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- Herausgeber: John Wiley & Sons
- Kategorie: Wissenschaft und neue Technologien
- Serie: TOPA Topics in Paleobiology
- Sprache: Englisch
The graptolites constitute one of the geologically most useful taxonomic groups of fossils for dating rock successions, understanding paleobiogeography and reconstructing plate tectonic configurations in the Lower Palaeozoic. Graptolites were largely planktic, marine organisms, and as one of the first groups that explored the expanses of the world’s oceans are vital for understanding Palaeozoic ecology. They are the best and often the only fossil group for dating Lower Palaeozoic rock successions precisely. Thousands of taxa have been described from all over the planet and are used for a wide variety of geological and palaeontological (biological) research topics. The recent recognition of the modern pterobranch Rhabdopleura as a living benthic graptolite enables a much better understanding and interpretation of the fossil Graptolithina.
In the decades since the latest edition of the Graptolite Treatise, the enormous increase of knowledge on this group of organisms has never been synthesised in a compelling and coherent way, and information is scattered in scientific publications and difficult to sort through. This volume provides an up-to-date insight into research on graptolites. Such research has advanced considerably with the use of new methods of investigation and documentation. SEM investigation and research on ultrastructure of the tubaria has made it possible to compare extant and extinct taxa in much more detail. Cladistic interpretation of graptolite taxonomy and evolution has advanced the understanding of this group of organisms considerably in the last two decades, and has highlighted their importance in our understanding of evolutionary processes. This book will show graptolites, including their modern, living relatives, in a quite new and fascinating light, and will demonstrate the impact that the group has had on the evolution of the modern marine ecosystem.
This book is aimed not only at earth scientists but also at biologists, ecologists and oceanographers. It is a readable and comprehensible volume for students at the MSc level, while remaining accessible to undergraduates and non-specialists seeking up-to-date information about this fascinating topic in palaeobiology.
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Veröffentlichungsjahr: 2017
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Table of Contents
Cover
Title Page
List of contributors
Preface
Acknowledgments
1 Graptolites
Biology
Evolution
Stratigraphy
Ecology
Paleogeography
Colony Shapes
History of Research
Outlook
2 Biological Affinities
Graptolites as Organisms
Hemichordata
Pterobranchia
Communality and Coloniality
Ontogeny and Astogeny
Cephalodiscida
Graptolithina
Outlook
3 Construction of Graptolite Tubaria
Naming the Tubarium Features
Construction Material
Tubarium Design
Thecal Tubes
Colony Growth
Extrathecal Developments
Outlook
4 Paleoecology of the Pterobranchia
Mode of Life
Benthic Graptolites
Planktic Graptolites
Synrhabdosomes
Locomotion
Graptolites and Sediments
Patterns of Occurrence
Population Structure
The Graptoloid Habitat
Spatial Distribution
Depth Distribution
Biogeography
Historical Biogeography
Graptolite Life History
Graptolites and the Food Chain
Feeding Style
The Diet
Parasitism
Tubarium Repair
Outlook
5 Graptolites as Rock Components
Graptolite Taphonomy and Preservation Potential
Death on the Sea Floor
In Situ
Death Assemblages
Transport of Graptolite Tubaria
Burial and Preservation
Diagenesis
Metamorphism and Organic Maturation
Mineral Replacement
Tectonic Deformation
Weathering
Outlook
6 Graptolites and Stratigraphy
Biostratigraphy and Graptolites
Types of Graptolite Zones
Graptolites and Chronostratigraphy
Graptolites and Absolute Ages
Graptolite Biozonations
Graphic Correlation
Graptolites and Exploration
Oil and Gas
Uranium Exploration
Outlook
7 Taxonomy and Evolution
Graptolites and Taxonomy
Nomenclature
Monophyly in Graptolite Taxonomy
Graptolite Cladistics
Relationships of the Major Graptolite Groups
Extinction Events and Radiations
Evolutionary Lineages
Convergent Evolution
Outlook
8 Bound to the Sea Floor
“Rooting” the Graptolite Colony
The Graptolithina
Family Rhabdopleuridae
Camaroids and Crustoids
The Cyclograptidae
The Dithecodendridae
The Dendroidea
The Mastigograptidae
The Dendrograptidae
The Acanthograptidae
The Extinction
Algae or Graptolites?
Outlook
9 The Planktic Revolution
Why Move into the Water Column?
What has Changed in Colony Development?
Attachment and the Free Nema
Increase in Symmetry
The “Dendroid” Bithecae
Increasing Diversity and Disparity
The Graptoloidea
Anisograptidae as Inventors
Proximal Development
Evolutionary Changes and Biostratigraphy
Turnover in the Late Tremadocian
Outlook
10 Early Ordovician Diversity Burst
The Great Ordovician Biodiversification Event
Change in the Colony Design
Suborder Sinograpta
Family Sigmagraptidae
Family Abrograptidae
Thecal Complexity
Suborder Dichograptina
Family Tetragraptidae
Family Didymograptidae
Family Pterograptidae
Symmetry and the Glossograptina
Family Isograptidae
The Manubrium
Scandency and Biserial Tubaria
Family Glossograptidae
Outlook
11 The Biserial Graptolites
The Axonophora Concept
Early Biserial Axonophorans
An Axis for Support
The Axonophoran Sicula
The Virgella
Proximal Development Types
The Thecal Styles
The Median Septum
The Diplograptina
Family Diplograptidae
Orthograptinae and the Antivirgellar Spine
Family Lasiograptidae
Family Climacograptidae
Family Dicranograptidae
The Neograptina
Outlook
12 The Retiolitid Graptolites
Tubarium Reduction?
Retiolitid Origins
Ancora Umbrella and Ancora Sleeve
Reticulum and Clathrium
Early Ancora Sleeve Development
The Retiolitinae
The Plectograptinae
Appendix and the Retiolitid Extinction
Outlook
13 The Monograptids
Monograptid Construction
Family Dimorphograptidae
Monograptid Thecal Styles
Tubarium Shapes
Cladia
Llandovery Diversification
Rastritid Monograptids
Streptograptids
Spirograptus, Cyrtograptus
and their Relatives
Pristiograptus
Clade
Aftermath of the
Lundgreni
Extinction
Cucullograptinae and Neocucullograptinae
Linograptinae
Kozlowskii
Event
Final Extinction
Outlook
14 Collection, Preparation and Illustration of Graptolites
Collecting Graptolites
Physical Preparation
Chemical Preparation
Methods of Illustration
Permanent Storage
Outlook
15 History of Graptolite Research
The First Collected Graptolite?
Foundation of Graptolite Research
The British Dominance
Early Graptolite Research in Scandinavia
Graptolites from “Down Under”
The 20th Century Graptolites
Graptolites in China and Russia
The Pterobranch Connection
Graptolite Reconstructions through Time
The Kirk Hypothesis – A Controversy
Outlook
References
Index
Supplemental Images
End User License Agreement
List of Tables
Chapter 07
Table 7.1 The taxonomy of the Pterobranchia (from Maletz 2014a).
List of Illustrations
Chapter 01
Figure 1.1 Images of well‐preserved graptolites, showing the complexity and beauty of their construction. (A)
Archiclimacograptus
sp., obverse view, SEM photo, Table Head Group, western Newfoundland, Canada. (B)
Dicranograptus irregularis
, obverse view, relief specimen, Scania, Sweden. (C)
Spirograptus turriculatus
(Barrande, 1850), proximal end, SEM photo, Kallholn Shale, Llandovery, Dalarna, Sweden. Scale indicated by 1 mm long bar in each photo.
Figure 1.2 Pterobranchs and their housing constructions (tubaria). Extant
Cephalodiscus
(A, B, F, G) and
Rhabdopleura
(C–E) to show the zooids (A–D) and their tubaria. Illustrations after Sars 1874 (C, D), Lester 1985 (B), Dilly et al. 1986 (A), Emig 1977 (F), and M’Intosh 1887 (G). Illustrations not to scale.
Figure 1.3 Large‐scale evolutionary changes in graptoloids. (A) Encrusting benthic graptolite,
Rhabdopleura normani
Allman, 1869. (B) Benthic dendroid,
Dictyonema cavernosum
Wiman, 1896. (C) Multiramous
Goniograptus thureaui
(M’Coy, 1876). (D) Two‐stiped, reclined
Isograptus mobergi
Maletz, 2011d. (E) Biserial graptolite,
Archiclimacograptus
sp. (F) Straight monograptid
Monograptus priodon
(Bronn, 1834). (G) Coiled monograptid
Demirastrites
sp. (H) Secondarily multiramous
Abiesgraptus
sp. Graptolite illustrations not to scale.
Figure 1.4 Graptolite biostratigraphy of the Floian, Lower Ordovician of Scandinavia, based on the evolution of
Baltograptus
species with subhorizontal (
B. geometricus
) to pendent (
B. minutus
) habit, as an example of biostratigraphical subdivision of an evolutionary lineage. Graptolite illustrations not to scale.
Figure 1.5 Organisms associated with graptolites. (A) Silicified ostracod carapace. (B–C) Phosphatic brachiopods. (D–E) Conodont elements. (F)
Obruchevella
, silicified microbial organism. (G) Radiolarian. (H) Chitinozoan. (I) Phyllocarid fragment in shale. Specimens from the Middle Ordovician of eastern Canada.
Figure 1.6 Paleogeography of graptolites. (A) Biogeography of Laurentia showing oceanic and shallow water, epicratonic (Laurentian) biofacies (based on Maletz et al. 2005a, Fig. 4). (B) Biogeographical distribution of the Floian dichograptid
Baltograptus
in high latitudes (from Goldman et al. 2013). The areas in grey show the distribution of
Baltograptus
species.
Figure 1.7 Colony variation in Ordovician and Silurian graptoloids. (A)
Cyrtograptus multibrachiatus
Bjerreskov, 1992, Arctic Canada. (B)
Expansograptus abditus
Williams and Stevens, 1988, Cow Head Group, western Newfoundland, Canada. (C)
Neogothograptus balticus
(Eisenack, 1951), glacial boulder, Northern Germany. (D)
Streptograptus galeus
Lenz & Kozłowska, 2006. (E)
Paradelograptus norvegicus
(Monsen, 1937), Fezouata Biota, Morocco. (F)
Amplexograptus maxwelli
(Decker, 1935). (G)
Dicranograptus nicholsoni
Hopkinson, 1870, Viola Limestone. (H)
Monograptus priodon
(Bronn, 1835), Arctic Canada. Illustrations not to scale.
Figure 1.8 (A, B) Reconstruction and interpretation of indeterminable biserial graptolite fragment from Richter (1853, pl. 12, Figs 12, 13). (C, D) Excellent illustration by Georg Liljevall (ca. 1895) for Gerhard Holm,
Monograptus priodon
(Bronn, 1835), proximal end in obverse (C) and reverse (D) views (published in Bulman 1932a, pl. 1, Figs 10, 11). (E) Erroneous reconstruction of biserial graptolite with complex float structure by Franziska Zörner‐Bertina (ca. 1950) for Rudolf Hundt, unpublished (original at Naturkundemuseum Gera, Germany; provided by Frank Hrouda, Gera).
