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Cryptic bivalves with descriptions of new species from the Rodrigues lagoon

Taylor & Francis
Journal of Natural History
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Abstract

This paper describes a number of new taxa in the Arcidae, Philobryidae, Mytilidae, Galeommatidae, Kelliidae, Cyamiidae, Carditidae, Thraciidae and Clavagellidae. A further 17 species are included as new to the malacofauna of Rodrigues. Most of the species included came from cryptic habitats, of which the most interesting are apparently cavernicolous and similar to those described from southern Japan. Other species were described from crevice or algal turf communities. The most understudied group is the Galeommatoidea with four species described as new and a further five with as yet unresolved taxonomy.
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... He also noted that one specimen "gave birth to a single young […] which was approximately 300 microns in length (and) immediately began to burrow", thus suggesting brooding up to the juvenile stage. Brooding was also recorded for a few other species from different genera, and some data on the soft parts were provided along conchological observations in faunistic or taxonomic studies (Bernard, 1896a;Bernard, 1896b, Bernard, 1898Waller, 1973;Salas & Rolán, 1990;Salas & von Cosel, 1991;Middelfart, 2002a;Middelfart, 2002b;Oliver & Holmes, 2004;Güller & Zelaya, 2013;Malchus & Sartori, 2013). ...
... In a small species of Astartidae (Goodallia triangularis (Montagu, 1803), with a maximum shell size of four mm), only a single demibranch (the inner) is present (Saleuddin, 1965); this condition also occurs in species of Crassinella (Crassatellidae) (Harry, 1966;Allen, 1968). For species classified as condylocardiids, reduction to a single demibranch was recorded by Middelfart (2002aMiddelfart ( , 2002b, Oliver & Holmes (2004) and Güller & Zelaya (2013) (in this latter work, a species of the Carditopsis was reported as having a single demibranch, whereas two species of Carditella were recorded as bearing both demibranchs, with the outer strongly reduced). Reduction of ctenidial elements such as demibranchs or filaments is a recurrent phenomenon in Bivalvia, occurring independently in many groups (e.g., Purchon, 1939;Bieler et al., 2014, MorphoBank matrix), and frequently associated with small size (as discussed for species classified as condylocardiids by Güller & Zelaya, 2013). ...
... For the condylocardiids, observations are mainly restricted to the information provided by Bernard (1896aBernard ( , 1896b, Moore (1961), Waller (1973), Salas & Rolán (1990), Salas & von Cosel (1991) and Middelfart (2002aMiddelfart ( , 2002b. Moore (1961) was the only author to record ovoviviparity in a living condylocardiid, and Salas & von Cosel (1991), Middelfart (2002a) and Oliver & Holmes (2004) observed chemically fixed animals with brooded juveniles, albeit in different places of the parent's body: in two specimens of Carditopsis gofasi Salas & von Cosel, 1991, the young appeared to be held among the gill filaments (Salas & von Cosel, 1991;figs. 42-47). ...
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‘Miniaturization’ is a widespread phenomenon among the Metazoa. In the molluscan class Bivalvia, records of miniaturization are numerous. Among the Archiheterodonta, Warrana besnardi (Klappenbach, 1963) has attracted attention for its tiny size, which does not exceed 1.5 mm in shell length, and because it belongs to a group with limited anatomical information and often-debated status, the “Condylocardiidae” (which recent molecular studies place deeply nested within the family Carditidae). All species of Warrana Laseron, 1953 are small-bodied, and so miniaturization presumably occurred from a large-bodied ancestor within the Carditidae sensu lato. South American W. besnardi is here studied in detail. Its small size and the enlargement of the anterodorsal region during growth, reflects (and likely led) to infaunal habit, living as a burrowing bivalve that passively feeds on deposit particles entering the pallial cavity anteriorly. Mantle glands, previously reported as a common feature of other archiheterodonts, are missing in W. besnardi, but spongiform tissue in the antero-ventral portion of the mantle lobes presumably represents a blood sinus that might compensate for the great reduction of the ctenidia. Lecithotrophy is reported, with yolky oocytes bearing a thick non-cellular capsule layer; brooding was not observed, and it is here hypothesized that the extreme miniaturization, with the great reduction of ctenidia, is responsible for a shift in the reproductive mode of condylocardiids, contrasting with the commonly reported ovoviviparity of the carditids.
