INDEX PENNATULACEA

ANNOTATED BIBLIOGRAPHY AND INDEXES OF THE SEA PENS
OF THE WORLD 1469-2002
(COELENTERATA: OCTOCORALLIA)

By
Gary C. Williams
Department of Invertebrate Zoology
California Academy of Sciences
Golden Gate Park
San Francisco, California 94118

A reasonably comprehensive bibliography of the living and fossil pennatulacean Octocorallia is presented with the goal of including all published accounts regarding the biology of the sea pens. This compilation of approximately 1000 citations represents 533 years of published research. Complete citations for periodicals are used throughout, rather than use of abbreviated notations for journal titles. Many of the citations listed here are annotated with brief descriptive notes in bold print. Taxonomic, geographic, and field-of-study indexes to the literature are included, as well as a synopsis of historical periods in the study of the Pennatulacea.

A B C D E F G H I J K L M N O P Q R S T U V W X Y Z

CONTENTS

Introduction

Acknowledgements

Historical Account

Pre-Linnean Period (1554-1757)
The Linnean Period (1758-1858)
The Darwinian Period (1859-1899)
The Early Twentieth Century (1900-1949)
The Late Twentieth Century (1950-present)

Synonyms and misspelled generic names of Pennatulacea

Taxonomic Index

Geographic Index

Africa - southern
Africa - western
Arctic Ocean
Australia - southern
Australia - western
Indo-Pacific (including the Red Sea)
Japan
New Zealand
North America - Pacific
North Atlantic and Mediterranean (Europe and eastern North America)
Pacific Ocean - northern
South America (Atlantic)
South America (Pacific)
Southern Oceans (Antarctica and the subantarctic)
Western Tropical Atlantic and Caribbean
Worldwide

Subject Index

Behaviour
Bibliography
Biography (Regarding Authors of Pennatulacean Research)
Bioluminescence
Corals (Pre-Linnean General Natural History)
Ecology and Biotic Interactions
General Natural History and General Biology
Histology
History of Science (Regarding Coelenterates and/or Sea Pens)
Molecular Biology and Genetics
Morphology and Ultrastructure
Natural Products Chemistry, Biochemistry, and Toxicology
Paleontology (Mesozoic and Cenozoic Fossil Taxa)
Paleontology (Vendian Frond-like Fossil Taxa)
Photography (Color Photographs of Living Sea Pens)
Phylogeny, Cladistics, Evolution, and Biogeography
Physiology and Cell Biology
Polymorphism, Phenotypic Variability, and Biodiversity
Quoted Passages
Reproductive and Developmental Biology
Taxonomy, Distributional & Bathymetric Records

INTRODUCTION

The pennatulaceans, commonly known as sea pens and sea pansies, are a highly specialized and distinct group of sessile benthic coelenterates. They are distributed throughout the world's oceans from the polar seas to the tropics, and at all depths from the intertidal to over 6200 meters. Morphologically, sea pens are highly diverse and exhibit morphological changes as evolutionary events within different lineages such as bilateral symmetry, concentration and localization of feeding polyps, the development of lateral processes as polyp leaves or ridges, and the reduction in the number and size of sclerites. As octocorallian coelenterates, pennatulaceans are characterized by having eight pinnate tentacles surrounding the mouth of each polyp, and eight mesenteries. Unlike other octocorals, however, mature colonies develop from a single large polyp (the oozooid) that produces secondary zooids by lateral budding of its body wall. Also unique to the Pennatualacea, is the character of an unbranched muscular peduncle, which anchors the animal by peristaltic contractions into soft substrata such as sand, mud, or abyssal ooze. The secondary polyps (larger autozooids for feeding and reproduction, and smaller siphonozooids for internal water circulation) are restricted to the distal rachis, which is also unbranched but may have laterally-produced polyp leaves.
The ability to inhabit soft substrata has allowed several abyssal-dwelling sea pens to have nearly cosmopolitan distributions. Despite these very widespread geographic ranges, pennatulacean species diversity of the deep-sea is relatively low, and may be attributable to a combination of factors including: relatively low energy input and depauperate productivity in the abyssal environment, coupled with a relative lack of ecological diversity.
Thirty-two genera in fifteen families of living pennatulaceans are currently recognized. Of the 436 nominal species names described in the literature, approximately one half are currently considered valid. Major monographic works on the Pennatulacea include Kükenthal (1915), Hickson (1916), and Williams (1990, 1995a).
The sparsely represented pennatulacean fossil record extends back to the Cretaceous Period and consists primarily of virgulariid fossil taxa in the genera Graphularia and Virgularia. Bayer (1956: 228) observed that fossils of Graphularia closely resemble the axial structure of the genus Stylatula, and the two may be congeneric. Problematic and controversial frond-like taxa that resemble pennatulid sea pens are described from the Vendian Period of the late Precambrian. In fact, the coming of age of Precambrian paleobiology in the 1980's has produced a wealth of publications dealing with the Ediacaran or Vendian biota. Many of the Vendian frond-like taxa have been aligned with the Pennatulacea by various authors, but these interpretations remain highly disputatious. Because of the importance of recent fossil discoveries, which have helped to bridge the gap between fossil biotas of the Precambrian and the Cambrian, many of the references on these Vendian pennatulacean-like organisms are included here, but the coverage should in no way be considered comprehensive.
The idealistic goal of this work is to produce a bibliography of all published works pertaining in part or in full to the pennatulacean octocoral coelenterates. Although no work of this kind can claim to be absolutely comprehensive, this bibliography can at least be considered reasonably complete, certainly more so than any previously published similar work. The fine bibliography of octocorals by Bayer (1996) concentrates on taxonomic works for all of the Octocorallia, but does not claim to be inclusive for Pennatulacea or fields of study outside of taxonomy.
The following annotated and alphabetical bibliography to the Pennatulacea of the world is derived from a variety of sources - primarily the Zoological Record (1864-1998), as well as the author's research library. Other sources include Bayer (1981), Bayer (1996), and Hickson (1916). My annotations are in square brackets
and bold print at the end of many of the citations = [sample annotation].
This work is intended for a variety of users including professional biologists, biology students, fisheries biologists and marine resource managers.
The intent of the presentation is to make this work as user friendly as possible. In order to facilitate this goal, full citations are given for the vast majority of the references listed, as abbreviations for periodical publications have proved in the past to be confusing, inconsistent, or often unusable.
The Melvyl System, library data base of the University of California, [http://www.melvyl.ucop.edu] was used extensively to acquire complete citations for periodicals. Other sources used for abbreviations and complete citations of periodicals include the List of Serial Publications in the BM(NH) Library, the World List of Scientific Periodicals 1900-1960, and Biosis Serial Sources.
For the most recently recognized valid species names of Pennatulacea, see Williams (1995a and 1997f).

ACKNOWLEDGMENTS

I am grateful to Larry Currie (Librarian, California Academy of Sciences, San Francisco), who was instrumental in providing expertise and in locating several references.

I thank Leen van Ofwegen (Nationaal Natuurhistorisch Museum, Leiden), Phil Alderslade (Museum and Art Gallery of the Northern Territory, Darwin), Frederick Bayer (National Museum of Natural History, Washington, D.C.), James Morin (Cornell University), Michael Ghiselin and Alan Leviton (California Academy of Sciences, San Francisco) for their valuable contributions and comments.

