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- Herausgeber: John Wiley & Sons
- Kategorie: Wissenschaft und neue Technologien
- Serie: TOPA Topics in Paleobiology
- Sprache: Englisch
Bryozoa are among the most abundant yet least understood of phyla in the fossil record. These exclusively colonial animals can be traced back to the Ordovician as fossils and are common elements of sediments deposited in shallow marine environments. On occasion their calcareous skeletons are sufficiently numerous to produce bryozoan limestones. The potential of bryozoans in facies analysis, and their use in macroevolutionary studies, have both been widely recognised, but to date have been incompletely exploited.
Bryozoan Paleobiology brings together the scattered research on living and fossil bryozoans in broad and profusely illustrated overview that will help students and researchers alike in understanding this fascinating group of animals. Beginning with the basics of bryozoan morphology, ecology and classification, the book progresses from the smallest scale of skeletal ultrastructure, to the largest of bryozoan distributions in time and space. On the way, topics such as the origin of zooidal polymorphism and macroevolutionary trends in colony forms are covered. Case studies illuminate these topics, and areas in which further research is particularly required are highlighted.
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Table of Contents
Cover
Preface
Acknowledgements
1 Introduction
1.1 Zooids
1.2 Colonies
1.3 Colony Propagation
1.4 Ecology
1.5 Taxonomy
1.6 Collecting and Studying Fossil Bryozoans
2 Biomineralization and Geochemistry
2.1 Skeletal Wall Types
2.2 Pores and Pseudopores
2.3 Skeletal Growth
2.4 Skeletal Ultrastructure
2.5 Spines
2.6 Mineralogy
2.7 Geochemistry
3 Zooid Morphology and Function
3.1 Autozooids
3.2 Ontogenetic and Astogenetic Variations
3.3 Ancestrulae
3.4 Polymorphism
3.5 Reproductive Polymorphs
3.6 Active Defensive Polymorphs
3.7 Structural Polymorphs
3.8 Spinozooids
3.9 Rhizooids
3.10 Cleaning Polymorphs
3.11 Locomotory Polymorphs
3.12 Microenvironmental Variability
3.13 Zooid‐level Skeletal Structures
3.14 Extrazooidal Structures
3.15 The Cormidial ‘Zooids’ of Advanced Cheilostomes
4 Colony Morphology and Function
4.1 Colony‐form Classifications
4.2 Growth and Colony‐form
4.3 Functional Morphology of Colony‐form
4.4 Colony Propagation in Lunulitiform Bryozoans
4.5 Multizooidal Feeding Morphologies
4.6 Life Histories
4.7 Colonial Integration
4.8 Endolithic and Etching Bryozoans
5 Biotic Interactions
5.1 Competition
5.2 Predation
5.3 Symbioses
5.4 Bryozoans as Habitat Providers
6 Ecology and Palaeoecology
6.1 Bryozoan Habitats
6.2 Bryozoans in Reefs and Mounds
6.3 Colony‐form and Palaeoenvironments
6.4 Depth Distributions and Palaeobathymetry
6.5 Bryozoans as Sediment Producers
6.6 Taphonomy
6.7 Palaeoclimatology and Zooid Size
7 Biogeography
7.1 Cosmopolitan vs. Endemic Distributions
7.2 Modes of Dispersal
7.3 Biogeography of Bryozoans at the Present Day
7.4 Latitudinal Diversity Gradient
7.5 Palaeobiogeography
8 Phylogeny
8.1 Relationships with Other Phyla
8.2 Inter‐relationships of Bryozoan Classes
8.3 Inter‐relationships of Bryozoan Orders
8.4 Morphological Phylogenies
8.5 Molecular Phylogenies
9 Evolution and Fossil History
9.1 Phanerozoic Bryozoan Diversity
9.2 Cambrian Bryozoans?
9.3 Great Ordovician Biodiversification Event
9.4 End‐Ordovician Extinction
9.5 Devonian Extinctions
9.6 Permian Mass Extinctions
9.7 Triassic Diversity and Mass Extinction
9.8 Jurassic Cyclostome Radiation
9.9 Cretaceous–Palaeogene Radiations
9.10 End Cretaceous and Danian Extinctions
9.11 Convergence
9.12 Palaeostomates and Post‐Palaeozoic Cyclostomes Compared
9.13 Frontal Shield Evolution in Ascophoran Cheilostomes
9.14 Cyclostomes vs. Cheilostomes
9.15 Colony‐forms Through Geological Time
9.16 Evolutionary Tempo in Bryozoans
10 Prospective Future Research
10.1 Biomineralization
10.2 Polymorphism
10.3 Environmental Distributions of Colony‐forms
10.4 Taphonomy
10.5 Small Palaeozoic Bryozoans
10.6 Phylogeny and Classification
References
Index
Supplemental Images
End User License Agreement
List of Tables
Chapter 1
Table 1.1 Morphological characteristics of the three classes of bryozoa...
