Bivalvia

Outer shell (above) and inner mould of Fordilla troyensis, from Walcott (1886).


Belongs within: Mollusca.
Contains: Daonella, Halobia, Polidevciidae, Sanguinolariidae, Ferganoconcha, Astartila, Myonia, Perrierinidae, Palaeomutelidae, Pleurophorus, Opponobrachia, Anatinidae, Leptonidae, Palaeoheterodonta, Anomalodesmata, Pteriomorphia, Heterodonta.

The Bivalvia are one of the main subgroups of the molluscs, readily distinguished from other classes by the division of the shell into two hinged lateral valves. Bivalves also lack a developed head and mostly feed by filtering organic particles from the water (with some notable exceptions). The valves of the shell are commonly connected at the hinge by a series of interlocking teeth. In members of the clade Autobranchia, the gills are enlarged and folded within the mantle cavity to function in filter-feeding. More basal bivalve lineages have the gills as simple non-reflected lamellae that function for respiration only, with the labial palps as the main food-collecting organs. Bivalves outside the clade Amarsipobranchia have at least the first few anterior filaments of the gills inserted unfused into a distal oral groove. In most members of the Amarsipobranchia, however, these filaments are fused or not inserted (Plazzi et al. 2011).

The earliest fossils that have been assigned to the bivalves come from the Early and Middle Cambrian and include the genera Fordilla, Pojetaia and Arhouriella (Budd & Jensen 2000). These are small forms, only a few millimetres in length, that may have had a burrowing lifestyle similar to that of the modern Nuculidae (Morton 2012). Fordilla and Pojetaia both had laterally compressed, subequally valved and suboval shells with few teeth in the hinge. Arhouriella had a veneriform shell with two teeth overhanging the hinge plate on the right valve (the left valve is unknown).

See also: Pig's toes and water nymphs.

Characters (from Giribet & Wheeler 2002): Body laterally compressed with an external bivalved shell that is hinged dorsally. Valves connected by a partially calcified elastic ligament and held together by 1 or 2 adductor muscles. Buccal or radular apparatus absent, mantle lobes either joined or free ventrally. Mantle cavity spacious, extending upwards on each side of visceral mass and containing a pair of laterally suspended ctenidia. Ctenidia may be enlarged, lamellate and plicate. Mouth and anus located at opposite ends of body, gut typically convoluted. Pair of ciliated labial palps connecting ctenidia to the mouth, directing food particles into it. Foot extensible, either elongated or laterally compressed.

<==Bivalvia [Acephala, Arcacea, Aviculae, Cryptodonta, Nuculiformii, Nuculoida, Palaeotaxodonta, Pelecypoda, Protobranchia]
    |--Fordillidae [Fordilloida] C96
    |    |--Neofordilla elegans Krasilova 1977 SB93
    |    |--Pojetaia Jell 1980 C96, M12
    |    |    `--P. runnegari Jell 1980 SB93
    |    `--Fordilla BJ00
    |         |--F. sibirica P08
    |         `--F. troyensis Barrande 1881 M12
    `--+--Opponobranchia PC11
       `--Autobranchia (see below for synonymy) PC11
            |  i. s.: Anatinidae W77
            |         Mactromyopsis [Mactromyidae] SB93
            |           `--M. (Mactromyella) inflata (Thévenin 1909) SB93
            |         Montanaria devonicus (Beushausen 1884) SB93
            |         Plesiocyprinella carinata Holdhaus 1918 SB93
            |         Leptonidae SB93
            |         Vokesella inopinata Chavan 1952 SB93
            |         Neoleptonidae SB93
            |           |--Goodalliopsis terminalis (Deshayes 1858) SB93
            |           |--Neolepton Monterosato 1875 F27a
            |           |    `--N. citrinum F27a
            |           `--Puyseguria Powell 1927 SB93, P61
            |                |--*P. cuneata Powell 1927 P61
            |                |--P. crenulifera Maxwell 1969 SB93
            |                |--P. prognata Powell 1927 P61
            |                `--P. tani Powell 1939 P61
            |         Cardiolarioidea C04
            |           |  i. s.: Cardiolaria beirensis C96
            |           |         Eritropis C04
            |           `--Praenuculidae SB93
            |                |--Ledopsis JB12
            |                |--Nuculodonta JB12
            |                `--Deceptrix C04
            |                     |--D. carinata Fuchs 1919 SB93
            |                     `--D. levata C96
            |--Palaeoheterodonta PC11
            `--Amarsipobranchia PC11
                 |--Anomalodesmata PC11
                 `--+--Pteriomorphia PC11
                    `--Heterodonta PC11