Chapter 02
Figure 2.1 Adults of Enteropneusta and Pterobranchia. (A) The enteropneust
Saccoglossus
sp. (based on Bulman 1970a). (B, C)
Cephalodiscus
sp. (B) Part of colony with zooids sitting at the openings (apertures) of the tubarium. (C) Single zooid at the apertural spine of a tubarium, showing the main body parts.
Figure 2.2 The phylogenetic relationships of the Hemichordata (based on Maletz 2014a).
Figure 2.3 Ontogeny of
Saccoglossus horsti
(after Burdon‐Jones 1952). (A) Burrowing larva. (B) Newly hatched planktic larva. (C–D) Settling larva in different views. (E) Post‐settlement creeping larva. (F–G) Burrowing larva in ventral (F) and lateral (G) views. (H)
Planctosphaera pelagica
(after van der Horst 1936). Illustrations not to scale.
Figure 2.4
Mazoglossus ramsdelli
Bardack, 1997. Counterparts, Mason Creek Biota. The white spots on the specimen may be an effect of recent weathering.
Figure 2.5 Enteropneust burrows and traces (after Hyman 1959, Fig. 50). (A)
Saccoglossus inhacensis
, spiral burrow. (B–C)
Balanoglossus clavigerus
, funnel opening and fecal coil as surface indication of burrow (B) and scheme of U‐shaped burrow (C). Illustrations not to scale.
Figure 2.6 Tubaria of extant and fossil Pterobranchia. (A)
Rhabdopleura compacta
Hincks, 1880, extant, part of small colony showing fuselli, SEM photo. (B)
Streptograptus sartorius
(Törnquist, 1881), fossil, infrared photo showing fuselli. (C)
Pseudoglyptograptus vas
(Bulman & Rickards 1968), fossil specimen in reverse view in shale, showing the median septum and the interthecal connections (common canal) of the two stipes of an axonophoran graptolite, vague striation indicates the fusellar construction. Scale bar indicates 0.1 mm in (A–B) and 1 mm in (C).
Figure 2.7 (A)
Rhabdopleura normani
, part of colony, showing the stolon (black line) at the base of the thecal tubes and the zooids retracted into the basal parts of their tubes (based on Schepotieff 1907a, pl. 22). (B)
Jenkinsograptus
spinulosus
, proximal end with sicula and a number of successively formed tubes; the stolon is not preserved Illustrations not to scale.
Figure 2.8 Ontogeny and astogeny in
Rhabdopleura compacta
. (A) Swimming larva. (B) Settled larva in cocoon. (C–D) Developing zooid. (E) Juvenile colony with dome and thecal tube of sicular zooid. (F) Larger colony with five thecal tubes. Illustrations not to scale.
Figure 2.9 Monopodial and sympodial budding. (A) Monopodial growth, fragment of
Rhabdopleura normani
colony with growing end and permanent terminal zooid (based on Lankester 1884, pl. 39). (B) Sympodial growth, reconstruction of dichograptid branch with zooids; each zooid is the terminal one at one time (zooids reconstructed from Schepotieff 1906, pl. 25).
Figure 2.10 The zooids of
Atubaria
and
Cephalodiscus
. (A)
Cephalodiscus
sp., pseudocolony of mature and associated juveniles (B)
Cephalodiscus dodecalophus
M’Intosh, showing internal anatomy (from Schepotieff 1907b, pl. 48). (C)
Atubaria heteroplopha
Sato, 1936 Illustrations not to scale.
Figure 2.11 Tubarium shapes in extant Cephalodiscidae. (A)
Cephalodiscus nigrescens
(B)
Cephalodiscus calciformis
(C)
Cephalodiscus
(
Orthoecus
)
rarus
(from Andersson 1907, pl. 2, Fig. 6). (D)
Cephalodiscus
(
Idiothecia
)
levinseni
Harmer, 1905 (after Harmer 1905, pl. 2, Fig. 11). (E)
Cephalodiscus sibogae
(from Ridewood 1907, Fig. 2). Illustrations not to scale.
Figure 2.12 (A) Graptolite zooids in a fragment of a biserial tubarium, surface showing cortical bandages (reconstruction based on Crowther & Rickards 1977). (B–C)
Rhabdopleura normani
zooids (based on Schepotieff 1906, pl. 25). Body of the zooids is ca. 1 mm long without arms.
Chapter 03
Figure 3.1 (A) Uniserial tubarium of
Neocolonograptus lochkovensis
(Přibyl, 1940) showing fusellar construction (based on Urbanek 1997a, Fig. 49). (B) Fuselli and cortex (based on Kozłowski 1938, Fig. 2). (C) Biserial tubarium of
Diplacanthograptus spiniferus
(Ruedemann, 1912), showing increasing width of fuselli towards thecal apertures (based on Mitchell 1987). Illustrations not to scale.
Figure 3.2 Ultrastructure of graptolite tubarium. (A)
Desmograptus micronematodes
(Spencer, 1884), SEM, fuselli, with thicker ectocortex below, and thinner endocortex above. (B)
Gothograptus nassa
(Wiman, 1895), SEM photo, bandages on genicular hood, with pustules. (C)
Dendrograptus
? sp., TEM section through two fuselli and bandages of the ectocortex. (D) Stacked fuselli with endo‐ and ectocortex (based on Bates & Kirk 1986a, Fig. 1).
Figure 3.3 (A)
Rhabdopleura normani
Allman, 1869, large encrusting colony, showing highly irregular placing of thecal tubes and branching points (from Ridewood 1907, Fig. 6). (B)
Clonograptus flexilis
(Hall, 1865), planktic graptoloid with dichotomous branching (after Lindholm & Maletz 1989, Fig. 2 F). (C)
Dendrograptus fruticosus
Hall, 1865; note the massive stem and slender stipes showing dichotomous branching (from Hall 1865, pl. 19, Fig. 8). Illustrations not to scale.
Figure 3.4 Cladial branching in
Cyrtograptus
. (A)
Cyrtograptus perneri
Bouček, 1933, SEM photo. (B–F) Several stages in the development of a cladium, showing the secondary nema as the leading rod of the cladial stipe (based on Bulman 1970a, Fig. 65). (G)
Cyrtograptus radians
Törnquist, 1887, Thuringia, specimen with numerous cladia and fragment of
Monoclimacis
sp. on top. Illustrations not to scale.
Figure 3.5 (A)
Palaeodictyota
sp. showing the typical anastomosing stipes of a conical, benthic graptolite colony (B)
Dictyonema
(
Dictyonema
)
elongatum
Bouček, 1957, fragment showing fine dissepiments in a benthic dendroid graptolite (based on Bouček 1957, Fig. 20C). Reconstructions not to scale.
Figure 3.6 (A) Pendent to scandent colony shapes (B)
Isograptus victoriae
, reclined two‐stiped colony. (C)
Pseudamplexograptus distichus
, scandent, dipleural colony. (D)
Paraglossograptus proteus
, scandent, monopleural colony. Shading is used in (B–D) to show the disposition of the two stipes in these taxa. Illustrations not to scale.
Figure 3.7 (A) Terminology of dichograptid thecae. (B)
Expansograptus holmi
(Törnquist, 1901), obverse view, showing thecal style of relief specimen, common canal and interthecal septae visible on the stipe. Scale indicated by 1 mm long bar in (B).
Figure 3.8 Thecal development. (A) Thecal diagram showing triad budding and the presence of a dicalycal theca (based on Cooper & Fortey 1983). (B) Stipe fragment with bithecae, dorsal view, showing lateral origin of thecae. (C) Stipe fragment with bithecae in lateral view. (D) Stipe fragment with plaited overlap, but lacking bithecae. (E) Dichograptid stipe fragment without bithecae and dorsal thecal origins. Illustrations not to scale.
Figure 3.9
Archiclimacograptus
sp., showing the thecal notation for biserial axonophorans.
Figure 3.10 Thecal styles in planktic graptolites. (A)
Monograptus priodon
Bronn, 1835, hooked thecae. (B)
Pseudostreptograptus williamsi
Loydell, 1991b, thecae with cupulae (arrows) and branched lateral apertural spines. (C)
Proteograptus opimus
(Lenz & Melchin, 1991), hooked thecae showing biform development, with distal thecae bearing paired lateral lobes. (D)
Paraorthograptus pacificus
(Ruedemann, 1947), geniculate thecae with spines. (E)
Neodiplograptus tcherskyi tcherskyi
(Obut & Sobolevskaya, 1967), biform thecae in biserial graptoloid, proximally thecae are geniculate, distally straight, outwards inclined. Various magnifications.
Figure 3.11 Thecal style. (A) Isolated thecal apertures in
Psigraptus arcticus
Jackson, 1967, isolated specimen, Erdaopu, Jilin, China (photo provided by Zhang Yuandong). (B)
Sinograptus typicalis
Mu, 1957, showing prothecal and metathecal folding. Scale indicated by 1 mm long bar in each photo.
Figure 3.12 Ontogeny and early astogeny. (A) Dome of
Rhabdopleura compacta
Hincks, 1880 (based on Stebbing 1970b, Fig. 3). (B–D)
Rectograptus gracilis
(Roemer, 1861), sicular development, showing the individual parts (based on Kraft 1926). (E)
Monograptus
sp., sicula with first theca (based on Kraft 1926). (F–G)
Rectograptus gracilis
(Roemer, 1861), proximal ends in reverse (F) and obverse (G) view (based on Kraft 1926). Illustrations not to scale.
Figure 3.13 The astogeny of
Dicranograptus nicholsoni
Hopkinson, 1870, Viola Limestone, JM26. (A) Sicula with indication of th1
1
origin. (B) Downward growth of th1
1
visible. (C) Th1
1
nearly complete. (D) Sicula with th1
1
before secretion of apertural spine. (E) sicula with incomplete first two thecae. (F) Proximal end with th1
1
and th1
2
complete. (G) Specimen with six thecal pairs and indication of distally diverging stipes. All specimens in reverse view, except for (D), which is in obverse view. Scale indicated by 0.5 mm long bar in each photo.
Figure 3.14 Nematularia. (A)
Pseudoclimacograptus scharenbergi
(from Bulman 1947, pl. 9). (B–C)
Archiclimacograptus decoratus
(Harris & Thomas, 1935), distal end of flattened colony with fragmented nematularium, Table Head Group, western Newfoundland. (D) Isolated complete nematularium of
Archiclimacograptus decoratus
(Harris & Thomas, 1935) showing growth lines. (E)
Cystograptus vesiculosus
(Nicholson), section through part of nematularium (based on Urbanek, Koren & Mierzejewski 1982, Fig. 5). Scale indicated by 1 mm long bar in each photo.
Figure 3.15 Proximal webs and membranes. (A)
Loganograptus kjerulfi
Herrmann, 1882, Christiania, Norway (from Herrmann 1882, pl. 2, Fig. 12), showing a large proximal membrane. (B) ?
Loganograptus kjerulfi
Herrmann, 1882, juvenile with initial membrane growth (from Herrmann 1882, pl. 1, Fig. 1). (C–D)
Didymograptus murchisoni
Beck in Murchison, 1839, inside views of specimen with extensive proximal membranes (identified as
Didymograptus pakrianus
in Jaanusson 1960, pl. 1, Figs 5–6). (E)
Cyrtograptus
sp., Cape Phillips Formation, Arctic Canada, dorsal view of proximal membrane. Illustrations not to scale.