Table 1.2 Main morphological features and geological ranges of the orde...
Chapter 4
Table 4.1 Bryozoan colony‐form nomenclature derived from exemplar cheilostome...
List of Illustrations
Chapter 1
Figure 1.1 Basic cheilostome bryozoan labelling some of the most important...
Figure 1.2 Schematic figure of polypide cycling in a bryozoan in which bro...
Figure 1.3 Comparative external skeletal morphology of the feeding zooids ...
Figure 1.4 Typically tubular zooidal skeletons of a stenolaemate compared ...
Figure 1.5 Cutaway diagram of a branch from a ramose trepostome colony (af...
Figure 1.6 Correlation between the proportion of brooding zooids within co...
Figure 1.7 Life history of the fenestrate bryozoan
Archimedes
(based on Mc...
Figure 1.8 Statoblasts produced by freshwater phylactolaemate bryozoans fo...
Figure 1.9 Dormancy and regrowth in the Cretaceous cheilostome ‘
Micropora’
...
Figure 1.10 Linear growth rates of bryozoan species using data compiled fr...
Figure 1.11 Immured fossil ctenostome bryozoans: (A) mould bioimmuration o...
Figure 1.12 Examples of fossil bryozoans belonging to the eight orders pos...
Chapter 2
Figure 2.1 Surface textures of skeletal walls in bryozoans: (A) interior wal...
Figure 2.2 Stylized vertical section through the base and stem of a hypothet...
Figure 2.3 Transversely broken branch of the cheilostome
Chiplonkarina campb
...
Figure 2.4 Pores and pseudopores in bryozoan skeletal walls: (A) round pseud...
Figure 2.5 Skeletal wall ultrastructural fabrics in some recent bryozoans: (...
Figure 2.6 Unusually pristine preservation of Palaeozoic bryozoans occurs in...
Figure 2.7 Cyclostome mural spines (A–D), and cheilostome spinules (E) and s...
Figure 2.8 Palaeostomate styles: (A) thin section of the esthonioporine
Ibex
...
Figure 2.9 Mineralogical compositions of cheilostome bryozoans in biotas fro...
Figure 2.10 Mineralogical composition of cheilostome bryozoan skeletons in s...
Figure 2.11 Frequency distributions of exozonal wall thickness in ramose tre...
Figure 2.12 Dissolution of the outer aragonitic skeleton that formed the fro...
Figure 2.13 Skeletal mineralogy of 1051 specimens of marine bryozoans plotte...
Chapter 3
Figure 3.1 Opercula closing the orifices of cheilostomes (A–E) and the apert...
Figure 3.2 Frequency distribution of zooid size, expressed as a proxy of fro...
Figure 3.3 Basic mechanism of lophophore protrusion and retraction in bryozo...
Figure 3.4 The four main modes of tentacle sheath eversion, and therefore lo...
Figure 3.5 Morphological features of the zooidal skeletons in some anascan (...
Figure 3.6 Longitudinal section of a polypide situated within the elongate l...
Figure 3.7 Schematic vertical and horizontal sections through a bryozoan zoo...
Figure 3.8 Hexagonally close‐packed arrays of lophophores with basic zooid o...
Figure 3.9 Diaphragms and peristomes in some cyclostome bryozoans: (A) longi...
Figure 3.10 Ontogenetic changes in autozooids of the ascophoran cheilostome
Figure 3.11 Anascan cheilostome
Wilbertopora
sp. with the opesiae of three a...
Figure 3.12 Zones of astogenetic change: (A) early astogeny in the uniserial...
Figure 3.13 Ancestrulae in stenolaemate (A–F) and cheilostome (G–J) bryozoan...
Figure 3.14 Cyclostome gonozooids for larval brooding showing variation in s...
Figure 3.15 Cheilostome ovicells used for larval brooding: (A) band of ovice...
Figure 3.16 Primitive ovicell comprising a cage of several spines formed by ...
