162:
192:
280:
268:
1679:
1673:
1667:
211:
20:
177:
117:
161:
448:
Klug, C., G. Schweigert, D. Fuchs & G. Dietl (2010). First record of a belemnite preserved with beaks, arms and ink sac from the
Nusplingen Lithographic Limestone (Kimmeridgian, SW Germany).
629:
Hunt, S. & M. Nixon (1981). A comparative study of protein composition in the chitin-protein complexes of the beak, pen, sucker disc, radula and oesophageal cuticle of cephalopods.
1326:
Uchikawa, K., M. Sakai, T. Wakabayashi & T. Ichii (2009). The relationship between paralarval feeding and morphological changes in the proboscis and beaks of the neon flying squid
140:. Cephalopod beaks gradually become less stiff as one moves from the tip to the base, a gradient that results from differing chemical composition. In hydrated beaks of the
191:
1349:
1028:
Cherel, Y. & K.A. Hobson (2005). Stable isotopes, beaks and predators: a new tool to study the trophic ecology of cephalopods, including giant and colossal squids.
588:
1069:
Clarke, M.R. & R.E. Young (1998). Description and analysis of cephalopod beaks from stomachs of six species of odontocete cetaceans stranded on
Hawaiian shores.
169:
1350:"Current status of using beaks to identify cephalopods: III International Workshop and training course on cephalopod beaks, Faial Island, Azores, April 2007"
1914:
1397:
Xavier, J.C., R.A. Phillips & Y. Cherel (2011). Cephalopods in marine predator diet assessments: why identifying upper and lower beaks is important.
1535:
1919:
136:, beaks are more-or-less indigestible and are often the only identifiable cephalopod remains found in the stomachs of predatory species such as
701:
1437:
1232:
1137:
Hobson, K.A. & Y. Cherel (2006). Isotopic reconstruction of marine food webs using cephalopod beaks: new insight from captively raised
1213:
425:
Tanabe, K., P. Trask, R. Ross & Y. Hikida (2008). Late
Cretaceous octobrachiate coleoid lower jaws from the north Pacific regions.
733:
Miserez, A., T. Schneberk, C. Sun, F.W. Zok & J.H. Waite (2008). The transition from stiff to compliant materials in squid beaks.
654:
1109:
517:
Morton, N. & M. Nixon (1987). Size and function of ammonite aptychi in comparison with buccal masses of modem cephalopods.
1233:"Cephalopod beak identification and biomass estimation techniques: tools for dietary studies of southern Australian finfishes"
1356:
779:"A beak key for eight eastern tropical Pacific cephalopod species with relationships between their beak dimensions and size"
279:
267:
599:
1722:
1062:
Clarke, M.R. & L. Maddock (1988). Beaks of living coleoid
Cephalopoda. In: M.R. Clarke & E.R. Trueman (eds.)
1430:
1678:
1672:
803:
1666:
1159:
360:
1382:
393:
Kanie, Y. (1998). New vampyromorph (Coleoidea: Cephalopoda) jaw apparatuses from the Late
Cretaceous of Japan.
69:(lower) mandible and together they function in a scissor-like fashion. The beak may also be referred to as the
1292:
Ogden, R.S., A.L. Allcock, P.C. Watts & J.P. Thorpe (1998). The role of beak shape in octopodid taxonomy.
84:
Fossilised remains of beaks are known from a number of cephalopod groups, both extant and extinct, including
1707:
540:
Lehmann, U. & C. Kulicki (1990). Double function of aptychi (Ammonoidea) as jaw elements and opercula.
348:
Tanabe, K., Y. Hikida & Y. Iba (2006). Two coleoid jaws from the Upper
Cretaceous of Hokkaido, Japan.
2034:
1423:
806:
Identification and estimation of size from the beaks of 18 species of cephalopods from the
Pacific Ocean
1727:
1717:
329:
1909:
1904:
758:
The identification of cephalopod "beaks" and the relationship between beak size and total body weight
698:
1045:
Clarke, M.R. & N. MacLeod (1974). Cephalopod remains from a sperm whale caught off Vigo, Spain.
409:
Tanabe, K. & N.H. Landman (2002). Morphological diversity of the jaws of
Cretaceous Ammonoidea.