Chapter 04
Figure 4.1 Lifestyle of graptolites. (A)
Dictyonema cavernosum
Wiman, 1896b, benthic, erect colony (based on Wiman 1896b, pl. 1) (from Erdtmann 1986a). (B)
Rhabdinopora flabelliformis
(Eichwald, 1840), planktic graptoloid. (C)
Tetragraptus
(?)
norvegicus
(Monsen, 1937), planktic graptoloid with free nema, chemically isolated specimen (from Maletz 2011d, Fig. 1A). (D)
Cephalodiscus
(
Orthoecus
)
rarus
Andersson, 1907, specimen attached to a piece of rock (from Andersson 1907, pl. 2, Fig. 7). Illustrations not to scale.
Figure 4.2 Early graptolites and their associates. (A)
Sphenoecium mesocambricus
(Öpik, 1933), holotype, showing poorly preserved phosphatic brachiopod in centre. (B)
Goniagnostus nathorsti
(Brögger, 1878), cranidium. (C)
Goniagnostus nathorsti
(Brögger, 1878), pygidium. (D)
Leiopyge calva
, cranidium. All specimens from Krekling, Norway, Middle Cambrian
Goniagnostus nathorsti
Biozone.
Figure 4.3 Misleading concepts of epiplanktic graptolites. (A) Colony of
Orthograptus quadrimucronatus
(Hall, 1865), showing synrhabdosome with floating vesicle (“pneumatophore”), gonangia and tubaria (“stipes” or “rhabdosomes”). (B)
Pterograptus elegans
Holm, 1881, attached to seaweed with long nema. Illustration modified from Maletz (2014a).
Figure 4.4 (A–B) Zooid model of Melchin and DeMont (1995). (C)
Limacina antarctica
, an Antarctic pteropod (after Woodward 1854, pl. 14, Fig. 41).
Figure 4.5 Vertical movement of graptolite colonies (after Rigby & Rickards 1989, Fig. 4).
Figure 4.6 The graptolite biofacies: schematic distribution of lithofacies and graptolite biofacies in a sedimentary basin bounded by a carbonate platform (after Podhalańska 2013).
Figure 4.7 (A) Length‐frequency relationship of
Orthograptus quadrimucronatus micracanthus
of sample SM X23260 (after Rigby & Dilly 1993, Fig. 14). (B) Survivorship curve for
Orthograptus quadrimucronatus micracanthus
(after Rigby 1993a, Fig. 9).
Figure 4.8 Graptolite paleobiogeography of the Middle Ordovician. Isograptid biofacies in black; shallow‐water graptolite biofacies in grey (based on Fortey & Cocks 1986, Fig. 3).
Figure 4.9 Transgressive succession in the Lower Ordovician of North Wales, showing early appearance of
Azygograptus
species in the most shallow facies (after Beckly & Maletz 1991, Fig. 1).
Figure 4.10 Depth distribution of graptolites (based on Egenhoff & Maletz 2007, Fig. 11).
Figure 4.11 Diabasbrottet section, Västergötland, Sweden, GSSP section for the base of the Floian Stage, Ordovician System, graptolite biostratigraphy and recognition of maximum flooding surfaces (mfs) (based on Egenhoff & Maletz 2007, Fig. 3).
Figure 4.12 Paleocontinent reconstruction map for the Dapingian (Middle Ordovician) showing graptolite localities and distribution of graptolite biofacies. Black dots are localities that have graptolite successions during the Dapingian. Grey shaded ovals indicate oceanic (isograptid) biofacies and striped ovals indicate shelf (didymograptid) biofacies (based on Goldman et al. 2013).
Figure 4.13 Example of historical biogeography and origination of graptoloid clades (based on Zhang & Chen 2007, Fig. 3).
Figure 4.14 Area cladogram illustrating the geographical regions occupied at the nodes (see Goldman et al. 2013 for details).
Figure 4.15 Graptoloids showing growth limitations. (A, B)
Archiclimacograptus decoratus
(Harris & Thomas, 1935). (A) NMVP 31932, two specimens with heart‐shaped nematularium (B)
Rectograptus intermedius
(Elles & Wood, 1907), Viola Limestone, Oklahoma, showing nema inside and growing end. (C)
Corynites divnoviensis
(Kozłowski, 1953), glacial boulder, Germany (coll. Kühne) with reduced and coiled third theca. (D)
Holoretiolites erraticus
(Eisenack, 1951) with appendix, Sellin, Rügen, glacial boulder. (E)
Neogothograptus eximinassa
Maletz, 2008, Spandau, near Berlin, Germany, glacial boulder, showing appendix with strong reticulum. Scale indicated by 1 mm long bar in each photo.
Figure 4.16 Feeding. (A) Ciliary currents in zooid of
Cephalodiscus
, rejection current in centre of arm basket, food particles moving downwards towards mouth (B) Postulated water flow in dendroid graptolite colony (after Kirk 1990, Fig. 1a). (C) Modelled water flow in dendroid colony according to Melchin and Doucet (1996).
Figure 4.17 Parasites in
Cephalodiscus
. (A)
Cephalodiscus gilchristi
Ridewood, 1907, showing the position of the parasite
Zanclopus gilchristi
Calman 1908 in its stomach. (B–E)
Zanclopus gilchristi
, several specimens. (B–C) Female in two views. (E) Male in 5th copepodid stage, appendages omitted. (D) Earliest known larval stage (after Calman 1908, pl. 18, 19). Illustrations not to scale.
Figure 4.18 Possible epibionts on graptolites. (A)
Helicotubulus dextrogyra
(Kozłowski, 1967), holotype on fragment of
Mastigograptus
sp. (after Kozłowski 1967, Fig. 7). (B)
Clistrocystis graptolithophilius
Kozłowski, 1959 (after Kozłowski 1965, Fig. 1). (C) Tubotheca on
Acanthograptus
sp. (after Kozłowski 1970, Fig. 5). Illustrations not to scale.
Figure 4.19 (A, B, E)
Anticostia lata
(Elles & Wood, 1907), specimens with round blisters on the surface of the tubarium and enlargement of blister (E). (C)
Hustedograptus teretiusculus
(Hisinger, 1837) with blister, relief specimen in shale. (D)
Geniculograptus typicalis
(Hall, 1865), proximal end with several short tubes on the surface (from Bates & Loydell 2000). (F)
Archiclimacograptus riddellensis
(Harris & Thomas, 1935), specimen with coiled structure. Scale indicated by 1 mm long bar in each photo.
Figure 4.20 (A)
Rhabdinopora flabelliformis anglica
(Bulman, 1927a), showing regrown branches in the distal part of the colony (from Bull 1996, Fig. 1, reproduced with permission from The Palaeontological Association). (B)
Normalograptus scalaris
(Hisinger, 1837), isolated specimen from Dalarna, Sweden, distally lacking second stipe (based on Maletz 2003). (C)
Cardiograptus
sp. with one distorted stipe (from Han & Chen 1994, reproduced with permission from John Wiley & Sons). (D)
Dicaulograptus hystrix
(Bulman, 1932b), specimen with aborted second stipe (from Bulman 1932b, pl. 9, Fig. 9). (E)
Rectograptus gracilis
, regeneration of theca (after Kraft 1926, pl. 12). (F)
Bohemograptus bohemicus bohemicus
(Barrande, 1850) with misdirected first theca (from Urbanek 1970, Fig. 3, reproduced under the terms of the Creative Commons Attribution Licence 4.0, CC‐BY‐4.0). (G)
Slovinograptus balticus
(Teller, 1966) with partially biserial colony (from Urbanek 1997a, Fig. 10, reproduced with permission from Instytut Paleobiologii PAN). Illustrations not to scale.
Chapter 05
Figure 5.1 The preservable organic housing construction of the graptolites. (A)
Rhabdopleura normani
Allman in Norman (1869), single transparent tube of an extant graptolite. (B)
Rhabdopleura
sp., tube of a Middle Ordovician graptolite, Öland, Sweden. (C)
Xiphograptus primitivus
Maletz, 2010, proximal end of a planktic Ordovician graptolite. (B) and (C) are chemically isolated specimens showing the fossil preservation of the organic tubarium.
Figure 5.2 Graptolite preservation. (A)
Sigmagraptus praecursor
Ruedemann, 1904, NMVP 320445B, Victoria, Australia, multiramous sigmagraptine in typical black shale. (B)
Parisograptus caduceus
(Salter, in Bigsby 1853), NMVP 319348A, Victoria, Australia; darker, organically preserved graptolite in strongly weathered (bleached) shale.
Figure 5.3 “Clingfilm” preservation of
Spirograptus turriculatus
(Barrande, 1850). (A–C) Preservation in turbidite depositional system. (D–F) Preservation in clingfilm mode (based on Jones et al. 2002, Fig. 4); see also Plate 16 for
Spirograptus turriculatus
preservation.
Figure 5.4 Graptolites and trace fossils at Spudgels Cove, western Newfoundland, Darriwilian shales of American Tickle Formation with pyrite‐stained trace fossils (
Chondrites
sp.) and
Archiclimacograptus
sp. (A) Field photo showing traces. (B) Photo of shale from same layer showing traces and graptolite (white box). (C)
Archiclimacograptus
sp., enlarged from (B), SP4.12.01b. (D)
Archiclimacograptus
sp., SP4.12.19, well‐preserved specimen from same layer, coated with ammonium chlorite to show details. Scale is 1 mm in (C–D).
Figure 5.5 (A) Current‐aligned specimens of
Orthograptus apiculatus
(Elles & Wood, 1907) from Laggan Burn, Ayrshire, Scotland (B)
Uncinatograptus uniformis
(Přibyl, 1940), Loitzsch quarry, Thuringia, Lower Devonian. Scale indicated by 1 cm long bar in each photo.
Figure 5.6 Different views of graptolite colonies filled with sand in turbidite layer,
Levisograptus primus
(Legg, 1976), St. Pauls Inlet, western Newfoundland, Lower Head Sandstone, Darriwilian. (A) Apertural (scalariform) view, coated with ammonium chlorite. (B) Oblique view, uncoated. (C) Reverse view, showing proximal development in partial relief. Scale indicated by 1 mm long bar in each photo.
Figure 5.7 Diagenesis in graptolites. (A–B)
Amplexograptus perexcavatus
(Lapworth, 1876), JM 79, Lebanon Limestone, Murphreesboro, Tennessee (see Goldman et al. 2002), chemically isolated specimen showing internal and external growth of small pyrite crystals. (C)
Cochlograptus veles
(Richter, 1871), LO 1071 t, showing colony surrounded by dispersed pyrite crystals. (D)
Cymatograptus bidextro
Toro & Maletz, 2008, specimen showing black fusellum and pink mineral fill (see also Pl. 8A). (E)
Rivagraptus bellulus
(Törnquist, 1890), LO 1128 t, polished section showing pyrite fill of tubarium, fusellum shown as black outline and in the interthecal septae. Scale indicated by 1 mm long bar in each photo.
