See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/233979427

Systema Porifera: A Guide to the Classification of Sponges

Book · January 2002

DOI: 10.1007/978-1-4615-0747-5

CITATIONS READS
758 4,195

2 authors:

John N.A. Hooper Rob van Soest

Queensland Museum Naturalis Biodiversity Center
416 PUBLICATIONS 7,799 CITATIONS 641 PUBLICATIONS 13,622 CITATIONS

SEE PROFILE SEE PROFILE

Some of the authors of this publication are also working on these related projects:

SpongeMaps Project View project

Sponge science in the Aegean Sea and adjacent areas View project

All content following this page was uploaded by John N.A. Hooper on 27 May 2014.

The user has requested enhancement of the downloaded file.

Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer Academic/Plenum Publishers, New York, 2002
Systema Porifera. A Guide to the Classification of Sponges
John N.A. Hooper1 & Rob W.M. Van Soest2
1
Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au).
2
Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT, Amsterdam, Netherlands. (soest@science.uva.nl)
“If I have seen further it is by standing on the shoulders of Giants” (Sir Isaac Newton, 1676 [Turnbull et al., 1959])
INTRODUCTION
Why write a book on sponge taxonomy; who would read such
a book; what is the relevance of sponges; and what are sponges
anyway?
The first two questions are not easily answered, except to
offer the explanation that over the past few centuries there have
been many ‘books’ written on various aspects of sponges, and pre-
sumably over the next few centuries there will be many more.
During the late 19th to the mid 20th centuries a great many
monographic publications on sponges were produced (e.g.,
Johnston, 1842; Duchassaing & Michelotti, 1864; Schmidt, 1862,
1864, 1868, 1870, 1879, 1880b; Bowerbank, 1864, 1866, 1874b,
1882; Haeckel, 1872; Pomel, 1872; Marshall & Meyer, 1877;
Zittell, 1877, 1879; Quenstedt, 1878b; Hinde, 1887–1893;
Saville Kent, 1880–1882; Lendenfeld, 1888, 1889a, 1897a, 1907,
1910a,b, 1915; Ridley, 1884a; Poléjaeff, 1883, 1884; Ridley &
Dendy, 1887; Sollas, 1888; Rauff, 1893–1894; Topsent, 1892a,
1900, 1901d, 1904b, 1913a, 1928c; Hall & Clarke, 1898;
Lundbeck, 1902, 1905, 1910; Schulze, 1904; Schrammen,
1910–1912, 1937; Dendy, 1922b; Wilson, 1925; Ijima, 1926; de
Laubenfels, 1936a, 1948, 1954; Reid, 1958a; Hill, 1965; Burton,
1963; Penney & Racek, 1968; and others). This trend is now sadly
in decline, although there are some notable exceptions among the
contemporary authors, especially the palaeontological literature
(e.g., Lévi, 1956a; Hartman, 1958b; Koltun, 1959, 1964b, 1967;
Finks, 1960; Vacelet, 1964; Wiedenmayer, 1977b, 1994; Rigby,
1986c; Rigby & Webby, 1988; Senowbari-Daryan, 1990; Boika
et al., 1991; Reitner, 1992; Rigby & Senowbari-Daryan, 1996; Pisera,
1997, and others). There are also several general texts, mono-
graphic taxonomic overviews, and a few popular books published
on sponges (e.g., Delage & Hérouard, 1899; Minchin et al., 1900;
Hentschel, 1923; Arndt, 1935, 1937; Hyman, 1940; Koltun, 1955;
Moore, 1955; Teichert, 1972; Brien et al., 1973; Bergquist, 1978;
Hartman, 1982; Hartman et al., 1980; Simpson, 1984; Zea, 1987;
Ackers et al., 1992; Rigby et al., 1993; Hooper & Wiedenmayer,
1994; Lévi et al., 1998). Since 1970 there has also been a growing
literature from the international sponge symposia, together repre-
senting a comprehensive treatment of sponge biology (Fry, 1970;
Harrison & Cowden, 1976; Lévi & Boury-Esnault, 1979; Jones,
1987; Vacelet & Boury-Esnault, 1987; Rützler, 1990; Reitner &
Keupp, 1991; Uriz & Rützler, 1993; Van Soest et al., 1994;
Willenz, 1996; Watanabe & Fusetani, 1998; Hooper, 1999).
Together with the many thousands of other existing publications on
sponges this literature has contributed substantially to advancing
the knowledge of ‘Life On Earth’ – past and present; the diverse
sponge morphologies; their structural, physiological and biochemical
1
mechanisms that have evolved to survive over Earth’s changing
chemical history; their role in the cycles of life and death in the
oceans; and their co-evolution with other symbiotic life forms.
This knowledge has also provided practical answers to funda-
mental biological questions: such as understanding the biosynthesis
of chemicals, the evolution of eukaryotic immunology, cellular