333:
316:
370:
Zakharov, Y.D. & T.A. Lominadze (1983). New data on the jaw apparatus of fossil cephalopods.
1239:
1086:
Hernańdez-García, V., U. Piatkowski & M.R. Clarke (1998). Development of the darkening of
838:
Jackson, G.D. (1995). The use of beaks as tools for biomass estimation in the deepwater squid
312:
2011:
1525:
1347:
Xavier, J.C., M.R. Clarke, M.C. Magalhães, G. Stowasser, C. Blanco & Y. Cherel (2007).
1999:
1275:(Cephalopoda: Ommastrephidae) throughout the Atlantic Ocean; by body and beak characters.
661:
8:
1983:
1748:
1638:
1623:
1602:
1587:
1455:
896:(Cephalopoda: Onychoteuthidae) from the Falkland Islands region of the Patagonian Shelf.
149:
471:
Tanabe, K. (2012). Comparative morphology of modern and fossil coleoid jaw apparatuses.
1562:
995:
Chen, X., H. Lu, B. Liu, Y. Chen, S. Li & M. Jin (2012). Species identification of
552:
529:
506:
382:
1618:
1852:
1787:
921:
641:
460:
45:
1120:
757:
483:
2007:
1978:
1888:
1777:
1767:
1692:
1687:
1597:
1572:
1405:
1338:
1317:
1300:
1283:
1203:
1149:
1098:
1077:
1053:
1036:
1019:
986:
965:
937:
904:
877:
850:
741:
683:
637:
571:
548:
525:
502:
479:
456:
433:
378:
356:
251:
58:
990:
969:
957:
Aldridge, A.E. (2009). Can beak shape help to research the life history of squid?
563:
Seilacher, A. (1993). Ammonite aptychi; how to transform a jaw into an operculum?
254:
length and total body weight of the original animal as well as the total ingested
1845:
1835:
1830:
1818:
1803:
1743:
1648:
1592:
1510:
1110:"Age determined from the daily deposition of concentric rings on common octopus (
705:
687:
54:
1940:
1762:
1753:
1712:
1577:
1567:
1304:
1102:
892:
Jackson, G.D., N.G. Buxton & M.J.A. George (1997). Beak length analysis of
247:
239:
141:
1342:
1321:
1287:
1207:
1081:
1057:
908:
865:
Jackson, G.D. & J.F. McKinnon (1996). Beak length analysis of arrow squid
2028:
1965:
1515:
1409:
1023:
210:
926:(Cephalopoda: Psychroteuthidae) and its use for size and biomass estimation"
745:
1924:
1823:
1582:
1482:
1309:
Roeleveld, M.A.C. (2000). Giant squid beaks: implications for systematics.
1040:
97:
31:
941:
778:
1878:
1633:
575:
181:
137:
121:
50:
24:
655:"Jumbo squid beaks: Inspiration for design of robust organic composites"
631:
Comparative
Biochemistry and Physiology Part B: Comparative Biochemistry
19:
2015:
1970:
1782:
1772:
1520:
1477:
1446:
881:
854:
227:
116:
35:
1170:. Master's thesis, National Sun Yat-sen University, Kaohsiung, Taiwan.
829:
NOAA Technical Report NMFS 17, NOAA/National Marine
Fisheries Service.
814:
176:
2003:
1945:
1643:
1492:
1487:
1415:
1179:
beaks and rostral length relationships with size and weight of squids
437:
93:
1153:
197:
Side view of the upper beak from the same specimen (2.7 mm URL)
1868:
1840:
1263:
Martínez, P., A. Sanjuan & Á. Guerra (2002). Identification of
1064:
The Mollusca. Volume 12. Paleontology and Neontology of Cephalopods
105:
101:
62:
1174:
1071:
Journal of the Marine Biological Association of the United Kingdom
1047:
Journal of the Marine Biological Association of the United Kingdom
898:
Journal of the Marine Biological Association of the United Kingdom
1758:
1530:
586:
Saunders, W.B., C. Spinosa, C. Teichert & R.C. Banks (1978).
255:
133:
89:
66:
494:
Morton, N. (1981). Aptychi: the myth of the ammonite operculum.