Figure 3.17 Spinose cheilostome ovicells: (A) crescent of circular bases on ...
Figure 3.18 Female polymorphs in cheilostomes: (A) gonozooid (centre) of the...
Figure 3.19 Possible larval brood chambers in palaeostomates: (A) shallow de...
Figure 3.20 Sexual polymorphism in the living cheilostome
Celleporella
sp., ...
Figure 3.21 Comparison of an autozooid and an avicularium in the anascan che...
Figure 3.22 Schematic diagram of some of the main types of avicularia found ...
Figure 3.23 Cheilostome avicularia: (A) ‘bird’s head’ avicularium with intac...
Figure 3.24 Polymorphic cheilostome zooids with modified opercula/mandibles:...
Figure 3.25 Variability in the avicularia of mid‐Cretaceous species of the e...
Figure 3.26 Cladistic phylogenetic tree of species of the early neocheilosto...
Figure 3.27 Eleozooids of eleid cyclostomes: (A)
Meliceritites
sp. with a ro...
Figure 3.28 Kenozooids in some cheilostomes (A–D) and cyclostomes (E–H): (A)...
Figure 3.29 Thin sections of palaeostomate bryozoans showing internal morpho...
Figure 3.30 Cheilostome spinozooids: (A) oblique view of long, paired, basal...
Figure 3.31 Underside of a fossil colony of
Schizorthosecos interstitia
with...
Figure 3.32 Nanozooids of cyclostomes: (A) primary nanozooids with tiny aper...
Figure 3.33 Cheilostome vibracula: (A) crescent‐shaped vibracula on the dors...
Figure 3.34 Some skeletal structures of bryozoan zooids: (A)–(B) cap‐like ap...
Figure 3.35 Thin section of the trepostome
Stenoporella romingeri
(Carbonife...
Figure 3.36 Trepostome living chambers of different proportions and morpholo...
Figure 3.37 Tangential thin section of dark, horseshoe‐shaped lunaria in aut...
Figure 3.38 Ascophoran cheilostome frontal shields: (A) spinocystal (cribrim...
Figure 3.39 Orificial characters in ascophoran cheilostomes: (A) paired cond...
Figure 3.40 Longitudinal thin section of the cystoporate
Fistulipora elegant
...
Figure 3.41 Surface of a branch of the cyclostome
Hornera erugata
showing au...
Figure 3.42 Cormidial skeleton of an ascophoran cheilostome zooid formed of ...
Chapter 4
Figure 4.1 Growth patterns in some encrusting bryozoan colonies: (A) runner ...
Figure 4.2 Small encrusting cheilostome colonies from the Pleistocene Red Cr...
Figure 4.3 Fungiform and conical bryozoan colony‐forms: (A) fungiform cyclos...
Figure 4.4 Character states for bryozoan colony‐forms related to the six cat...
Figure 4.5 Small colonies of encrusting stenolaemate bryozoans with circumfe...
Figure 4.6 Cheilostome row bifurcation showing change in zooid width along r...
Figure 4.7 Spiral budding around the ancestrula (a) in the cheilostome
Bisel
...
Figure 4.8 Axial structures visible in transverse sections of some cryptosto...
Figure 4.9 Variation in colony shape between sites experiencing different hy...
Figure 4.10 The distinctive cyclostome
Terebellaria ramosissima
(Jurassic, B...
Figure 4.11 Superstructures developed above the colony surface: (A) in the f...
Figure 4.12 Extrazooidal calcification obscuring the zooids at base of a col...
Figure 4.13 Variation in the shapes of colonies according to microhabitat in...
Figure 4.14 Articulated colonies of cyclostomes (A–C) and cheilostomes (D–F)...
Figure 4.15 (A) holdfast of unidentified ptilodictyine cryptostomes with two...
Figure 4.16 Reconstruction of a colony of the Carboniferous cystoporate
Meek
...
Figure 4.17 Two living colonies of the lunulitiform cheilostome
Selenaria ma
...
Figure 4.18 Small free‐living colonies of the cheilostome
Volviflustrellaria
...
Figure 4.19 Conescharelliniform cheilostome colonies figured upside down in ...
Figure 4.20 Comparison of sexual (larvally recruited) and asexual (clonal) c...
Figure 4.21 Cycle of clonal propagation by peripheral fragmentation in the f...
Figure 4.22 Schematic vertical section through a bryozoan with monticules. T...
