1578:
2050:
139:
1469:
165:
1627:
560:) and different arrangements of the theca, features which are important in the identification of graptolite fossils. Colonies can be classified by their total number of theca rows (biserial colonies have two rows, uniserial have one) and the number of initial stipes per colony (multiramous colonies have many stipes, pauciramous colonies have two or one). Each thecal tube is mostly made up by two series of stacked semicircular half-rings, known as
501:
1316:
568:
1441:, the colonies bear male and female zooids but fertilized eggs are incubated in the female tubarium, and stay there until they become larvae able to swim (after 4–7 days) to settle away to start a new colony. Each larva surrounds itself in a protective cocoon where the metamorphosis to the zooid takes place (7–10 days) and attaches with the posterior part of the body, where the stalk will eventually develop.
613:
1520:, it is likely that asymmetrical states in hemichordates are not under a strong developmental or evolutionary constraint. The origin of this asymmetry, at least for the gonads, is possibly influenced by the direction of the basal coiling in the tubarium, by some intrinsic biological mechanisms in pterobranchs, or solely by environmental factors.
1564:(G). This mutation decreases the efficiency of the autoproteolytic cleavage and therefore, the signalling function of the protein. It is not clear how this unique mechanism occurred in evolution and the effects it has in the group, but, if it has persisted over millions of years, it implies a functional and genetic advantage.
583:
fall into this category, with an overall encrusting colony form combined with erect, vertical theca. Most of the erect, dendritic or bushy/fan-shaped graptolites are classified as dendroids (order
Dendroidea). Their colonies were attached to a hard substrate by their own weight via an attachment
1381:
developed from the cephalic shield or feeding tentacles. In some species, the thecal aperture was probably so restricted that the appendages hypothesis is not feasible. On the other hand, buoyancy is not supported by any extra thecal tissue or gas build-up control mechanism, and active swimming
1385:
There are still many questions regarding graptolite locomotion but all these mechanisms are possible alternatives depending on the species and its habitat. For benthic species, that lived attached to the sediment or any other organism, this was not a problem; the zooids were able to move but
690:
is considered to be a sister subclass of
Graptolithina. One of the main differences between these two groups is that Cephalodiscida species are not colonial organisms. In Cephalodiscida organisms, there is no common canal connecting all zooids. Cephalodiscida zooids have several arms, while
564:(sing: fusellum). The fuselli resemble growth lines when preserved in fossils, and the two stacks meet along a suture with a zig-zag pattern. Fuselli are the major reinforcing component of a tubarium, though they are assisted by one or more additional layers of looser tissue, the cortex.
828:, groups that had great development during the Ordovician. Old taxonomic classifications consider the orders Dendroidea, Tuboidea, Camaroidea, Crustoidea, Stolonoidea, Graptoloidea, and Dithecoidea but new classifications embedded them into Graptoloidea at different taxonomic levels.
1362:, active swimming, and so forth) are not clear yet. One proposal, put forward by Melchin and DeMont (1995), suggested that graptolite movement was analogous to modern free-swimming animals with heavy housing structures. In particular, they compared graptolites to "sea butterflies" (
1694:
distorted rocks. The fossil can also appear stretched or distorted. This is due to the strata that the graptolite is within, being folded and compacted. They may be sometimes difficult to see, but by slanting the specimen to the light they reveal themselves as a shiny marking.
620:
A mature zooid has three important regions, the preoral disc or cephalic shield, the collar and the trunk. In the collar, the mouth and anus (U-shaped digestive system) and arms are found; Graptholitina has a single pair of arms with several paired tentacles. As a
1623:, but generally these rocks were deposited in conditions which were more favorable for bottom-dwelling life, including scavengers, and undoubtedly most graptolite remains deposited here were generally eaten by other animals.
1350:
zone, from inshore to open ocean. Living rhabdopleura have been found in deep waters in several regions of Europe and
America but the distribution might be biased by sampling efforts; colonies are usually found as
342:
share a colony structure of interconnected zooids housed in organic tubes (theca) which have a basic structure of stacked half-rings (fuselli). Most extinct graptolites belong to two major orders: the bush-like
1734:
with species from the boundary
Ordovician-Silurian. Since the group had a wide distribution, fossils are also abundant in several parts of the United States, Canada, Australia, Germany and China, among others.
2418:"New insights on the Silurian graptolite biostratigraphy of the La Chilca Formation, Poblete Norte section, Central Precordillera of San Juan, Argentina: faunal replacement and paleoenvironmental implications"
1786:) influenced changes in the morphology of the colonies and thecae, giving rise to new groups like the planktic Graptoloidea. Later, some of the greatest extinctions that affected the group were the
1338:
Inferring by analogy with modern pterobranchs, they were able to migrate vertically through the water column for feeding efficiency and to avoid predators. With ecological models and studies of the
596:, drifting freely through the water column. They were a successful and prolific group, being the most important and widespread macroplanktonic animals until they died out in the early part of the
2269:
579:
animals, with a colony attached to the sea floor. Several early-diverging families were encrusting organisms, with the colony developing horizontally along a substrate. Extant
1905:
533:. The colony structure has been known from several different names, including coenecium (for living pterobranchs), rhabdosome (for fossil graptolites), and most commonly
633:
of chordates. Proper fossils of the soft parts of graptolites have yet to be found, and it is not known if they had pharyngeal gill slits or not, but based on extant
2372:
Bapst, D., Bullock, P., Melchin, M., Sheets, D. & Mitchell, C. (2012) Graptoloid diversity and disparity became decoupled during the
Ordovician mass extinction.
1612:, and had no scavengers. The dead planktic graptolites, having sunk to the sea floor, would eventually become entombed in the sediment and were thus well preserved.
1935:
1335:
because the most abundant and diverse species were planktonic. Graptolites were most likely suspension feeders and strained the water for food such as plankton.
