the surviving genera as well as the Triassic recovery of this group. Two new species (Laschmaspira lirata sp. nov. and Altadema hausmannae sp. nov.) and one new subfamily (Cheilotomoninae) are introduced. Murchisonia-like caenogastropods, chiefly Goniasmatidae, were diverse and abundant until the Permian, barely survived the end-Permian extinction, regained a certain generic diversity within the Triassic with the evolution of several new genera but failed by far to regain their Permian generic diversity. This once successful and diverse group shares a similar fate (surviving the end-Permian extinction, a reduced Triassic diversity and extinction during Late Triassic crises) as conodonts, orthoceratids, conulariids, and others. This diversity pattern does not qualify for the “Dead Clade Walking”
phenomenon, i.e., the extinction shortly after a major mass extinction event (survival without recovery) because they have survived for ca. 30 Ma (at least until the Norian) and even produced a number of new genera. The exact time of their extinction is unknown but there are no safe Rhaetian occurrences. Their extinction is part of a long-term selective trend against the character “shell-slit”.
... Toward the mid-Darriwilian (late Middle Ordovician), Macluritoidea generic diversity dropped rapidly and the superfamily became extinct toward the latest Ordovician (Frýda and Rohr, 2004). According to Wagner (2002) and Frýda et al. (2008), species within this superfamily typically are diagnosed by a base formed from a posteriorly projected inner margin, nearly planispiral to visually dextral coiling, open-coiled and heterostrophic early shell, a shallow V-shaped sinus, and a sharp thin peripheral band located on top of the aperture. Another diagnostic character is the change in shape of the aperture during ontogeny. ...
... Wagner (2002) also pointed out that a horn-shaped calcified operculum with a handle-like knob diagnoses Teiichispira and more-derived macluritid species. The Family Macluritidae is mostly represented by genera restricted to the Ordovician (Frýda et al., 2008) with a cosmopolitan paleobiogeographical distribution. In the Argentinean Precordillera, the genera Teiichispira and Maclurites are known probably as a result of the extensive record and good preservational conditions in which their opercula are found. ...
Two species of Teiichispira Yochelson and Jones reported from the Early/Middle Ordovician marine deposits of the San Juan Formation in the Argentine Precordillera are described. The new species Teiichispira teresae n. sp. is a component of the Early/Middle Ordovician marine gastropod assemblage in the studied region; Teiichispira argentina (Kayser), previously known from the San Juan Formation, is described with a complete teleoconch and associated operculum. The opercula of both Teiichispira species are complete and preserved in life position associated with the shell of T . argentina . The unguiculate morphology of the operculum is here interpreted as a mechanism for increasing the shell weight and ensuring anchoring to the substrate in a more or less fixed mode of life for Teiichispira , and as protection. The new occurrence of Teiichispira provides new taxonomic data on early Paleozoic marine gastropods in Argentina and testifies to a wide paleobiogeographical distribution for the genus, restricted to tropical and subtropical regions during the Early/Middle Ordovician.
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... 485 Ma (Wagner 2002) and ca. 420 Ma (Fryda et al. 2008), 2) its position in some phylotranscriptomic trees contradicting the monophyly of the Orthogastropoda (Zapata et al. 2014;Cunha and Giribet 2019), and 3) the difficulty of defining its members (Bouchet and Rocroi 2005;Bouchet et al. 2017;Ponder et al. 2019). Vetigastropoda is exclusively marine and includes more than 4064 living species (MolluscaBase 2021). ...
Gastropods have survived several mass extinctions during their evolutionary history resulting in extraordinary diversity in morphology, ecology, and developmental modes, which complicate the reconstruction of a robust phylogeny. Currently, gastropods are divided into six subclasses: Caenogastropoda, Heterobranchia, Neomphaliones, Neritimorpha, Patellogastropoda, and Vetigastropoda. Phylogenetic relationships among these taxa historically lack consensus, despite numerous efforts using morphological and molecular information. We generated sequence data for transcriptomes derived from twelve taxa belonging to clades with little or no prior representation in previous studies in order to infer the deeper cladogenetic events within Gastropoda and, for the first time, infer the position of the deep-sea Neomphaliones using a phylogenomic approach. We explored the impact of missing data, homoplasy, and compositional heterogeneity on the inferred phylogenetic hypotheses. We recovered a highly supported backbone for gastropod relationships that is congruent with morphological and mitogenomic evidence, in which Patellogastropoda, true limpets, are the sister lineage to all other gastropods (Orthogastropoda) which are divided into two main clades (i) Vetigastropoda s.l. (including Pleurotomariida + Neomphaliones) and (ii) Neritimorpha + (Caenogastropoda + Heterobranchia). As such, our results support the recognition of five subclasses (or infraclasses) in Gastropoda: Patellogastropoda, Vetigastropoda, Neritimorpha, Caenogastropoda and Heterobranchia.