Bivalvia incertae sedis:
  Crytodonta saffordi PR79
  Pyrenomoeus EB01
  Batolites Montfort 1808 D-DP03
    `--B. organisans Montfort 1808 D-DP03
  Lapeirousiinae D-DP03
    |--Lapeirousia Bayle 1878 D-DP03
    |--Praelapeirousia D-DP03
    `--Lapeirousella D-DP03
  Plicatomytilus AV03
  Cataceramus HS03
    |--C. barabini (Morton 1834) HS03
    `--C. subcircularis (Meek 1876) HS03
  Horiopleura lamberti HS02
  Polyconites verneuili HS02
  Pseudotoucasia santanderensis HS02
  Sphaerulites HS02
  Chondrodonta HS02
  Nodipecten nodosus TL03
  Callithaca adamsii L10
  Swiftopecten swifti GAS03
  Liocyma XZ10
    |--L. chinensis XZ10
    `--L. fluctuosa GAS03
  Alveinus ojianus GAS03
  Panomya arctica KBC03
  Sertifer keenae KBC03
  Gouldia M62
    |--G. cerina WE63
    |--G. minima M62 [=Circe minima C64]
    `--G. pacifica C64
  Carolia placunoides G92
  Cubitostrea G92
  Nicaisolopha G92
  Devonia perrieri (Malard 1903) (see below for synonymy) BK77
  Halobiidae [Halobioidea] S04
    |--Daonella S04
    `--Halobia KI70
  Natasia boliviensis SV03
    |--N. b. boliviensis SV03
    `--N. b. fragilis SV03
  Arisaigia postornata P98
  Potamocorbula XZ10
    |--P. amurensis (Schrenck 1867) XZ10
    |--P. laevis (Hinds 1843) XZ10
    `--P. ustulata (Reeve 1865) XZ10
  Polidevciidae SB93
  Pseudophytina rugifera GW02
  Mactromeris polynyma GR98
  Sarepta Adams 1860 [Sareptidae] DK08
    |--S. obolella (Tate 1886) [=Leda obolella) F27a
    `--S. planiuscula (Tate 1886) [=Leda planiuscula, Ovaleda planiuscula] DK08
  Saxicava [Saxicavidae] F27a
    |--S. antarctica F27a
    |--S. australis F27a
    |--S. norvegica N79
    |--S. pholadis Middendorff 1851 N79 (see below for synonymy)
    |--S. rugosa [incl. S. rugosa var. praecisa] U79
    `--S. subalata F27a
  Barcoona Finlay 1927 [=Pachydomella Etheridge 1907 non Ulrich 1891] F27b
    `--*B. chutus (Etheridge 1907) [=*Pachydomella chutus] F27b
  Modiolarca impacta HS01
  Pseudoarcopagia disculus HS01
  Varinucula gallinacea HS01
  Melina cumingii (Reeve 1858) [=Perna cumingii] H09
  Crenatula H09
    |--C. flammea Reeve 1858 H09
    `--C. picta (Gmelin 1791) [=Ostrea picta] H09
  Meleagrina H09
    |--M. anomioides (Reeve 1857) [=Avicula anomioides] H09
    |--M. fimbriata [incl. M. mazatlanica] C64
    |--M. flexuosa (Reeve 1857) [=Avicula flexuosa] H09
    |--M. lucunata (Reeve 1857) [=Avicula lucunata] H09
    |--M. margaritifera (Linnaeus 1758) [=Mytilus margaritiferus] H09
    |--M. maxima (Jameson 1901) [=Margaritifera maxima] H09
    |--M. sugillata (Reeve 1857) [=Avicula sugillata] H09
    `--M. tegulata (Reeve 1857) [=Avicula tegulata] H09
  Aenigma aenigmatica [=Tellina aenigmatica] H09
  Placenta H09
    |--*P. placenta (Linnaeus 1758) [=Anomia placenta] H09
    `--P. lobata Soerby 1871 H09
  Myrina coppingeri Smith 1885 H09
  Coelodon elongatus Carpenter 1846 H09
  Crassatellites H09
    |--C. janus Hedley 1906 H09
    |--C. rhomboides (Smith 1885) [=Crassatella rhomboides] H09
    |--C. torresi (Smith 1885) [=Crassatella torresi] H09
    `--C. ziczac [=Crassatella ziczac] H09
  Carditella H09
    |--C. infans Smith 1885 H09
    `--C. torresi Smith 1885 H09
  Opisocardium subretusum (Sowerby 1841) [=Cardium subretusum] H09
  Glaucomya H09
    |--G. desmoyersii H09
    `--G. rugosa Hanley 1828 [=Glauconomya rugosa] H09
  Metis H09
    |--M. meyeri (Philippi 1846) [=Tellina meyeri] H09