... The underlying processes are not understood. However, there is sufficient evidence to correlate these growth patterns with extended brooding (hayaMi & kase, 1993;MiddelFart, 2002a;oliver & holMes, 2004;this study). Because the distinction of shell stages is important for homology hypotheses and for a morphologybased classification, shells with indistinct boundaries (or where a clear boundary could represent either P-1/P-2 or P/N) are presently called cryptoconch rather than prodissoconch or metaconch. ...
... 2;MiddelFart, 2002aMiddelFart, , 2002b. Larger pits and ridges may be traced on the shell interior ( Fig. 15-16 liNse, 2002;MoraN, 2004a;oliver & holMes, 2004). Conical elevations often possess distinct sculptural characteristics ( Fig. 15.2), and the underlying microstructure is antimarginal granular-prismatic. ...
... Examples include numerous arcoids (e.g., Philobryidae, Barbatia Gray, 1842); Cyclochlamydidae (Cyclochlamys FiNlay, 1926, right valves);and Condylocardiidae (Austrocardiella MiddelFart, 2002a;Benthocardiella PoWell, 1930), among others (Fig. 37.3-37.8;dell, 1964MiddelFart, 2002a;oliver & holMes, 2004;Malchus, 2006). chaParro and others (2011) illustrated a cryptoconch (probably P-1) of Gaimardia bahamondei osorio & arNaud, 1984, which represents a transitional shell type between ST-2D (overall outline) and ST-3B (domed central portion with distinct microsculpture and differentiation of ears). ...
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This chapter provides a state-of-the-art review (as of 2013) of the ontogeny of the early shell in bivalve molluscs written for a revised Bivalve Treatise. All bivalves go through a succession of two or more shell-forming phases before the adult valves take shape. These early, larval to juvenile portions of the shell, and their morphology, morphogenetic patterns, taxonomic distribution, and evolutionary interpretations are the subjects of this chapter. Some aspects of the processes of shell formation at the level of cell lineages and control genes are briefly summarized. Most research conducted on these topics focuses on phases preceding the veliger and pericalymma larvae. However, from a morphological perspective, the most instructive modifications occur around the metamorphic window, where shell features are most diverse and record differences in reproductive modes and heterochronic changes. These changes provide clues to the evolution of autobranchs and protobranchs. The following sections are dedicated to establishing a theoretical framework for interpretation and defining the essential terms.
... The underlying processes are not understood. However, there is sufficient evidence to correlate these growth patterns with extended brooding (hayaMi & kase, 1993;MiddelFart, 2002a;oliver & holMes, 2004;this study). Because the distinction of shell stages is important for homology hypotheses and for a morphologybased classification, shells with indistinct boundaries (or where a clear boundary could represent either P-1/P-2 or P/N) are presently called cryptoconch rather than prodissoconch or metaconch. ...
... 2;MiddelFart, 2002aMiddelFart, , 2002b. Larger pits and ridges may be traced on the shell interior ( Fig. 15-16 liNse, 2002;MoraN, 2004a;oliver & holMes, 2004). Conical elevations often possess distinct sculptural characteristics ( Fig. 15.2), and the underlying microstructure is antimarginal granular-prismatic. ...
... Examples include numerous arcoids (e.g., Philobryidae, Barbatia Gray, 1842); Cyclochlamydidae (Cyclochlamys FiNlay, 1926, right valves);and Condylocardiidae (Austrocardiella MiddelFart, 2002a;Benthocardiella PoWell, 1930), among others (Fig. 37.3-37.8;dell, 1964MiddelFart, 2002a;oliver & holMes, 2004;Malchus, 2006). chaParro and others (2011) illustrated a cryptoconch (probably P-1) of Gaimardia bahamondei osorio & arNaud, 1984, which represents a transitional shell type between ST-2D (overall outline) and ST-3B (domed central portion with distinct microsculpture and differentiation of ears). ...