... hemocyanin) in immunological applications (Harris and Markl 1999;Mora Román et al. 2019). Vetigastropods were the dominant clade of gastropods throughout the Paleozoic and most of the Mesozoic eras, with fossils going back at least to the Silurian (Fryda et al. 2008), and the divergence of crown groups being estimated as in the Devonian (Zapata et al. 2014). Despite their diversity, evolutionary importance, and applications for human food and health, the phylogeny of vetigastropods remains contentious, likely due to basal divergences being ancient and the small amount of molecular data that has been available thus far. ...
... Compared to analyses based on site-homogeneous models, site-heterogeneous models have been shown to more accurately infer the true species tree even in the presence of ILS (Wang et al. 2019). These deep vetigastropod divergences are between 200 and 400 Myr old (Zapata et al. 2014), if not older (Fryda et al. 2008), therefore we could alternatively hypothesize high heterogeneity in the evolution of sequences to be the cause of topological discordance. In this case, the more accurate inference could be that from the concatenated methods using profile mixture models. ...
Phylogenetic analyses may suffer from multiple sources of error leading to conflict between genes and methods of inference. The evolutionary history of the mollusc clade Vetigastropoda makes them susceptible to these conflicts, their higher level phylogeny remaining largely unresolved. Originating over 350 million years ago, vetigastropods were the dominant marine snails in the Paleozoic. Multiple extinction events and new radiations have resulted in both very long and very short branches and a large extant diversity of over 4000 species. This is the perfect setting of a hard phylogenetic question in which sources of conflict can be explored. We present 41 new transcriptomes across the diversity of vetigastropods (62 terminals total), and provide the first genomic-scale phylogeny for the group. We find that deep divergences differ from previous studies in which long branch attraction was likely pervasive. Robust results leading to changes in taxonomy include the paraphyly of the order Lepetellida and the family Tegulidae. Tectinae subfam. nov. is designated for the clade comprising Tectus, Cittarium and Rochia. For two early divergences, topologies disagreed between concatenated analyses using site heterogeneous models vs. concatenated partitioned analyses and summary coalescent methods. We investigated rate and composition heterogeneity among genes, as well as missing data by locus and by taxon, none of which had an impact on the inferred topologies. We also found no evidence for ancient introgression throughout the phylogeny. We further tested whether uninformative genes and over-partitioning were responsible for this discordance by evaluating the phylogenetic signal of individual genes using likelihood mapping, and by analyzing the most informative genes with a full multispecies coalescent model. We find that most genes are not informative at the two conflicting nodes, but neither this nor gene-wise partitioning are the cause of discordant results. New method implementations that simultaneously integrate amino acid profile mixture models and the multispecies coalescent might be necessary to resolve these and other recalcitrant nodes in the Tree of Life.
... As a result, a few key genera control the observed pattern of distribution and the biogeographical inferences that can be deduced from this. In general terms, gastropod diver sity increases steadily throughout the Ordovician until it drops at the end of the period, but with different patterns of dispersal, diversity and ecological distribution among gastropod groups, palaeocontinents and facies (Novack Gottshall and Miller 2003;Frýda and Rohr 2004;Frýda et al. 2008;Ebbestad et al. 2013), but also among members of the planktic and benthic realms (Nützel et al. 2006;Servais et al. 2010). Gastropod and tergomyan diversity and dis persal trends follow suite of other major benthic organisms during the complex and dynamic biodiversity increase seen throughout the Ordovician, constituting the socalled Great Ordovician Biodiversity Event (Servais and Harper 2018). ...
... Classification above family level is un certain (see Wagner 2002: 21), and therefore left tentative. See also discussions in Frýda and Rohr (2004), Frýda et al. (2008), andFrýda (2012). ...
... Findings of early Mimospira species in Avalonia, the Franconian Forest, Bohemia and Baltica may reflect a wide dispersal due to planktotrophy. It has been argued that the protoconch of Mimospira consist of an embryonic shell (protoconch I) and a larval shell (protoconch II) which is present in gastropods with planktotrophic larval dispersal (Frýda et al. 2008;Frýda 2012). This would allow the larvae to stay longer in the water column and facilitate a wider dispersal. ...
... Horný, 1964 [pars.]; Dzik, 1983Dzik, , 1999Frýda, 1999Frýda, , 2001Frýda, , 2012Rohr, 2004, 2006;Frýda et al., 2008). Current classifications regard them as basal gastropods of uncertain systematic position (Bouchet et al., 2005(Bouchet et al., , 2017. ...