    `--M. spectabilis (Hanley 1844) [=Tellina spectabilis] H09
  Phylloda foliacea (Linnaeus 1758) [=Tellina foliacea] H09
  Sanguinolariidae P61
  Psammosolen H09
    |--P. australis (Dunker 1861) [=Macha australis] H09
    |--P. coarctatus (Gmelin 1791) [=Solen coarctatus] H09
    |--P. deshayesii (Dunker 1861) [=Macha deshayesii] H09
    |--P. minutus (Dunker 1861) [=Azor minutus] H09
    `--P. sulcatus (Dunker 1861) [=Macha sulcatus] H09
  Labiosa grayi (Adams 1872) [=Raeta grayi] H09
  Davila plana (Hanley 1843) [=Mesodesma plana] H09
  Roccellaria pupina (Deshayes 1855) [=Gastrochaena pupina] H09
  Nausitoria H09
    |--N. manni Wright 1865 H09
    `--N. thoracites (Gould 1856) [=Calobates thoracites] H09
  Margarita Leach 1814 BR05
    |--M. maculata [incl. M. elegantula Jeffreys 1864 (n. n.)] J64
    `--M. minutissima Mighels 1843 J49
  Peloris Poli 1791 BR05
  ‘Venilia’ Morton 1833 non Rafinesque 1815 BR05
  Fasciculiconcha guangxiensis ZP86
  Aloidis [Aloididae] M54
    `--A. iredalei M54
  Conradilla caelata USDI77
  Cyrtonaias tampicoensis USDI77
  Plethobasis USDI77
    |--P. cicatricosus USDI77
    |--P. cooperianus USDI77
    `--P. cypha B66
  Eotrigonia semiundulata Q72
  Papyridea BD86
    |--P. australe (Sowerby 1834) BD86
    `--P. papyraceum (Gmelin 1791) BD86
  Bathymodiolus Kenk & Wilson 1985 AJH07
    `--B. brevior R01
  Mesosaccella Chavan 1946 SY85
  Danlengiconcha Lu 1978 K84
  Volsellina zhesiensis Liang in Ding, Xia et al. 1984 DX84
  Ferganoconcha ZZ88
  Nakamuranaia ZZ88
    |--N. chingshanensis ZZ88
    |--N. subrotunda ZZ88
    `--N. zhejiangensis ZZ88
  Sibireconcha ZZ88
    |--S. anodontoides ZZ88
    `--S. jenissejensis ZZ88
  Solenaia menyinensis ZZ88
  Pseudocardiniidae SB93
    |--Tutuella chachlovi Ragozin 1938 SB93
    |--Utschamiella tungussica Ragozin 1938 SB93
    `--Kija SB93
         |--K. kibetenensis Lebedev 1958 SB93
         `--K. tjazhinensis Lebedev 1958 SB93
  Actinoceramus KK08
    |--A. concentricus KK08
    `--A. salomoni (d’Orbigny 1850) KK08
  Actinopterella F71
    |--A. lamellosa Sherrard 1959 F71
    `--A. minuta Sherrard 1959 F71
  Actinopteria F71 [incl. Clathrocoelia Hall 1879 F62]
    |--A. australis Dun 1896 F71
    `--A. decussata [incl. *Clathrocoelia eborica Hall 1879] F62
  Astartila F71
  Aucella F71
    |--A. hausmanni G31
    `--A. hughendensis F71
  Girtypecten tessellatus [=Aviculopecten tessellatus] F71
  Burrier dunii Laseron 1910 F71
  Stutchburia F71
    |--S. compressa F71
    |--S. costata [=Cardinia costata] F71
    |--S. farleyensis Etheridge 1900 F71
    |--S. obliqua Etheridge 1900 F71
    |--S. praerupta (Dana 1849) [=Cypricardia praerupta] F71
    |--S. recta (Dana 1849) [=Cardinia recta] F71
    |--S. simplex (Dana 1849) [=Cypricardia simplex] F71
    `--S. veneris [=Cypricardia veneris] F71
  Cleobis F71
    |--C. grandis (Dana 1849) (see below for synonymy) F71
    `--C. gryphoides [=Cardiomorpha gryphoides, Megadesmus (Megadesmus) gryphoides] F71
  Corimya primula F71
  Cypricardinia Hall 1859 SB93
    |--C. gregarius (Etheridge 1900) (see below for synonymy) F71
    |--C. sinensis G31
    `--C. subelegans G31
  Cytheria moorei F71
  Diaphragmella merismopteroides Etheridge 1918 F71
  Permopecten aviculatum [=Entolium aviculatum] F71
  Glyptodesma bundanoonensis Etheridge 1917 F71
  Glyptoleda F71
    |--G. glomerata Fletcher 1945 F71
    `--G. reidi Fletcher 1945 F71