HISTORICAL ACCOUNT

Pre-Linnean Period (1554 - 1757)
E. Newton Harvey (1952: 170), writing about the history of research regarding pennatulacean bioluminescence, relates the following, "Although the Romans knew of sea pens, referring to them as 'Penna marina', the sea feather, or as 'Mentula alata', the winged penis, the ability to luminesce does not appear to have been recorded. Probably the first mention of luminous sea pens comes to us in the famed 'De lunariis' and the 'Historia Animalium' of Conrad Gesner, published in the middle sixteenth century."
The earliest published accounts describing aspects of the natural history of octocorals including the sea pens are Pliny the Elder (1469), Guillaume Rondelet (1554-1555), Conrad Gesner (1555 and 1558), Ferrante Imperato (1599), and Captain Lancaster (1601). Other noteworthy pre-Linnean works dealing at least in part with the Octocorallia are those of Francisci Erasmi (1668), Paolo Boccone (1674), Hans Sloane (1707), Jacques Barrelier (1714), Herman Boerhaave (1720 and 1727), Christlob Mylius (1753, 1754), and John Ellis (1753, 1755).
Pennatulacean bioluminescence is at least mentioned by François Boussuet (1558), Caspar Bauhin (1620, 1671), Ulisse Aldrovandi (1642, 1648), Johann Bauhin (1650-51), and Thomas Shaw (1738-46).
Louis Agassiz (1860) praised the perspicacity of early workers such as Rondelet and Gesner, who first established the animal nature of the "acalephs" (many of the taxa that we now regard as coelenterates), departing from the earlier views of Aristotle and Pliny, who compared them to plants (the "zoöphytes").
Pertaining to Rondelet, Agassiz (1860: 8) states, "...we are chiefly indebted to Rondelet for contributions to the natural history of the Acalephs. He was, indeed, not only better acquainted with the inhabitants of the Mediterranean than all his predecessors, but he knew them even more accurately than any naturalist that lived before the present century."
In his praise of sixteenth century naturalists, Agassiz (1860: 10) continues, "It has been a source of constant delight for me, while perusing the works of the earlier naturalists, to sympathize with the genial spirit and the ernestness that pervade their writings, so free from egotism, and animosity against their fellow-students. Their devotion to their studies is equal to the spirit of reverence with which they look upon nature; and it is disgraceful to our age, that we must contrast width such dispositions the ill-will, the jealousies, the quarrels for priority, and the profanation, which pervade the discussions of certain modern authors. Moreover, in a systematic point of view, the great naturalists of the sixteenth century deserve to be studied more fully than they have been thus far. It is astonishing, for instance, to see how near Rondelet, in discussing the views of Aristotle upon the affinities of animals, came to perceiving their true affinities, and their natural classification..."
A good example of the evolution of thought in natural history is exemplified by various views regarding the nature of corals in the seventeenth and eighteenth centuries. The concept of the "zoophyte", or transitional nature of corals, was established by the Roman scholars Sextus Empiricus and Pliny the Elder (Caius Plinius Secundus), based on Aristotle's notion of the gradation from plants to higher animals. This undoubtedly served to propagate a state of confusion for nearly two thousand years --- the false view that corals as living beings share the natures of both animals and plants (Agassiz, 1860: 6-7). Paolo Boccone (1670) and John Woodward (1695) emphasized the inanimate aspect of corals and regarded them as stones, while L.F. Marsigli (1725) considered them to be plants, and the coral polyps as "flowers of the coral". It was the perceptive observer, Jean-André Peyssonnel (1753), that first recognized the animal nature of corals and their affinities to other coelenterates, after observing the contraction, expansion and movement of the tentacles of coral polyps (Hyman, 1940: 365).

The Linnean Period (1758 - 1858)
The publication of Systema Naturae by Linnaeus (Carl von Linné, 1758) is accepted as the starting point for our contempory system of binominal nomenclature. Linnaeus named the genus Pennatula, from which the ordinal name Pennatulacea is derived. Other contributors during the Linnean Period include William Borlase (1758), Job Baster (1759-65), Joannes Baptist Bohadsch (1761), John Ellis (1764), Peter Simon Pallas (1766 --- first monographer of the "zoophytes"), Pieter Boddaert (1771), I. Lepechin (1781), John Ellis and Daniel Solander (1786), Lazzaro Spallanzani (1796), Georges Cuvier --- originally Jean Léopold Nicolas Frédéric Cuvier (1797 --- who named the genera Umbellula and Veretillum), Jean Baptiste Pierre Antoine de Monet de Lamarck (1816 --- who named the genera Funiculina, Renilla, and Virgularia), Tilesius von Tilenau (1819), S. della Chiaje (1827, 1841-44), M.W. Rapp (1827), H.-M. Ducrotay de Blainville (1834), Christian Gottfried Ehrenberg (1834a --- who introduced the family name Pennatulina, later corrected by Dana, 1846 to Pennatulidae), Fredrich Sigismund Leuckart (1841), James Dwight Dana (1846 --- who corrected the familial name of Ehrenberg to Pennatulidae), George Johnston (1847), John Edward Gray (1840-1873 --- who named the genus Sarcoptilus), Henri Milne Edwards and Jules Haime (1850 ---who named the fossil genus Graphularia), Achille Valenciennes (1850 --- who named the veretillid genus Lituaria), P. Chr. Asbjørnsen (1856--- who named the genus Kophobelemnon), and Jan Adrianus Herklots (1858 --- who named the genera Scytalium and Pteroeides).

The Darwinian Period (1859 - 1899)
The publication of the Origin of Species by Charles Darwin initiated what is here referred to as the Darwinian Period.
The pre-eminent author of the Pennatulacea during this period was Rudolf Albert von Kölliker, who produced major descriptive taxonomic works between 1865 and 1880. Kölliker described the sea pens of the H.M.S. Challenger Expedition, in which thirty species were described. Kölliker also named the genera Acanthoptilum, Anthoptilum, Halipteris, Protoptilum, Sclerobelemnon, Scleroptilum, and Stachyptilum. Other noteworthy contributions during this period include Pieter Bleeker (1859 --- who described several species of Pteroeides), Jan Adrianus Herklots (1863), Josua Lindahl (1874), Johan Koren & Daniel C. Danielssen (1847-1884), Addison Emery Verrill (1865-1882 --- who named the genera Distichoptilum, Ptilosarcus, and Stylatula), Sir Charles Wyville Thomson (1874 --- chief scientist of the H.M.S. Challenger Expedition), Ambrosius Arnold Willem Hubrecht (1885 --- who named the genus Echinoptilum), G. Herbert Fowler (1888 and 1894), A. Milnes Marshall (1883a and 1883b), A. Milnes Marshall & G. Herbert Fowler (1887 and 1889), and A. Milnes Marshall & William P. Marshall (1882), T.H. Tizard, et al (1885), Théophile Studer (1891 --- who named the genus Gyrophyllum), and Addison Emery Verrill (1864-1922 --- who introduced the name Pennatulacea in 1865 as a subordinal name, later elevated to ordinal status by Studer, 1887a) .
The Darwinian Period produced numerous studies of evolutionary relationships for many groups of organisms. Important works pertaining to pennatulacean phylogeny during this time include R.A. von Kölliker (1870 and 1880), Gottlieb von Koch (1878), and A. Milnes Marshall (1887).

The Early Twentieth Century (1900 - 1949)
Significant contributions in octocorallian systematics (including pennatulaceans) during the early part of the Twentieth Century include Hector F.E. Jungersen (1904, 1907, and 1917), Monsieur Ch. Gravier (1906 --- who named the genus Scytaliopsis), Charles Cleveland Nutting (1908 --- who named the genus Calibelemnon), Heinrich Balss (1909 and 1910), Hjalmar Broch (1910-1961), Willy Kükenthal (1902-1925 --- who named the genera Actinoptilum, Amphiacme, and Chunella), W. Kükenthal & H. Broch (1911 --- who named the genus Cavernulina), Sydney John Hickson (1883-1940), Elisabeth Deichmann (1936 and 1941), and J. Arthur Thomson (1905-1931). The latter published in collaboration with several co-authors including George Crane, Doris L. Mackinnon, Nita I. Rennet, W. D. Henderson, James Ritchie, and James J. Simpson. F.M. Bayer (pers. comm.) states, "Deichmann...is the 20th Century link with the 19th Century in octocoral systematics. Her Blake report (1936) is pivotal for the western Atlantic."
It was during this period that the argument was introduced by Libbie Henrietta Hyman (1940: 365) to abandon the name "Coelenterata"of Rudolph Leuckart in favor of "Cnidaria" introduced by Berthold Hatschek. This argument has apparently convinced many zoologists to substitute a name long in use with a more recently created one. Hyman states, "Leuckart (1847) clearly grasped the fundamental differences between the two great radiate groups, the coelenterates and the echinoderms, and separated them, creating the name Coelenterata. Leuckart's Coelenterata, however, included the sponges and the ctenophores...The proper splitting of Leuckart's Coelenterata was achieved by Hatscheck (1888), who recognized three phyla: Spongiaria, Cnidaria, and Ctenophora. We therefore consider Cnidaria to be the most suitable name for the phylum..."
The acceptance of this line of reasoning is both unnecessary and inconsistent, however. As an example, Ernst Haeckel's "Scyphozoa" originally included both the Scyphozoa and the Anthozoa, yet the name Scyphozoa is still retained by most zoologists. This is just one example of a major animal group in which the name has long usage and is still recognized today, even though it orginally encompassed more taxa than are now accepted. On the other hand, Land (2003) [Zoologische Verhandelingen 345: 209-213] argues that because of modern common usage, the name Coelenterata should no longer be used.