... Table 1). It has been argued that the sinistral asymmetry of the embryonic protoconch 1 shows that they could not be hyperstrophic (Dzik, 1983(Dzik, , 1999, implying sinistral orthostrophy, and that other compelling evidence, e.g., an operculum, to demonstrate this condition is lacking (Frýda, 1995(Frýda, , 2012Frýda and Blodgett, 2001;Rohr, 2004, 2006;Frýda et al., 2008). The Clisospiroidea, as proposed by Bouchet et al. (2005Bouchet et al. ( , 2017, is largely a summary of the proposals by Knight et al. (1960) and Horný (1964) (see Table 1). ...
... Lack of conclusive evidence, such as an operculum, and the general problem of establishing hyperstrophy vs sinistral orthostrophy based on the shell alone, also preclude firm conclusions about the nature of shell coiling in mimospirids. Furthermore, the modified concept of a presumed untorted class Paragastropoda, now only including the mimospirids, has undermined the validity of that concept (Frýda et al., 2008;Frýda, 2012). Nützel (2014) expected higher disparity and different forms and clades among early gastropods, and mimospirids could very well represent part of a mainly late Cambrian and Ordovician clade of sinistral mollusks that later went extinct. ...
A new sinistrally coiled univalved mollusk Catalanispira n. gen. is described with two species; Catalanispira reinwaldti (Öpik, 1930) from the Middle Ordovician Kõgekallas Formation (Darriwilian) of Estonia and Catalanispira plattevillensis n. gen. n. sp. from the Upper Ordovician Platteville Formation (Sandbian) of northern Illinois, USA. Morphological features include a large, low-trochiform shell, a narrow lenticular aperture, a deep funnel-like umbilicus, a falcate inner lip and a large (1.4 mm wide) protoconch. Ornamentation consists of fine commarginal growth lines or ribs but superimposed on a slightly irregular shell surface. Catalanispira n. gen. is placed within the sinistrally coiled order Mimospirida and the family Onychochilidae, and Catalanispirinae n. subfam. is proposed. The large Lower Ordovician (Tremadocian) Pelecyogyra Ebbestad and Lefebvre, 2015 from Morocco and France is transferred to this new subfamily. The well-preserved initial growth stage of Catalanispira plattevillensis n. gen. n. sp. is cap-shaped, slightly asymmetrical, unusually large, and smooth, and represents either an unusually large embryonic shell (protoconch 1) or a larval shell (protoconch 2). It differs from the smaller protoconch described for the clisospirine Mimospira Koken in Koken and Perner, 1925, which might include a multiwhorled larval shell (protoconch 2). Mimospirids are dominantly Ordovician, and have been classified as untorted mollusks (only distantly related to gastropods), dextral hyperstrophic gastropods, or sinistral orthostrophic gastropods. Sinistral asymmetry already in the embryonic shell and lack of conclusive evidence for coiling direction, e.g., an operculum, could suggest that Catalanispira n. gen. or similar mimosprids were sinistral orthostrophic gastropods. Currently the group is therefore classified as a group of sinistral orthostrophic gastropods, unranked within the Gastropoda.
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... However, this is in contrast to the results from the study of Dinapoli (2009), where the point of origin of Euthyneura was traced back to the upper Triassic period. The present result defies the proposal of the origin of Euthyneura 'not before Mesozoic' (Fryda et al. 2008;Dinapoli 2009). ...
The phylogenetic relationships among the major groups of Pulmonata were studied by the information derived from a concatenated dataset consisting of mitochondrial (16S and COI) and nuclear (18S and 28S) markers. Heterobranchia are recovered as monophyletic. Euthyneura as paraphyletic due to the emergence of taxa from Opisthobranchia and lower Heterobranchia. The major groups of Pulmonata, namely Stylommatophora, Veronicellidae, Onchidiidae, Otinoidea, Siphonarioidea and Hygrophila are recovered as monophyletic. Monophyly of Basommatophora was not confirmed due to the variable position of Siphonarioidea and Amphiboloidea. Evolutionary divergence times for different taxa were also estimated using a relaxed molecular clock method in Bayesian evolutionary analysis by sampling trees (BEAST). The common ancestor of Heterobranchia and Caenogastropoda was originated in the Silurian period and the common ancestors of Euthyneura and Pulmonata were originated in the Carboniferous and lower Triassic periods, respectively.