  Grammatodon daintreei Etheridge 1907 F71
  Grammysioides declivis Sherrard 1959 F71
  Maccoyella F71
    |--M. barklyi F71
    |--M. corbiensis F71
    |--M. mariaeburnensis C16
    |--M. reflecta F71
    `--M. umbonalis F71
  Myonia F71
  Nucundata darwini [=Nuculana darwini, Phestia darwini, Tellinomya darwini] F71
  Pachydomus sacculus M’Coy 1847 F71
  Protovirgus F71
    |--P. dunstani (Etheridge 1888) [=Unio dunstani] F71
    `--P. flemingi McMichael 1956 F71
  Solenomya translineata Laseron 1910 F71
  Tatella aptiana F71
  Nototrigonia cinctuta (Etheridge 1902) [=Trigonia cinctuta] F71
  Undulomya rugulata Fletcher 1946 F71
  Unionella F71
    |--U. bowralensis Etheridge 1888 F71
    |--U. carnei Etheridge 1888 F71
    `--U. wianamattensis (Etheridge 1888) [=Unio wianamattensis] F71
  Mysidioptera Salomon 1895 [=Latemaria Wilckens 1909] W60
    |--*M. ornata Salomon 1895 (see below for synonymy) W60
    |--M. exima Diener 1913 W60
    |--M. globosa Broili 1930 [=M. (Latemaria) globosa] W60
    |--M. globularis (Trechmann 1918) [=Megalodon globularis] W60
    |--M. kittlii Bittner 1895 [=M. (*Latemaria) kittlii] W60
    |--M. obliqua Broili 1903 W60
    |--M. riceae Waterhouse 1960 W60
    |--M. vixcostata W60
    `--M. woehrmanni Salomon 1895 W60
  Hokonuia Trechmann 1918 W60
    `--H. limaeformis Trechmann 1918 (see below for synonymy) W60
  Cardiomorpha nuggetensis Trechmann 1918 W60
  Foramelina exempla MG-H11
  Fissilunula clarkei H75
  Apidopholas Fischer 1887 H75
  Karadjalia venusta H79
  Volviceramus K79
    |--V. exogyroides K79
    `--V. involutus K79
  Cremnoceramus K79
    |--C. inconstans K79
    |--C. koeneni K79
    `--C. wandereri K79
  Endocostea K79
    |--E. baltica K79
    `--E. typica K79
  Enantiostreon difforme (Schlotheim 1820) KL04
  Eurynia D56 [=Euryma (l. c.) B64]
    `--E. iris D56
  Camya Hinz-Schallreuter 1995 M12
  Arhouriella Geyer & Streng 1998 M12
  Mixtipecten amuriensis (Woods 1917) S98
  Pectenoproductus Likharev 1930 M65
    `--*P. proprius Likharev 1930 M65
  Halophiala Koken 1925 KC60
  Shaanxiconcha cliovata HH09
  Perrierinidae P61
  Obovaria V16
    |--O. levigata (Rafinesque 1820) [=Unio levigata, Elliptio levigata; incl. Obovaria lens Lea 1831] V16
    |--O. olivaria (Rafinesque 1820) (see below for synonymy) V16
    |--O. retusa Lam. 1819 [incl. O. torsa Rafinesque 1820, Unio torsa] V16
    `--O. subrotunda (Rafinesque 1820) (see below for synonymy) V16
  Symphynota viridis (Rafinesque 1820) [=Unio viridis, Elliptio viridis] V16
  Hemimactra elongata H86
  Hiatula H86
    |--H. compacta C64
    |--H. diphos F66
    |--H. incerta H86
    `--H. nuttalli C64
  Mysia neozelanica H86
  Solenella H86
    |--S. australis [incl. Nucula ornata] H86
    `--S. funiculata H86
  Macrodon H86
    |--M. australis H86
    |--M. kingianum G31
    |--M. naliwkini G31
    |--M. striatum G31
    `--M. verneuilianum G31
  Mactrinula dolabrata (Reeve 1854) XZ10
  Raetellops pulchella (Adams & Reeve 1850) XZ10
  Coelomactra antiquata (Spengler 1802) XZ10
  Nitidotellina iridella (Mastensi 1868) XZ10
  Cyclina XZ10
    |--C. sinensis (Gmelin 1791) XZ10
    `--C. subquadrata C64
  Trigonothracia XZ10
    |--T. jinxingae Xu 1980 XZ10
    `--T. pusilla (Gould 1861) XZ10
  Avella sinica (Wang & Qi 1984) XZ10
  Talonostrea talonata Li & Qi 1994 XZ10
  Tugonia XZ10
    |--T. anatina PP64
    `--T. huanghaiensis Xu 1987 XZ10