The Late Twentieth Century (1950 - present)
Previous to this period, publications on the Pennatulacea dealt predominantly with descriptive taxonomy or general natural history (particularly bioluminescence). This most recent period has produced a flourishing of works in fields of study other than taxonomy - including behaviour, ecology, morphology, physiology, cell biology, natural products biochemistry, and evolutionary biology. Important contributions in systematics during this period include Frederick M. Bayer (1955-1996), Huzio Utinomi (1956-1982); Manfred Grasshoff (1972-1991), F.M. Bayer, M. Grasshoff, & Jakob Verseveldt (1983), F.M. Bayer & M. Grasshoff (1997), Marie-José d'Hondt (1984), Andrée Tixier-Durivault (1954-1987 --- who named the genera Crassophyllum and Malacobelemnon), and Gary C. Williams (1986-1997). The latter author introduced cladistic methodology to the study of evolutionary relationships in the Pennatulacea.

SYNONYMS AND MISSPELLED GENERIC NAMES OF PENNATULACEA

(Revised from Williams, 1995a: 101; synonyms are listed on the left with the valid genera to which they belong shown in parentheses; * = misspelled names or transcription errors; ** = see Bayer & Grasshoff, 1997)

*Acanthoptilon Traquair in Zoological Record 7, 1870 (Acanthoptilum)
Actinoptilon Kükenthal, 1910 (Actinoptilum)
*Actinoptinum Day et al., 1970 (Actinoptilum)
Amphianthus Kükenthal, 1902 (Amphiacme)
Argentella Gray, 1870 (Pteroeides)
Balticina Gray, 1870 (Halipteris)
Bathypenna Marion, 1906 (possibly synonymous with Gyrophyllum according to Kükenthal, 1915)
Bathyptilum Kölliker, 1872 (Kophobelemnon)
Benthoptilum Verrill, 1885 (Anthoptilum)
*Benthoptillum Haddon in Zoological Record 22, 1885 (Anthoptilum)
Cladiscus Koren & Danielssen, 1877 (Virgularia)
Clavella Gray, 1870 (Lituaria)
Crinillum Harting, Miquel & Hoeven, 1861 (Umbellula)
Crispella Gray, 1870 (Pteroeides)
Deutocaulon Marshall & Fowler, 1888 (Virgularia)
Dübenia Danielssen & Koren, 1884 (Stylatula)
Encrinus Lamarck, 1801 (Umbellula)
Fusticularia Simpson, 1905 (probably synonymous with Cavernularia according to Kükenthal, 1915)
*Godefroyia Leuckart, 1872 (Pteroeides)
Godeffroyia Kölliker, 1870 (Pteroeides)
Göndul Koren & Danielssen, 1883 (Halipteris)
Gunneria Danielssen & Koren, 1884 (Kophobelemnon)
Halisceptrum Herklots, 1863 (Virgularia)
Helicoptilum Nutting, 1912 (Distichoptilum)
Herklotsia Gray, 1860 (Renilla)
Juncoptilum Thomson & Henderson, 1905 (Distichoptilum)
*Leioptilum Verrill, 1865 (Ptilosarcus)
*Leioptillum Verrill, 1868 (Ptilosarcus)
Leioptilus Gray, 1860 (Pennatula)
Leioptilus of authors other than Gray, 1860 (Ptilosarcus)
Leptoptilum Kölliker, 1880 (Funiculina)
*Lioptilum Kölliker, 1872 (Ptilosarcus)
Lygomorpha Koren & Danielssen, 1877 (Halipteris)
Lygus Herklots, 1858 (Virgularia)
Mesobelemnon Gravier, 1907 (considered synonymous with Sclerobelemnon by Hickson, 1916)
Microptilum Kölliker, 1880 (Halipteris)
Norticina Gray, 1870 (Halipteris)
Ombellulaires Cuvier, 1817 (Umbellula)
**Ombellula Cuvier, 1797 (Umbellula)
Ombellularia Lamarck, 1836 (Umbellula)
Osteocella Gray, 1870 (Halipteris)
Parabelemnon Thomson & Simpson, 1909 (possibly synonymous with Cavernularia according to Kükenthal, 1915)
Pavonaires Cuvier, 1817 (Funiculina)
Pavonaria Schweigger, 1820 (Funiculina)
Pavonaria Kölliker, 1869 (Halipteris)
Penna Bohadsch, 1761 (Pennatula, in part) (name unavailable as the work was suppressed by ICZN)
Phosphorella Gray, 1870 (Pennatula)
Policella Gray, 1870 (Veretillum)
Prochunella Balss, 1909 (Calibelemnon)
Protocaulon Kölliker, 1880 (Virgularia)
*Pteroides Pfeffer, 1886 (Pteroeides)
Pteromorpha Herklots, 1858 (Pteroeides)
Ptilella Gray, 1870 (Pennatula)
*Renila Schweigger, 1820 (Renilla)
Renillina Gray, 1860 (possibly a young example of the alcyoniid soft coral Sarcophyton according to Kükenthal, 1915)
Sarcobelemnon Herklots, 1858 (Cavernularia)
Sarcophyllum Kölliker, 1870 (Sarcoptilus)
Sceptonidium Richiardi, 1869 (Virgularia)
Stephanoptilum Roule, 1905 (Anthoptilum)
Stichoptilum Grieg, 1887 (Halipteris)
Struthiopteron Broch, 1910 (Pteroeides)
Stylobelemnon Kölliker, 1872 (Cavernularia)
Stylobelemnoides J.A. Thomson & Henderson, 1905 (possibly synonymous with Cavernularia according to Kükenthal, 1915)
Svava Danielssen & Koren, 1884 (Virgularia)
Svavopsis Roule, 1908 (Virgularia)
Thesioides J.A. Thomson & Henderson, 1906 (Anthoptilum)
Trichoptilum Kölliker, 1880 (Funiculina)
Umbellaria Schweigger, 1820 (Umbellula)
Umbellularia Lamarck, 1801 (Umbellula)
*Verrilia Lütken in Zoological Record 10, 1873 (Halipteris)
Verrillia Stearns, 1873 (Halipteris)
Vorticella Linnaeus, 1767 (Umbellula, in part)

(Original descriptions and other selected taxonomic references - restricted primarily to taxa considered valid by Williams,1995a, 1997f, and the present work; + = fossil taxa; ++ = taxa with both living and fossil representation)