Figure 5.8 Coalification of graptolite fusellum. (A)
Pseudophyllograptus densus
(Törnquist, 1879), CN 2234, Skattungbyn, Dalarna, remains of black fusellum with some weathering. (B)
Baltograptus
sp., LO 10582 t, Lerhamn drillcore, Scania, Sweden, low coalification. (C)
Expansograptus latus
(Hall, 1907), Diabasbrottet, Sweden, relief specimen, fusellum highly coalified through contact metamorphosis. (D)
Archiclimacograptus wilsoni
(Lapworth, 1876), SM A 19619, highly coalified, Dob’s Linn, Scotland. Scale indicated by 1 mm long bar in each photo.
Figure 5.9 Coalification and contact metamorphosis. (A)
Pendeograptus simplex
(Törnquist, 1904), CN 1801,
Tetragraptus phyllograptoides
Biozone, high coalification, silvery fusellum. (B)
Baltograptus geometricus
(Törnquist, 1901), LO 1585 T,
Cymatograptus protobalticus
Biozone, flattened specimen with patches of surrounding mineral growth partly obscuring and distorting the specimen. (C, E)
Baltograptus
sp., coated latex cast (C) and mould (E) of specimen, showing crystal growth. (D)
Baltograptus jacksoni
Rushton, 2011,
Baltograptus vacillans
Biozone, originally pyrite‐filled parts preserved only. All specimens from Hunneberg, Sweden. Scale indicated by 1 mm long bar in each photo.
Figure 5.10 (A, B) Stipe fragment of
Didymograptus
sp., showing pattern of sediment fill and chlorite pressure shadow distributions (based on Mitchell et al. 2008, Fig. 3). (C) Multiramous graptolite on cleaved black shale, largely showing preservation of pressure shadow minerals (light colour), JOS 23.1a, Sandia Region Peru (Maletz et al. 2010).
Figure 5.11 Tectonic deformation.
Arienigraptus zhejiangensis
Yu & Fang, 1981. (A) JOS 21.1, original specimen, tectonically deformed. (B) Retro‐deformed specimen, vertically shortened by about 50%. (C) JOS 21.2, original specimen. (D) Retro‐deformed specimen, horizontally shortened to about 75%. (E) Partial relief specimen, latex cast for comparison.
Didymograptus murchisoni
(Beck). (F) Proximal end, flattened, showing tectonic lineation, preserved largely as pressure shadow minerals. (G) Fragment, fusellum in dark red, strong tectonic lineation visible. (A–D, F–G) Sandia Region, southern Peru. (E) Lerhamn drillcore, Scania, Sweden. Scale indicated by 1 mm long bar in each photo.
Figure 5.12 Graptolites in weathered (yellow) shale from the Ordovician of China. (A)
Prorectograptus uniformis
(Chen, in Mu et al. 1979), flattened specimen with fusellum preserved as small carbon flakes. (B)
Undulograptus formosus
(Mu & Lee, 1958), relief specimen in obverse view, weathered pyritic cast with partial cover of black fusellum. (C)
Azygograptus
sp., flattened, flakes of fusellum perserved. (D)
Baltograptus geometricus
(Törnquist, 1901), weathered pyritic cast with cover of black fusellum. (E)
Arienigraptus
sp., mainly yellow‐stained pressure shadow minerals. Scale indicated by 1 mm long bar in each photo.
Chapter 06
Figure 6.1 Fossil correlation. The presence of
Nicholsonograptus fasciculatus
can be used to correlate Middle Ordovician sections in Norway and western Newfoundland, the basis for biostratigraphical correlation of lithological successions on two different continents (based on data in Maletz et al. 2011).
Figure 6.2 Correlation of graptolite faunas across the Iapetus Ocean using biofacies overlap and transitional faunas (based on Maletz et al. 2011).
Figure 6.3 Devonian graptolite biozonation, showing the most important taxa and their actual ranges (compiled from data in Jaeger 1988; Loydell 2012; Lenz 2013).
Figure 6.4 The Ordovician/Silurian boundary interval at Dob’s Linn, Scotland, showing ranges of important graptolite taxa (based on https://engineering.purdue.edu/Stratigraphy/gssp/ordsil.htm).
Figure 6.5 Examples of biozone types as discussed in the text. FAD (first appearance datum), LAD (last appearance datum).
Figure 6.6 Biostratigraphy of the Aeronian/Telychian boundary, showing the evolution and biostratigraphy of the genus
Spirograptus
, length of zones and subzones not to scale (based on Loydell 1992; Loydell et al. 1993).
Figure 6.7 Lower Paleozoic GSSPs based on graptolites. The right side shows the “golden spike” for the base of the Floian at Diabasbrottet, Västergötland, Sweden, and a specimen of
Tetragraptus approximatus
, the graptolite species defining the boundary.
Figure 6.8 Ordovician biostratigraphy of Laurentia (North America) as an example of Ordovician biostratigraphy. A number of typical graptolite genera for the Ordovician are shown. (A)
Rhabdinopora
(Lower Tremadocian). (B)
Adelograptus
(Upper Tremadoian). (C)
Clonograptus
(Floian). (D)
Expansograptus
(Floian to Lower Darriwilian). (E)
Tetragraptus
, reclined (Floian to Lower Darriwilian). (F)
Isograptus
(Dapingian to Lower Darriwilian). (G)
Arienigraptus
(Upper Dapingian to Lower Darriwilian). (H)
Archiclimacograptus
(Darriwilian to Katian). (I)
Nemagraptus gracilis
(Sandbian). (J)
Normalograptus
(upper Darriwilian to Silurian). (K)
Amplexograptus
(Sandbian to Katian). (L)
Dicellograptus
(Uppermost Darriwilian to Katian). Graptolite illustrations from various sources, not to scale.
Figure 6.9 Silurian graptolite biostratigraphy (based on Koren et al. 1996; Loydell 2012), showing some of the characteristic Silurian graptolites. (A)
Akidograptus ascensus
. (B)
Petalolithus
. (C)
Dimorphograptus
. (D)
Demirastrites
. (E)
Rastrites
. (F)
Stimulograptus
. (G)
Spirograptus
. (H)
Streptograptus
. (I)
Monoclimacis
. (J)
Oktavites
. (K)
Retiolites
. (L)
Cyrtograptus
. (M)
Pristiograptus
. (N)
Heisograptus
. (O)
Saetograptus
. (P)
Bohemograptus
. Graptolite illustrations from various sources, not to scale.
Figure 6.10 Graphic correlation of Lower Ordovician graptolite sequences, composite standard sequence of FA (first appearance) events plotted against Australasian sequence (modified from Cooper & Lindholm 1990, Fig. 2).
Chapter 07
Figure 7.1 The virgellar spine. (A) Sigmagraptine indet., virgellar spine not present. (B)
Xiphograptus lofuensis
(Lee, 1961), dorsal virgellar spine. (C)
Archiclimacograptus
sp., juvenile with ventral virgellar spine. (D)
Archiclimacograptus
sp., proximal end with two complete thecae. (E)
Archiclimacograptus
sp., longer specimen. (F)
Saetograptus leintwardinensis
(Lapworth, 1880a), specimen showing ventral virgellar spine and branched dorsal apertural tongue. V = virgellar spine, N = nema. Scale indicated by 1 mm long bar close to each specimen.
Figure 7.2 Cladistic interpretation of
Rhabdopleura
, strict consensus of 12 equally parsimonious trees obtained from the full 17 taxon set, showing the position of extant
Rhabdopleura
inside the clade of the Graptolithina (based on Mitchell et al. 2013).
Figure 7.3 The evolution of the
Pristiograptus dubius
lineage (based on Urbanek et al. 2012). Illustrated specimen is
Pristiograptus dubius frequens
.
Figure 7.4 Extinction events in graptolite history, based on Cooper et al. (2014).
Figure 7.5 The evolution of the
Demirastrites triangulatus
group, modified from Sudbury (1958) and Urbanek (1960).
Figure 7.6 Convergent evolution of uniserial, single‐stiped Ordovician and Silurian graptoloids. (A)
Jishougraptus novus
Beckly & Maletz, 1991. (B)
Azygograptus suecicus
Moberg, 1892. (C)
Nicholsonograptus fasciculatus
(Nicholson, 1869). (D)
Pseudazygograptus incurvus
(Ekström, 1937). (E)
Atavograptus ceryx
(Rickards & Hutt, 1970). Bars show the ranges of genera, not the species shown as examples. Reconstructions not to scale.
Figure 7.7 The micro‐evolutionary changes of
Uncinatograptus
in the Ludfordian, Upper Ludlow, Silurian, from the Mielnik‐1 borehole section of Poland (based on Urbanek 1995).
Chapter 08
Figure 8.1 Cambrian Series 3, biostratigraphy and pterobranch faunas. (A)
Sphenoecium robustus
(Maletz et al., 2005b), Luh, Czech Republic. (B)
Yuknessia simplex
Walcott, 1919, Burgess Shale, holotype. (C)
Sphenoecium wheelerensis
Maletz & Steiner, 2015, Spence Shale, Utah. (D)
Sphenoecium mesocambricus
(Öpik, 1933), Alum Shale, Sweden (from Bengtson & Urbanek 1986). (E)
Sphenoecium discoidalis
(Chapman & Thomas, 1936), Heathcote fauna, Tasmania. (F)
Sphenoecium wheelerensis
Maletz & Steiner, 2015, Wheeler Shale, Utah. (G)
Sphenoecium robustus
(Maletz et al., 2005b), Konicek, Czech Republic. (H)
Sphenoecium wheelerensis
Maletz & Steiner, 2015, Marjum Fm., Utah. (I)
Sphenoecium mesocambricus
(Öpik, 1933), Krekling, Norway. Illustration of specimens not to scale.
Figure 8.2 The Middle Cambrian graptolite
Sphenoecium wheelerensis
Maletz & Steiner, 2015. (A–C) KUMIP 204381, Spence Shale, Utah, USA, complete specimen and details showing impressions of fusellar construction. (D–E) WHE‐001, Wheeler Shale, Utah, USA (see Maletz et al. 2005b). B–E are SEM‐BSE photos showing the fusellum in black and providing evidence of the fusellar construction.
Figure 8.3 Colony shape and attachment in erect, benthic graptolites. (A)
Dictyonema
sp., conical colony without stem. (B)
Dictyonema
sp. without stem. (C)
Dendrograptus
sp., colony with robust stem, thecae not visible. (D)
Dictyonema
sp., specimen with long, thecate stem. (E)
Dendrograptus
sp., bushy colony without stem. (F)
Dictyonema cavernosum
Wiman, specimen with irregular attachment disc. Reconstructions based on Bulman (1928) and Chapman et al. (1996), not to scale.
Figure 8.4 Middle Cambrian (Series 3) rhabdopleurids from Monegeeta, Australia. (A)
Archaeolafoea longicornis
Chapman, 1919, NMVP 13112, holotype. (B) Growing end of same. (C) NMVP 13114, holotype of
Archaeocryptolaria skeatsi
Chapman, 1919, growing end. (D) Central part of another colony from same slab. Scale indicated by 1 mm long bar in each photo.