238:, respectively. These are the standard measures of beak size in
1883:
129:
1218:(Cephalopoda: Octopoda) in the Thracian Sea (NE Mediterranean)
869:(Cephalopoda: Ommastrephidae) in southern New Zealand waters.
1628:
85:
473:
Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen
214:
Giant squid beak and associated muscles with hand for scale
78:
1191:
Lalas, C. (2009). Estimates of size for the large octopus
1107:
Hernández-López, J.L. & J.J. Castro-Hernández (2001).
762:
Bulletin of the British Museum (Natural History), Zoology
974:
Bolstad, K.S. (2006). Sexual dimorphism in the beaks of
1030:
Proceedings of the Royal Society B: Biological Sciences
978:
Smith, 1881 (Cephalopoda: Oegopsida: Onychoteuthidae).
1311:
Journal of the Marine Biological Association of the UK
1168:(Cephalopoda: Octopodidae) based on landmarks of beaks
842:(Cephalopoda: Onychoteuthidae) in New Zealand waters.
699:
Organic composite is exceptionally robust: jumbo squid
81:
because they crush bone while most bird beaks do not.
1197:
New Zealand Journal of Marine and Freshwater Research
1011:(Ommastrephidae) using beak morphological variables.
959:
New Zealand Journal of Marine and Freshwater Research
919:
Gröger, J., U. Piatkowski & H. Heinemann (2000).
722:
A Handbook for the Identification of Cephalopod Beaks
328:
Young, R.E., M. Vecchione & K.M. Mangold (2000).
311:
Young, R.E., M. Vecchione & K.M. Mangold (1999).
184:
glasses are recommended to view this image correctly.
652:
Miserez, A., Y. Li, J.H. Waite & F. Zok (2007).
1214:Analysis of beak morphometry of the horned octopus
1090:beaks and its relation to growth and reproduction.
922:"Beak length analysis of the Southern Ocean squid
1394:British Antarctic Survey, Cambridge. 129 pp.
411:Abhandlungen der Geologischen Bundesanstalt, Wien
2026:
1183:Revista de Investigación y Desarrollo Pesquero
1066:. Academic Press, San Diego. pp. 121–131.
1431:
1385:Cephalopod Beak Guide for the Southern Ocean
173:(3.6 mm LRL, 160 mm ML (estimate))
1173:Ivanovic, M.L. & N.E. Brunetti (1997).
716:
714:
395:Bulletin of Gumma Museum of Natural History
1438:
1424:
124:, surrounded by the buccal mass and limbs
1195:from measures of beaks in prey remains.
711:
209:
115:
104:– paired plate-like structures found in
18:
1294:South African Journal of Marine Science
1092:South African Journal of Marine Science
2027:
1445:
1230:Lu, C.C. & R. Ickeringill (2002).
593:and its palaeontological implications"
344:
342:
1419:
1380:Xavier, J.C. & Y. Cherel (2009).
1372:Arquipélago: Life and Marine Sciences
1212:Lefkaditou E. & P. Bekas (2004).
307:
305:
303:
1718:Flotation devices ("secondary fins")
148:) this stiffness gradient spans two
339:
250:. They can be used to estimate the
108:– may also have been jaw elements.
13:
2010:) → Juvenile → Subadult → Adult •
950:
724:. Oxford University Press, Oxford.