Figure 4.23 Monticule formed of non‐feeding kenozooids surrounded by radial ...
Figure 4.24 Schematic transverse section through two branches of a cribrate ...
Figure 4.25 Schematic transverse section through three branches of a fenestr...
Figure 4.26 Inferred multizooidal feeding currents in cross sections of the ...
Figure 4.27 Ctenostome borings (A–D) and cheilostome etchings (E–F); (A) par...
Chapter 5
Figure 5.1 Marginal overgrowth and fouling. Arrows show centrifugal growth d...
Figure 5.2 The three possible outcomes – overgrowth, stand‐off, and reciproc...
Figure 5.3 Encrusting cheilostomes with different budding processes recogniz...
Figure 5.4 Some encrusting cyclostomes are able to elevate their growing edg...
Figure 5.5 Examples of overgrowths involving bryozoans in fossil communities...
Figure 5.6 Putative predatory drillholes: (A) eleid cyclostome
Meliceritites
...
Figure 5.7 Intramural buds, possibly indicating reparative growth after pred...
Figure 5.8 Bryozoan symbionts: (A) scleractinian coral
Culicia parasitica
on...
Figure 5.9 Pseudoborings (bioclaustrations) and borings in palaeostomates: (...
Figure 5.10 Thin section of the inferred paguroid symbiont
Reptomultisparsa
...
Figure 5.11 Bryozoan‐associated microendoliths and microbial threads: (A) mi...
Chapter 6
Figure 6.1 Abiotic (A–B) and biotic (C–I) fossil substrates colonized by bry...
Figure 6.2 Sheet‐like growth of the cheilostome
Tamanicella lapidosa
from a ...
Figure 6.3 Relative importance of bryozoan colony‐forms, based on combined a...
Figure 6.4 Depth gradients in ratio of encrusting to erect species (black sq...
Figure 6.5 Positive relationship between bryozoan species diversity and dept...
Figure 6.6 Distribution of seven bryozoan colony‐forms according to inferred...
Figure 6.7 Contrasting morphologies of shallow‐ and deep‐water colonies of t...
Figure 6.8 Contrast between the palaeolatitudinal distributions of bryozoan‐...
Figure 6.9 Taphonomy of bryozoans with different colony‐forms and substrates...
Figure 6.10 Latitudinal changes in autozooid size in species belonging to tw...
Figure 6.11 Decline in the size of avicularia with increasing summer tempera...
Figure 6.12 Correlation in 29 species of Recent cheilostomes between within‐...
Figure 6.13 Mean annual range in temperature at the present day (square boxe...
Chapter 7
Figure 7.1 Beta diversity of North Atlantic bryozoan species (data from Clar...
Figure 7.2 Paratethyan Middle Miocene biogeographical links showing numbers ...
Chapter 8
Figure 8.1 Phylogenetic relationships between the lophophorate phyla Brachio...
Figure 8.2 Basal encrusting bryozoans: (A) dried colony of the ctenostome
Ar
...
Figure 8.3 (A) Phylogeny of stenolaemate orders (Timanodictyina excluded)...
Figure 8.4 Palaeozoic crownoporid cyclostomes that may belong in the stem‐gr...
Figure 8.5 Phylogenetic tree of the cheilostome
Macropora
based on morpholog...
Figure 8.6 Molecular phylogeny of cyclostome bryozoans with traditional subo...
Figure 8.7 Fixed and free‐walled organizations in stenolaemates as exemplifi...
Figure 8.8 Molecular phylogeny of selected cheilostome bryozoans based on th...
Chapter 9
Figure 9.1 Generic diversity of fossil bryozoans through time.
Figure 9.2 Mean species diversity in bryozoan assemblages through geological...
Figure 9.3 Phanerozoic species‐level diversity of bryozoans based on the dat...
Figure 9.4 Diversity of bryozoan genera and species through the stages of th...
Figure 9.5 Rates of first appearance of major traits in cyclostome (pale gre...
Figure 9.6 Homeomorphy is striking between the Palaeozoic cryptostome
Worthe
...
Figure 9.7 Gonozooid of the eleid cyclostome
Reptomultelea oceani
with the o...
Figure 9.8 Homoemorphy between the cheilostome
Chiplonkarina
and cerioporine...
Figure 9.9 Palaeozoic fenestellid bryozoan and two recent homeomorphs, all w...
Figure 9.10 Homeomorphic lunulitiform cheilostomes belonging to Lunulitidae ...
