Graptolites

BGS fossils and geological time

Didymograptus 'bifidus' from the Ordovician of South Wales. About 2 cm long.
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Didymograptus ‘bifidus’ from the Ordovician of South Wales. About 2 cm long. BGS ©UKRI. All rights reserved.

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Fossil graptolites are thin, often shiny, markings on rock surfaces that look like pencil marks, and their name comes from the Greek for ‘writing in the rocks’. Graptolites are the remains of intricate colonies, some of which accommodated up to 5000 individual animals; these individuals lived in a skeleton of collagen, similar to the material from which our finger nails are made. We focus on the two main groups of graptolites:

  • dendroids
  • planktonic graptoloids

They lived between the Cambrian and Carboniferous periods, about 520 to 350 million years ago.

The animal

Didymograptus, a tuning folk graptoloid, showing the main parts of the rhabdosome.
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Didymograptus, a tuning folk graptoloid, showing the main parts of the rhabdosome. BGS ©UKRI. All rights reserved.

Graptolites were colonial animals that lived in an interconnected system of tubes. From an initial ’embryonic’ cone-like tube (the sicula), subsequent tubes (thecae) are arranged in branches (stipes) to make up the whole colony (rhabdosome).

A close-up of the thecae of Climacograptus partly cut away to show the internal structure of the interconnecting tubes and one of the zooids. The thread-like, central nema (or virgula), which may protrude some distance beyond the stipes, may have been used for attachment when juvenile, for strengthening or to attach a floatation device or vane.
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A close-up of the thecae of Climacograptus partly cut away to show the internal structure of the interconnecting tubes and one of the zooids. The thread-like, central nema (or virgula), which may protrude some distance beyond the stipes, may have been used for attachment when juvenile, for strengthening or to attach a floatation device or vane. BGS ©UKRI. All rights reserved.

Environment

The earliest graptolites lived on the sea bed, see below for full description.
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The earliest graptolites lived on the sea bed, see below for full description. BGS ©UKRI. All rights reserved.

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The earliest graptolites lived on the sea bed, attached to boulders (a) or forming upright cones rooted into soft mud (b). This is still the lifestyle of the living relatives of graptolites. At the beginning of the Ordovician period graptolites became free floating. They were amongst the first animals to colonise the open sea and were able to exploit enormous untapped reserves of food (single celled organisms) in the upper layers of the oceans. Different species evolved rapidly in order to exploit these food reserves and in response to the new challenge of a floating life.

Some graptolite forms; including those fixed to the sea bed (a, b) and floating species (c–l). Graptolites were common where food was abundant, especially in upwelling zones where deep water is forced to the surface with its load of nutrients. Some were deep-water specialists (c) whilst others were opportunists seeking out temporary supplies of food (e, d). Using the distribution of fossil graptolites, we can begin to reconstruct the oceans in which they lived.

Feeding: graptolites were most common and diverse where major upwelling currents brought nutrients from deep water into the shallows, in areas such as the tropics and at the edge of the continental shelf. Some colonies evolved into enormous harvesting arrays, capable of living to a significant age, perhaps up to 20 years (c). Others evolved into slim, short colonies (d, e) or into gently curved forms which rotated through the water as they fed (f, g).

Floating: a floating life is a challenge, and graptolites responded to this by evolving a range of hydrodynamic strategies. Long nemas evolved to retard sinking (e, h) and hooks, spines and net forms appeared which would have had a high drag so that they moved slowly (i, j, k). The zooids may have helped the colony to move and possibly secreted gas or low-density fat to help the colony to rise through the water.

Graptolites: modern relatives

Cephalodiscus gracilis, a living pterobranch thought to be related to graptolites. Two zooids have extended their feathery filtering nets to feed.
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Cephalodiscus gracilis, a living pterobranch thought to be related to graptolites. Two zooids have extended their feathery filtering nets to feed. BGS ©UKRI. All rights reserved.

Although graptolites are now extinct, living marine animals called pterobranchs appear to be closely related. Pterobranchs do not grow their tube-like skeleton in the same way as we grow our bones or an oyster makes its shell. Rather, the pterobranch zooids actively construct them, much as a spider weaves its web or termites build their nest. Graptolites probably did the same: the criss-crossing bandages in the outer layer of a theca, which look rather like the bandages wrapping an Egyptian mummy, were apparently ‘trowelled’ on by a secretory organ akin to that of pterobranchs.

The geologists’ tool

Graptolites are excellent geological time-keepers, for they can be used to date the rocks in which they are found. They evolved quickly and assumed a wealth of easily recognisable shapes. Many of these evolutionary steps, which can be traced around much of the world, define periods of time. Some of these time-slices are only a few hundred thousand years long, which to a geologist, amounts to pinpoint accuracy given that all this took place hundreds of millions of years ago.