Figure 8.5 Wimanicrustidae and Cysticamaridae. (A)
Bithecocamara
, thigmophylic, sheet‐like colony, one theca highlighted, reconstruction based on Bulman (1970a), Fig. 31, with permission from the Paleontological Institute. (B)
Bulmanicrusta
? sp. runner‐type colony, specimen with irregular basal sheet, stolon and single partly preserved autotheca. (C)
Bulmanicrusta
, reconstruction, showing thecal style and graptoblast (after Urbanek, unpublished). (D)
Bulmanicrusta
? sp., graptoblast. (B, D) adapted from Mitchell et al. (1993) with permission from Cambridge University Press. Illustrations not to scale.
Figure 8.6 The Cyclograptidae. (A)
Galeograptus nicholasi
Bulman & Rickards, 1966, dorsal view. Adapted from Bulman and Rickards (1966). (B)
Discograptus
, dorsal view. (C)
Tubidendrum bulmani
Kozłowski, 1949, fragment showing coiled thecae. (D)
Kozlowskitubus erraticus
(Kozłowski,1963), proximal end with erect stipes. (E)
Galeograptus
, lateral view. (F)
Dendrotubus wimani
Kozłowski, 1949, initial coiled part of thecal tube. (B), (E) and (F) adapted from Bulman (1970a) with permission from Paleontological Institute, and from Bulman (1970b) with permission from Société Belge de Géologie de Paléontologie et d’Hydrologie. Illustrations not to scale.
Figure 8.7 Dithecodendridae. (A)
Ovetograptus gracilis
Sdzuy, 1974, SMF 30028, holotype. (B)
Tarnagraptus cristatus
Sdzuy, 1974, SMF 30021, holotype. (C–D)
Tarnagraptus palma
Sdzuy, 1974, SMF 30000, holotype. All specimens from the Cantabrian Mountains, Spain (see Sdzuy 1974).
Figure 8.8 Examples of Mastigograptidae. (A–B)
Mastigograptus tenuiramosus
(Walcott, 1883), large fragment and detail showing thecae. (C–E)
Mastigograptus
sp., chemically isolated thecae showing triad budding. (A–D) adapted from Bulman (1970a) with permission from Paleontological Institute, and from Bulman (1970b) with permission from Société Belge de Géologie de Paléontologie et d’Hydrologie. Illustrations not to scale. (E) adapted from Andres (1977) with permission from Springer.
Figure 8.9 Dendrograptidae. (A)
Dendrograptus hallianus
(Prout, 1851), fragment of bushy colony showing thecal style on distal stipes, based on Hall (1865, Fig. on p. 127). (B)
Dictyonema retiformis
(Hall, 1851), conical colony with numerous dissepiments forming a typical meshwork (after Hall 1865, Fig. 10). (C)
Dictyonema estlandicum
Bulman, 1933, fragment showing thecal development and dissepiments (after Bulman 1933, pl. 7, Fig. 5). Illustrations not to scale.
Figure 8.10 Planktic dendroids. (A–C)
Calyxdendrum graptoloides
Kozłowski, 1960 (after Kozłowski 1960). (A) Distal fragment showing thecal development. (B) Proximal end with nematophorus sicula. (C) Isolated sicula with initial part of first theca. (D)
Dictyonema ghodisiae
Rickards, Hamedi & Wright, 1994, showing nematophorus sicula and vesicular bodies or nematularium(?), Kerman District, Iran (after Rickards et al. 1994).
Figure 8.11 Acanthograptidae. (A)
Acanthograptus sinensis
Hsü and Ma, 1948, fragment preserved in partial relief, showing thecal tubes, Tremadocian, China. (B–C).
Acanthograptus divergens
Skevington, 1963, fragments in relief, SEM photos, Darriwilian, Middle Ordovician, Öland, Sweden. The scale bar indicates 1 mm in each photo.
Figure 8.12 The Carboniferous graptolite
Ptiograptus fournieri
Ubaghs, 1941, holotype, Molignée Formation, Visean, Denée, Belgium (photo provided by B. Mottequin, 2015). Scale indicates 10 cm.
Figure 8.13 Algae and graptolites. (A)
Inocaulis plumulosa
Hall, 1852 (B)
Diplospirograptus goldringae
Ruedemann, 1925 (after Ruedemann 1947, pl. 41). (C)
Medusaegraptus mirabilis
Ruedemann, 1925 (after Ruedemann 1947, pl. 42). (D)
Medusaegraptus graminiformis
(Pohlmann, 1887) (after Ruedemann 1947, pl. 41). Reconstructions not to scale.
Chapter 09
Figure 9.1 (A)
Dictyonema
sp., benthic graptoloid, Burgberg section, Germany, Middle Devonian. (B)
Rhabdinopora parabola
(Bulman), Dayangcha, Jilin, China, details of stipes showing thecae. (C)
Rhabdinopora parabola
(Bulman), planktic graptoloid, Dayangcha, Jilin, China, complete specimen showing colony shape and irregularly placed dissepiments.
Figure 9.2 (A)
Rhabdinopora parabola
(Bulman), Dayangcha, fragment showing irregular dissepiments. (B)
Sagenograptus macgillivrayi
(Hall, 1897), NMVP 13096, top view of proximal end with regular dissepiments. (C)
Callograptus elegans
(Hall, 1865), GSC 956a, cotype, latex cast, showing anastomosis (arrows). (D)
Callograptus salteri
(Hall 1865, pl. 19, Fig. 7), GSC 955, cotype, showing short bridges or dissepiments (1) and anastomosis (2). Scale in each photo 1 mm, except for (B) in which it is 10 mm.
Figure 9.3 (A)
Epigraptus
sp., juvenile with dome and initial (sicular) tube (B)
Dendrotubus
sp. with a distal helical line in the prosicula (based on Kozłowski 1971, Fig. 5). (C)
Dendrograptus communis
Kozłowski, 1949, tube‐like prosicula with helical line (based on Kozłowski 1949, Fig. 1). (D) Conus and cauda (based on Hutt 1974a). (E–G):
Adelograptus tenellus
(Linnarsson, 1871), based on Hutt (1974a). (E) Conus and cauda in incomplete sicula, showing resorption foramen of first theca in prosicula. (F) Juvenile with complete sicula, part of th1
1
and crossing canal of th1
2
, reverse view. (G) Sicula with complex nema. (D–G): adapted from Hutt (1974a) with permission from John Wiley & Sons. Illustrations not to scale.
Figure 9.4 (A, B)
Rhabdinopora proparabola
(Lin, 1986), Dayangcha section, Jilin, China, showing nematularium with possible fusellar increments (B). (C)
Staurograptus dichotomous
Emmons, 1855, St Paul’s Bridge section, western Newfoundland, coll. Erdtmann. (D, E)
Rhabdinopora parabola
(Bulman, 1954), Dayangcha section, Jilin, China, specimen with multiple branched nemata. (F)
Rhabdinopora parabola
(Bulman, 1954), Shangsonggang section, Jilin, China, specimen with distally branched nema.
Figure 9.5 Increase in symmetry and regularity. (A)
Anisograptus matanensis
Ruedemann, 1937, Matane, Quebec, Canada, adapted from Bulman (1950) from Geological Society London. (B)
Clonograptus rigidus
(Hall, 1858), Levis, Quebec, Canada, after Lindholm and Maletz (1989). (C)
Expansograptus grandis
(Monsen, 1937). (D)
Isograptus lunatus
Harris, 1933. (E)
Archiclimacograptus
sp. All illustrations are reconstructions, not to scale.
Figure 9.6 (A, B)
Hunnegraptus copiosus
Lindholm, Hunneberg, Sweden, proximal end in obverse (A) and reverse (B) views, showing sicular bitheca (arrow) and stipe development. (C) ?
Paratemnograptus magnificus
(Pritchard, 1892), MG 1967, latex cast, stipe fragment showing plaited thecal overlap without bithecae, Fezouata Biota, Morocco. (D)
Kiaerograptus kiaeri
(Monsen, 1925), PMO 72833, fragment showing long bithecae (arrows). Scale indicated by 1 mm long bar in each photo.
Figure 9.7 (A) Quadriradiate (
Rhabdinopora
,
Staurograptus
), (B) triradiate (
Anisograptus
,
Triograptus
) and (C) biradiate (
Adelograptus
,
Kiaerograptus
) proximal development in dorsal view (based on Maletz 1992a). Distal dicalycal thecae labelled DD.
Figure 9.8 Lower Tremadocian graptolite biostratigraphy (based on Cooper et al. 1998). A general succession of quadri‐, tri‐ and biradiate horizontal taxa can be differentiated in addition to the more precise biostratigraphical zonation. The
Rhabdinopora
lineage remains quadriradiate through the whole Lower Tremadocian. Graptolite illustrations from various sources, not to scale.
Figure 9.9 Upper Tremadocian psigraptids. (A)
Chigraptus supinus
Jackson & Lenz, 1999, GSC 117666, Yukon Territory, Canada. (B)
Ancoragraptus bulmani
(Spjeldnaes, 1963), GSC 123191, Yukon Territory, Canada. (C–D)
Psigraptus jacksoni
Rickards & Stait, 1984, TM 01, Yeongwol area, Korea, one juvenile and one mature specimen associated on a slab. (E–F)
Psigraptus arcticus
Jackson, 1967, JM 45, Erdaopu Section, Jilin, China. Scale indicated by 1 mm long bar in each photo.
Figure 9.10 Transitional graptoloids. (A)
Sagenograptus macgillivrayi
(Hall, 1897), Victoria, Australia. (B)
Hunnegraptus copiosus
Lindholm, 1991, Hunneberg, Sweden. (C)
Hunnegraptus novus
(Berry, 1960a), juvenile, Texas, USA. (D)
Paratemnograptus magnificus
(Pritchard, 1892), Victoria, Australia. (E)
Paradelograptus mosseboensis
Erdtmann, Maletz and Gutierrez‐Marco, 1987, Hunneberg, Sweden. (F)
Paradelograptus
sp., Cow Head Group, western Newfoundland, Canada. (G)
Aorograptus victoriae
(Hall, 1899), Cow Head Group, western Newfoundland, Canada. Illustrations not to scale.
Chapter 10
Figure 10.1 Ordovician graptolite diversity diagrams, modified from Sadler et al. (2011, Fig. 14). Three main events are shown by horizontal lines. (1) Origin of Dichograptina in the late Tremadocian and diversity burst in late Floian. (2) Origin of Isograptidae in basal Dapingian with subsequent increase in diversity. (3) Origin of Axonophora (Normalograptidae, Diplograptidae, Lasiograptidae) at the base of the Darriwilian and subsequent diversification.
Figure 10.2 Loss of stipes in the Dichograptina. (A)
Paratemnograptus magnificus
(Pritchard, 1892), large multiramous graptoloid, Victoria, Australia. (B)
Tetragraptus amii
(Elles & Wood, 1902), side view showing three out of four stipes, latex cast, Hunneberg, Sweden. (C)
Expansograptus validus
(Törnquist, 1901), two‐stiped, flattened, Hunneberg, Sweden. (D)
Expansograptus
sp., latex cast, Tøyen, Norway. Magnification indicated by 1 mm long bar in (B–D) and 10 cm long bar in (A).