theory and totipotency, understanding physiological adaptive
strategies to cope with extreme environments, gene function and
evolution of the Metazoa, and some realistic assessments of
the magnitude of marine biodiversity, including impacts by
humans on the marine environment. It is significant that many of
these contemporary studies have been collaborative, multidiscipli-
nary efforts that seek a broader understanding of the role of
sponges in time and space.
It is also obvious from the existing sponge literature that this
present book is unique. Apart from the pioneering attempt by
de Laubenfels (1936a) to make the sponge systematics ‘widely avail-
able’ at the practical level of classification (families and genera), no
other publication has attempted to achieve what we propose here.
The latter questions are perhaps easier to answer. Sponges are
among the most highly diverse and successful of the 28 aquatic
invertebrate phyla, both in terms of the numbers of species and
their range of morphological characters. About 7,000 extant
species are currently described (but these undoubtedly represent
only a small fraction of those that have ever lived). Sponges have
survived largely unchanged in their fundamental bauplans since the
Late Cambrian (509MYA). They were important reef builders dur-
ing the Phanerozoic – a role subsequently usurped by the faster
growing scleractinarian corals during the late Mesozoic. Although
most of the older groups became extinct during the Late Devonian
crisis (373MYA), sponges have radiated and diversified in Recent
seas, with representatives found in all aquatic habitats – from
ephemeral (quasi-terrestrial) ones to the more stable marine
abyssal zones. Consequently, sponges have demonstrated a huge
capacity to adapt and survive in marginal life situations – and it has
been suggested that they are clearly set to outsurvive those who
study them (Vacelet, 1999).
Sponges continue to have an economic potential beyond their
fundamental roles in the marine ecological processes. These
include commercial ‘bath sponges’, important since early Greek
civilization, and more recently as potential sources of therapeutic
drugs. This renewed interest in sponges has accelerated the discov-
ery and documentation of species in all the oceans, with recent
estimates from the various museum collections that the extant
fauna may be twice as diverse as that currently described (e.g.,
Hooper & Lévi, 1994). As a consequence, it was necessary
to revisit the existing systematics in order to accommodate these
2 Systema Porifera. A Guide to the Classification of Sponges
substantial new collections, which was the initial impetus to
produce this book.
HISTORY OF SPONGE TAXONOMY
A long and tortuous taxonomic history
Johnston (1842), an ‘amateur’ sponge taxonomist, remarked
long ago that sponges were a “comparatively limited and isolated
class of organized beings, obscure in character and possessed of
less interest than attaches every other. The class may be said to
occupy at present the confines of debateable land, lying between
the confines of the two organic Kingdoms, – too poor and barren to
be an object of contest with the subjects of either … This is not an
easy task, for there is so much that is in common to them, and each
adapts itself so readily to circumstances and assumes a new mask,
that it requires a tact, to be gained only by some experience, to rec-
ognize them under their guises; while we labour, perhaps in vain,
to devise phrases which shall aptly pourtray to others the charac-
teristics of objects that have no fixed shape, and whose distinctive
peculiarities almost cheat the eye.”
Over recorded human history the taxonomy of sponges has
been long and tortuous, gaining an unenviable reputation among
zoologists as presenting appalling difficulties (Vacelet, 2000).
Homer made numerous references to them (Lendenfeld, 1888) as
regular household commodities in early Greek society (referring to
the so-called ‘bath sponges’, which apparently served in diverse
roles, from cushioning body armour to cleansing bodies). He also
coined the name ‘sponges’, “to squeeze”: “Then with a sponge he
drest, his face all over, necke and hands, and all his hairie breast”
(translation by Chapman, quoted in Johnston, 1842). Aristotle was
apparently the first to describe sponges as objects of scientific
inquiry (Johnston, 1842), differentiating three ‘species’ within the