Dendroids

Graptoloids

Graptoloids were all planktonic, had only one kind of theca and may have been hermaphrodites. Dichograptids flourished in the early Ordovician. Their stipes were few and arranged in various ways: e.g. stretched out sideways (horizontal), hanging down (pendant) or turned upwards (reclined). Tetragraptus had four horizontal or pendant stipes (a). Didymograptus had two pendant stipes (b). Isograptus had reclined stipes (and a long nema extending above the sicula) (c).
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Graptoloids were all planktonic, had only one kind of theca and may have been hermaphrodites. Dichograptids flourished in the early Ordovician. Their stipes were few and arranged in various ways: e.g. stretched out sideways (horizontal), hanging down (pendant) or turned upwards (reclined). Tetragraptus had four horizontal or pendant stipes (a). Didymograptus had two pendant stipes (b). Isograptus had reclined stipes (and a long nema extending above the sicula) (c). © The British and Irish Graptolite Group (BIG G).

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Diplograptids

Diplograptids evolved from dichograptids in the Ordovician. Orthograptus and Dicranograptus are from the Ordovician of southern Scotland.

a) The stipes of Orthograptuswere fused ‘back-to-back’ (i.e. scandent). b) Some later types of diplograptids became partly ‘unzipped’, like the Y-shaped Dicranograptus (the biserial part is 5mm long). c) Spirograptus turriculatuswas coiled or spring-like d) Rastriteshad a curved stipe with long thecae up to 3mm long
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a) The stipes of Orthograptus were fused ‘back-to-back’ (i.e. scandent).

b) Some later types of diplograptids became partly ‘unzipped’, like the Y-shaped Dicranograptus (the biserial part is 5mm long).

c) Spirograptus turriculatus was coiled or spring-like

d) Rastrites had a curved stipe with long thecae up to 3mm long

© The British and Irish Graptolite Group (BIG G).

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One-stiped monograptids appeared at the beginning of the Silurian and evolved rapidly into many coiled, curved and straight species, with elongate, hooked or lobed thecae. Spirograptus turriculatus, Rastrites and Monograptus lobiferus are from the Silurian (Llandovery).

Graptolites through time

The history of graptolites (dendroids and graptoloids) and their diversity related to global events. The geological position of illustrated specimens are shown in italics.
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The history of graptolites (dendroids and graptoloids) and their diversity related to global events. The geological position of illustrated specimens are shown in italics. BGS ©UKRI. All rights reserved.

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Graptolites in Britain

Palaeozoic rocks of the UK and some good places to look for graptolites (see black spots). The colour code for geological periods is the same as for the history diagram.
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Palaeozoic rocks of the UK and some good places to look for graptolites (see black spots). The colour code for geological periods is the same as for the history diagram. BGS ©UKRI. All rights reserved.

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On death graptolites fell to the sea floor, became entombed into the sea floor muds and, if conditions were right, preserved as fossils for us to find today. They are found only in Palaeozoic rocks such as those in Scotland, Wales and north-western England. The oldest dendroids occur in Middle Cambrian rocks, but they can be found in rocks as young as the Carboniferous. Planktonic graptolites are particularly common in Ordovician and Silurian shales and mudstones.

Fun facts

Graptolites are thought to have been hermaphroditic — with both male and female sexual organs. To avoid self-fertilisation, or inbreeding, zooids could have had the potential to be temporarily male, female or neuter. Zooids maturing at the near end of a colony might initially have been males whilst those occupying the larger, distal thecae may have manifested the female phase. But nobody knows for sure!

3D fossil models

Orthograptus truncatus var. corrisensis. (SM A 10007 – Holotype). Ashgill Series (Ordovician Period) (443.8 – 449 Ma B.P.) [Closest ICS interval: Late Ordovician Epoch]. See 3D fossils online. GB3D Type Fossils.
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Orthograptus truncatus var. corrisensis Davies. (SM A 10007 – Holotype). Ashgill Series (Ordovician Period) (443.8 – 449 Ma B.P.) [Closest ICS interval: Late Ordovician Epoch]. See 3D fossils online. GB3D Type Fossils. BGS ©UKRI. All rights Reserved.

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Many of the fossils in the BGS palaeontology collections are available to view and download as 3D models. To view this fossil, or others like it, in 3D visit GB3D Type Fossils.

Reference

Wilkinson, I P, Rigby, S, and Zalasiewicz, J A. 2001. Graptolites: fossil focus. Nottingham, British Geological Survey.

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