Figure 10.3 The loss of bithecae. (A)
Paradelograptus antiquus
(T.S. Hall, 1899), Yukon, Canada, showing sicular bitheca. (B)
Kiaerograptus supremus
Lindholm 1991, Sweden, specimen in reverse view with regular bithecae along stipes. (C)
Baltograptus vacillans
(Tullberg, 1880), latex cast in reverse view, Hunneberg, Sweden, bithecae are absent throughout the stipes, and the origin of thecae is from the dorsal side of stipes. Bithecae indicated by white arrows in (A–B).
Figure 10.4 Revised interpretation of the Maletz et al. (2009) analysis, using preferred taxon names. Insets show proximal development of the (A) Anisograptidae (
Adelograptus
). (B) Sigmagraptidae (
Paradelograptus
). (C) Didymograptidae (
Didymograptellus nitidus
). (D) Isograptidae (
Isograptus
). Graptolite illustrations not to scale.
Figure 10.5 The proximal development of the Anisograptidae (A), Sinograpta (B–C) and Dichograptina (D–F). (A)
Anisograptus matanensis
Ruedemann, 1937, NGPA 216/07, oblique sicula and asymmetrical development. (B) Sigmagraptine indet., SPI 63, vertical sicula and asymmetrical development. (C).
Sigmagraptus
sp. with elongated, slender sicula, CHN 11.4E, nearly symmetrical development. (D)
Xiphograptus
sp., GSC 133392. (E–F)
Expansograptus hirundo
(Salter, 1863) in obverse (E) and reverse (F) views, Tøyen Shale, Slemmestad, Norway. (A–D) flattened specimens, Cow Head Group, western Newfoundland. Magnification for all specimens provided by 1 mm long bar in each photo.
Figure 10.6 Colony design of the Sigmagraptidae. (A)
Paradelograptus smithi
(Harris & Thomas, 1938a). (B)
Paradelograptus mosseboensis
Erdtmann, Maletz and Gutierrez‐Marco, 1987, proximal end. (C)
Yushanograptus
sp. (D)
Goniograptus
sp., GSC 125786, proximal end. (E)
Sigmagraptus praecursor
Ruedemann, 1904, GSC 79889, western Newfoundland. (F)
Etagraptus tenuissimus
(Harris & Thomas, 1942), holotype. (G) ?
Goniograptus
sp., GSC 125768, proximal end in reverse view. (H) Sigmagraptine indet with single stipe, GSC 125815. (I)
Trichograptus dilaceratus
Herrmann, 1885. (J)
Goniograptus
sp., GSC 125788, juvenile. (K) Sigmagraptine indet, GSC 125806. Various magnifications.
Figure 10.7 Specialized tubaria in Abrograptidae. (A)
Jiangshanites
(?)
dubius
Maletz, 1993, reconstruction (based on Maletz 1993). (B)
Jiangshanites
(?)
dubius
, GSC 102774. (C)
Jiangshanites
(?)
dubius
Maletz, 1993, GSC 102779, holotype. (D)
Abrograptus formosus
Mu, 1958. (E)
Dinemagraptus warkae
Kozłowski, 1951. (F)
Parabrograptus tribrachiatus
Mu & Qiao, 1962. (B, C) flattened, isolated specimens, digitally cleaned, see also Maletz (1993); (D–F) modified from Finney (1980, Fig. 11). Arrows indicate preservation of part of first theca in
Jiangshanites
(?)
dubius
. Various magnifications.
Figure 10.8 Sinograptidae. (A)
Anomalograptus reliquus
Clark, 1924, NIGP 8847, small specimen with comparably few stipes, thecal details not available. (B)
Allograptus mirus
Mu, NIGP 8868, paratype. (C)
Anomalograptus reliquus
Clark, 1924, NIGP 8852, large specimen with typical dichotomous branching and long funicle. (D)
Anomalograptus reliquus
Clark, 1924, wb2.34‐42b. (E)
Anomalograptus reliquus
Clark, 1924, wb2.34.29a. (D, E) are flattened isolated specimens from shales of the Cow Head Group, western Newfoundland, Canada. Scale indicates 5 mm (A–C) and 1 mm (D–E).
Figure 10.9 Sinograptidae. (A–B)
Holmograptus callotheca
(Bulman, 1932), holotype, isolated material, Öland, Sweden. (C)
Holmograptus lentus
(Törnquist, 1892), LO 3260 t, holotype, Scania, Sweden. (D)
Holmograptus
sp., apertural view of theca (from Kozłowski 1954). (E)
Holmograptus bovis
Williams & Stevens, 1988, reverse view (from Bulman 1936). (F–G)
Nicholsonograptus fasciculatus
(Nicholson, 1869), Table Head Group, western Newfoundland. Scale indicates 1 mm in each photo.
Figure 10.10 Complex thecal structure of the Sinograptidae. (A)
Sinograptus typicalis
Mu, 1957, holotype, weathered pyritic internal cast (right stipe) and high relief imprint in light coloured shale (left stipe). (B)
Sinograptus
, thecal reconstruction with prothecal and metathecal folds. (C)
Holmograptus
, thecal reconstruction with prothecal folds. Reconstructions not to scale.
Figure 10.11 Various branching patterns in Dichograptina (A–D, F) and Sinograpta (E). (A)
Clonograptus
, progressive, dichotomous branching. (B)
Schizograptus
, dichotomous branching with lateral origin of stipes. (C)
Holograptus
, dichotomous, lateral branching with fairly irregular branching. (D)
Triaenograptus
, branching in triads. (E)
Goniograptus
, monoprogressive branching. (F)
Dichograptus
, dichotomous branching proximally, specimen with large proximal web. Illustrations not to scale.
Figure 10.12 Tetragraptidae. (A)
Tetragraptus amii
Elles and Wood, 1902, horizontal. (B)
Pseudophyllograptus
sp., cross‐section. (C)
Tetragraptus serra
(Brongniart, 1828), reclined. (D)
Pseudophyllograptus densus
(Hall, 1865). (E)
Tetragraptus phyllograptoides
Strandmark, 1902, showing three out of four stipes. (F)
Phyllograptus
sp., showing central columella. (G)
Tetragraptus phyllograptoides
Strandmark, 1902, showing proximally united stipes. Scale indicates 1 mm in all photos, except (A), where it indicates 5 mm.
Figure 10.13 Tetragraptidae. (A–B)
Tetragraptus bigsbyi
(Hall, 1865). (C–D)
Tetragraptus phyllograptoides
Strandmark, 1902. (E–F)
Tetragraptus cor
(Strandmark, 1902). (A–F from Strandmark, 1902). (G–H)
Tetragraptus archaios
(Braithwaite, 1976), Trail Creek, Idaho (see Maletz et al., 2005), showing distally united stipes. (A, C, E, G) in a‐b orientation; (B, D, F, H) in 1‐2 orientation (see Chapter 3). Illustrations not to scale.
Figure 10.14 Didymograptidae. (A)
Expansograptus holmi
(Törnquist, 1901), latex cast, showing the high (prosicular) origin of the first theca (th1
1
) from the sicula, Diabasbrottet, Hunneberg, Sweden. (B)
Baltograptus vacillans
(Tullberg, 1880), latex cast, showing the origin of the th1
1
from middle part of the sicula, Diabasbrottet, Hunneberg, Sweden. (C)
Expansograptus praenuntius
(Törnquist, 1901), LO 1611 t, latex cast, Flagabro, Scania, Sweden. (D)
Expansograptus grandis
(Monsen, 1937), latex cast, Slemmestad, Norway. (E)
Didymograptus murchisoni
(Beck in Murchison, 1839),
artus
type proximal development typified by th1
1
as the dicalycal theca, Ebbe anticline, Germany. (F–G)
Jenkinsograptus spinulosus
(Perner, 1895), isograptid‐type proximal development typified by th1
2
as the dicalycal theca, Krapperup, Scania, Sweden. All specimens in reverse view, except for (C–D) in obverse view. Bar indicates 1 mm in each photo.
Figure 10.15 Growth stages and adults of some species in the Pterograptidae.
Pterograptus elegans
Holm, 1881: (A) reconstruction of colony.(C) Fragmented proximal end showing cladial origin of theca (arrow). (D) Proximal end showing artus type development and metasicular origin of th1
1
.
Xiphograptus formosus
(Bulman, 1936): (B) Proximal end. (E–F) Juveniles showing construction and growth of virgellar spine (G)
Didymograptellus bifidus
(Hall, 1865), juvenile showing strong virgellar spine.
Yutagraptus mantuanus
Riva, 1994: (H) Holotype (Riva 1994). (I) Isolated juvenile showing virgellar spine attached to ventral wall of theca 1
2
(arrow). Illustrations not to scale.
Figure 10.16 Different symmetries in Tetragraptidae and Isograptidae. The maeandrograptid (A) and isograptid (B–D) symmetry. (A)
Tetragraptus reclinatus
. (B)
Isograptus victoriae
. (C)
Parisograptus caduceus
. (D)
Arienigraptus zhejiangensis
. Sicula, dicalycal theca (th1
2
) and downward growing part of manubriate thecae (in D) is shaded. Reconstructions based on Maletz & Zhang (2003); Maletz (2011d).
Figure 10.17 Middle Ordovician biostratigraphy based on isograptids and pseudisograptids (based on Cooper 1973; Cooper & Ni 1986). Graptolite specimens from various sources, not to scale.
Figure 10.18 The manubriate isograptids. (A)
Pseudisograptus manubriatus janus
Cooper and Ni, 1986, GSC 82977, flattened specimen, Melville Island, Arctic Canada. (B)
Arienigraptus
sp., LO 12244 t, relief specimen in reverse view, coated with ammonium chlorite, Krapperup drillcore, Scania, Sweden. (C)
Arienigraptus
sp., GSC 82979, flattened, Melville Island, Arctic Canada. (D)
Arienigraptus dumosus
Harris, 1933, GSC 82978, flattened, Melville Island, Arctic Canada. (E)
Pseudisograptus manubriatus
ssp., KR‐5b‐2, latex cast in reverse view, Krapperup drillcore, Scania, Sweden. Magnification indicated by 1 mm long bar in each photo.
Figure 10.19 Evolution of scandency in the Glossograptina. (A)
Tetragraptus reclinatus
. (B)
Isograptus lunatus
. (C)
Isograptus mobergi
. (D)
Oncograptus
. (E)
Cardiograptus
. (F)
Pseudisograptus
. (G)
Exigraptus
. (H)
Bergstroemograptus
. (I)
Cryptograptus
. (J)
Glossograptus
. (K)
Kalpinograptus
. Scandency seen in (E) (Isograptidae, through
Isograptus
), (G) (Isograptidae, through
Pseudisograptus
, leading to the Axonophora), (H–J) (Glossograptidae, through
Isograptus
; (K) lost scandency secondarily). Illustrations not to scale, based on various sources.
Figure 10.20
Parisograptus caduceus
(Salter in Bigsby, 1853), (A) NIGP 126527, obverse view; (B, D) NIGP 12523, reverse view. (C) NIGP 126522, obverse view. Scale indicates 1 mm in each illustration.