  Sinolyonsia sinica Xu 1992 XZ10
  Porterius dalli (Smith 1885) XZ10
  Asthenothaerus XZ10
    |--A. huaughaensis Xu 1989 XZ10
    `--A. villosior C64
  Crassithracia ovata (Xu 1989) XZ10
  Bulluniella borealis P08
  Pseudomyonia Runnegar 1983 P08
  Gastrana fragilis PP64
  Alba tenuis PP64
  ‘Pleurodonta’ Conrath 1887 non Herrmanssen 1847 BR17
  Caesariella JB12
  Paleostraba JB12
  Chesapecten middlesexensis P12
  Mansfieldostrea geraldjohnsoni P12
  Dualina T08
    |--‘Nautilus’ desideratus Barrande 1865–1867 T08
    `--D. longiuscula (Barrande 1881) T08
  Algerina algena Kříž 2008 T08
  Sainshandia V-RR93
    |--S. bainshirensis V-RR93
    |--S. dushiulensis V-RR93
    |--S. mongolica V-RR93
    `--S. ongonulensis V-RR93
  Pseudohyria V-RR93
    |--P. cardiiformis V-RR93
    |--P. debilicostata V-RR93
    `--P. marginodentata V-RR93
  Plicatotrigonioides V-RR93
    |--P. gobiaensis V-RR93
    `--P. khamariensis V-RR93
  Gogiola multicostata V-RR93
  Buginella V-RR93
    |--B. buginica V-RR93
    `--B. pseudoanodonta V-RR93
  Plicatella transaltica V-RR93
  Carnalpia rostrata Křiž 1974 D94
  Thoralia [Thoraliidae] SB93
    `--T. languedociana (Thoral 1935) SB93
  Pristiglomidae SB93
  Siliculidae SB93
  Lametilidae SB93
  Tindaria [Tindariidae] SB93
    `--T. arata Bellardi 1875 SB93
  Pachymytilus Zittel 1881 SB93
  Pseudopachymytilus Krumbeck 1923 SB93
  Boiomytilus Růžička & Prantl 1961 SB93
  Glendella dickinsi Runnegar 1970 SB93
  Sichuania marwicki Waterhouse 1980 SB93
  Carditomantea spinata Quenstedt 1929 SB93
  Cypricardella Hall 1858 SB93
  Tellidorella Berry 1962 SB93
  Rhaetidia zittellii Bittner 1895 SB93
  Ceromyella miotaurina Sacco 1901 SB93
  Roxoa intrigans (Mendes 1944) SB93
  Prilukiellidae SB93
    |--Amnigeniella kumsassiana (Ragozin 1960) SB93
    `--Microdontella problematica Lebedev 1944 SB93
  Palaeomutelidae SB93
  Acharax Dall 1908 AJH07
    `--A. cretacea Kanie & Nishida 2000 AJH07
  Linga De Gregorio 1884 AJH07
  Thyella lamellosa [incl. Cumingia elegans] M01
  Ciliatocardium ciliatum HG98
  Serratina eugonia PG98
  Stagnestesta Tasch 1961 T61
    `--*S. solitaria Tasch 1961 T61
  Rostrotortus Tasch 1961 T61
    `--*R. dissimilis Tasch 1961 T61
  Productae Tasch 1961 T61
    `--*P. dunbaris Tasch 1961 T61
  Elliptoidea Tasch 1961 T61
    `--*E. vulgaris Tasch 1961 T61
  Harvella elegans C64
  Gratelupia hydeana C64
  Mactrella C64
    |--M. carinata [incl. M. alata] C64
    `--M. exoleta C64
  Pseudobakewellia krasnowidowoensis G31
  Netschajewia G31
    |--N. elongata G31
    `--N. globosa G31
  Pleurophorina simplex G31
  Pleurophorus G31
  Alula kutorgi G31
  Bakewellia G31
    |--B. antiqua G31
    |--B. bicarinata G31
    |--B. ceratophaga G31
    |--B. ladina G31
    |--B. nikitowkensis G31
    `--B. sulcata G31
  Cyrtodontarca bakewellioides G31
  Clinopristha radiata G31
  Angustella praeangusta [=Gervilleia praeangusta] G31
  Palaeostrea sinensis G31
  Cucullopsis quadrata G31
  Acanthopecten shansiensis G31
  Posidoniella G31
    |--P. elongata G31
    |--P. laevis G31
    |--P. pyriformis G31
    |--P. variabilis G31
    |--P. vetusta G31
    `--P. yuani G31
  Vola praecursor G31
  Posidonomya becheri G31
  Jousseaumiella concharum Knudsen 1944 H64
  Trigonicardia antillarum WE63
  Canthyria N65b
    |--C. collina N65b
    `--C. spinosa N65b
  Schistodesmus lampreyanus N65b
  Dozierella gouldii N65a