553:10.1111/j.1502-3931.1990.tb01365.x
530:10.1111/j.1502-3931.1987.tb02043.x
507:10.1111/j.1502-3931.1981.tb01074.x
383:10.1111/j.1502-3931.1983.tb02000.x
300:
14:
2046:
77:. These beaks are different from
1754:Hepatopancreas (digestive gland)
1677:
1671:
1665:
461:10.1111/j.1502-3931.2009.00203.x
278:
266:
190:
175:
160:
1723:Funnel–mantle locking apparatus
1255:Museum Victoria Science Reports
913:
886:
859:
832:
795:
770:
750:
727:
692:
646:
623:
580:
557:
534:
511:
488:
205:
167:Side view of the lower beak of
53:and surrounded by the muscular
1910:Nuchal folds (occipital folds)
1905:Nuchal crest (occipital crest)
1399:ICES Journal of Marine Science
980:New Zealand Journal of Zoology
465:
442:
419:
403:
387:
364:
322:
111:
1:
991:10.1080/03014223.2006.9518459
970:10.1080/00288330.2009.9626529
589:"The jaw apparatus of Recent
293:
2016:Protoconch (embryonic shell)
1222:Mediterranean Marine Science
688:10.1016/j.actbio.2006.09.004
642:10.1016/0305-0491(81)90071-7
7:
1143:Canadian Journal of Zoology
565:American Journal of Science
484:10.1127/0077-7749/2012/0243
330:Cephalopod Beak Terminology
10:
2051:
1305:10.2989/025776198784126476
1103:10.2989/025776198784126485
1005:Sthenoteuthis oualaniensis
361:10.1666/0022-3360(2006)080
1994:
1958:
1933:
1897:
1861:
1811:
1800:
1736:
1700:
1686:
1663:
1611:
1555:
1548:
1503:
1470:
1463:
1454:
1343:10.1007/s12562-008-0036-2
1322:10.1017/S0025315499001769
1288:10.1007/s00227-002-0796-7
1208:10.1080/00288330909510029
1160:Geomorphometric study of
1082:10.1017/S0025315400041667
1058:10.1017/S0025315400057684
909:10.1017/S0025315400038765
1024:10.3989/scimar.03408.05B
924:Psychroteuthis glacialis
334:Tree of Life Web Project
317:Tree of Life Web Project
16:Body part of cephalopods
746:10.1126/science.1154117
427:Journal of Paleontology
350:Journal of Paleontology
1410:10.1093/icesjms/fsr103
1328:Ommastrephes bartramii
1041:10.1098/rspb.2005.3115
997:Ommastrephes bartramii
215:
128:Composed primarily of
125:
27:
1997:Developmental stages:
1889:Spadix and antispadix
1773:Nephridia ("kidneys")
942:10.1007/s003000050009
756:Clarke, M.R. (1962).
720:Clarke, M.R. (1986).
226:are commonly used in
213:
119:
22:
1269:I. illecebrosus
1088:Todarodes sagittatus
801:Wolff, G.A. (1984).
776:Wolff, G.A. (1981).
576:10.2475/ajs.293.A.20
313:Cephalopoda Glossary
285:Upper rostral length
273:Lower rostral length
236:upper rostral length
232:lower rostral length
170:Chiroteuthis picteti
1984:Squid giant synapse
1158:Hsu, C.-C. (2002).
1035:(1572): 1601–1607.
867:Nototodarus sloanii
740:(5871): 1816–1819.
150:orders of magnitude
2035:Cephalopod zootomy
1915:Occipital membrane
1788:Pericardial glands
1483:Argonautid eggcase
1273:I. argentinus
1193:Macroctopus maorum
976:Moroteuthis ingens
894:Moroteuthis ingens
882:10.1007/BF02329211
855:10.1007/BF00236118
840:Moroteuthis ingens
704:2012-01-06 at the
677:Acta Biomaterialia
218:The abbreviations
216:
126:
49:, situated in the
28:
2022:
2021:
1954:
1953:
1853:Suckers and hooks
1796:
1795:
1778:Nidamental glands
1708:Dermal structures
1661:
1660:
1657:
1656:
1544:
1543:
1332:Fisheries Science
1139:Sepia officinalis
246:is preferred for
185:
132:and cross-linked
2042:
2008:Doratopsis stage
1979:Squid giant axon
1898:Occipital region
1809:
1808:
1749:Ctenidia (gills)
1744:Branchial hearts
1737:Internal anatomy
1701:External anatomy
1698:
1697:
1681:
1675:
1669:
1553:
1552:
1468:
1467:
1461:
1460:
1440:
1433:
1426:
1417:
1416:
1404:(9): 1857–1864.
1393:
1391:
1370:
1368:
1367:
1361:
1355:. Archived from
1354:
1253:
1251:
1250:
1244:
1238:. Archived from
1237:
1216:Eledone cirrhosa
1177:Illex argentinus
1129:Fishery Bulletin
1127:
1125:
1119:. Archived from
1118:
1112:Octopus vulgaris
1009:Illex argentinus
964:(5): 1061–1067.