Acanthopitilum Kölliker, 1870
[See also: Balss (1910); Bayer (1957); Kükenthal (1915); Williams (1995a)]
Acanthoptilum album Nutting, 1909
Acanthoptilum agassizi Kölliker, 1872
Acanthoptilum annulatum Nutting, 1909
Acanthoptilum gracile (Gabb, 1863)
Acanthoptilum oligacis Bayer, 1957
Acanthoptilum pourtalesii Kölliker, 1870
Acanthoptilum scalpelifolium Moroff, 1902
Actinoptilum Kükenthal in: Kükenthal & Broch, 1911
[See also: Kükenthal (1910, 1915); Williams (1990, 1995a)]
Actinoptilum molle (Kükenthal, 1910)
Amphiacme Kükenthal, 1903
[See also: Hickson (1916); Kükenthal (1902a, 1915); Kükenthal & Broch (1911); Williams (1990, 1995a)]
Amphiacme abysorrum (Kükenthal,1902)
Anthoptilum Kölliker, 1880
[See also: Grasshoff (1982a, 1982b); Hickson (1916); Kükenthal (1915); Kükenthal & Broch (1911); Williams (1990, 1995a)]
Anthoptilum grandiflorum (Verrill, 1879)
Anthoptilum murrayi Kölliker, 1880
+Bensonularia Hamilton, 1958
+Bensonularia spatulata Hamilton, 1958 [Eocene of New Zealand]
Calibelemnon Nutting, 1908
[See also: Balss (1910); Hickson (1916); Kükenthal (1915); Williams (1990, 1995a)]
Calibelemnon hertwigi (Balss, 1909)
Calibelemnon indicum (Thomson & Henderson, 1906)
Calibelemnon symmetricum Nutting, 1908
Cavernularia Valenciennes in Milne Edwards & Haime, 1850
[See also: Hickson (1916); Hondt (1984b); Kükenthal (1915); Lopez-Gonzalez, Gili, & Williams (in press); Williams (1989b, 1990, 1995a)]
Cavernularia capitata Williams, 1989
Cavernularia clavata Kükenthal & Broch, 1911
Cavernularia chuni Kükenthal & Broch, 1911
Cavernularia dayi Tixier-Durivault, 1954
Cavernularia dedeckeri Williams, 1989
Cavernularia elegans (Herklots, 1858)
Cavernularia glans Kölliker, 1872
Cavernularia habereri Kölliker, 1872
Cavernularia luetkeni Kölliker, 1872 (also spelled C. lütkeni and C. lutkeni)
Cavernularia malabarica Fowler, 1894
Cavernularia mirifica Tixier-Durivault, 1963
Cavernularia obesa Valenciennes in Milne Edwards & Haime, 1850
Cavernularia pusilla (Philippi, 1835)
Cavernulina Kükenthal & Broch, 1911
[See also: Hickson (1916); Hondt (1984b); Kükenthal (1915); Williams (1989b, 1990, 1995a)]
Cavernulina cylindrica Kükenthal & Broch, 1911
Cavernulina darwini (Hickson, 1921)
Cavernulina grandiflora Hondt, 1984b
Cavernulina orientalis Thomson & Henderson, 1909
Chunella Kükenthal, 1902
[See also: Kükenthal (1915); Kükenthal & Broch (1911); Hickson (1916); Williams (1990, 1995a, 1997c)]
Chunella gracillima Kükenthal, 1902
Crassophyllum Tixier-Durivault, 1961
[See also: Williams (1995a, 1995d)]
Crassophyllum cristatum Tixier-Durivault, 1961
Crassophyllum thessalonicae Vafidis & Koukouras, 1991
Distichoptilum Verrill, 1882
[See also: Hickson (1916); Kükenthal (1915); Kükenthal & Broch (1911); Williams (1990, 1995a)]
Distichoptilum gracile Verrill, 1882
Echinoptilum Hubrecht, 1885
[See also: Balss (1910); Hickson (1916); Kükenthal (1915); Kükenthal & Broch (1911); Sachs (1913); Williams (1990, 1995a)]
Echinoptilum asperum Hickson, 1916
Echinoptilum echinatum (Kükenthal, 1910)
Echinoptilum elongatum Hickson, 1916
Echinoptilum macintoshi Hubrecht, 1885
Echinoptilum minimum Hickson, 1916
Echinoptilum roseum Hickson, 1916
Funiculina Lamarck, 1816
[See also: Balss (1910); Herklots (1858); Hickson (1916); Jungersen (1904); Kölliker (1872); Kükenthal (1915); Kükenthal & Broch (1911); Williams (1990, 1995a)]
Funiculina armata Verrill, 1879
Funiculina parkeri Kükenthal, 1909
Funiculina quadrangularis (Pallas, 1766)
+Graphularia Milne Edwards & Haime, 1850
+Graphularia ambigua (Morton, 1830) [Cretaceous/Tertiary of southeastern North America]
[See also: Shapiro & Ramsdell (1965)]
+Graphularia badenia Strand, 1928
+Graphularia belgica Vincent (1893) [Eocene of Belgium]
+Graphularia crecelii Andrée, 1912 [Oligocene of Germany]
+Graphularia desertorum Kuhn, 1949
+Graphularia groenwalli Nielsen , 1914 [Cretaceous of Denmark]
+Graphularia incerta Malaroda, 1951 [Oligocene of Italy]
+Graphularia irregularis Nielsen, 1914 [Cretaceous of Denmark]
+Graphularia kalimnae Chapman & Crespin, 1928 [Tertiary of Australia]
+Graphularia longissima Squires, 1958 [Cretaceous/Tertiary of New Zealand]
+Graphularia meijeri Voight, 1958
+Graphularia nigra Malaroda, 1951 [Oligocene of Italy]
+Graphularia quadrata Voigt, 1958
+Graphularia salisburgensis Traub, 1938 [Paleocene of Austria]
+Graphularia sp. [Eocene of Hungary]
+Graphularia sulcata Nielsen, 1914 [Cretaceous of Denmark]
[See also: Kolosvary (1949)]
+Graphularia wetherelli Milne-Edwards & Haime, 1850 [Eocene of England]
[See also: Davis (1936)]
+Graphularia ? yamakawai Yabe & Sugiyama, 1937 [Pleistocene of Japan]
Gyrophyllum Studer, 1891
[See also: Hickson (1916); Kükenthal (1915); Roule (1905); Studer (1901); Williams (1995a, 1995d)]
Gyrophyllum hirondellei Studer, 1891
Gyrophyllum sibogae Hickson, 1916
Halipteris Kölliker, 1869
[See also: Willams (1990, 1995a)]
Halipteris africana (Studer, 1879)
Halipteris californica (Moroff, 1902)
Halipteris christii (Koren & Danielssen, 1847)
Halipteris finmarchica (Sars, 1851)
Halipteris heptazooidea Acuña & Zamponi, 1992
Halipteris willemoesi Kölliker, 1870
Kophobelemnon Asbjørnsen, 1856
[See also: Balss (1910); Danielssen & Koren (1884); Herklots (1858); Hickson (1916); Jungersen (1904); Kölliker (1872); Kükenthal (1915); Kükenthal & Broch (1911); Thomson & Simpson (1909); Williams (1990, 1995a)]
Kophobelemnon affine Studer, 1894
Kophobelemnon heterospinosum Kükenthal, 1910
Kophobelemnon hispidum Nutting, 1912
Kophobelemnon irregulatus Keller, Pasternak & Naumov, 1975
Kophobelemnon leucharti Cecchini, 1917
Kophobelemnon macrospinosum J.A. Thomson, 1927
Kophobelemnon molanderi Pasternak, 1975
Kophobelemnon pauciflorum Hickson, 1916
Kophobelemnon stelliferum (Müller, 1776)
Lituaria Valenciennes in Milne Edwards & Haime, 1850
[See also: Balss (1910); Gray (1870); Hickson (1916); Hondt (1984b); Light (1921); Kölliker (1872); Kükenthal (1915); Thomson & Simpson (1909); Williams (1990, 1995a)]
Lituaria amoyensis Koo, 1935
Lituaria australasiae (Gray, 1860)
Lituaria breve Light, 1921
Lituaria habereri Balss, 1910
Lituaria hicksoni Thomson & Simpson, 1909
Lituaria kuekenthali Light, 1921
Lituaria molle Light, 1921
Lituaria phalloides (Pallas, 1766)
Lituaria philippinesis Light, 1921
Lituaria valenciennesi Hondt, 1984
Malacobelemnon Tixier-Durivault, 1965
[See also: Williams (1995a)]
Malacobelemnon stephensoni Tixier-Durivault, 1965
*Octobasis Malecki, 1982 [Upper Cretaceous of Poland]
+Pavonaria portisi Angelis, 1895? [Tertiary of Italy]
+Pavonaria ? singularis Malaroda, 1951 [Oligocene of Italy]
Pennatula Linnaeus, 1758
[See also: Balss (1910); Gray (1870); Herklots (1858); Kölliker (1869); Lamarck (1816); Leuckart (1872); ); Hickson (1916); Kükenthal (1915); Kükenthal & Broch (1911); Williams (1990, 1995a, 1995d)]
Pennatula aculeata Danielssen, 1860
Pennatula argentina Acuña & Zamponi, 1992
Pennatula delicata Tixier-Durivault, 1966
Pennatula fimbriata Herklots, 1858
Pennatula grandis Ehrenberg, 1834
Pennatula indica Thomson & Henderson, 1906
Pennatula inflata Kükenthal, 1910
Pennatula mollis Alder, 1867 (considered a junior synonym of Pennatula phosphorea by Cornelius & Garfath, 1980, after examination of the two well preserved syntypes)
Pennatula phosphorea Linnaeus, 1758
Pennatula prolifera Jungersen, 1904
Pennatula rubra (Ellis, 1764)