Figure 9.11 Intermediate morphologies in ascophoran cheilostome showing the ...
Figure 9.12 Generic diversity from the Jurassic to the Holocene in cyclostom...
Figure 9.13 Overgrowths between cheilostome and cyclostome bryozoans recorde...
Figure 9.14 Moving average trend of changes in the total number of bryozoan ...
Figure 9.15 Average proportions of species in bryozoan assemblages through g...
Figure 9.16 Parallel increase through time in the proportion of unilaminate ...
Figure 9.17 Punctuated pattern of morphological evolution in the cheilostome...
Supplemental Images
Plate 1 Recent bryozoans in their natural habitats. (A)–(B) Rottnest Island,...
Plate 2 Fossil bryozoans in the field. (A) Palaeozoic sequences of interbedd...
Plate 3 Bryozoan‐rich carbonates. (A) Bedding plane of Late Ordovician limes...
Plate 4 Bryozoans in thin section. (A) Brown deposits in the zooidal chamber...
Plate 5 Lophophores. The tentaculate feeding structures of some living bryoz...
Plate 6 Bryozoan colony‐forms. (A)–(C) Encrusting: (A) sheet‐like cheilostom...
Plate 7 Bryozoan colony disparity 1. (A) Flabellate colony of the cheilostom...
Plate 8 Bryozoan colony disparity 2. (A) Fenestrate
Reteporina reticulata
sh...
Plate 9 Cheilostome frontal wall types. (A) Anascan with a wide opesial open...
Plate 10 Indicators of multizooidal current systems, substrates, and bryozoa...
Plate 11 Conchicole symbiont bryozoans. Bryozoans forming symbioses with kno...
Plate 12 Bryozoan colony disparity 3. (A)–(C) Three distantly related cheilo...
Guide
Cover
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Bryozoan Paleobiology
Paul D. Taylor
Natural History MuseumLondon, UK
This edition first published 2020© 2020 Natural History Museum
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Library of Congress Cataloging‐in‐Publication Data
Names: Taylor, Paul D., author.Title: Bryozoan paleobiology / Paul D. Taylor, Natural History Museum, London, UK.Description: Hoboken, NJ : Wiley‐Blackwell, 2020. | Series: Topa topics in paleobiology | Includes bibliographical references and index.Identifiers: LCCN 2020012771 (print) | LCCN 2020012772 (ebook) | ISBN 9781118455005 (paperback) | ISBN 9781118454985 (adobe pdf) | ISBN 9781118454992 (epub)Subjects: LCSH: Bryozoa, Fossil. | Bryozoa–Biology. | Bryozoa–Ecology.Classification: LCC QE798 .T39 2020 (print) | LCC QE798 (ebook) | DDC 564/.67–dc23LC record available at https://lccn.loc.gov/2020012771LC ebook record available at https://lccn.loc.gov/2020012772
Cover Design: WileyCover Image: © Screw axes of the bryozoan Archimedes from the Carboniferous of Alabama© Trustees of the Natural History Museum, London
Preface
Until the early nineteenth century, natural historians were puzzled by organisms at the time known as zoophytes: were they animals (zoo‐), plants (‐phyte), or something in between? Perhaps they were even the common ancestors of animals and plants? Zoophytes as then conceived included sponges, corals, and coralline algae, as well bryozoans, the subject of this book. The so‐called ‘zoophyte problem’ greatly engaged Charles Darwin when he set sail from Plymouth Sound on board HMS Beagle in December 1831. Indeed, Darwin’s first scientific paper, which was read by his mentor Robert Grant before both the Wernerian and Plinian societies when Darwin was a medical student at the University of Edinburgh, had concerned species of zoophytes we now know to be the bryozoans Flustra and Carbasea. And he made detailed observations of the intriguing behaviour of the peculiar ‘bird‐head’ structures in bryozoans dredged off Patagonia during the Beagle voyage (Keynes 2003).
Zoophyta has long been abandoned as a taxonomic group and we now know much more about the biology of the diverse animals formerly grouped together under this name. Nevertheless, our anthropocentric view of life still makes it difficult to comprehend these peculiar plant‐like, colony‐forming animals that are so very different from the dogs, spiders, and other animals we encounter daily. In their immobility and growth, colonial animals resemble plants but they are not autotrophs that photosynthesize but are instead heterotrophs that must obtain their nutrition by consuming other plants or animals. The resemblance in form between benthic colonial animals and higher terrestrial plants reflects not only their sessile lifestyles but also a shared modular construction (Hughes 2005).