1846:
1647:
Fossils are often found flattened along the bedding plane of the rocks in which they occur, though may be found in three dimensions when they are infilled by
1947:
1412:
in shallow waters; it is assumed that graptolite fossils had a similar development as their extant representatives. The life cycle comprises two events, the
1532:
as a pterobranch representative. It was found that hedgehog gene in pterobranchs is expressed in a different pattern compared to other hemichordates as the
1390:
but is particularly useful for feeding. Using their arms and tentacles, which are close to the mouth, they filter the water to catch any particles of food.
1983:
1887:
1456:. It is possible that in graptolite fossils the terminal zooid was not permanent because the new zooids formed from the tip of latest one, in other words,
2417:
2012:
1953:
1211:
2032:
2022:
1798:
1154:
1771:
at the end of the
Ordovician eliminated most graptolites except the neograptines. Diversification from the neograptines that survived the Ordovician
1490:. For example, graptolites are used to study asymmetry in hemichordates, especially because their gonads tend to be located randomly on one side. In
1262:
736:
549:. In some colonies, there are two sizes of theca, the larger autotheca and smaller bitheca, and it has been suggested that this difference is due to
2017:
1275:
1224:
1116:
1107:
1050:
2007:
2547:
1235:
1163:
1089:
1003:
695:, and the size of the zooids. In the fossil record, where mostly tubaria (tubes) are preserved, it is complicated to distinguish between groups.
691:
Graptolithina zooids have only one pair of arms. Other differences include the type of early development, the gonads, the presence or absence of
147:
2027:
1917:
1253:
1176:
484:
2250:
961:
467:
cephalopods. By the mid-20th century, graptolites were recognized as a unique group closely related to living pterobranchs in the genera
1448:, and the larvae are ciliated and pigmented, with a deep depression on the ventral side. Astogeny happens when the colony grows through
1965:
1959:
2225:. (2nd Edition). Geological Society of America and University of Kansas Press, Boulder, Colorado and Lawrence, Kansas, XXXII + 163 pp.
2359:
Sato, A., White-Cooper, H., Doggett, K. & Holland, P. 2009. Degenerate evolution of the hedgehog gene in a hemichordate lineage.
1779:
1541:. An important conserved glycine–cysteine–phenylalanine (GCF) motif at the site of autocatalytic cleavage in hh genes, is altered in
362:. Due to their widespread abundance, planktonic lifestyle, and well-traced evolutionary trends, graptoloids in particular are useful
1386:
restricted within the tubarium. Although this zooid movement is possible in both planktic and benthic species, it is limited by the
677:. According to recent phylogenetic studies, rhabdopleurids are placed within the Graptolithina. Nonetheless, they are considered an
2560:
1759:
rocks as they evolved rapidly with time and formed many different distinctive species. Geologists can divide the rocks of the
2237:
Sato, A., Bishop, J. & Holland, P. (2008). Developmental
Biology of Pterobranch Hemichordates: History and Perspectives.
2220:
2565:
2285:
1452:
from the tip of a permanent terminal zooid, behind which the new zooids are budded from the stalk, a type of budding called
2649:
2674:
1416:
and the astogeny, where the main difference is whether the development is happening in the individual organism or in the
2679:
2346:
Sato, A. & Holland, P. (2008). Asymmetry in a
Pterobranch Hemichordate and the Evolution of Left-Right Patterning.
1790:
in the
Ordovician and the Lundgreni in the Silurian, where graptolite populations were dramatically reduced (see also
2659:
2199:
1971:
1831:
1830:
2639:
2606:
1528:(hh), a highly conserved gene implicated in neural developmental patterning, was analyzed in Hemichordates, taking
1358:
Their locomotion was relative to the water mass in which they lived but the exact mechanisms (such as turbulence,
584:
disc. Graptolites with relatively few branches were derived from the dendroid graptolites at the beginning of the
513:
Each graptolite colony originates from an initial individual, called the sicular zooid, from which the subsequent
455:
Graptolite fossils were later referred to a variety of groups, including other branching colonial animals such as
2669:
2654:
2644:
529:. These zooids are housed within an organic structure comprising a series of tubes secreted by the glands on the
2664:
1989:
1512:
being a sister group of hemichordates, and therefore, the asymmetry might be a feature that developed early in
2386:
Urbanek, Adam (1993). "Biotic Crises in the
History of Upper Silurian Graptoloids: A Palaeobiological Model".
1844:
1843:
1839:
1837:
1825:
1824:
1342:, it was observed that, at least for Ordovician species, some groups of species are largely confined to the
1767:
periods into graptolite biozones; these are generally less than one million years in duration. A worldwide
164:
1881:
17:
2101:
Mitchell, C.E., Melchin, M.J., Cameron, C.B. & Maletz, J. (2013) Phylogenetic analysis reveals that
1840:
1821:
541:. The composition of the tubarium is not clearly known, but different authors suggest it is made out of
1524:
556:
Early in the development of a colony, the tubarium splits into a variable number of branches (known as
446:, he included a figure of a "fossil or graptolite of a strange kind" currently thought to be a type of
329:, is known from the Middle Cambrian. Recent analyses have favored the idea that the living pterobranch
2189:
1838:
1836:
1835:
1834:
1829:
1828:
1827:
1826:
1823:
1822:
2611:
2575:
2330:
Röttinger, E. & Lowe, C. (2012) Evolutionary crossroads in developmental biology: hemichordates.
2063:
1842:
1841:
1743:
Graptolite fossils have predictable preservation, widespread distribution, and gradual change over a
648:
281:
625:, graptolites have a simple layer of fibers between the epidermis and the basal lamina, also have a
1833:
1832:
1537:
1460:
budding. These new organisms break a hole in the tubarium wall and start secreting their own tube.
425:
1730:
also yield rich and well-preserved graptolite faunas. A famous graptolite location in Scotland is
1600:
Graptolites are common fossils and have a worldwide distribution. They are most commonly found in
2528:
2479:
1893:
1660:
1636:
320:
2588:
816:
graptolites). The latter is the most diverse, including 5 suborders, where the most assorted is
2490:
575:
The earliest graptolites appeared in the fossil record during the Cambrian, and were generally
2583:
2467:
1474:
1417:
1399:
1382:
requires a lot of energetic waste, which would rather be used for the tubarium construction.