+Pennatulites Nelli, 1903
+Pennatulites manzonii Nelli, 1903 [Miocene of Italy]
+Prographularia Frech, 1890
+Prographularia tradica Frech, 1890 [Triassic]
Protoptilum Kölliker, 1872
[See also: Balss (1910); Hickson (1916); Jungersen (1904); Kölliker (1880); Kükenthal (1915); Kükenthal & Broch (1911); Williams (1995a)]
Protoptilum carpenteri Kölliker, 1872
Protoptilum celebense Hickson, 1916
Protoptilulm cyaneum Kükenthal, 1910
Protoptilum denticulatum Jungersen, 1904
Protoptilum smitti Kölliker, 1872
Protoptilum thomsoni Kölliker, 1872
Pteroeides Herklots, 1858
[See also: Gray (1870); Hickson (1916); Hondt (1984a); Kölliker (1872); Kükenthal (1915); Kükenthal & Broch (1911); Leuckart (1872); Williams (1990, 1995a, 1995d)]
Pteroeides acutum Tixier-Durivault, 1966
++Pteroeides argenteum (Ellis & Solander, 1786)
[See also: Bayer (1955a) - Tertiary of Kei Islands]
Pteroeides bankanense Bleeker, 1859
Pteroeides bestae Hondt, 1984
Pteroeides breviradiatum Kölliker, 1869
Pteroeides caledonicum Kölliker, 1869
Pteroeides carnosum Tixier-Durivault, 1972
Pteroeides crossieri Tixier-Durivault, 1966
Pteroeides densum Tixier-Durivault, 1966
Pteroeides duebeni Kölliker, 1869
Pteroeides dofleini Balss, 1909
Pteroeides durum Kölliker, 1872
Pteroeides esperi Herklots, 1858
Pteroeides flexuosum Tixier-Durivault, 1966
Pteroeides humesi Tixier-Durivault, 1966
Pteroeides hymenocaulon Bleeker, 1859
Pteroeides isosceles J.S. Thomson, 1915
Pteroeides jungerseni Broch, 1910
Pteroeides laboutei Hondt, 1984
Pteroeides latissimum Kölliker, 1869
Pteroeides lusitanicum Broch, 1910
Pteroeides malayense Hickson, 1916
Pteroeides morbusus Tixier-Durivault, 1961
Pteroeides oblongum Gray, 1860
Pteroeides sagamiense Moroff, 1902
Pteroeides sparmanni Kölliker, 1869
Pteroeides spicatum Tixier-Durivault, 1972
Pteroeides spinosum (Ellis, 1764)
Pteroeides tenerum Kölliker, 1869
Pteroeides timorense Hickson, 1916
Pteroeides triangulum Tixier-Durivault, 1972
Ptilosarcus Verrill, 1865
[See also: Leuckart (1872); Nutting (1909); Verrill (1865); Williams 1995a, 1995d)]
Ptilosarcus gurneyi (Gray, 1860)
Ptilosarcus undulatus (Verrill, 1865)
Renilla Lamarck, 1816
[See also: Ehrenberg (1834); Eisen (1876); Gray (1860); Herklots (1858); Kölliker (1871b); Kükenthal (1915); Kükenthal & Broch (1911); Müller (1864); Williams (1995a); Zamponi & Pérez (1996); Zamponi, Pérez, & Capitoli (1997)]
Renilla koellikeri (also spelled R. kollikeri or R. köllikeri) Pfeffer, 1886
Renilla muelleri (also spelled R. mulleri or R. mülleri) Kölliker, 1872
Renilla musaica Zamponi & Pérez, 1995
Renilla octodentata Zamponi & Pérez, 1995
Renilla reniformis (Pallas, 1766)
Renilla tentaculata Zamponi, Pérez, & Capitoli, 1997
Sarcoptilus Gray, 1848
[See also: Williams (1995a, 1995c, 1995d)]
Sarcoptilus bollonsi (Benham, 1906)
Sarcoptilus grandis Gray, 1848
Sarcoptilus nullispiculatus Williams, 1995
Sarcoptilus rigidus Williams, 1995
Sarcoptilus shaneparkeri Williams, 1995
Sclerobelemnon Kölliker, 1872
[See also: Balss (1910); Hickson (1916); Kölliker (1872); Kükenthal (1915); Kükenthal & Broch (1911); Thomson & Henderson (1906b); Williams (1995a, 1995d)]
Sclerobelemnon burgeri (Herklots, 1858)
Sclerobelemnon elongatum Hickson, 1916
Sclerobelemnon gracile (Gravier, 1908)
Sclerobelemnon gravieri Hickson, 1916
Sclerobelemnon koellikeri Thomson & Henderson, 1906
Sclerobelemnon magniflorum Hickson, 1916
Sclerobelemnon schmeltzi Kölliker, 1872
Sclerobelemnon theseus Bayer, 1959
Scleroptilum Kölliker, 1880
[See also: Balss (1910); Kükenthal (1915); Kükenthal & Broch (1911); Williams (1990, 1995a)]
Scleroptilum grandiflorum Kölliker, 1880
Scytaliopsis Gravier, 1906
[See also: Gravier (1908); Kükenthal (1915); Williams (1990, 1995a)]
Scytaliopsis djiboutiensis Gravier, 1906
Scytaliopsis ghardagensis (?Gravenhorst, 1821) (also spelled S. ghardaqana; a probable nomen nudem since it is unclear whether this citation represents an original description of a new taxon, or merely the use of an unsubstantiated name)
[See also: Atiya (1994); Schuhmacher & Hinterkircher (1996)]
Scytalium Herklots, 1858
[See also: Balss (1910); Hickson (1916); Kölliker, 1870; Kükenthal (1915); Kükenthal & Broch (1911); Thomson & Simpson (1909); Williams (1995a)]
Scytalium balssi Hickson, 1916
Scytalium martensi Kölliker, 1870
Scytalium sarsii Herklots, 1858
Scytalium tentaculatum Kölliker, 1880
Stachyptilum Kölliker, 1880
[See also: Kükenthal (1915); Kükenthal & Broch (1911); Williams (1995a)]
Stachyptilum dofleini Balss, 1909
Stachyptilum macleari Kölliker, 1880
Stachyptilum superbum Studer, 1894
Stylatula Verrill, 1864
[See also: Balss (1910); Jungersen (1904); Kölliker (1870); Kükenthal (1915); Kükenthal & Broch (1911); Williams (1995a)]
Stylatula antillarum Kölliker, 1870
Stylatula brasiliensis (Gray, 1870)
Stylatula darwini Kölliker, 1870
Stylatula diadema Bayer, 1959
Stylatula elegans (Danielssen, 1860)
Stylatula elongata (Gabb, 1862)
Stylatula gracilis (Gabb, 1864)
Stylatula kinbergi Kölliker, 1870
Stylatula lacazi Kölliker, 1870
Stylatula polyzoidea Zamponi & Pérez, 1996
Umbellula Cuvier, 1797 (see Bayer & Grasshoff, 1997)
[See also: Broch (1957, 1958a); Cuvier (1800); Ellis (1753, 1755); Grasshoff (1971, 1973, 1982a, 1982b); Heezen & Hollister (1971); Hickson (1916); Kölliker (1875, 1880); Kükenthal (1902b, 1915); Kükenthal & Broch (1911); Lamarck (1801); Mylius (1755); Pasternak (1975); Williams (1990, 1993b, 1995a, 1997a, 1997c); Williams & Rogers (1989)]
Umbellula durissima Kölliker, 1880
Umbellula encrinus Linnaeus, 1758
Umbellula hemigymna Pasternak, 1975
Umbellula huxleyi Kölliker, 1880
Umbellula lindahli Kölliker, 1874
Umbellula monocephalus Pasternak, 1975
Umbellula pellucida Kükenthal, 1902
Umbellula spicata Kükenthal, 1902
Umbellula thomsoni Kölliker, 1874
Veretillum Cuvier, 1797
[See also: Balss (1910); Gray (1870); Herklots (1858); Hickson (1916); Kükenthal (1915); Kükenthal & Broch (1911); Williams (1990, 1995a)]
Veretillum australis (Gray, 1870)
Veretillum cynomorium (Pallas, 1766)
Veretillum leloupi Tixier-Durivault, 1960
Veretillum malayense Hickson, 1916
Veretillum manillensis (Kölliker, 1872)
Veretillum tenuis (Marshall & Fowler, 1889)
Veretillum vanderbilti Boone, 1938
Virgularia Lamarck, 1816
[See also: Herklots (1858); Kölliker (1880); Koren & Danielssen (1877); Kükenthal (1915); Kükenthal & Broch (1911); Richiardi (1869); Williams 1990, 1995a)]
Virgularia abies Kölliker, 1870
Virgularia brochi Kükenthal, 1915
Virgularia bromleyi Kölliker, 1880
Virgularia densa Tixier-Durivault, 1966
Virgularia galapagensis Hickson, 1930
Virgularia glacialis Kölliker, 1870
Virgularia gracillima Kölliker, 1880
Virgularia gustaviana (Herklots, 1863)
Virgularia halisceptrum Broch, 1910
Virgularia juncea (Pallas, 1766)
Virgularia kophameli May, 1899
Virgularia loveni Utinomi, 1971
Virgularia mirabilis (Müller, 1776)
Virgularia patagonica (a probable nomen nudem since an original description cannot be found and a type specimen has not been located)
[See also: Barrattini & Ureta, 1960; Darwin, 1860]
Virgulaira patachonica (see Virgularia patagonica)
++Virgularia presbytes Bayer, 1955a
[See also: Bayer (1957); Belém & Figueiredo Alvarenga (1973) - Tertiary of Trinidad]
Virgularia reinwardti Herklots, 1858
Virgularia rumphi Kölliker, 1870
Virgularia schultzei Kükenthal, 1910
Virgularia tuberculata Marshall, 1883