Autobranchia [Anatinacea, Autolamellibranchiata, Dimyaria, Eulamellibranchia, Heteroconchia, Lamellibranchia, Lucinata, Lucinoida, Mesosyntheta, Metabranchia, Mytilacea, Mytiliformii, Prionodesmacea] PC11

Cleobis grandis (Dana 1849) [=Maeonia grandis, Megadesmus (Cleobis) grandis; incl. Pachydomus gigas M’Coy 1847, Maeonia gigas, Myonia gigas, Ma. gracilis Dana 1849, Megadesmus (Cleobis) gracilis, Myonia gracilis, Globicarina grossula Waterhouse 1965, Cleobis robusta Laseron 1910] F71

Cypricardinia gregarius (Etheridge 1900) [=Pleurophorus gregarius; incl. P. gregarius var. inflata Laseron 1910] F71

Devonia perrieri (Malard 1903) [=Synapticola perrieri, Entovalva perrieri; incl. Montacuta donacina Bateson 1923] BK77

Hokonuia limaeformis Trechmann 1918 [incl. Nuggetia morganiana Wilckens 1927, H. parki Wilckens 1927, H. rotundata Trechmann 1918] W60

*Mysidioptera ornata Salomon 1895 [incl. M. ornata var. laevigata Bittner 1895, M. ornata var. lombardica Bittner 1895] W60

Obovaria olivaria (Rafinesque 1820) [=Amblema olivaria, Unio olivaria; incl. Obovaria ellipsis Lea 1828] V16

Obovaria subrotunda (Rafinesque 1820) [=Obliquaria subrotunda, Unio subrotunda; incl. Obovaria circulus Lea 1829, O. striata Rafinesque 1820, Unio striata] V16

Saxicava pholadis Middendorff 1851 N79 [incl. S. clava C64, S. distorta C64, S. gastrochaenoidea C64, S. legumen C64]

*Type species of generic name indicated

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