SCIENTIFIC PORTAL FOR ALL USERS INTERESTED IN GRAPTOLITES AND PTEROBRANCHS
What Are Graptolites?

For many years it was believed that the graptolites are an
extinct group of colonial marine invertebrates that lived
during Paleozoic Era, from the Middle Cambrian to the Middle
Carboniferous, or even Permian. However, Professor Noel P.
Dilly suggested in 1993 that the Recent pterobranch
hemichordate
Cephalodiscus graptolitoides is probably
classifiable as a living graptolite. It is important to note
that in the past, a similar conclusion was made by an eminent
zoologist, Professor Vladimir N. Beklemishev (1890- -1962),
author of the famous "
Principles of Comparative Anatomy of
Invertebrates
". He proposed the class Graptolithoidea... >>>
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Epigraptus sp. (Tuboidea).
Ordovician erratic boulder from Poland.
SEM micrograph of an  incomplete
rhabdosome composed of four
autothecae and thecorhiza.
Back-Scattered Electron Microscopy (BSEM) -
a tool for the study of graptolites

The 1970' saw the aplication of the scanning electron
microscopy (SEM) to graptolite studies. First SEM studies of
graptolitic material were published in Poland by
R. Kozłowski
(1971) and in Great Britain by B. Rickards et. al. (1971). The
conventional mode of SEM use is an invaluable tool...
>>>
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Phylogenetic relationships among benthic
graptolites
Since the classic work of Kozłowski (1949), the phylogenetic
relationships among primitive (i.e. non-graptoloid) benthic
orders of the Graptolithoidea have only rarely been discussed
(e.g. Skevington 1963; Obut 1964;
Kozłowski, 1962, 1966;
Bulman 1968; Mierzejewski 1985; Urbanek 1986); Chapman et
al. 1996). Bulman (1968, 1970) accepted the validity of only
five orders of sessile graptolites (
Camaroidea, Crustoidea,
Stolonoidea, ... >>>
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Professor Roman Kozłowski
(1889-1977)
An eminent graptolite researcher
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Selected web pages of Graptolite Net

 Graptolite researchers

Dietmar Andres - Doctor's disseratation (1976)
Bedrich Boucek (1902-19??)
Oliver M.B. Bulman (1902 -1974)
Alfred Eisenack (1891-1982)
Edvard J.G. Holm (1853-1926)
Hermann Jaeger (1929 - 1992)
Roman Kozlowski (1899 -1977)
-    
Obituary - written by Krystyna Pożaryska
-    
Biographical note - written By M. Gonera and K. Miśkiewicz
-   
Graptolite, pterobranch and related taxa erected by Roman Kozlowski
-    
Taxa named in honour of Roman Kozlowski
Rudolf Ruedemann (1864 -1960)
Carl Wiman (1867-1944)

  Graptolite classification

On the structure and relationships of graptolites (Kozlowski 1966)
Evolution of graptolites   
Graptolite classification by V.N. Beklemishev (1951) - Class Graptolithoidea
Graptolite classification by K. E. Kinman (1994) - Class Pterobranchea

Fine structure of graptolites

Back-scattered electron microscopy (BSEM) - a tool for the study of graptolites
Cortical bandages in Orthograptus gracilis (Roemer)
Cortical bandage-like structure in Dictyonema sp.
Cortical fibrils in Recent species of Rhabdopleura
Ultrastructure of the tuboid graptolite tubotheca
Reticulate fabric in graptolites
Verrucose fabric in graptolites

               Varia

A giant graptolite!
Lower Ordovician graptolites of North China
Large sized graptolites
Collagen in pterobranch skeletons
Dormancy or diapause in graptolites
Discoveries on graptolites by X-ray studies
Evolutionary changes in graptolite larval stages (siculae)
The nature and mode of the graptoloid zooid
Graptolites and early vascular land plants
Soft tissues in Silurian graptolites from Latvia
Graptolite-bearing sites in Poland - concept of protection. In memory of Professor Roman Kozłowski
Helicotubulus Mierzejewski and Kulicki, 2003, an epibiont on graptolites   [Polish]
Clistrocystis Kozlowski, 1959, i.e. cephalopod eggs on Mastigograptus sp.
Graptolites - A Geology Song

GRAPTOLITES sensu lato, i.e. Graptolithina + Pterobranchia = Graptolithoidea

order Rhabdopleuroidea
 Beklemishev, 1951 (= Rhabdopleurida Fowler, 1892)

Stolon system in Rhabdopleura compacta
Classification of the Rhabdopleuroidea   
Discovery of the Rhabdopleuroidea in the Silurian
Ultrastructure of Palaeozoic Rhabdopleuroidea
Rhabdopleuran hemichordates: new fossil forms and review
The discovery of
Rhabdopleura (Pterobranchia) in the Jurassic of Poland
Middle Cambrian pterobranchs and the Question: What is a graptolite?
Diplohydra szaniawskii Mierzejewski & Kulicki, 2002 from the late Permian (Kazanian) of Barents Sea
Rhabdopleura compacta (Recent)
Rhabdopleura graysoni Chapman, Durman, Rickards, 1995 from the Carboniferous
Rhabdoplura kozlowskii Kulicki, 1969 from the Jurassic of Poland
Rhabdopleura normani (Allman) in Dietmar Andres'  drawings
"Rhabdopleura" obuti Durman & Sennikov, 1994 from the Middle Cambrian of Siberia
Rhabdopleura sinica Chapman, Durman & Rickards, 1995 from the Ordovician of China
Rhabdopleurites primaevus from Öland (Lower Ordovician)
Rhabdopleuroides expectatus from the Ordovician of Poland
Rhabdotubus johanssoni Bengtson and Urbanek, 1986 (Middle Cambrian)
Rhabdopleura-like fossil from the Silurian of Gotland

order Cephalodiscoidea Beklemishev, 1951  (= Cephalodiscida Fowler, 1892)