... With > 70,000 living species, they are adapted to marine (any depth), brackish, freshwater, terrestrial and even arboreal habitats (Aktipis et al., 2008). They also show a rich fossil record and a wide variety of morphologies (Fryda et al., 2008). Therefore, they could constitute a sound model system for evolutionary biology studies, once a robust phylogeny for the group is available. ...
... ribosomal protein genes) have also been shown to bias phylogenetic inference [19,20], while genes with fast rates and high levels of saturation can cause longbranch attraction [15,21]. An additional model violation comes from gene tree discordance, not accounted for by concatenation methods, that can be caused by incomplete lineage sorting and be particularly relevant in areas of the tree with short internal branches [22][23][24], such as the radiation of crown gastropods during the Ordovician [16,25]. More commonly considered issues include rate heterogeneity between sites and missing data. ...
... Other traits shared by neritimorphs and apogastropods include complex reproductive anatomy, internal fertilization and encapsulated eggs, which hatch into a feeding veliger larva or directly into a juvenile [6,25,60 -62]. By contrast, vetigastropods and patellogastropods are mostly broadcast spawners, with embryos that develop in the plankton into non-feeding larvae, first as a trochophore that later gives rise to a veliger [6,25,60,62,63]. Character states shared by patellogastropods and vetigastropods have historically been interpreted as plesiomorphic based on the phylogenetic hypothesis in which patellogastropods were the sister group to all other gastropods, or due to a misguided notion that these are 'primitive' taxa [5,55]. ...
Gastropod molluscs are among the most diverse and abundant animals in the oceans, and are successful colonizers of terrestrial and freshwater environments. Past phylogenetic efforts to resolve gastropod relationships resulted in a range of conflicting hypotheses. Here, we use phylogenomics to address deep relationships between the five major gastropod lineages-Caenogastropoda, Heterobranchia, Neritimorpha, Patellogastropoda and Vetigastropoda-and provide one congruent and well-supported topology. We substantially expand taxon sampling for outgroups and for previously underrepresented gastropod lineages, presenting new transcriptomes for neritimorphs and patellogastropods. We conduct analyses under maximum-likelihood, Bayesian inference and a coalescent-based approach, accounting for the most pervasive sources of systematic errors in large datasets: compositional heterogeneity, site heterogeneity, heterotachy, variation in evolutionary rates among genes, matrix completeness, outgroup choice and gene tree conflict. We find that vetigastropods and patellogastropods are sister taxa, and that neritimorphs are the sister group to caenogastropods and heterobranchs. We name these two major unranked clades Psilogastropoda and Angiogastropoda, respectively. We additionally provide the first genomic-scale data for internal relationships of neritimorphs and patellogastropods. Our results highlight the need for reinterpreting the evolution of morphological and developmental characters in gastropods, especially for inferring their ancestral states.
... Even though the Permian-Triassic mass extinction severely affected caenogastropods, it is generally agreed that at least some cerithioids (or stem group cerithioids) were present in the late Palaeozoic (e.g., Nützel, 1998Nützel, , 2002Nützel and Bandel, 2000;Bandel et al., 2002;Nützel and Pan, 2005;Frýda et al., 2008;Ponder et al., 2008). The phylogenetic relationships and timing of appearance of various cerithioids in the Mesozoic, however, remain unclear. ...
Turritellid gastropods are important components of many Cretaceous–Recent fossil marine faunas worldwide. Their shell is morphologically simple, making homoplasy widespread and phylogenetic analysis difficult, but fossil and living species can be recognized based on shell characters. For many decades, it has been the consensus that the oldest definite representatives of Turritellidae are from the Lower Cretaceous, and that pre-Cretaceous forms are homeomorphs. Some morphological characters of the present turritelline species resemble those of mathildoids, but many diagnostic characters clearly separate these two groups. We here describe and/or redescribe—based on examination of more than 2600 near complete specimens—four species from the Upper Jurassic Dhosa Oolite Member of the Chari Formation in Kutch, western India, and demonstrate that they are members of Turritellidae, subfamily Turritellinae, on the basis of diagnostic characters including apical sculptural ontogeny (obtained from SEM study), spiral sculpture, and growth line patterns. The four species are in order of abundance, Turritella jadavpuriensis Mitra and Ghosh, 1979; Turritella amitava new species; Turritella jhuraensis Mitra and Ghosh, 1979, and Turritella dhosaensis new species. The turritelline assemblages occur only on the northeastern flank of the Jhura dome (23°24’47.57”N, 69°36’09.26”E). Age of the Dhosa Oolite has recently been confirmed based on multiple ammonite species. All these points indicate that these fossils are the oldest record of the family Turritellidae—by almost 30 million years—in the world.
