Fragments of Stolonodendrum, from Bulman (1970).

Belongs within: Hemichordata.
Contains: Graptoblasti, Dendroidea, Graptoloida, Wimanicrustidae.

The Graptolithina, graptolites, are a group of colonial organisms known from the Middle Cambrian to the Carboniferous that produced linear, branched or encrusting colonies.

Characters (from Bulman 1970): Colonial organisms secreting a sclerotised exoskeleton with characteristic growth bands (fuselli) and growth lines. Thecae usually arranged in single or double row along branches (stipes) of colony (rhabdosome), rarely in irregular aggregates; thecae may be polymorphic, may be related to internal sclerotised stolon system. Rhabdosomes originating by single bud from initial zooid housed in conical sicula, producing simple, branched or rarely encrusting colonies. Sessile or pelagic.

    |--+--Dendroidea B70
    |  `--Graptoloida B70
    |--Mastigograptida [Mastigograptidae] BU02
    |    |--Micrograptus Eisenack 1974 BU02
    |    `--Mastigograptus Ruedemann 1908 BU02
    |         |--*M. tenuiramosus (Walcott 1879) [=Dendrograptus tenuiramosus] BU02
    |         `--M. gracillimus (Lesquereux 1878) BU02
    |--Crustoidea B70
    |    |--Hormograptus Öpik 1930 [=Thallograptus Öpik 1928 non Ruedemann 1925; Hormograptidae] B70
    |    |    `--*H. sphaericola (Öpik 1928) [=*Thallograptus sphaericola] B70
    |    `--Wimanicrustidae B70
    `--Stolonoidea B70
         |--Stolonofolliculis Zessin & Puttkamer 1994 [Stolonofolliculidae] MU04
         |    `--*S. signum (Öpik 1930) (n. d.) [=Melanostrophus signum] MU04
         `--Stolonodendrum Kozłowski 1948 [Stolonodendridae] B70
              |--*S. uniramosum Kozłowski 1948 B70
              `--S. parasiticum Kozłowski 1948 K48