505:
2598:
616:
A hypothetical graptolite zooid inside its theca, reconstructed based on living pterobranchs
463:. The term Graptolithina was established by Bronn in 1849, who considered them to represent
2634:
2552:
1929:
1783:
1589:
1492:
1449:
1408:
576:
344:
335:
represents an extant graptolite which diverged from the rest of the group in the Cambrian.
152:
1500:
is located asymmetrically, and possibly other structures such as the oral lamella and the
8:
2055:
1875:
1806:
1744:
1504:. The significance of these discoveries is to understand the early vertebrate left-right
654:
316:
1805:. This event has been attested in locations such as today's Canada, Libya as well as in
1911:
1683:
by the casual observer, as it has been the case for the first graptolite descriptions.
159:
2085:
Maletz, J. (2014). Hemichordata (Pterobranchia, Enteropneusta) and the fossil record.
2001:
1608:
deposited in relatively deep water that had poor bottom circulation, was deficient in
509:
colony with creeping and erect tubes, showing the zigzag suture where the fuselli meet
2195:
2151:
1715:
1687:
1284:
1198:
941:
550:
2461:
1655:
1577:
400:' rather than true fossils, though later workers supposed them to be related to the
2431:
2395:
2141:
1869:
1719:
1189:
1145:
1136:
923:
914:
879:
253:
1604:
and mudrocks where sea-bed fossils are rare, this type of rock having formed from
800:
Graptolithina includes several minor families as well as two main extinct orders,
1791:
1516:. Since the location of the structures is not strictly established, also in some
1068:
1021:
905:
870:
852:
530:
430:
428:
and incrustations which resembled actual fossils. In 1768, in the 12th volume of
304:
289:
246:
234:
537:(for both). The individual tubes, each occupied by a single zooid, are known as
2422:
2265:
1899:
1482:
In recent years, living graptolites have been used as a hemichordate model for
1445:
1127:
1098:
1039:
1030:
1012:
974:
950:
888:
861:
845:
724:
687:
679:
622:
448:
378:
260:
240:
2473:
2399:
1468:
442:, respectively a possible plant fossil and a possible graptolite. In his 1751
2628:
2513:
2455:
2436:
2155:
2146:
2129:
1752:
1731:
1727:
1723:
1703:
1674:
1517:
1513:
1363:
1293:
992:
983:
667:
601:
475:
421:
405:
393:
363:
325:
296:
292:
206:
138:
87:
2130:"The classification of the Pterobranchia (Cephalodiscida and Graptolithina)"
1977:
1631:
1584:
from the Bendigonian Australian Stage (Lower Ordovician; 477–474 mya) near
1533:
1437:
1244:
1059:
764:
673:
662:
657:, graptolites have generally been thought to be most closely allied to the
635:
588:
period. This latter major group, the graptoloids (order Graptoloidea) were
469:
397:
358:. These orders most likely evolved from encrusting pterobranchs similar to
355:
331:
196:
126:
2539:
1626:
2522:
2223:. Part V. Graptolithina, with sections on Enteropneusta and Pterobranchia
1941:
1669:
1487:
1347:
1332:
658:
630:
308:
62:
31:
2317:, Loydell, D. & Bates, D. (2012) Palaeoecology of the Graptoloidea.
2271:
Life: A Natural History of the First Four Billion Years of Life on Earth
1865:
The following is a selection of graptolite and pterobranch researchers:
1315:
1995:
1923:
1787:
1760:
1756:
1546:
1453:
1378:
1343:
1079:
749:
692:
593:
585:
401:
367:
347:
107:
72:
567:
2314:
1696:
1691:
1616:
1593:
1549:
1509:
1505:
1457:
1367:
1328:
1320:
464:
176:
112:
56:
2484:
500:
2507:
1810:
1802:
1764:
1665:
1605:
1501:
1497:
1483:
1413:
1402:
of Graptholitina has been possible by the discovery of the species
1359:
1352:
813:
639:, it is likely that the grapotlite zooids had the same morphology.
597:
542:
460:
389:
371:
351:
102:
97:
82:
77:
67:
49:
2472:
What are Fossil Graptolites and why are they useful in geology? -
1654:
or some other minerals. They vary in shape, but are most commonly
1772:
1768:
1748:
1640:
1585:
1561:
1424:
805:
589:
518:
456:
312:
117:
92:
661:, a rare group of modern marine animals belonging to the phylum
1651:
1609:
1557:
1387:
1374:
1339:
546:
514:
300:
186:
1820:
1323:, with swimming appendages developed from the cephalic shield.
1085:(biserial graptolites, and also retiolitids and monograptids)
1711:
1707:
1680:
1620:
1601:
1428:
1370:
612:
1702:
A well-known locality for graptolite fossils in Britain is
1648:
1463:
1377:
or swimming via an undulatory movement of paired muscular
629:
that gives rise to several nerve branches, similar to the
1431:
produced by sexual reproduction, which later becomes the
2379:
2251:
Fundamentals of Invertebrate Palaeontology: Macrofossils
1596:. There are two overlapping, three-stiped rhabdosomes.
665:. Comparisons are drawn with the modern hemichordates
483:, as a genus, was officially abandoned in 1954 by the
1690:
on the rock's surface or as light grey clay films in
479:, which had been described in the late 19th century.
2045:
1797:
Graptolite diversity was greatly reduced during the
600:
period. The dendroid graptolites survived until the
388:
meaning "rock", as many graptolite fossils resemble
151:
from the Silurian of South America. Specimen at the
1373:. Under this suggestion, graptolites moved through
27:
Subclass of Pterobranchs in the phylum Hemichordata
2416:Lopez, Fernando Enrique; Kaufmann, Cintia (2023).