Classificaton of the Cephalodiscoidea
Larval coccon of Cephalodiscus densus Andersson, 1907
Ciliary currents in Cephalodiscus zooid
Cephalodiscus from Bermuda
Cephalodiscus densus Andersson from Ross Sea
Cephalodiscus fumosus John, 1931
Cephalodiscus from  Admiralty Bay, King George Island, West Antarctica
Cephalodiscus graptolitoides Dilly, 1993
Cephalodiscus lutetianus Abrard et al., 1950 from the Eocene of France
Melanostrophus fokini - a Cephalodiscus-like hemichordate
Pterobranchites antiquus Kozlowski, 1949 from the upper Tremadoc of Poland
Pterobranchites antiquus Kozlowski, 1949 - an aberrant graptolite?

Stolonoidea Kozlowski, 1938
General remarks

                     order Crustoidea Kozlowski, 1962

Cellurar tissue in graptoblasts
Classification of the order Crustoidea
Discovery of the Crustoidea in the Silurian
Sicula of a crustoid graptolite?
Bulmanicrusta latialata Kozlowski, 1962 (Lower Ordovician-Upper Silurian)
Holmicrusta Kozlowski, 1962
Graptoblasti or graptoblasti
Hormograptus Öpik, 1930 from the Ordovician of Estonia
New observations on
the fine structure of graptoblasts

                                     order Tuboidea Kozlowski, 1938

Tuboidea - general remarks, list of genera
Tuboid graptolites from erratic boulders of Poland
Siculae of the Tuboidea
Camarotubus graptocamaraeformis Mierzejewski, 2001   [Polish text]
Cyclograptus Spencer, 1884
Epigraptus bidens  Eisenack, 1941 from the Silurian of Baltoscandia
Epigraptus kozlowskii Mierzejewski, 1978
Kozlowskitubus  Mierzejewski, 1978 from the Ordovician and Silurian of Poland

                   order Camaroidea Kozlowski, 1938

Erecticamara maennili Mierzejewski, 2000
Graptocamara Kozlowski, 1949 from the upper Tremadocian of Poland
Xenotheka klinostoma Eisenack, 1937 from the Ordovician of Baltoscandia
Tubicamara coriacea Kozłowski, 1949 from the upper Tremadocian of Poland

                                  order Dendroidea Nicholson, 1872

Graptolodendrum mutabile n.gen. [...] an aberrant dendroid graptolite
On anisograptid affiliation of 'Dictyonema' flabelliforme
Airograptus Ruedemann, 1916
Callograptus indianensis Erdtmann and Adams, 1975 - the latest North American dendroid graptolite
Dendrograptus grabaui Sun, 1935 from the Lower Ordovician of China
Dictyonema - diagrammatic illustrations
Dictyonema altayense Sennikov, 1976 fro the Silurian
Dictyonema crassibasale Gurley from the Silurian of North America
Dictyonema flabelliforne polonicum Tomczyk, 1962 from the Ordovician of Poland
Dictyonema kozlowskii Boucek, 1957 from the Silurian of Bohemia
Graptolodendrum mutabile Kozlowski, 1966 from the Ordovician erratic boulders
Ophigraptus hercyniae Jaeger, 1992 from the Lower/Middle Devonian boundary in Harz, Germany
Pseudodictyonema heyi Chapman, Rickards & Grayson, 1993

                   order Graptoloidea Lapworth, 1875

Virgular apparatus (nematularium) in Cystograptus Hundt, 1942
Affinities of Isograptus, Glossograptus, Cryptograptus, Corynoides, and allied graptolites
Cucullograptus Urbanek, 1954 from the Silurian erratic boulders     
Cystograptus Hundt, 1942 emend. Jones & Rickards, 1967
Diplograptacea - evolution and phylogenetic classification
Didymograptus fearnsidesi Sun, 1935 - Lower Ordovician of North China
Diplograptus penna Hopkinson, 1869
Enigmagraptus Rickards and Wright, 2004 (Pridoli, Silurian)
Lobograptus Urbanek, 1958 from the Upper Silurian
Monoclimacis zawadanensis Teller, 1986 from the Upper Wenlock of Poland
Neocucullograptus Urbanek, 1970 from the Upper Silurian
Neolobograptus Urbanek, 1970 from the Upper Silurian
Saetograptus Pribyl, 1942

                   order Dithecoidea Obut, 1960
General remarks

                        order Mastigograptida Bates and Urbanek, 2002  

General remarks
Mastigograptus Ruedemann, 1908 -  incomplete list of species
Micrograptus Eisenack, 1974 from the Ordovician erratic boulders of Baltoscandia

                               informal group Graptovermida Kozlowski, 1949

Nature of graptovermids

Graptolite-like fossilis

Chaunograptus confertus Boucek from the Silurian of Bohemia
Chaunograptus - an enigmatic encruster from the Ordovician of U.S.A. by J.M.St. John and M.A. Wilson
Diplospirograptus Ruedemann, 1925 - a green alga (Chlorophyceae)
Haplograptus Ruedemann, 1933
Haplograptus canadensis Ruedemann, 1947 from the Ordovician of Canada and China
Medusaegraptus sp. from the Silurian of Gotland, Sweden
Medusaegraptus from the Ordovician of China
Melanostrophus signum Öpik from the Ordovician of Estonia
Inocaulis and the graptolite order Inocaulida
under reconstruction! sorry...