Major advances have been made in recent years in our knowledge of some types of colonial animals, especially corals reflecting their importance in reef ecosystems. But bryozoans have been far less intensively studied and remain poorly understood, particularly by non‐specialists. This belies the fact that the Bryozoa are a diverse phylum, with more than 6000 named species living today and a predicted 5000 more yet to be described (Gordon and Costello 2016), and have ecological importance in many marine and some freshwater habitats at the present day (Plate 1). Their rich fossil record – comprising more than 1300 genera – gives them considerable geological significance as well: bryozoans are common fossils in Ordovician–Holocene rocks deposited in shallow marine environments (Plates 2–3). Sometimes bryozoan colonies inhabiting the seafloor have supplied sufficient carbonate skeletal material to the sediment to form bryozoan limestones.
While the utility of bryozoans in applied geology as zonal fossils or palaeoenvironmental indicators has been limited, which is one of the reasons they have attracted too little attention from geologists, Bryozoa are the best phylum in which to study the evolution of coloniality. Furthermore, their skeletons can preserve key aspects of their life histories such as the timing of sexual reproduction, and overgrowths between encrusting bryozoans provide rare instances in which competition is ‘frozen’ in the fossil record. Bryozoans living today and preserved as fossils are also intriguing and frequently enigmatic creatures that offer great opportunities for making new discoveries.
This book brings together information from the scattered literature on living and fossil bryozoans with the intention of providing a broad overview of the palaeobiology (and biology) of these fascinating animals. It updates the standard general text on bryozoans written by Ryland (1970), and the excellent book of McKinney and Jackson (1989) focusing on the adaptive morphology of bryozoans. Beginning with an introduction covering the basics of bryozoan morphology, ecology, and systematics, Bryozoan Paleobiology progresses from the smallest to the largest scale: from the skeleton and its microstructure, via zooid‐ and colony‐level features and functions to biotic interactions, ecology, biogeography, phylogeny, and evolution. Important topics are highlighted – such as zooidal polymorphism – and critical areas for future research are identified.
Our understanding of bryozoan palaeobiology inevitably depends on making comparisons with the biology of living bryozoans, hence the large number of references to neontological studies. In common with other synthesis of this kind, personal experience has played a leading role on what is included and conversely what is excluded. While this undoubtedly colours many of the interpretations presented, I hope to have avoided unjustifiably strong biases.
Paul D. Taylor
London
November 2019
Acknowledgements
Serendipity invariably has a role in the career pathways of scientists and academics. I was fortunate in finding a Jurassic bryozoan during my independent geological mapping project while an undergraduate at the University of Durham in the early 1970s, and equally fortunate when, unbeknownst to me at the time, my main lecturer in palaeontology, Gilbert Larwood, just happened to be a bryozoan specialist. Gilbert took me under his wing, became my first mentor, and went on to supervise my doctoral research. I then enjoyed two years as a postdoc at the University College of Swansea where I was attracted by the presence of John Ryland’s group studying living bryozoans, including Peter Hayward and John Thorpe to whom I owe a major debt for teaching me so much about bryozoan biology. Another influence in Swansea was Derek Ager. Derek was a highly original thinker about the fossil record and palaeoecology who did not receive the credit he deserved.
After Swansea I was appointed to a research post on bryozoans in the Department of Palaeontology at the then British Museum (Natural History). Pat Cook, my counterpart in the Department of Zoology, taught me never to be too dogmatic when making statements about bryozoans – there are many surprises waiting around the corner to embarrass the unwary in the study of these diverse, complex, and often enigmatic animals.
I will forever be grateful to Daphne Lee at the University of Otago who encouraged me to apply for a William Evans Fellowship there in the late 1980s. This proved to be my personal Beagle voyage. Keith Probert at the Portobello Marine Laboratory introduced me to the wonderful diversity of animals on the Otago Shelf that interact with bryozoans, not just the hermit crabs I was there to study, while Doug Campbell and Dave MacKinnon (Canterbury University) took me in the field to see the rich Cenozoic bryozoan faunas of New Zealand (Doug’s son Hamish was later to guide me around the glorious Chatham Islands on two memorable geological fieldtrips). My collaborations with Dennis Gordon (NIWA, Wellington), the global authority on cheilostome bryozoans, describing the taxonomy of fossil and Recent bryozoans of New Zealand, began at this time. While at Portobello I was contacted by Richard Boardman (Smithsonian Institution) and F. Ken McKinney (Appalachian State University) who were partway through a study of the peculiar cyclostome bryozoan Cinctipora, one of the bioconstructional species on the Otago Shelf. Their invitation to join them in this research marked the beginning of my interest in bryozoan biomineralization, the skeleton of Cinctipora showing a striking ultrastructure (Boardman, McKinney, and Taylor 1992).