2264:
1751:of rocks throughout the world. They are important
820:(biserial graptolites, etc.). This group includes
2087:Palaeogeography, Palaeoclimatology, Palaeoecology
2626:
1327:Graptolites were a major component of the early
935:Lapworth, 1875 in Hopkinson & Lapworth, 1875
416:The name "graptolite" originates from the genus
2374:Proceedings of the National Academy of Sciences
2361:Proceedings of the National Academy of Sciences
1686:Graptolites are normally preserved as a black
2415:
2309:
2307:
2305:
2123:
2121:
2119:
2117:
2115:
517:will develop. They are all interconnected by
2466:Graptolites gallery by Michael P. Klimetz -
2233:
2231:
1679:). Their remains may be mistaken for fossil
831:Taxonomy of Graptolithina by Maletz (2014):
653:Since the 1970s, as a result of advances in
420:("writing on the rocks"), which was used by
124:Survive to the present via the living genus
2302:
2183:
2181:
2179:
2177:
2175:
2173:
2171:
2169:
2167:
2165:
2112:
2097:
2095:
137:
2435:
2411:
2409:
2274:. New York: Alfred A. Knopf. p. 129.
2228:
2145:
1780:Great Ordovician Biodiversification Event
1615:These colonial animals are also found in
1486:studies, as have their sister group, the
2460:Podcast on Graptolites by David Bapst -
2454:Classification of the Graptolithoidea -
2353:
2342:
2340:
2260:
2258:
2081:
2079:
1625:
1576:
1467:
1423:The life cycle begins with a planktonic
1314:
611:
566:
499:
2385:
2162:
2092:
1567:
1464:Graptolites in evolutionary development
14:
2627:
2406:
2187:
2127:
1747:. This allows them to be used to date
1319:Hypothetical zooid inspired by modern
2489:
2488:
2337:
2255:
2221:Treatise on Invertebrate Paleontology
2208:
2076:
1714:, where they occur in rocks from the
2576:da7516e5-8192-4cd8-8f5c-4f1898933218
1775:began around 2 million years later.
571:Diversity of graptolite colony forms
404:; now they are widely recognized as
1699:graptolite fossils are also found.
521:, a true colonial system shared by
495:
452:(a genus of biserial graptolites).
377:The name graptolite comes from the
24:
25:
2691:
2448:
1972:Vladimir Nikolayevich Beklemishev
299:organisms are known chiefly from
2048:
1635:from the middle Silurian of the
1444:The development is indirect and
945:Mu & Lin, 1981 in Lin (1981)
323:). A possible early graptolite,
163:
60:
2366:
2324:
1738:
1572:
1331:ecosystems, especially for the
2278:
2244:
1860:
1552:(T) in the N-terminal, and in
13:
1:
2480:Stephen Hui Geological Museum
2069:
1472:Left and right gonads (g) in
1393:
490:
396:originally regarded them as '
2456:Graptolites and Pterobranchs
1815:
7:
2650:Carboniferous invertebrates
2041:
1990:William Carmichael M'Intosh
1809:of Argentina (then part of
1193:Ĺ torch & Serpagli, 1993
909:Roemer, 1897 in Frech, 1897
701:Phylogeny of Pterobranchia
642:
553:of zooids within a colony.
398:pictures resembling fossils
10:
2696:
2675:Cambrian first appearances
2037:Hermann Jaeger (1929–1992)
1853:Ranges of Graptolite taxa.
1556:there is a replacement of
1310:
646:
411:
284:, members of the subclass
29:
2680:Carboniferous extinctions
2497:
2400:10.1080/10292389309380442
2105:is an extant graptolite.
2064:List of graptolite genera
1818:
978:Cooper & Fortey, 1982
927:Bates & Urbanek, 2002
762:
747:
740:
722:
715:
649:List of graptolite genera
607:
231:
226:
160:Scientific classification
158:
145:
136:
41:
2660:Ordovician invertebrates
2437:10.5027/andgeov50n2-3617
2147:10.3140/bull.geosci.1465
1582:Pendeograptus fruticosus
1538:Saccoglossus kowalevskii
1435:who starts a colony. In
1266:BouÄŤek & MĂĽnch, 1952
30:Not to be confused with
2640:Paleozoic invertebrates
2478:Writing on the rocks -
2191:Graptolite Paleobiology
2134:Bulletin of Geosciences
1906:Sven Leonhard Törnquist
1894:Henry Alleyne Nicholson
1637:Cape Phillips Formation
1632:Cyrtograptus canadensis
1545:by an insertion of the
937:(planktic graptolites)
384:meaning "written", and
338:Fossil graptolites and
2670:Silurian invertebrates
2655:Devonian invertebrates
2645:Cambrian invertebrates
2348:Developmental Dynamics
1936:Alexander Robert Keble
1658:or branching (such as
1644:
1597:
1479:
1420:growth of the colony.
1324:
1288:Elles & Wood, 1908
617:
572:
510:
424:in 1735 for inorganic
2665:Permian invertebrates
2584:Paleobiology Database
2319:Earth-Science Reviews
2290:samnoblemuseum.ou.edu
2188:Maletz, Jörg (2017).
2128:Maletz, Jörg (2014).
1629:
1580:
1475:Rhabdopleura compacta
1471:
1400:developmental biology
1318:
647:Further information:
615:
570:
506:Rhabdopleura compacta
503:
459:("moss animals") and
392:written on the rock.
2363:, 106(18):7491-7494.
2218:Teichert, C. (ed.).
1930:Thomas Sergeant Hall
1656:wiktionary:dendritic
1568:Geological relevance
1493:Rhabdopleura normani
1450:asexual reproduction
933:Order †Graptoloidea
153:Royal Ontario Museum
2376:, 109(9):3428-3433.
2194:. Wiley-Blackwell.
2056:Paleontology portal
1948:William John Harris
1876:Hanns Bruno Geinitz
1807:La Chilca Formation
1745:geologic time scale
686:On the other hand,
655:electron microscopy
317:Lower Carboniferous
34:, a genus of moths.
2388:Historical Biology
2266:Fortey, Richard A.