GEOGRAPHIC INDEX

Africa - eastern (see Indo-Pacific)
Africa - northern (see North Atlantic and Mediterranean)
Africa - southern

Branch, et al (1994); Broch (1939, 1940); Day (1974b); Day, et al (1970); Molander (1929); J.S. Thomson (1915, 1917, 1924); Tixier-Durivault (1954, 1960); Williams (1987, 1989a, 1989b, 1990, 1992, 1993a); Williams & Rogers (1989).

Africa - western

Broch (1910a; 1914b; 1958b); Buchanan (1955); Dollfus (1938); López- González et al (in press); Molander (1929); Pax & Müller (1954); Studer (1878); Tixier-Durivault (1961b, 1963); Williams (1987, 1989b, 1990, 1992).

Africa - Atlantic (see: Africa - western)
Antarctica (see Southern Oceans)
Arctic Ocean

Broch (1913c, 1955, 1956); Jungersen (1907, 1915, 1917); Lindahl (1874c); Madsen (1948); May (1900); Mylius (1753, 1754); Pasternak (1980); Verrill (1922); Yashnov (1948);

Asia - northeastern (see Pacific Ocean - northern; or Japan)
Atlantic - eastern (see North Atlantic and Mediterranean or Africa - western)
Atlantic - northern (see North Atlantic and Mediterranean)
Atlantic - western temperate (see North Atlantic and Mediterranean or South America - Atlantic)
Atlantic - western tropical (see Western Tropical Atlantic and Caribbean)
Australia - eastern (see Indo-Pacific)
Australia - northern (see Indo-Pacific)
Australia - southern

Broch (1910c); Briggs (1915); Godfrey (1943); Gray (1848); McCoy (1892); Pasternak (1966); Utinomi (1971); Utinomi & Shepherd (1982); Williams (1995c).

Australia - western

Broch (1910c); Williams (1995c).

Caribbean Sea (see Western Tropical Atlantic & Caribbean)
Cosmopolitan (see Worldwide)
East Africa (see Africa - eastern)
East Asia (see Indo-Pacific)
Eastern Australia (see Indo-Pacific)
Eastern Pacific - western North America (see North America - Pacific)
Eastern Pacific - western South America (see South America - Pacific)
Europe (see North Atlantic and Mediterranean)
Indo-Pacific (including Red Sea*)

Alcock (1902); Allen & Steene (1994); *Atiya (1994); Briggs (1915); Clastres, et al (1984); Colin & Arneson (1995); Dinamani (1965); Fowler (1894); Gosliner, et al (1996); Gravely (1941); *Gravenhorst (1821); *Gravier (1906a, 1906b, 1906c, 1907a, 1908, 1912a); Hickson (1905); Hondt (1984a, 1984b); Hornell (1922); Humes (1978); Imahara (1991); Koo (1935, 1940); *Kükenthal (1913b); Light (1921); *Magnus (1966); A.M. Marshall & Fowler (1889); Mather & Bennet (1993); Nakasone & Yu (1987); Nutting (1908); Pasternak (1964); Ridley (1883); Roule (1908); Sankolli & Neelakantan (1971); *Schuhmacher & Hinterkircher (1996); Simpson (1905); J.A. Thomson (1905); J.A. Thomson & Crane (1909a, 1909b); J.A. Thomson & Henderson (1905a, 1905b, 1906a, 1906b); Tixier-Durivault (1960, 1965, 1966, 1972); Tixier-Durivault & Hondt (1974a); Utinomi (1956b).

Japan

Balss (1910); Fisher (1874); Hubrecht (1885); Kumano (1937); Moroff (1902b); Nutting (1912); Okutani (1969); Stearns (1883); Stimpson (1855); J.A. Thomson & Rennet (1927); Utinomi (1956a, 1958, 1961, 1964).

Mediterranean Sea (see North Atlantic and Mediterranean)
New Zealand

Benham (1906, 1907); Dendy (1897); Hamilton (1958); Hutton (1904); Powell (1947); Williams (1995c).

North America - temperate Atlantic (see North Atlantic and Mediterranean)
North America - Pacific

Allen (1969); Anonymous (1898); Batie (1972); Belcik (1977); Blake (1872, 1873); Brusca (1980); Dawson (1872); Fautin, et al (1987); Flora & Fairbanks (1966); Gabb (1862, 1864); Gotshall (1987, 1994); Gotshall & Laurent (1979); Hartman (1960); Hickson (1930a); Hochberg & Ljubenkov (1998); Human (1973); Johnson & Snook (1935); Kerstitch (1989); Kozloff (1974, 1983, 1987); Kükenthal (1913a); Lam, et al (1982); Nutting (1909); Ricketts, et al (1985); Sclater (1872, 1873); Shimek (1998); Stearns (1873a); Strathmann (1988); Studer (1894); Verrill (1865, 1868a, 1868b).

 

SUBJECT INDEX

Anatomy (see Morphology and Ultrastructure)

Behaviour
P.A.V. Anderson (1976); P.A.V. Anderson & Case (1975); Buisson (1974, 1976, 1980, 1988); Darwin (1860); Couet (1979); Dickinson (1978); Hoare & Wilson (1977); Kastendiek (1975a, 1975b, 1976); Lancaster (1601); Langton, et al (1990); Mori (all citations); Mori & Ondo (1957); Mori & Tanase (1973); Pantin (1950); Pavans de Ceccatty & Buisson (1965); Takada & Mori (1956, 1957).

Bibliography
Bayer (1981a, 1996); Hickson (1916); Kükenthal (1915); D.W. Thompson (1885); Williams (1990).
Biography (Regarding Authors of Pennatulacean Research)
Boeseman (1973); Forest (1996); Fransen, et al (1997); Gardiner (1940); Holthuis (1993, 1995); Smit (1979); Zimmer (1925).

Biography (Regarding Authors of Pennatulacean Research)
Boeseman (1973); Forest (1996); Fransen et al. (1997); Gardiner (1940); Holthuis (1993, 1995); Smit (1979); Zimmer (1925).

Bioluminescence
Agassiz (1850); Aldrovandi (1642, 1648); Anctil et al (1982); J.M. Anderson & Cormier (1978); J.M. Anderson et al (1974, 1978); P.A.V. Anderson & Case (1975); Awad & Anctil (1993a, 1993b); Batie (1972); C. Bauhin (1620, 1671); J. Bauhin (1650-51); Bilhaut (1975a, 1975b); Blainville (1834); Boussuet (1558); Buck (1973); Chiaje (1827); Charbonneau (1981); Cormier (1978); Davenport & Nicol (1956); DeLuca, et al (1976); Dittrich (1888); Ehrenberg (1834c); Ellis (1764); Forbes (1847); Forbes & Goodsir (1851); Germain & Anctil (1988); Gesner (1555, 1558); Grant (1827, 1829); Grober (1990a, 1990b); Hart, et al (1979); Harvey (1917, 1920, 1940, 1952); Hastings (1968, 1983); Hastings & Morin (1969); Herdman (1913a); Herring (1978, 1991); Hori & Cormier (1973); Hori, et al (1972, 1973); Imperato (1599); Korotneff (1887); Kreiss & Cormier (1967); Krukenberg (1887, 1888); Mangold (1910); N.B. Marshall (1979); Matern (1984); Matthews, et al (1977); Morin (1974, 1976, 1998); Nealson, et al (1986); Nicol (1955a, 1955b, 1955c, 1958); Panceri (1871b, 1871c, 1872a, 1872b, 1872c); Parker (1920c); Péron (1804); Pratje (1923); Rapp (1827); Royal Society (1870); Satterlie & Case (1979); Shaw (1838-46); Shimomura & Johnson (1975, 1979); Soares & Sawaia (1975); Spallanzani (1784, 1796); Spurlock & Cormier (1975); C.W. Thomson (1874); Tilesius von Tilenau (1819); Titschack (1965, 1966); Tizard et al (1885); Wampler, et al (1971, 1973); Ward (1979); Ward & Cormier (1978a, 1978b); Williams (1990).

Corals (Pre-Linnean General Natural History)
Boccone (1670); Marsigli (1725); Peyssonnel (1753); Rondelet (1554-1555); Woodward (1695).