944:
929:
917:
911:
903:(4): 1235–1238.
890:
884:
863:
857:
836:
830:
828:
826:
825:
819:
813:. Archived from
812:
799:
793:
787:Fishery Bulletin
785:
783:
774:
768:
754:
748:
731:
725:
718:
709:
696:
690:
675:
673:
672:
666:
660:. Archived from
659:
650:
644:
627:
621:
613:
611:
610:
604:
598:. Archived from
597:
584:
578:
561:
555:
538:
532:
515:
509:
492:
486:
469:
463:
446:
440:
438:10.1666/07-029.1
423:
417:
407:
401:
391:
385:
368:
362:
346:
337:
326:
320:
309:
282:
270:
258:of the species.
194:
180:
179:
164:
39:have a two-part
2050:
2049:
2045:
2044:
2043:
2041:
2040:
2039:
2025:
2024:
2023:
2018:
1990:
1975:Nervous system
1950:
1929:
1920:Olfactory organ
1893:
1857:
1802:
1792:
1732:
1691:
1682:
1676:
1670:
1653:
1607:
1540:
1511:Belemnoid guard
1499:
1488:Nautiloid shell
1450:
1444:
1414:
1389:
1381:
1365:
1363:
1359:
1352:
1348:
1265:Illex coindetii
1248:
1246:
1242:
1235:
1231:
1175:Description of
1154:10.1139/z06-049
1123:
1116:
1108:
1013:Scientia Marina
1001:Dosidicus gigas
953:
951:Further reading
948:
947:
920:
918:
914:
891:
887:
864:
860:
837:
833:
823:
821:
817:
810:
802:
800:
796:
781:
777:
775:
771:
755:
751:
732:
728:
719:
712:
706:Wayback Machine
697:
693:
670:
668:
664:
657:
653:
651:
647:
628:
624:
608:
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581:
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388:
369:
365:
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289:
286:
283:
274:
271:
208:
203:
202:
201:
198:
195:
186:
174:
165:
146:Dosidicus gigas
114:
55:head appendages
17:
12:
11:
5:
2048:
2038:
2037:
2020:
2019:
1995:
1992:
1991:
1989:
1988:
1987:
1986:
1981:
1973:
1968:
1966:Chromatophores
1962:
1960:
1956:
1955:
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1951:
1949:
1948:
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1937:
1935:
1931:
1930:
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1901:
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1876:
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1856:
1855:
1850:
1849:
1848:
1843:
1838:
1828:
1827:
1826:
1815:
1813:
1812:Brachial crown
1806:
1798:
1797:
1794:
1793:
1791:
1790:
1785:
1780:
1775:
1770:
1765:
1756:
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1740:
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1733:
1731:
1730:
1725:
1720:
1715:
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1702:
1695:
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1683:
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1662:
1659:
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1600:
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1518:
1513:
1507:
1505:
1501:
1500:
1498:
1497:
1496:
1495:
1485:
1480:
1478:Ammonoid shell
1474:
1472:
1465:
1458:
1452:
1451:
1443:
1442:
1435:
1428:
1420:
1413:
1412:
1395:
1378:
1345:
1337:(2): 317–323.
1324:
1316:(1): 185–187.
1307:
1290:
1282:(1): 131–143.
1277:Marine Biology
1261:
1228:
1210:
1202:(2): 635–642.
1189:
1171:
1156:
1148:(5): 766–770.
1135:
1126:on 2013-05-13.
1105:
1097:(1): 363–373.
1084:
1076:(2): 623–641.
1067:
1060:
1052:(4): 959–968.
1043:
1026:
993:
985:(4): 317–327.
972:
954:
952:
949:
946:
945:
912:
885:
876:(3): 227–230.
858:
831:
794:
769:
767:(10): 419–480.
749:
726:
710:
691:
682:(1): 139–149.
645:
636:(4): 535–546.
622:
579:
556:
533:
524:(3): 231–238.
510:
487:
464:
455:(4): 445–456.
441:
432:(2): 398–408.
418:
402:
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142:Humboldt squid
120:The beak of a
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1531:Spirula shell
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1526:Gladius (pen)
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1516:Cirrate shell
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792:(2): 357–370.