2214:Bulman, M. (1970)
1984:George Ossian Sars
1912:Sven Axel Tullberg
1672:"-shaped (such as
1645:
1598:
1480:
1366:), small swimming
1325:
898:Order †Dendroidea
618:
573:
511:
2622:
2621:
2491:Taxon identifiers
2350:, 237:3634 –3639)
2013:Denis E. B. Bates
1954:David Evan Thomas
1858:
1857:
1854:
1722:of Scotland, the
1716:Ordovician Period
1398:The study of the
1298:
1289:
1285:Dimorphograptidae
1280:
1267:
1258:
1249:
1240:
1229:
1220:
1212:Neodiplograptinae
1207:
1199:Neodiplograptidae
1194:
1185:
1168:
1159:
1150:
1141:
1132:
1121:
1112:
1103:
1094:
1084:
1073:
1064:
1055:
1044:
1035:
1026:
1017:
1008:
997:
988:
979:
970:
955:
946:
942:Graptodendroidina
936:
928:
919:
910:
901:
893:
884:
875:
866:
857:
840:
808:graptolites) and
798:
797:
793:
792:
784:
783:
775:
774:
551:sexual dimorphism
275:
274:
222:
131:
16:(Redirected from
2687:
2615:
2614:
2602:
2601:
2592:
2591:
2579:
2578:
2569:
2568:
2556:
2555:
2543:
2542:
2533:
2532:
2531:
2518:
2517:
2516:
2486:
2485:
2442:
2441:
2439:
2413:
2404:
2403:
2383:
2377:
2370:
2364:
2357:
2351:
2344:
2335:
2334:, 139:2463-2475.
2328:
2322:
2311:
2300:
2299:
2297:
2296:
2282:
2276:
2275:
2262:
2253:
2248:
2242:
2235:
2226:
2212:
2206:
2205:
2185:
2160:
2159:
2149:
2125:
2110:
2099:
2090:
2083:
2058:
2053:
2052:
2051:
2033:David K. Loydell
2023:Chris B. Cameron
1870:Joachim Barrande
1852:
1816:
1799:Sedgwickii Event
1720:Southern Uplands
1296:
1287:
1278:
1265:
1256:
1247:
1238:
1227:
1214:
1201:
1192:
1190:Normalograptidae
1179:
1166:
1157:
1155:Dicranograptinae
1148:
1146:Dicranograptidae
1139:
1137:Climacograptidae
1130:
1119:
1110:
1101:
1092:
1082:
1071:
1062:
1053:
1042:
1033:
1024:
1015:
1006:
995:
986:
977:
964:
953:
944:
934:
926:
924:Mastigograptidae
917:
915:Acanthograptidae
908:
899:
891:
882:
880:Dithecodendridae
873:
864:
855:
838:
743:
742:
718:
717:
708:
707:
698:
697:
496:Colony structure
354:, free-floating
282:colonial animals
254:Dithecodendridae
220:
168:
167:
141:
123:
122:
59:
53:
45:Temporal range:
39:
38:
21:
2695:
2694:
2690:
2689:
2688:
2686:
2685:
2684:
2625:
2624:
2623:
2618:
2610:
2605:
2597:
2595:
2587:
2582:
2574:
2572:
2564:
2559:
2551:
2546:
2538:
2536:
2527:
2526:
2521:
2512:
2511:
2506:
2493:
2451:
2446:
2445:
2414:
2407:
2384:
2380:
2371:
2367:
2358:
2354:
2345:
2338:
2329:
2325:
2321:, 112(1):23-41.
2312:
2303:
2294:
2292:
2284:
2283:
2279:
2263:
2256:
2249:
2245:
2236:
2229:
2213:
2209:
2202:
2186:
2163:
2126:
2113:
2100:
2093:
2084:
2077:
2072:
2054:
2049:
2047:
2044:
2002:Roman Kozłowski
1888:Frederick M'Coy
1863:
1847:
1845:
1792:Lilliput effect
1741:
1718:. Sites in the
1575:
1570:
1554:S. kowalesvskii
1466:
1396:
1313:
1263:Plectograptinae
1131:Lapworth, 1880e
1069:Glossograptidae
1054:Jaanusson, 1960
1022:Didymograptidae
906:Dendrograptidae
900:Nicholson, 1872
871:Wimanicrustidae
853:Rhabdopleuridae
794:
785:
776:
737:Eugraptolithina
651:
645:
627:collar ganglion
610:
531:cephalic shield
498:
493:
431:Systema Naturae
426:mineralizations
414:
305:Middle Cambrian
303:found from the
280:are a group of
247:Wimanicrustidae
235:Rhabdopleuridae
219:
162:
132:
121:
120:
115:
110:
105:
100:
95:
90:
85:
80:
75:
70:
65:
54:
47:
46:
43:
35:
28:
23:
22:
15:
12:
11:
5:
2693:
2683:
2682:
2677:
2672:
2667:
2662:
2657:
2652:
2647:
2642:
2637:
2620:
2619:
2617:
2616:
2603:
2593:
2580:
2570:
2557:
2544:
2534:
2519:
2503:
2501:
2495:
2494:
2483:
2482:
2476:
2470:
2464:
2458:
2450:
2449:External links
2447:
2444:
2443:
2423:Andean Geology
2405:
2378:
2365:
2352:
2336:
2323:
2301:
2277:
2254:
2243:
2227:
2207:
2200:
2161:
2140:(3): 477–540.