... The same is not true of the free-living 'gastropodlike' species with ornate/colourful mantles. Many presumably freeliving species with this probable general appearance were described in the 19th century by, for example, Deshayes (1856) who named no fewer than 39 species of ' Scintilla ' based on empty shells collected by Cuming in the Philippines and elsewhere in the tropical Pacific, an impediment also lamented by Oliver & Holmes (2004) . Deshayes emphasized ' animal ignotum ' and many of these nominal species have never been treated in the recent literature or, when they have, in a manner that does not add to the original description. ...
Article
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Galeommatoidea are bivalves with small (usually 1–15 mm), fragile, sometimes vestigial, shells, occurring free-living or in association with other benthic invertebrates from the intertidal to depths in excess of 7,000 m. They are the most diverse but least studied and most underrepresented group of bivalves in biodiversity surveys and museum collections. The majority of commensal species of Galeommatoidea have dull, frequently uncoloured, unsculptured shells, but free-living species have papillate, sometimes coloured, mantles; they can be gregarious, and superficially resemble crawling gastropods. Many species of Galeommatoidea remain to be discovered, and we argue that future descriptions of free-living new species should ideally include photos of living animals, while many of the historically known species described based on their shells will probably be best treated as nomina dubia.
... Condylocuna tricosa Middelfart, 2002 ( Figure 5C) was described from Qld and synonymised with Condylocuna io (Barsch, 1915) by Graham and Holmes (2004) despite Middelfart (2002) comparing C. tricosa with the South African type of C. io (United States National Museum (USNM 251043) and stating the dissimilarity. Oliver and Holmes (2004) did not examine the type of C. io, and compared it only to material from Rodrigues Island, which is C. tricosa and not C. io. Hence, the synonymy is here revoked and the distribution of C. tricosa is extended across the tropical Indo-Pacific region from Qld to Rodrigues Island and the Kimberley region. ...
... However, A. plicata may comprise a complex of species (Oliver and Holmes, 2004). The holotype of B. ...
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The bivalve molluscan fauna of Jeju Island (Jeju-do), the largest island of Korea, is reviewed based on original and literature data. An annotated and illustrated catalogue of species with indication of local distribution and taxonomic comments is prepared, supplemented also with data on general distribution, habitats, and primary synonyms. This first part of the paper includes 105 species belonging to 21 families (Nuculidae through Limidae). The history of Jeju molluscan research, a bibliography on all aspects of mollusk biology, and the general ecology of Jeju bivalves in various habitats are also included. Five species are reported as new for Jeju Island, two species as new for Korea, and 11 species have been reported in literature but were not included in the first bivalve catalogue of Jeju by Noseworthy et al. (2007); in total 16 species are added to the fauna.
... A study of H. strangei by Morton (2002b), however, has shown to the contrary that Humphreyia is most closely linked to the Brechites (or Penicillus) "lineage" (as Savazzi, 1999 and2005 puts it). Moreover, Dianadema multangularis (Tate, 1887) has hitherto been placed in the genus Clavagella (B.J. Smith, 1971(B.J. Smith, , 1976 but Morton (2003a) has shown that it is distinct in that it possesses a crown of tubules arising from the apex of the crypt, as also seen in the Indian Ocean D. mascarenensis recently described by Oliver and Holmes (2004) and the fossil Ascaulocardium armatum (Pojeta and Soh!, 1987). 3. Morton (2005Morton ( , 2006a has studied the structure and method of formation of the adventitious tubes of the fossil clavagellid Stirpulina coronata (Deshayes, 1824) and the extant Stirpulina ramosa and showed that the process involved is fundamentally different from that of the penicillid Brechites vaginiferus, as elucidated by Harper and Morton (2004). ...
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