Mediterranean fauna – although it is highly likely that recognition
of more than one species of ‘sponge’ substantially predates his writ-
ings: Tragos, Manon, Achilleum. Pliny (translation by Holland,
1634) also mentioned a fourth species: Aplysiae. More significantly,
however, Aristotle was apparently also the first author to recognize
that sponges might be animals (Johnston, 1842), although Pliny
provided conflicting statements that they were “neither living crea-
tures nor yet plants … intermediate between vegetable and animal
kingdoms”, and again, “sponges have a life … yea and a sensible
life … for there is found of their bloud settled within them”.
The ensuing scientific void during the ‘dark ages’ contributed
little to advancing biological or taxonomic knowledge of the phy-
lum. Occasionally ‘bestialities’ included reference to sponges (e.g.,
Entzel, 1551), and their medicinal properties (e.g., Pomet, 1694),
sometimes reiterating Aristotle’s assertion of their animality (e.g.,
Wotton, 1552), although with little comprehension of their origin
and development – “fermentation of the sea’s scum or its sponta-
neous pullulations … certaine matter wrought together, of the fome
and froth of the sea, which we call spunges” (Gerarde, 1578), and
sometimes asserting their vegetable nature – “cryptogamous veg-
etables and nearly allied to the Fungi” (Imperato, 1599). Indeed,
Linnaeus (1759) initially arranged Spongia amongst the cryptoga-
mous algae, but later (1767) included it in Animalia. This debate
continued for well over a century (see Johnston, 1842; Lévi, 1999),
with Pallas (1766) and Ellis (1755, 1786) being the first to provide
an informed scientific opinion as to their animal nature, which was
finally settled under the influential gaze of Grant (1826b).
Despite a number of important publications during the late 18th
and early 19th centuries (Linnaeus, Cuvier, Lamarck, Lamouroux,
Pallas, Esper, Nardo etc.; see bibliography of Vosmaer, 1928),
Lendenfeld (1888) suggested that our earliest reliable taxonomic
knowledge of sponges stemmed from the mid-19th century works
of Bowerbank (e.g., 1841) and Schmidt (e.g., 1862) on the British
and Adriatic faunas, respectively. While it is true that these authors
introduced new suites of characters to sponge taxonomy, raising the
standard of contemporary systematic debates, there were certainly
many other significant contributions to the growing body of knowl-
edge at that time, including works of Carter, Duchassaing,
Gray, Hyatt, Johnston, Lieberkuhn, Norman, Thompson, Wyville
Thompson and Verrill, to name only the more prominent, and many
of whose descriptions are still recognizable today. Towards the end
the 19th and beginning of the 20th centuries there was a phenome-
nal proliferation of sponge research that impacted on the systemat-
ics of Porifera, with the most prominent contributions by:
Annandale, Arndt, Arnesen, Bidder, Brondsted, Dendy, Döderlein,
Fristedt, Haeckel, Hallmann, Hanitsch, Hentschel, Hinde, Ijima,
Jenkin, Keller, Kieschnick, Kirk, Kirkpatrick, Lambe, Lendenfeld,
Lindgren, Lundbeck, Maas, Marshall, Minchin, Moret, Okada,
Poléjaeff, Rauff, Ridley, Row, Saville Kent, Schlüter, Schrammen,
Schulze, Sollas, Thiele, Topsent, Vosmaer, Weltner, Whitelegge,
Wilson and Zittel. It was also during this period that most of the
7,000 currently known species were documented, and the frame-
work of the existing classification developed (e.g., see statistics on
history of Australian sponge collections on the web at http://
www.environment.gov.au/abrs/abif-fauna/volswww.htm).
In contrast with this earlier halcyon era of scientific endeavour
only a few authors studied sponges during the middle of the
20th century (e.g., Brien, Burton, de Laubenfels, Tanita, Topsent,
Tuzet), followed by several others over the next few decades
(e.g., Bergquist, Borojevic, Cuif, Gautret, Hartman, Jones, Koltun,
Lévi, Pulitzer-Finali, Rasmont, Sarà, Simpson, Termier, Vacelet,
Wendt, Wiedenmayer), with expertise gradually escalating to number
several hundreds of researchers by the end of the 20th century, includ-
ing the 30 or so contributors to this present book. In comparison to
this escalating number of neontological authors our palaeontological
capabilities continue to decline, with many of the more prominent