2111:
2091:
2074:
2073:
2071:
2068:
2067:
2066:
2060:
2059:
2043:
2040:
2039:
2038:
2035:
2030:
2025:
2020:
2018:Alfred C. Lenz
2015:
2010:
2005:
1999:
1993:
1987:
1981:
1975:
1969:
1963:
1957:
1951:
1945:
1939:
1933:
1927:
1921:
1915:
1909:
1903:
1900:John Hopkinson
1897:
1891:
1885:
1879:
1873:
1862:
1859:
1856:
1855:
1849:
1848:
1819:
1740:
1737:
1574:
1571:
1569:
1566:
1465:
1462:
1446:lecithotrophic
1395:
1392:
1312:
1309:
1308:
1307:
1306:
1305:
1304:
1303:
1302:
1301:
1300:
1299:
1297:Lapworth, 1873
1290:
1279:Lapworth, 1873
1276:Monograptoidea
1272:
1271:
1270:
1269:
1268:
1259:
1257:Lapworth, 1873
1248:Lapworth, 1873
1239:Lapworth, 1873
1232:
1231:
1230:
1225:Petalolithinae
1221:
1195:
1173:
1172:
1171:
1170:
1169:
1167:Lapworth, 1873
1160:
1158:Lapworth, 1873
1149:Lapworth, 1873
1142:
1133:
1128:Lasiograptidae
1124:
1123:
1122:
1120:Mitchell, 1987
1117:Orthograptinae
1113:
1111:Lapworth, 1873
1108:Diplograptinae
1102:Lapworth, 1873
1099:Diplograptidae
1093:Lapworth, 1880
1076:
1075:
1074:
1072:Lapworth, 1873
1065:
1051:Glossograptina
1047:
1046:
1045:
1040:Tetragraptidae
1036:
1031:Pterograptidae
1027:
1018:
1016:Lapworth, 1873
1013:Dichograptidae
1007:Lapworth, 1873
1000:
999:
998:
989:
980:
975:Sigmagraptidae
958:
957:
956:
951:Anisograptidae
931:
930:
929:
920:
911:
896:
895:
894:
889:Cyclograptidae
885:
876:
867:
862:Cysticamaridae
858:
846:Incertae sedis
796:
795:
791:
790:
787:
786:
782:
781:
778:
777:
773:
772:
769:
768:
761:
758:
757:
754:
753:
746:
741:
739:
733:
732:
729:
728:
725:Rhabdopleurida
721:
716:
714:
706:
703:
702:
688:Cephalodiscida
680:incertae sedis
644:
641:
623:nervous system
609:
606:
497:
494:
492:
489:
449:Climacograptus
436:G. sagittarius
434:, he included
413:
410:
315:) through the
297:filter-feeding
273:
272:
271:
270:
267:
264:
261:Cyclograptidae
257:
250:
243:
241:Cysticamaridae
237:
229:
228:
224:
223:
214:
210:
209:
204:
200:
199:
194:
190:
189:
184:
180:
179:
174:
170:
169:
156:
155:
143:
142:
134:
133:
116:
111:
106:
101:
96:
91:
86:
81:
76:
71:
66:
61:
44:
26:
9:
6:
4:
3:
2:
2692:
2681:
2678:
2676:
2673:
2671:
2668:
2666:
2663:
2661:
2658:
2656:
2653:
2651:
2648:
2646:
2643:
2641:
2638:
2636:
2633:
2632:
2630:
2613:
2608:
2604:
2600:
2594:
2590:
2585:
2581:
2577:
2571:
2567:
2562:
2558:
2554:
2549:
2545:
2541:
2535:
2530:
2529:Graptolithina
2524:
2520:
2515:
2509:
2505:
2504:
2502:
2500:
2499:Graptolithina
2496:
2492:
2487:
2481:
2477:
2475:
2471:
2469:
2465:
2463:
2459:
2457:
2453:
2452:
2438:
2433:
2429:
2425:
2424:
2419:
2412:
2410:
2401:
2397:
2393:
2389:
2382:
2375:
2369:
2362:
2356:
2349:
2343:
2341:
2333:
2327:
2320:
2316:
2310:
2308:
2306:
2291:
2287:
2286:"Graptolites"
2281:
2273:
2272:
2267:
2261:
2259:
2252:
2247:
2241:, 46:587-591.
2240:
2234:
2232:
2224:
2222:
2217:
2211:
2203:
2201:9781118515617
2197:
2193:
2192:
2184:
2182:
2180:
2178:
2176:
2174:
2172:
2170:
2168:
2166:
2157:
2153:
2148:
2143:
2139:
2135:
2131:
2124:
2122:
2120:
2118:
2116:
2108:
2104:
2098:
2096:
2088:
2082:
2080:
2075:
2065:
2062:
2061:
2057:
2046:
2036:
2034:
2031:
2029:
2026:
2024:
2021:
2019:
2016:
2014:
2011:
2009:
2006:
2003:
2000:
1997:
1994:
1991:
1988:
1985:
1982:
1979:
1976:
1973:
1970:
1967:
1964:
1961:
1958:
1955:
1952:
1949:
1946:
1943:
1940:
1937:
1934:
1931:
1928:
1925:
1922:
1919:
1916:
1913:
1910:
1907:
1904:
1901:
1898:
1895:
1892:
1889:
1886:
1883:
1880:
1877:
1874:
1871:
1868:
1867:
1866:
1851:
1850:
1817:
1814:
1812:
1808:
1804:
1800:
1795:
1793:
1789:
1785:
1781:
1776:
1774:
1770:
1766:
1762:
1758:
1754:
1753:index fossils
1750:
1746:
1736:
1733:
1729:
1728:Welsh Borders
1725:
1724:Lake District
1721:
1717:
1713:
1709:
1705:
1704:Abereiddy Bay
1700:
1698:
1693:
1689:
1684:
1682:
1678:
1676:
1675:Didymograptus
1671:
1667:
1663:
1662:
1657:
1653:
1650:
1642:
1638:
1634:
1633:
1628:
1624:
1622:
1618:
1613:
1611:
1607:
1603:
1595:
1591:
1587:
1583:
1579:
1565:
1563:
1559:
1555:
1551:
1548:
1544:
1540:
1539:
1535:
1531:
1527:
1526:
1521:
1519:
1518:enteropneusts
1515:
1514:deuterostomes
1511:
1507:
1503:
1499:
1495:
1494:
1489:
1485:
1477:
1476:
1470:
1461:
1459:
1455:
1451:
1447:
1442:
1440:
1439:
1434:
1433:sicular zooid
1430:
1426:
1421:
1419:
1415:
1411:
1410:
1405:
1401:
1391:
1389:
1383:
1380:
1376:
1372:
1369:
1365:
1361:
1356:
1354:
1349:
1345:
1341:
1336:
1334:
1330:
1322:
1317:
1295:
1294:Monograptidae
1291:
1286:
1282:
1281:
1277:
1274:Superfamily â€
1273:
1264:
1260:
1255:
1251:
1250:
1246:
1242:
1241:
1237:
1236:Retiolitoidea
1234:Superfamily â€
1233:
1226:
1222:
1218:
1213:
1209:
1208:
1205:
1200:
1196:
1191:
1187:
1186:
1183:
1178:
1174:
1165:
1164:Nemagraptinae
1161:
1156:
1152:
1151:
1147:
1143:
1138:
1134:
1129:
1125:
1118:
1114:
1109:
1105:
1104:
1100:
1096:
1095:
1091:
1090:Diplograptina
1087:
1086:
1081:
1077:
1070:
1066:
1061:
1057:
1056:
1052:
1048:
1041:
1037:
1032:
1028:
1023:
1019:
1014:
1010:
1009:
1005:
1004:Dichograptina
1001:
994:
993:Abrograptidae
990:
985:
984:Sinograptidae
981:
976:
972:
971:
968:
963:
959:
952:
948:
947:
943:
939:
938:
932:
925:
921:
916:
912:
907:
903:
902:
897:
890:
886:
881:
877:
872:
868:
863:
859:
854:
850:
849:
848:
847:
843:
842:
841:
837:
836:Graptolithina
832:
829:
827:
823:
822:Diplograptids
819:
815:
811:
807:
803:
789:
788:
780:
779:
771:
770:
767:
766:
760:
759:
756:
755:
752:
751:
745:
744:
738:
735:
734:
731:
730:
727:
726:
720:
719:
713:
712:Graptolithina
710:
709:
705:
704:
700:
699:
696:
694:
689:
684:
682:
681:
676:
675:
670:
669:
668:Cephalodiscus
664:
660:
656:
650:
640:
638:
637:
632:
628:
624:
614:
605:
603:
602:Carboniferous
599:
595:
591:
587:
582:
578:
569:
565:
563:
559:
554:
552:
548:
544:
540:
536:
532:
528:
527:Cephalodiscus
524:
520:
516:
508:
507:
502:
488:
486:
482:
478:
477:
476:Cephalodiscus
472:
471:
466:
462:
458:
453:
451:
450:
445:
441:
437:
433:
432:
427:
423:
419:
409:
407:
406:hemichordates
403:
399:
395:
391:
387:
383:
380:
375:
373:
369:
365:
364:index fossils
361:
357:
353:
349:
346:
341:
336:
334:
333:
328:
327:
326:Chaunograptus
322:
321:Mississippian
318:
314:
310:
306:
302:
298:
294:
293:Pterobranchia
291:
287:
286:Graptolithina
283:
279:
269:†Graptoloidea
268:
265:
262:
258:
255:
251:
248:
244:
242:
238:
236:
233:
232:
230:
225:
218:
217:Graptolithina
215:
212:
211:
208:
207:Pterobranchia
205:
202:
201:
198:
195:
192:
191:
188:
185:
182:
181:
178:
175:
172:
171:
166:
161:
157:
154:
150:
149:
148:Cryptograptus
144:
140:
135:
129:
128:
119:
114:
109:
104:
99:
94:
89:
84:
79:
74:
69:
64:
58:
51:
40:
37:
33:
19:
2498:
2427:
2421:
2391:
2387:
2381:
2373:
2368:
2360:
2355:
2347:
2331:
2326:
2318:
2313:Cooper, R.,
2293:. Retrieved
2289:
2280:
2270:
2246:
2238:
2219:
2215:
2210:
2190:
2137:
2133:
2106:
2103:Rhabdopleura
2102:
2089:, 398:16-27.
2086:
2028:Adam Urbanek
1978:Michael Sars
1918:Gerhard Holm
1864:
1796:
1777:
1742:
1739:Stratigraphy
1701:
1692:tectonically
1685:
1673:
1659:
1646:
1630:
1614:
1599:
1581:
1573:Preservation
1553:
1542:
1536:
1534:enteropneust
1530:Rhabdopleura
1529:
1523:
1522:
1491:
1481:
1473:
1443:
1438:Rhabdopleura
1436:
1432:
1422:
1407:
1403:
1397:
1384:
1364:Thecostomata
1357:
1337:
1326:
1254:Retiolitinae
1245:Retiolitidae
1228:Bulman, 1955
1216:
1203:
1181:
1175:Infraorder â€
1088:Infraorder â€
1063:Harris, 1933
1060:Isograptidae
966:
954:Bulman, 1950
918:Bulman, 1938
892:Bulman, 1938
874:Bulman, 1970
865:Bulman, 1955
856:Harmer, 1905
844:
835:
833:
830:
825:
821:
817:
810:Graptoloidea
809:
801:
799:
765:Graptoloidea
763:
748:
723:
711:
685:
678:
674:Rhabdopleura
672:
666:
663:Hemichordata
659:pterobranchs
652:
636:Rhabdopleura
634:
626:
619:
581:Rhabdopleura
580:
574:
561:
557:
555:
538:
534:
526:
523:Rhabdopleura
522:
512:
504:
481:Graptolithus
480:
474:
470:Rhabdopleura
468:
454:
447:
444:SkĂĄnska Resa
443:
439:
435:
429:
418:Graptolithus
417:
415:
385:
381:
376:
360:Rhabdopleura
359:
356:Graptoloidea
340:Rhabdopleura
339:
337:
332:Rhabdopleura
330:
324:
285:
277:
276:
216:
197:Hemichordata
146:
127:Rhabdopleura
125:
36:
2635:Graptolites
2523:Wikispecies
2468:Graptolites
2332:Development
2109:, 46:34–56.
2008:Jörg Maletz
2004:(1889–1977)
1998:(1916–2005)
1992:(1838–1931)
1986:(1837–1927)
1980:(1805–1869)
1974:(1890–1962)
1968:(1928–2013)
1962:(1917–1987)
1956:(1902–1978)
1950:(1886–1957)
1944:(1885–1957)
1942:Noel Benson
1938:(1884–1963)
1932:(1858–1915)
1926:(1867–1944)
1920:(1853–1926)
1914:(1852–1886)
1908:(1840–1920)
1902:(1844–1919)
1896:(1844–1899)
1890:(1817–1899)
1884:(1811–1898)
1878:(1814–1900)
1872:(1799–1883)
1861:Researchers
1755:for dating
1688:carbon film
1670:tuning fork
1668:-like, or "
1543:R. compacta
1488:acorn worms
1404:R. compacta
1355:of shells.