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730:
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708:. Ask Nature.
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667:on 2016-03-05
663:
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620:(1): 129–141.
619:
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615:Palaeontology
605:on 2016-10-05
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1996:
1925:Nuchal organ
1873:
1824:Hectocotylus
1713:Fins (wings)
1619:Body chamber
1583:Periostracum
1401:
1398:
1384:
1374:
1371:
1364:. Retrieved
1357:the original
1334:
1331:
1327:
1313:
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1299:(1): 29–36.
1296:
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1247:. Retrieved
1240:the original
1224:
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1196:
1192:
1185:
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662:the original
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607:. Retrieved
600:the original
590:
582:
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501:(1): 57–61.
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397:
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389:
377:(1): 67–78.
374:
371:
366:
352:
349:
324:
243:
235:
231:
230:to refer to
223:
219:
217:
206:Measurements
168:
145:
138:sperm whales
127:
83:
74:
70:
44:
40:
34:
29:
2012:Egg fossils
1971:Photophores
1934:Other parts
1879:Odontophore
1801:Head &
1634:Phragmocone
849:(1): 9–14.
547:: 325–331.
478:(1): 9–18.
244:hood length
182:3D red cyan
122:giant squid
112:Composition
51:buccal mass
36:cephalopods
25:giant squid
1946:Statocysts
1783:Osphradium
1521:Cuttlebone
1447:Cephalopod
1366:2013-04-07
1249:2013-04-07
1188:: 135–144.
824:2013-02-03
671:2012-01-08
609:2016-09-12
416:: 157–165.
294:References
228:teuthology
94:belemnites
79:bird beaks
2004:Paralarva
1831:Tentacles
1644:Siphuncle
1603:Umbilicus
1588:Sculpture
1493:Orthocone
570:: 20–32.
106:ammonites
90:octopuses
71:mandibles
2029:Category
1869:Aptychus
1841:Dactylus
1612:Internal
1563:Aperture
1556:External
1549:Features
1504:Internal
1471:External
1377:: 41–48.
1166:Cistopus
1114:) beaks"
702:Archived
591:Nautilus
400:: 23–34.
134:proteins
63:mandible
61:(upper)
1959:General
1759:Ink sac
1624:Camerae
1598:Sutures
1449:anatomy
1260:: 1–65.
1162:Octopus
735:Science
542:Lethaia
519:Lethaia
496:Lethaia
450:Lethaia
372:Lethaia
256:biomass
102:Aptychi
67:ventral
46:rostrum
1884:Radula
1836:Carpus
1693:funnel
1690:&
1688:Mantle
1649:Whorls
1573:Callus
1536:Stylet
252:mantle
130:chitin
96:, and
86:squids
59:dorsal
57:. The
32:extant
2000:Spawn
1846:Manus
1804:limbs
1639:Septa
1629:Nacre
1593:Spire
1578:Lirae
1464:Types
1456:Shell
1390:(PDF)
1360:(PDF)
1353:(PDF)
1243:(PDF)
1236:(PDF)
1124:(PDF)
1117:(PDF)
818:(PDF)
811:(PDF)
782:(PDF)
665:(PDF)
658:(PDF)
603:(PDF)
596:(PDF)
43:, or
1941:Eyes
1874:Beak
1819:Arms
1761:and
1728:Tail
1568:Apex
1271:and
1164:and
1007:and
234:and
222:and
75:jaws
41:beak
30:All
1763:ink
1406:doi
1339:doi
1318:doi
1301:doi
1284:doi
1280:141
1204:doi
1150:doi
1099:doi
1078:doi
1054:doi
1037:doi
1033:272
1020:doi
987:doi
966:doi
938:doi
905:doi
878:doi
851:doi
742:doi
738:319
684:doi
638:doi
572:doi
568:293
549:doi
526:doi
503:doi
480:doi
476:266
457:doi
434:doi
379:doi
357:doi
224:URL
220:LRL
73:or
2031::
2014:•
2002:→
1402:68
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1335:75
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1267:,
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934:23
901:77
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790:80
760:.
713:^
634:68
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545:23
522:20
499:14
453:43
430:82
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341:^
332:.
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302:^
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