Graptolithina incertae sedis:
  Protistograptus McLearn 1915 (n. d.) B70
  Haplograptus Ruedemann 1933 B70
    `--*H. wisconsinensis Ruedemann 1933 B70
  Graptovermis Kozłowski 1949 [Graptovermida] MU04
    |--*G. spiralis Kozłowski 1948 B70
    `--G. intestinalis Kozłowski 1948 K48
  Graptoblasti B70
  Aellograptus Obut 1964 B70
    `--*A. savitskyi Obut 1964 B70
  Archaeocryptolaria Chapman 1919 B70
    `--*A. skeatsi Chapman 1919 B70
  Ascograptus Ruedemann 1925 B70
    `--*A. similis Ruedemann 1925 B70
  Cactograptus Ruedemann 1908 B70
    `--*C. crassus Ruedemann 1908 B70
  Ceramograptus Hudson 1915 B70
    `--*C. ruedemanni Hudson 1915 B70
  Coelograptus Ruedemann 1947 B70
    `--*C. problematicus (Spencer 1878) [=Inocaulis problematica] B70
  Crinocaulis Obut 1960 B70
    `--*C. flosculus Obut 1960 B70
  Diplospirograptus Ruedemann 1925 B70
    `--*D. goldringae Ruedemann 1925 B70
  Dithecodendrum Obut 1964 B70
    `--*D. sibericum Obut 1964 B70
  Estoniocaulis Obut & Rytsk 1958 B70
    `--*E. jaervensis (Rosenstein in Obut & Rytsk 1958) [=Inocaulis jaervensis] B70
  Leveillites Foerste 1923 B70
    `--*L. hartnageli Foerste 1923 B70
  Medusaegraptus Ruedemann 1925 B70
    `--*M. mirabilis Ruedemann 1925 B70
  Palmatophycus Bouček 1941 B70
    `--*P. kettneri Bouček 1957 B70
  Polygonograptus Bouček 1957 B70
    `--*P. sokolowi (Obut 1953) [=Palaeodictyota sokolowi] B70
  Protohalecium Chapman & Thomas 1936 B70
    `--*P. hallianum Chapman & Thomas 1936 B70
  Rhadinograptus Obut 1960 B70
    `--*R. jurgensonae Obut 1960 B70
  Ruedemannograptus Termier & Termier 1948 [=Streptograptus Ruedemann 1947 non Yin 1937] B70
    `--*R. tenuis (Ruedemann 1947) [=*Streptograptus tenuis] B70
  Siberiodendrum Obut 1964 B70
    `--*S. robustum Obut 1964 B70
  Siberiograptus Obut 1964 B70
    `--*S. kotujensis Obut 1964 B70
  Sphenoecium Chapman & Thomas 1936 [=Sphenothallus Chapman 1917 non Hall 1848] B70
    `--*S. filicoides (Chapman 1917) [=Sphenothallus filicoides] B70
  Birastrites Geinitz 1866 (n. d.) B70
  Buthograptus Hall 1861 (n. d.) B70
  Cameragraptus Hundt 1951 (n. d.) B70
  Cardograptus Hundt 1965 (n. d.) B70
  Conograptus Ruedemann 1947 (n. d.) B70
  Ctenograptus Nicholson 1876 (n. d.) B70
  Cystoturriculograptus Hundt 1952 (n. d.) B70
  Demicystograptus Hundt 1942 (n. d.) B70
  Dibranchiograptus Hundt 1949 (n. d.) B70
  Didymograptoides Hundt 1951 (n. d.) B70
  Eiseligraptus Hundt 1965 (n. d.) B70
  Geminograptus Hundt 1951 (n. d.) B70
  Labrumograptus Hundt 1952 (n. d.) B70
  Limpidograptus Khaletskaya 1962 (n. d.) B70
  Nereitograptus Hundt 1951 (n. d.) B70
  Paradimorphograptus Hundt 1951 (n. d.) B70
  Phycograptus Gurley 1896 (n. d.) B70
  Planktograptus Yakovlev 1933 (n. d.) B70
  Procrytograptus Poulsen 1943 (n. d.) B70
  Protograptus Matthew 1886 (n. d.) B70
  Spinosidiplograptus Hundt 1951 (n. d.) B70
  Stelechograptus Ruedemann 1947 (n. d.) B70
  Strophograptus Ruedemann 1904 (n. d.) B70
  Thecocystograptus Hundt 1947 (n. d.) B70
  Thuringiagraptus Hundt 1935 (n. d.) B70
  ‘Triplograptus’ Hundt 1965 non Richter 1871 (n. d.) B70
  Undograptus Hundt 1949 (n. d.) B70
  Triramograptus atavus FEZ05
  Parakidograptus acuminatus P98
  Amansites Brongniart 1849 (n. d.) H62
    `--*A. dentatus (Brongniart 1828) (n. d.) [=Fucoides dentatus] H62
  Stimulograptus sedgwicki K-D04
  Lituigraptus convolutus K-D04

*Type species of generic name indicated


[BU02] Bates, D. E. B., & A. Urbanek. 2002. The ultrastructure, development, and systematic position of the graptolite genus Mastigograptus. Acta Palaeontologica Polonica 47 (3): 445–458.

[B70] Bulman, O. M. B. 1970. Graptolithina with sections on Enteropneusta and Pterobranchia. In: Teichert, C. (ed.) Treatise on Invertebrate Paleontology pt V 2nd ed. pp. V1–V149. The Geological Society of America, Inc.: Boulder (Colorado), and the University of Kansas: Lawrence (Kansas).

[FEZ05] Feng, H., B.-D. Erdtmann & Y. Zhang. 2005. Taxonomy and phylogeny of Staurograptus Emmons from the base of the Ordovician. Palaeontology 48 (5): 1007–1019.

[H62] Häntzschel, W. 1962. Trace fossils and problematica. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt W. Miscellanea: Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils and Problematica pp. W177–W245. Geological Society of America, and University of Kansas Press.

[K-D04] Kozłowska-Dawidziuk, A. 2004. Evolution of retiolitid graptolites—a synopsis. Acta Palaeontologica Polonica 49 (4): 505–518.

[K48] Kozłowski, R. 1948. Les graptolithes et quelques nouveaux groupes d’animaux du Tremadoc de la Pologne. Palaeontologica Polonica 3: i–xii, 1–235.

[MU04] Mierzejewski, P., & A. Urbanek. 2004. The morphology and fine structure of the Ordovician Cephalodiscus-like genus Melanostrophus. Acta Palaeontologica Polonica 49 (4): 519–528.

[P98] Prothero, D. R. 1998. Bringing Fossils to Life: An introduction to paleobiology. WCB McGraw-Hill: Boston.

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