Figure 10.21 The Glossograptidae. (A)
Cryptograptus schaeferi
Lapworth, 1880, Table Head Group, western Newfoundland. (B–C)
Nanograptus lapworthi
Hadding, 1915. (B) Specimen on GSM 5495 with lectotype of
Rogercooperia phylloides
(Elles & Wood, 1908). (C) LO 2743 t (paratype). (D)
Cryptograptus
sp., SPS 28, western Newfoundland. (E)
Paraglossograptus tentaculatus
(Hall, 1865), showing partially preserved lacinia. (F)
Glossograptus hincksii
(Hopkinson, 1872), LO 2370 t, Scania, Sweden (Hadding 1913, pl. 2, Fig. 6). Scale indicated by 1 mm long bar in each photo.
Figure 10.22 Proximal development of the Glossograptidae. (A)
Cryptograptus insectiformis
Ruedemann, 1908 (based on Maletz & Mitchell 1996). (B) Reconstruction of sicula and first theca in
Cryptograptus schaeferi
(based on Maletz & Mitchell 1996). (C)
Cryptograptus schaeferi
Lapworth, 1880, Table Head Group, western Newfoundland, Canada, juvenile showing complete sicula with lateral apertural spines and part of first theca; note the thickened rim around the sicular aperture that will become part of the list structure of younger cryptograptids. (D)
Glossograptus acanthus
Elles & Wood, 1908, proximal end in relief. Adapted from Ni and Cooper (1994, Fig. 1), with permission from Taylor & Francis. (E) Thecal diagram of
Glossograptus
, showing monopleural arrangement (after Maletz & Mitchell 1996). Illustrations not to scale.
Figure 10.23 Tubarium reconstruction of the Glossograptidae. (A)
Bergstroemograptus
. (B)
Cryptograptus
. (C)
Glossograptus
. (D)
Paraglossograptus
. (E)
Corynoides
. (F)
Corynites
. (G)
Kalpinograptus
. Reconstructions (JM) not to scale.
Chapter 11
Figure 11.1 The graptolite tubarium and the nema. (A)
Isograptus victoriae
Harris, 1933, Vinini Formation, Nevada, USA, completely free nema between two reclined stipes. (B)
Rectograptus intermedius
(Elles & Wood, 1907), nema incorporated in tubarium, visible through the colony in this infrared photo. (C)
Saetograptus leintwardinensis
(Hopkinson in Lapworth, 1880), glacial boulder, Nienhagen northern Germany, nema visible on dorsal side of stipe and distally of thecae. Scale indicated by 1 mm long bar in each photo.
Figure 11.2 Early biserial axonophorans and their biostratigraphy (based on Maletz 2011a, Fig. 4).
Figure 11.3 Unusual development of the nema. (A–C)
Climacograptus
(?)
uncinatus
Keble & Harris, 1934, all specimens from Trail Creek section, Idaho (see Goldman et al. 2007), drawings by Kristen Paris (UB Buffalo, 2005). (D)
Climacograptus ensiformis
Mu and Zhang in Mu, 1963 with paired spines and lateral membranes (after Mu 1963, Fig. 12d). Scale indicates 1 mm in each illustration.
Figure 11.4 Development of the axonophoran sicula. (A) Immature sicula with prosicula showing longitudinal rods and spiral line, several metasicular fusellar rings and development of virgellar spine. (B) Monograptid with complete sicula and five sicular annuli (labelled 1–5). (C) Section through sicular annulus showing construction from cortical layers. (D) Section through sicula showing development of sicular annuli on the inside. (A–B) Monograptid indet., ?
Heisograptus micropoma
(after Kraft 1926). (C–D)
Pseudomonoclimacis dalejensis
(Bouček, 1936) (after Urbanek 1958). Illustrations not to scale.
Figure 11.5 The virgellar spine development. (A) Rutellum. (B) Lamelliform rutellum. (C) Lanceolate virgella. (D) Virgella (after Maletz 2010a, Fig. 2). Specimens: (E) Isograptid indet., sicula with lamelliform rutellum, wb1 34 22. (F)
Isograptus
sp., chs 13 1 73, flattened proximal end with extended rutelli. (G)
Levisograptus sinicus
(Mu & Lee, 1958), GSC 133381, complete sicula with lanceolate virgella and part of first theca. (H)
Levisograptus sinicus
(Mu & Lee, 1958), GSC 133378, specimen with two thecal pairs, some parts broken. (I)
Levisograptus sinicus
(Mu & Lee, 1958), GSC 133386, small specimen with five thecae and lanceolate virgella. (J)
Archiclimacograptus
sp., bas 123, West Bay Centre Quarry, western Newfoundland. All specimens are flattened, chemically isolated from shales. Scale indicated by 1 mm long bar in each photo.
Figure 11.6 Examples of the proximal development types of biserial axonophorans. Patterns A to I are shown together with specimens as examples demonstrating the development (based on Mitchell 1987).
Figure 11.7 The transition from the reclined
Pseudisograptus
to the biserial, axonophoran
Archiclimacograptus
. The manubrium, sicula and th1
1
are highlighted to show the changes more clearly. (A)
Pseudisograptus
. (B–C)
Exigraptus
, showing prosicular origin of th1
1
(arrow in B). (D)
Archiclimacograptus
, juvenile, showing metasicula rorigin of th1
1
(arrow). (E)
Levisograptus
. (F)
Archiclimacograptus
. Reconstruction of specimens from various sources, not to scale.
Figure 11.8 Comparison of the Neograptina (Normalograptidae) (A–D) and the Diplograptina (Diplograptidae and Lasiograptidae) (E–L) in the Ordovician. (A)
Undulograptus formosus
(Mu & Lee, 1958). (B)
Undulograptus novaki
(Perner, 1895). (C)
Skanegraptus janus
Maletz, 2011c. (D)
Normalograptus antiquus
(Ge, in Ge et al. 1990). (E)
Orthograptus quadrimucronatus
(Hall, 1865). (F)
Levisograptus primus
(Legg, 1976). (G)
Diplograptus pristis
(Hisinger, 1837). (H)
Exigraptus clavus
Mu in Mu et al. 1979. (I)
Paraorthograptus pacificus
(Ruedemann, 1947). (J)
Pipiograptus
sp. (K)
Brevigraptus quadrithecatus
Mitchell, 1988. (L)
Amplexograptus
sp. Graptolite specimens and reconstructions from various sources, not to scale.
Figure 11.9 Median septum development. (A)
Levisograptus
sp., showing intrathecal folds with undulating median septum and long, double sigmoid thecae, Krapperup drill core, Scania, Sweden. (B)
Haddingograptus oliveri
(Bouček, 1973), showing a strongly zigzag median septum, Table Head Group, western Newfoundland. (C) (?)
Archiclimacograptus
sp., showing short thecae with nearly straight median septum, Table Head Group, western Newfoundland. (D)
Petalolithus minor
Elles, 1897, LO 1115 t, showing alternating thecae, median septum lacking, Scania, Sweden. (1) indicates the interthecal septum, (2) the intrathecal septum in (A–B). All specimens coated with ammonium chlorite. Scale indicated by 1 mm long bar in each photo.
Figure 11.10 The differentiation and evolution of the Axonophora. Graptolite examples: (A)
Normalograptus kukersianus
(Neograptina). (B)
Archiclimacograptus
sp. (Diplograptina). Diversities and ranges based on Sadler et al. (2011).
Figure 11.11 Examples of the Diplograptidae (A–D, F–I, K–L) and Lasiograptidae (E–F, J, M). (A)
Pseudamplexograptus
Mitchell, 1987. (B)
Urbanekograptus retioloides
(Wiman, 1895). (C)
Amplexograptus
sp. (D)
Hustedograptus uplandicus
(Wiman, 1895). (E)
Nymphograptus velatus
Elles & Wood, 1908. (F)
Orthoretiolites hami
Whittington, 1954. (G)
Diplograptus pristis
(Hisinger, 1837). (H)
Rectograptus gracilis
(Roemer, 1861), neotype. (I)
Rectograptus intermedius
(Elles & Wood, 1907). (J)
Lasiograptus harknessi
(Nicholson, 1867a) (K)
Geniculograptus typicalis
(Hall, 1865). (L)
Peiragraptus fallax
Strachan, 1954. (M)
Yinograptus disjunctus
(Yin & Mu, 1945). Illustrations not to scale, based on various sources.
Figure 11.12 The Climacograptidae. (A)
Undulograptus formosus
(Mu & Lee, 1958). (B)
Undulograptus clabavensis
Bouček, 1973. (C)
Oelandograptus oelandicus
(Bulman, 1963). (D)
Haddingograptus eurystoma
(Jaanusson, 1960), reverse view. (E)
Haddingograptus oliveri
(Bouček, 1973). (F)
Pseudoclimacograptus scharenbergi
(Lapworth, 1876). (G)
Proclimacograptus angustatus
(Ekström, 1937). (H, J)
Climacograptus bicornis
(I)
Styracograptus tubuliferus
(Lapworth, 1876), reverse view (based on Mitchell 1987). (K)
Diplacanthograptus spiniferus
(Ruedemann, 1908), reverse view (L)
Appendispinograptus venustus
(Hsü, 1959), showing complex parasicular development, Wufeng Formation, China. (M)
Appendispinograptus longispinus
(Hall, 1902), flattened (N)
Climacograptus hastatus
(Hall, 1902), flattened specimen with long parasiculaIllustrations are largely reconstructions, based on various sources, not to scale.
Figure 11.13 (A)
Haddingograptus eurystoma
(Jaanusson, 1960), reverse view, showing zigzag median septum and intrathecal folding, Elnes Formation, Slemmestad, Norway. (B)
Pseudoclimacograptus scharenbergi
(Lapworth, 1876), showing zigzag median septum and intrathecal folding (based on Bulman 1945, pl. 8, Fig. 6). (C)
Proclimacograptus angustatus
Ekström, 1937, showing slightly undulating median septum, no intrathecal folds, Elnes Formation, Slemmestad, Norway. (D)
Appendispinograptus supernus
? (Elles & Wood, 1906), NIGP 139881, internal cast of proximal end showing sicula (S) with parasiculae, sub‐scalariform view, Wufeng Formation, Guizhou, China. (E–F)
Climacograptus cruciformis
VandenBerg, 1990, Viola Limestone Formation, Oklahoma, showing prosicula reduced to two rods uniting distally to form the nema (photos provided by D. Goldman). Bar indicates 1 mm in each photo.
Figure 11.14 The Dicranograptidae. (A)
Levisograptus sinicus
(Mu & Lee, 1958). (B)
Dicaulograptus hystrix
(Bulman, 1932b). (C)
Levisograptus dicellograptoides
(Maletz, 1998b), reconstruction. (D)
Dicellograptus
sp., reconstruction. (E)
Jiangxigraptus alabamensis
(Ruedemann, 1908). (F)
Jiangxigraptus
sp. (G)
Dicellograptus caduceus
Lapworth, 1876 (after VandenBerg & Cooper 1992, Fig. 9 V). (H)
Tangyagraptus typicus
Mu, 1963 (from Mu 1963, Fig. 3e). (I)
Dicranograptus clingani
Carruthers, 1868, reconstruction. (J)
Neodicellograptus dicranograptoides
Mu and Wang in Wang & Jin, 1977, reconstruction (JM). Illustrations not to scale.