1348:mesopelagic
1333:zooplankton
1261:Subfamily â€
1252:Subfamily â€
1223:Subfamily â€
1210:Subfamily â€
1177:Neograptina
1162:Subfamily â€
1153:Subfamily â€
1140:Frech, 1897
1115:Subfamily â€
1106:Subfamily â€
1083:Frech, 1897
1043:Frech, 1897
839:Bronn, 1849
826:Neograptids
631:neural tube
440:G. scalaris
390:hieroglyphs
309:Miaolingian
288:within the
278:Graptolites
266:†Dendroidea
221:Bronn, 1849
55:510–0
42:Graptolites
32:Graptolitha
18:Graptolites
2629:Categories
2462:Palaeocast
2430:(2): 201.
2295:2018-12-28
2070:References
1996:Nancy Kirk
1924:Carl Wiman
1882:James Hall
1788:Hirnantian
1773:glaciation
1761:Ordovician
1757:Palaeozoic
1732:Dob's Linn
1677:murchisoni
1661:Dictyonema
1617:limestones
1547:amino acid
1454:monopodial
1409:R. normani
1394:Life cycle
1379:appendages
1344:epipelagic
1080:Axonophora
1078:Suborder â€
1049:Suborder â€
1002:Suborder â€
962:Sinograpta
960:Suborder â€
940:Suborder â€
883:Obut, 1964
818:Axonophora
802:Dendroidea
750:Dendroidea
693:gill slits
594:planktonic
586:Ordovician
491:Morphology
465:orthoconic
461:hydrozoans
402:hydrozoans
368:Ordovician
352:planktonic
348:Dendroidea
227:Subgroups
213:Subclass:
52:to Recent.
2394:: 29–50.
2315:Rigby, S.
2156:1214-1119
1697:Pyritized
1594:Australia
1550:threonine
1510:chordates
1506:asymmetry
1458:sympodial
1353:epibionts
1329:Paleozoic
1321:pteropods
834:Subclass
457:bryozoans
374:periods.
183:Kingdom:
177:Eukaryota
2537:BioLib:
2508:Wikidata
2268:(1998).
2042:See also
1966:Li Jijin
1960:Mu Enzhi
1811:Gondwana
1803:Aeronian
1765:Silurian
1666:sawblade
1606:sediment
1590:Victoria
1560:(S) for
1525:Hedgehog
1502:gonopore
1498:testicle
1484:Evo-Devo
1414:ontogeny
1368:pteropod
1360:buoyancy
1292:Family â€
1283:Family â€
1243:Family â€
1215:Melchin
1202:Melchin
1197:Family â€
1188:Family â€
1144:Family â€
1135:Family â€
1126:Family â€
1097:Family â€
1067:Family â€
1058:Family â€
1038:Family â€
1034:Mu, 1950
1029:Family â€
1025:Mu, 1950
1020:Family â€
1011:Family â€
996:Mu, 1958
991:Family â€
987:Mu, 1957
982:Family â€
973:Family â€
949:Family â€
922:Family â€
913:Family â€
904:Family â€
887:Family â€
878:Family â€
869:Family â€
860:Family â€
814:planktic
683:family.
643:Taxonomy
604:period.
598:Devonian
543:collagen
535:tubarium
525:but not
422:Linnaeus
394:Linnaeus
372:Silurian
366:for the
350:and the
295:. These
193:Phylum:
187:Animalia
173:Domain:
50:Cambrian
2514:Q332403
2474:Youtube
2239:Genesis
2107:Lethaia
1801:in the
1769:ice age
1641:Nunavut
1586:Bendigo
1562:glycine
1508:due to
1425:planula
1418:modular
1311:Ecology
1180:Ĺ torch
965:Maletz
851:Family
806:benthic
590:pelagic
577:sessile
562:fuselli
519:stolons
412:History
382:graptos
345:sessile
313:Wuliuan
301:fossils
203:Class:
2612:380352
2599:423307
2596:uBio:
2589:319610
2573:NZOR:
2566:993363
2540:165767
2198:
2154:
1749:strata
1681:plants
1652:pyrite
1621:cherts
1610:oxygen
1602:shales
1558:serine
1496:, the
1427:-like
1388:stolon
1375:rowing
1371:snails
1340:facies
1219:, 2011
1217:et al.
1206:, 2011
1204:et al.
1184:, 2011
1182:et al.
969:, 2009
967:et al.
608:Zooids
558:stipes
547:chitin
515:zooids
386:lithos
2607:WoRMS
2548:IRMNG
1712:Wales
1708:Dyfed
1429:larva
539:theca
379:Greek
290:class
2561:ITIS
2553:1175
2196:ISBN
2152:ISSN
1784:GOBE
1778:The
1763:and
1726:and
1649:iron
1619:and
1406:and
1346:and
824:and
671:and
592:and
485:ICZN
473:and
438:and
370:and
63:Preęž’
48:Mid
2432:doi
2396:doi
2142:doi
1813:).
1794:).
1664:),
545:or
2631::
2609::
2586::
2563::
2550::
2525::
2510::
2428:50
2426:.
2420:.
2408:^
2390:.
2339:^
2304:^
2288:.
2257:^
2230:^
2216:In
2164:^
2150:.
2138:89
2136:.
2132:.
2114:^
2094:^
2078:^
1710:,
1706:,
1643:.
1639:,
1592:,
1588:,
487:.
408:.
311:,
113:Pg
57:Ma
2440:.
2434::
2402:.
2398::
2392:7
2298:.
2204:.
2158:.
2144::
1782:(
1478:.
812:(
804:(
319:(
307:(
263:?
259:â€
256:?
252:â€
249:?
245:â€
239:â€
130:.
118:N
108:K
103:J
98:T
93:P
88:C
83:D
78:S
73:O
68:ęž’
20:)
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