Ecology and Biotic Interactions
P.K. Anderson (1989); Atkinson (1989); Bertsch (1968, 1982); Best (1988); Birkeland (1969, 1971, 1974); Brafield (1969); Creed & Coull (1984); Davis (1978); Davis, et al (1982); Dawydoff (1930, 1938); Day (1974a); Dube & Ball (1971); Fager (1968); Fujita & Ohta (1988); ); Gili, et al (1987); Gilluly (1970); Gosliner, et al (1996); Hornell (1922); Howson & Davies (1991); Humes (1978); Jones (1960); Kastendiek (1975a, 1975b, 1976, 1982); Lam, et al (1982); Laubier (1972); Magnus (1966); Mariscal & Bigger (1977); McDonald & Nybakken (1978); Miyajima (1897, 1900); Mori (all citations); Mori & Ondo (1957); Mori & Tanase (1973); Nakasone & Yu (1987); Okutani (1969); Ramesh, et al (1985); Rittschof, et al (1986, 1988); Rowe (1971); Sankolli & Neelakantan (1971); Shepherd (1983); Shimek (1998); Short & Trower (1986); Standing, et al (1984); Thompson, et al (1988); Tyler, et al (1995).

General Natural History and General Biology
Alcock (1902); Allen (1969); Arculeo et al (1990); Arndt (1912); R.D. Barnes (1987); R.S.K. Barnes, et al (1988); Bayer (1973, 1981a); Bayer, et al (1983); Bayer & Owre (1968); Bellomy (1974); Bourne (1900); Branch, et al (1994); Brusca & Brusca (1990); Colin & Arneson (1995); Coleman & Teague (1973); Cooke (1889); Dakin (1953); Dalyell (1848); Darwin (1860); Delage & Hérouard (1901); Donovan (1995); Dunn (1982); Faulkner & Chesher (1979); Fosså & Nilsen (1995); Frische (1991); Gage & Tyler (1991); Gesner (1565); Hadi (1994); Hardy (1967); Gosliner, et al (1996); Gotshall (1987, 1994); Gotshall & Laurent (1979); Gravier (1912b); Haeckel (1904); Hardy (1965, 1967); Heezen & Hollister (1971); Hickson (1906, 1909, 1930b); Huxlely (1907); Hyman (1940); Johnson & Snook (1935); Kaplan (1982, 1988);Kerstitch (1989); Koren & Danielssen (1877); Kozloff (1974,1983, 1987, 1990); Kükenthal (1923-1925); Lancaster (1601); Luther & Fiedler (1961); Lutz (1986); Williams (1986, 1993a, 1995a, 1997c); Lydekker (no date); MacGinitie (1938); MacGinitie & MacGinitie (1968); N.B. Marshall (1979); W.P. Marshall (1895); Mather & Bennet (1993); Meglitsch (1972); Milne & Milne (no date); Milne Edwards & Haime (1857); Ming (1993); Modeer (1786); H.B. Moore (1937); Moseley (1872); Moss (1878); Mylius (1753, 1754); Nordgaard (1905); Omori (1991); Pearse, et al (1987); Philippi (1835); Pimentel (1967); Ricketts, et al (1985); Riedl (1963); Ricketts, et al (1985); Rondelet (1554-1555); Rumphius (1705); Schechter (1959); Schömann (1949); Schuhmacher & Hinterkircher (1996); Sclater (1872, 1873); Smith (1964); Smith & Carlton (1975); Sphon (1964); Sprung & Delbeek (1997); C.W. Thomson (1874); Thurston (1890); Tilesius (1812); Tizard et al (1885); Tixier-Durivault (1987); A. Trembley (1744); Tyler & Zibrowius (1992); Utinomi (1956a, 1964); Voss (1976); Wilkins & Birkholz (1986); Zim & Ingle (1955).

Histology
Hasama (1944); Kölliker (1865); Korotneff (1887); Lyke (1965); Niedermeyer (1913, 1914); Titschack (1970).

History of Science (Regarding Coelenterates and/or Sea Pens)
Agassiz (1860); Bayer (1981a); Hickson (1916); Hyman (1940); Shapiro & Ramsdell (1965); Williams (1993b, 1995a, 1997c).

Molecular Biology and Genetics
Lorenz, et al (1991).

Morphology and Ultrastructure
Alonso (1979); Beklemishev (1969a, 1969b); Buisson (1970); Buisson & Franc (1969); Bujor (1901); Chia & Crawford (1977); Crawford & Chia (1974); Bullough (1950); Dunkelberger & Watabe (1972, 1974); Fautin & Mariscal (1991); S. Franc (1979); S. Franc, et al (1971, 1974, 1985); Franzén (1967); Germain & Anctil (1988); Hickson (1883); Ivester & Dunkelberger (1971); Jungersen (1888a, 1888b); Koch (1878, 1889, 1890); Kölliker (1865, 1871b, 1872, 1874); Korotneff (1887); Ledger & Franc (1978); Lenhoff, et al (1971); Lightbown (1918); Lyke (1965); Mariscal (1974, 1979); Mariscal & Bigger (1977); Marks, et al (1949); Niedermeyer (1911, 1912); Panceri (1870); Parker (1919); Roule (1907); Satterlie, et al (1976); Shapeero (1969); Spurlock & Cormier (1975); Titschack (1966, 1968, 1970); P. Trembley (1941, 1942); Watabe & Dunkelberger (1979); Wilbur (1976); Wilson (1884).

Natural Products Chemistry, Biochemistry, and Toxicology
Anctil (1987, 1989a, 1989b); Anctil et al (1982, 1984, 1991); J.M. Anderson & Cormier (1978); J.M. Anderson, et al (1974, 1978); P.A.V. Anderson (1976); Awad & Anctil (1993a, 1993b); Bernheimer & Arigad (1981); Bullock (1970); Charbonneau (1981); Clastres, et al (1984); Coan & Travis (1970); Datta, et al (1990); DeLuca, et al (1976); Fu, et al (in press); Goswami, et al (1995); Grimmelikhuijzen & Groeger (1987); Grimmelikhuijzen, et al (1987); Harmon, et al (1984); Hart, et al (1979); Hori & Cormier (1973); Hori, et al (1972, 1973); Huang & Mir (1972); Jones, et al (1979); Karkhanis & Cormier (1971); Keifer, et al (1986); Kittredge, et al (1962); Kreiss & Cormier (1967); Morin (1976); Kumar, et al (1990); Lorenz, et al (1991); Mackie (1987); Pani & Anctil (1994); Shapeero (1969); Shimomura & Johnson (1975, 1979); Standing, et al (1984); Thompson, et al (1988); Tillet-Barret, et al (1992); Vanderah & Djerassi (1977); Vidal, et al (1992); Waele, et al (1987); Wampler, et al (1971); Wekell (1974,
1978); Wratten, et al (1977a, 1977b).

Paleontology (Mesozoic and Cenozoic Fossil Taxa)
Andrée (1912); Angelis (1895?); Baluk & Pisera (1984); Bayer (1955a, 1955b, 1956); Bradley (1980, 1981); Branco (1885); Chapman & Crespin (1928); Davis (1936); Fowler (1911); Frech (1890); Gabb (1859, 1861); Gregorio (1890); Hamilton (1958); Häntzschel (1958); Howell (1947); Kolosvary (1949); Kuhn (1949); Malaroda (1951); Malecki (1982); Milne Edwards & Haime (1850); Morton (1830, 1834); R.C. Moore (1956); Nelli (1903); Nielsen (1914); Shapiro & Ramsdell (1965); Roemer (1880); Squires (1958); Strand (1928); Traub (1938); P. Trembley (1941, 1942); Valenciennes (1850); Vincent (1893); Voigt (1958); Yabe & Sugiyama (1937).

Paleontology (Vendian Frond-like Fossil Taxa)
Arthur (1997); Bergström (1989, 1991); Buss & Seilacher (1994); Conway Morris (1991, 1993); Deng & Chen (1981); Dzik (1991); Fedonkin (1992, 1996); Glaessner (1958a, 1958b, 1959, 1961, 1984); Glaessner & Daily (1959); Glaessner & Wade (1966); Jenkins (1985, 1992); Jenkins & Gehling (1978); Lewin (1984); Liu (1981, 1983); Richter (1955); Runnegar (1992); Seilacher (1989); Simonetta & Conway Morris (1991); Weiguo (1986); Williams (1995b, 1997c, 1997d); Wright (1997).