Figure 11.15 Torsion in the Dicranograptidae. (A)
Dicellograptus
with left‐handed torsion showing axial angle (B)
Dicellograptus bispiralis
(Ruedemann, 1947) (C)
Nemagraptus gracilis
(Hall, 1847), OSU 32962 (based on Finney 1985, Fig. 19‐1). (D)
Dicranograptus zigzag
Lapworth, 1876, reconstruction showing independent spiralling of stipes (E)
Dicellograptus caduceus
Lapworth, 1876, reconstruction (after Bulman 1964, Fig. 3. Illustrations not to scale.
Figure 11.16 The Nemagraptinae. (A)
Nemagraptus linmassiae
Finney, 1985, holotype. (B)
Amphigraptus
sp. (C)
Nemagraptus gracilis
(Hall, 1847), proximal end in reverse view, showing aperturally free sicula. (D) Proximal and distal thecae of
Nemagraptus gracilis
. (E) Proximal and distal thecae of
Nemagraptus linmassiae
, compare with shorter thecal overlap of thecae in
Nemagraptus gracilis
. Illustrations not to scale.
Figure 11.17 The Neograptina radiation (based on Melchin et al. 2011, Fig. 7). Note that the Retiolitoidea is defined differently in Maletz (2014a). HME, Hirnantian mass extinction interval.
Figure 11.18 Ordovician and Silurian Normalograptidae. (A)
Normalograptus brevis
(Elles & Wood, 1906), Scania, Sweden. (B)
Normalograptus scalaris
(Hisinger, 1837), Dalarna, Sweden. (C)
Rhaphidograptus toernquisti
(Elles & Wood, 1906), Röstånga drill core, Scania, Sweden. (D)
Pseudoglyptograptus vas
Bulman & Rickards, 1968, Röstånga drill core, Scania, Sweden. (E)
Metaclimacograptus
sp., reverse view, Röstånga drill core, Scania, Sweden. (F)
Metaclimacograptus undulatus
(Kurck, 1882), two proximal ends in obverse views, Röstånga drill core, Scania, Sweden. Scale indicated by a 1 mm long bar in each photo.
Figure 11.19 Examples of the Neodiplograptidae. (A)
Metabolograptus persculptus
(Elles & Wood, 1907), Röstanga, Sweden. (B)
Parapetalolithus
sp., Röstanga, Sweden. (C)
Rivagraptus bellulus
(Törnquist, 1890), obverse view, showing alternating thecae and no median septum, Röstanga, Sweden. (D)
Rivagraptus bellulus
(Törnquist, 1890), Dalarna, Sweden, proximal end showing apertural spines on thecae. (E)
Paraclimacograptus innotatus
(Nicholson, 1869), Southern Urals, Russia (from Koren & Rickards 2004, reproduced with permission from The Palaeontological Association). (F)
Hirsutograptus
sp. cf.
H. villosus
Koren & Rickards, 1996, Arctic Canada (infrared photo by Jason Loxton). Scale indicated by 1 mm long bar in each photo.
Chapter 12
Figure 12.1 (A)
Pipiograptus
sp., a lasiograptid species with the thecal series in the centre and the lacinia outside, thecal row outline of tubarium digitally enhanced. (B)
Retiolites geinitzianus
Barrande, 1850, showing partially preserved thecal fusellum covered by the ancora sleeve. (C)
Spinograptus spinosus
Wood, 1900, showing ancora sleeve with sparse reticulum (modified from Maletz 2010b, Fig. 5A). (D)
Spinograptus tubothecalis
Kozłowska, Dobrowolska & Bates, 2013, reconstruction (based on Kozłowska et al. 2013, Fig. 5), showing hypothetical development of ancora sleeve membranes. Illustrations not to scale.
Figure 12.2 The ancora umbrella in the Petalolithinae. (A)
Petalolithus minor
(Elles, 1897), sicula with four‐pronged ancora (from Bates & Kirk 1992, Fig. 32). (B)
Pseudorthograptus inopinatus
(Bouček, 1944), showing fusellum of sicula and thecae and the circular ancora umbrella; note the paired apertural spines on first theca. (C).
Pseudorthograptus
cf.
obuti
Koren & Rickards, 1996 with spiral ancora (first illustrated in Bates & Kirk 1984, pl. 5). (D)
Hercograptus introversus
Melchin, 1999, holotype, showing preservation of fusellum (photo by M.J. Melchin). Figures not to scale.
Figure 12.3 (A)
Retiolites
sp., showing connection between thecal framework and ancora sleeve (based on Bates et al. 2005, Fig. 5A, D). (B). Thecal development in
Spinograptus
, ancora sleeve not shown (based on Bates et al., 2005, Fig. 5D).
Figure 12.4 (A) Cross‐section through thickened list with seam (arrow), showing the cortical bandages of a plectograptine retiolitid. (B)
Paraplectograptus
sp., lists with seams (arrows) indicating the presence of a membrane. (C)
Quattuorgraptus muenchi
(Eisenack, 1951), pustular ornamentation of plectograptine retiolitid.
Figure 12.5 Early ancora sleeve development. (A)
Petalolithus folium
(Hisinger, 1837) with ancora reaching to aperture of th1. (B)
Petalolithus ovatoelongatus
(Kurck, 1882) with short ancora. (C)
Petalolithus ovatoelongatus
(Kurck, 1882), showing ancora sleeve lists in proximal end. (D)
Pseudorthograptus
? sp. C, specimen with preserved thecal walls and ancora sleeve. (E)
Pseudorthograptus mutabilis
(Elles & Wood, 1907), showing thecal outlines, ancora and proximal ancora sleeve membrane. (F)
Pseudorthograptus obuti
Rickards & Koren, 1974, showing extensive proximal membranes. All from Koren & Rickards (1996), reproduced with permission from The Palaeontological Association.
Figure 12.6 (A, C–D)
Retiolites geinitzianus
Barrande, 1850. (A) Distal fragment, from Tullberg (1883). (C) Cross‐section, from Holm (1890). (D) Incomplete proximal end, from Holm (1890). (B, E–H)
Stomatograptus toernquisti
Tullberg, 1883 (=
Stomatograptus grandis
Barrande, 1850). (B) Proximal end, from Tullberg (1883). (E) Fragment, from Tullberg (1883). (F) Cross‐section showing stomata, after Bates and Kirk (1997). (G) Fragment showing ventral thecal wall and reticulum with stomata, from Holm (1890). (H) Fragment. (I)
Pseudoretiolites perlatus
(Nicholson, 1868b), from Štorch (1998b). (J)
Pseudoplegmatograptus obesus
(Lapworth, 1877), from Blumenstengel et al. (2006). (K)
Eiseligraptus cystifer
Hundt, 1959, Weinberg Hohenleuben, Thuringia, Germany. Illustrations not to scale.
Figure 12.7 The ancora development in the Retiolitinae. (A)
Rotaretiolites exutus
Bates & Kirk, 1992, reconstruction. (B)
Rotaretiolites exutus
, ancora umbrella from outside (based on Bates & Kirk 1992, Fig. 88). (C)
Pseudoretiolites perlatus
(Nicholson, 1868b), spiral ancora umbrella, side view (based on Bates & Kirk, 1992, Fig. 122: identified as
Pseudoretiolites
sp. cf.
P. decurtatus
Bouček & Münch, 1944). (D)
Pseudoretiolites
sp., clearly spiralled ancora umbrella from below (based on Bates & Kirk 1992, Fig. 167). (E)
Retiolites geinitzianus
Barrande, 1850, proximal end in side view, showing shallow ancora umbrella with hexagonal meshes (based on Bates & Kirk 1986a, Fig. 28, reproduced with permission from The Royal Society). (F)
Retiolites angustidens
(Elles & Wood, 1908), reconstructed ancora umbrella from below (based on Bates & Kirk 1997, Fig. 91). Illustrations not to scale.
Figure 12.8 Biostratigraphy of plectograptine graptolites from German glacial boulders, specimens based on line drawings by Hermann Jaeger (modified from Maletz 2008).
Figure 12.9 Examples of the Plectograptinae. (A)
Neogothograptus balticus
(Eisenack, 1951). (B)
Neogothograptus eximinassa
Maletz, 2008. (C)
Gothograptus nassa
(Wiman, 1895), proximal end of long specimen. (D)
Papiliograptus
sp. (
P. regimarginatus
in Maletz, 2010b, Fig. 2). (E)
Holoretiolites erraticus
(Eisenack, 1951). (F)
Spinograptus spinosus
(Wood, 1900). (G)
Neogothograptus alatiformis
Lenz & Kozłowska‐Dawidziuk, 2004. (H)
Neogothograptus reticulatus
Kozłowska et al., 2009. (I)
Gothograptus nassa
(Holm, 1890). (J)
Spinograptus latespinosus
Kozłowska‐Dawidziuk, 1997. (K)
Spinograptus spinosus
(Wood, 1900). SEM photos of material from north German glacial boulders (Maletz 2008, 2010b), Poland (Kozłowska‐Dawidziuk 1997) and Arctic Canada (Lenz & Kozłowska‐Dawidziuk 2004). The scale indicates 1 mm in (A–G) and 200 µm in (H–K).
Figure 12.10 Comparison of large (A) and small retiolitid colonies (B, C). (A)
Stomatograptus
sp, fragment of distal part of tubarium with four pairs of thecae. (B)
Holoretiolites helenaewitoldi
Kozłowska‐Dawidziuk, 2004, finite tubarium with four pairs of thecae. (C)
Plectodinemagraptus gracilis
Kozłowska‐Dawidziuk, 1995 with strongly reduced ancora sleeve. 1 mm scale for all specimens.
Chapter 13
Figure 13.1 The monograptid tubarium. (A)
Atavograptus ceryx
Rickards & Hutt, 1970, reconstruction. (B)
Azygograptus validus
Törnquist, 1901, reconstruction. Arrows in (A, B) indicate direction of stipe growth. (C)
Sinodiversograptus lientanensis
(Mu, 1948), multiramous streptograptid with numerous cladial branches (D) Resorption foramen for th1 in amplexograptid sicula. (E–G) Sinus and lacuna stages of primary foramen for th1 in monograptids. (D–G) adapted from Bulman 1970a, with permission from The Paleontological Institute, and also from Bulman 1970b. Illustrations not to scale.
Figure 13.2 Diagram showing the concept of the Dimorphograptidae based on the analysis of Melchin et al. (2011, Fig. 2). (A)
Normalograptus brevis
. (B)
Avitograptus avitus
. (C)
Akidograptus ascensus
. (D)
Dimorphograptus swanstoni
. (E)
Monograptus priodon
. Graptolite specimens from various sources, not to scale.
Figure 13.3 (A)
Parakidograptus acuminatus
(Nicholson, 1867b), LO 1284 t, reverse view, low relief, latex cast, Tomarp, Scania (Törnquist 1897, pl. 2, Fig. 7). (B)
Dimorphograptus
sp. cf.
D. swanstoni
Lapworth, LO 476 t, latex cast, showing delay of median septum on reverse side, Bollerup, Scania (Kurck 1882, Figs 5, 6). (C)
Rhaphidograptus toernquisti