Before migrating to the world of ornithology, Mike Weedon undertook two postdocs with me at the NHM on bryozoan skeletal ultrastructures. Other scientists to whom I am indebted for various collaborative biomineralization projects include Chiara Lombardi (ENEA, La Spezia), Piotr Kuklinski (Polish Institute of Oceanology, Sopot), Noel James (Kingston University, Ontario), and Bill Schopf (UCLA).
The late F. Ken McKinney, who has already been mentioned, deserves my special thanks. He offered so many fresh insights and was a wonderful person to work with, as well as a great friend. To Dennis Gordon and Ken McKinney, I must add a third long‐standing collaborator, Mark Wilson (College of Wooster, Ohio). It has been a delight working with Mark over the years and benefitting from his knowledge of hard substrate palaeoecology combined with a clarity of thought and expression that makes him such a superb teacher.
More recently, I have been fortunate to collaborate with two of the rising stars of bryozoology. Andrea Waeschenbach (Life Sciences, NHM) undertook a postdoc with Tim Littlewood and me on bryozoan molecular phylogeny, ferreting out the false data in GenBank and adding a lot of new data of her own to show the power of molecular sequences in understanding bryozoan evolution. Lee Hsiang Liow (University of Oslo) has been employing her analytical and modelling prowess to investigate biotic interactions through geological time, in particular competition for substrate space, a field where bryozoans have the potential to make a wider impact in macroecology and evolutionary ecology.
Supervision of doctoral students has served to broaden my perspectives. In chronological order, I thank them all: Richard Carthew, Julian Hammond, Caroline Buttler, Jon Todd, Kevin Tilbrook, Sian Evans, Phil Watts, Jo Snell, Lais Ramalho, Tanya Knowles, Scott Tompsett, Caroline Sogot, Emanuela Di Martino, and Peter Batson.
Others with whom I have been privileged to collaborate and learn from include Ehrhard Voigt, Eckart Håkansson, Andrei Grischenko, Andrej Ernst, Roger Cuffey, Alan Cheetham, Aaron O’Dea, Eckart Håkansson, Andrew Ostrovsky, Al MacGowan, Ma Junye, Seo Ji Yun, Shun Mawatari, Kamil Zágoršek, Noel James, Jeremy Jackson, Antonietta Rosso, Joachim Scholz, Beth Okamura, Tim Palmer, Andrew Smith, Silviu Martha, Urszula Hara, Patrick Wyse Jackson, Hans Arne Nakrem, Matt Dick, Scott Lidgard, David Jablonski, Abby Smith, Seabourne Rust, Kjetil Voje, Loïc Villier, Leandro Vieira, Francoise Bigey, Helen Jenkins, Consuelo Sendino, and Mary Spencer Jones.
I am grateful to the photographers of the NHM, particularly Harry Taylor, Phil Crabb, and Phil Hurst, for their skilled macrophotography. Most of the SEM images were taken by me during countless productive hours spent in the NHM’s imaging and analysis laboratories where Alex Ball, Chris Jones, and several others were always at hand when help was needed. Piotr Kuklinski, Andrej Ernst, Caroline Buttler, and Thomas Schwaha are thanked for generously providing additional images used in this book.
The study of bryozoans has never attracted the funding it deserves – as Sir David Attenborough once remarked, you can’t care about something unless you know it exists. Alas, far too few people are aware of the existence of these perenielly ‘unfashionable’ animals, let alone have any inkling of their fascinating natural history. Nevertheless, I have benefitted over the years from grants awarded by the Natural Environment Research Council (NERC), Leverhulme Trust, European Union, Royal Society, British Council, and the Japan Society for the Promotion of Science (JSPS), all of which I gratefully acknowledge.
Finally, I would like to thank Caroline Buttler, Andrea Waeschenbach, Lee Hsiang Liow, and Emanuela Di Martino for their comments on the manuscript, and Louise Spencely for her skilled copy editing.