Photography (Color Photographs of Living Sea Pens)
Allen & Steen (1994); Branch, et al (1994); Colin & Arneson (1995); Faulkner & Chesher (1979); Gosliner, et al (1996); Fosså & Nilsen (1995); Gotshall (1987, 1994); Gotshall & Laurent (1979); Kerstitch (1989); Ming (1993); Nishimura (1992); Schumacher & Hinterkircher (1996); Shimek (1998); Sprung & Delbeek (1997); Weinberg (1996); Williams (1990, 1996); Wilkins & Birkholz (1986).

Phylogeny, *Cladistics, Evolution, and Biogeography
Altuna-Prados (1994); Bergström (1989, 1991); Bourne (1900); Broch (1913b); Conway Morris (1991); Grasshoff (1973, 1991); Hickson (1916, 1930b); Kinoshita (1912); Koch (1878); Kölliker (1869-72, 1872, 1880); Kükenthal (1912b, 1914, 1915a, 1921); Kükenthal & Broch (1910, 1911); A.M. Marshall (1883b); Niedermeyer (1913); Patterson (1986); Williams (1992, *1993b, 1995a, *1995b, *1995d, 1997a, 1997b, *1997c, 1997d, 1997e).

Physiology and Cell Biology
Anctil (1987, 1989a, 1989b); Anctil et al (1982, 1984, 1991); J.M. Anderson & Cormier (1978); J.M. Anderson et al (1974); P.A.V. Anderson (1976); P.A.V. Anderson & Case (1975); Awad & Anctil (1993a, 1993b); Bilhaut & Pavans de Ceccatty (1971a, 1971b); Brafield (1969); Brafield & Chapman (1967); Case & Morin (1966); Chapman (1972); Child (1951); Buck & Hanson (1967); Buisson (1964, 1969, 1971a, 1971b, 1973, 1976, 1979, 1988); Carlgren (1940); Charbonneau (1981); Crawford & Chia (1974); Dickinson (1978); J.M. Franc (1979); S. Franc (1970, 1973, 1979, 1980); Hagiwata, et al (1981); Honjo (1940); Imafuku (1973, 1975, 1976); Ivester (1977); Korotneff (1887); Krukenberg (1887); Ledger & Franc (1978); A.R. Moore (1926); Mori (all citations); Mori & Ondo (1957); Mori & Tanase (1973); Musgrave (1909); Nicol (1955b, 1955c); Parker (1920a, 1920b); Pavans de Ceccatty & Buisson (1964a, 1964b, 1965); Pavans de Ceccatty, et al (1963); Pratt (1909); Satterlie, et al (1976, 1980); Senut & Franc (1985); Takada & Mori (1956, 1957); Titschack (1965, 1966, 1968, 1970); Umbriaco, et al (1990); Waele, et al (1987).

Polymorphism, Phenotypic Variability, and Biodiversity
Hickson (1903b, 1903c); Human (1973); Jaworski (1939); A.M. Marshall (1883); Pasternak (1989); Williams (1992, 1993b, 1997b).

Quoted Passages
Agassiz (1860); Harvey (1952); Hyman (1940); Lankaster (1601); N.B. Marshall (1979); Péron (1804); C.W. Thomson (1874); Shapiro & Ramsdell (1965) quoted under Gregorio (1890); Tizard et al (1885), Williams (1993b).

Reproductive and Developmental Biology (including Growth Stages and Regeneration)
Birkeland (1969, 1971); Chia and Crawford (1973); Dalyell (1839); Delage and Hérouard (1901); Eckelbarger et al. (1998); Franzén (1967); Jungersen (1888b); Korschelt (1936); Lacaze-Duthiers (1865, 1887); Mori and Tanase (1973); Roule (1932); Satterlie and Case (1979); Strathmann (1988); Tarent and Tarent (1980); Torrey (1901); Tyler et al. (1995); Willemoes-Suhm (1875); Wilson (1880, 1881, 1882a, 1882b, 1883b, 1903).

Taxonomy, Distributional & Bathymetric Records
Abel (1963); Acuña & Zamponi (1992); Alder (1861, 1862, 1863, 1867); Allen (1969); Anonymous (1898); Arculeo et al (1990); Arndt (1912); Asbørnsen (1856); Atiya (1994); Atkinson (1989); Balss (1909, 1910, 1911); Barattini & Ureta (1960); Barreira y Castro (1990); Batie (1972); Bayer (1955b, 1956, 1957, 1959, 1961); Bayer & Grasshoff (1997); Belcik (1977); Belem & Alvarenga (1973); Belyaev (1972); Benham (1906, 1907); Boone (1933, 1938); Branch, et al (1994); Branch & Williams (1993); Broch (all citations); Buchanan (1955); Cairns, Hartog, & Arneson (1986); Carpine & Grasshoff (1985); Castro (1981); Colin & Arneson (1995); Costa Soares (1979); Cuvier (1797, 1800); Dana (1846); Danielssen & Koren (1884); Dawson (1966); Deichmann (1936a, 1936b, 1941); Ellis & Solander (1786); Fautin, et al (1987); Ehrenberg (1834a, 1834b), Erhardt & Moosleitner (1995); Fowler (1888, 1894); Fu, et al (in press); Gabb (1862, 1864); Gili (1986); Gili & Pagès (1987); Gosliner, et al (1996); Gotshall (1987, 1994); Gotshall & Laurent (1979); Grasshoff (all citations); Gravenhorst (1821); Gravier (all citations); Gray (all citations); Hartman (1960); Herklots (1858, 1863); Hickson (1890, 1894, 1900, 1903a, 1904, 1905, 1907, 1911, 1914,1916, 1921, 1922, 1930a, 1936, 1937, 1940); Hoare & Wilson (1977); Hochberg & Ljubenkov (1998); Hondt (1984a, 1984b); Hubrecht (1885); Imahara (1991); Jahn (1970); Johnston (1847); Jungersen (1904, 1905, 1907, 1915, 1917); Kölliker (all citations); Koo (1935, 1940); Koren & Danielssen (1847, 1856, 1874, 1877, 1883, 1884); Kükenthal (all citations); Kramp (1932, 1933, 1950); Kumano (1937); Lacaze-Duthiers (1891); Lamarck (1816, 1836); Langton, et al (1990); Lepechin (1781); Leuckart (1841); Leunis (1886); Light (1921); Lindahl (1874a, 1874b, 1874c); Linnaeus (1758, 1767); Lopez-Gonzalez, et al (in press); Madsen (1948); Manuel (1981); A.M. Marshall (1883b); A.M. Marshall & Fowler (1888); A.M. Marshall & Marshall (1882); May (1899, 1900); Milne Edwards & Haime (1850, 1857); Molander (1929); Moroff (1902a, 1902b); O.F. Müller (1776); F. Müller (1866); Naumov (1955); Nobre (1931); Norman (1867); Nutting (1908, 1909, 1912); Pallas (1766, 1787); Panceri (1871a); Pasternak (all citations); Pax (1936); Pax & Müller (1955b, 1955c, 1962); Pérez (1996); Pfeffer (1886); Poche (1914, 1915a, 1915b); Quoy & Gaimard (1827); Rho & Song(1976, 1977); Richiardi (1869); Richmond (1997); Ridley (1883); Riedl (1963, 1983); Riveros Zunica (1948); Robertson (1887); Rossi (1971); Roule (1905, 1906, 1908); Sachs (1913); Sars (1846,1851); M. Schultze (1871); F.E. Schulze (1875); Simpson (1905); Stearns (1873a, 1873b, 1873c, 1874, 1882, 1883); Stephens (1909); Stiasny (1937, 1938); Stimpson (1855); Studer (all citations); J.A. Thomson (1905, 1927); J.A. Thomson & Crane (1909a, 1909b); J.A. Thomson & Henderson (1905a, 1905b, 1906a, 1906b); J.A. Thomson & Mackinnon (1911); J.A. Thomson & Rennet (1927, 1931); J.A. Thomson & Ritchie (1906); J.A. Thomson & Simpson (1909); J.S. Thomson (1915, 1917, 1924); Tilesius (1826); Tixier-Durivault (all citations); Tixier-Durivault & Hondt (1974a, 1974b); Tixier-Durivault & Lafargue (1968); Utinomi (all citations); Utinomi & Shepherd (1982); Vafidis & Koukouras (1991); Vafidis et al. (1994); Van Soest (1977); Verrill (all citations); Waterman (1950); Williams (all citations); Wiktor (1974); Yashnov (1948); Zamponi & Pérez (1995a, 1995b,1996); Zamponi, et al (1997); Zuñiga (1948).