palaeontological authors now retired or approaching the end of their
careers (Debrenne, Finks, Pickett, Reid, Rigby, Zhuralev).
“Standing on the shoulders of giants”
In 1928 Geo P. Bidder wrote “Some younger biologists incline
now to cut themselves loose from the lengthening chain of literature,
and to read nothing that has appeared more than twenty years ago.
Whatever they gain in time for research, they lose much … They lose
knowledge because forgotten observations of great value lie hidden
in papers which excited little interest among contemporaries, and
consequently have not been passed through the generations on the
stream of second-hand quotation. … Traditional beliefs without rea-
sonable foundation … are at their strongest if we believe that our
own generation has made all its own thought, and that no brave
man lived before Agamemnon” (Bidder, in Vosmaer, 1928).
Unfortunately, the sponge literature is now so diverse, and some
works are nearly impossible to access in libraries, that the older
literature continues to rapidly recede into the past. Lévi (1999)
provides some pertinent examples of knowledge that has been lost
over time, or distorted during its transmission to the present, such
that sometimes we seem to repeatedly ‘reinvent the wheel’.
 Systema Porifera. A Guide to the Classification of Sponges
We take heed of Bidder’s warning, and consequently have
endeavored here to build on the accumulated knowledge of our
forebears to provide (within the rules of the ICZN; Anon, 1999): a
stable nomenclature; a sound contemporary classification; a rem-
edy for the nomenclatural mistakes of the past; and incorporating
more recent non-morphometric data into the contemporary classifi-
cation. We hope that these efforts will provide a solid platform for
the future, and that any shortcomings of this Systema Porifera will
mostly concern the phylogenetic controversies and unanswered
biological questions, which we anticipate will be solved by
advances in our biological knowledge and our abilities to better
sample the sponge genome.
AIMS OF THE ‘SYSTEMA PORIFERA’
A Taxonomic Revision
The primary aim of Systema Porifera is to summarise, and
revise where necessary the supraspecific classification of the Phylum
Porifera (including spongiomorphs such as ‘Sphinctozoans’,
Archaeocyatha and Heteractinida), based firstly on re-evaluation of
type material for each genus when possible, and subsequently incor-
porating any recent biological evidence. The Systema focuses on the
living fauna, and in most cases provides only a cursory treatment of
the fossil fauna and palaeontological literature. It is anticipated that
the forthcoming revision of the Treatise on Invertebrate Paleontology
will balance this bias (Rigby et al., in prep.).
A Sponge Guide
It is intended that the Systema Porifera will serve as a
practical guide, providing accessibility to the supra-specific
classification of sponges and spongiomorphs; providing a clear
explanation of morphological characters, their importance to the
taxonomy and systematics of each higher taxon, and to illustrate
the major characters used to classify each group.
Solving Nomenclatural Problems
The Systema Porifera provides a sound baseline for future
debate on sponge taxonomy and to address many long-outstanding
nomenclatural problems (such as those inadequately treated or
overlooked completely by de Laubenfels, 1936a).
ABOUT THIS BOOK
Firstly, an apology
It is obvious that we have committed a fundamental faux pas
in the title of this book, which of course should be Systema
Poriferorum. However, from the outset it was recognised that this
classic title was both ponderous and ambiguous in identifying the
Phylum Porifera as the target taxon, and consequently we chose to
ignore classic nomenclature for pragmatic reasons. Our other
endeavours in this book, however, have been more stringent.
Structure of the ‘Systema’
The structure of this book is aimed primarily at those faced with
the daunting task of identifying sponges – providing keys to diagnose
View publication stats
3
classes, subclasses, orders, suborders, families and genera – and
within each genus, listing the pertinent literature (where this exists) to
potentially identify species. This structure, however, is fundamentally
at odds with how most sponge taxonomists operate – which is from
the species level upwards (a ‘gestalt’ approach). Anyone familiar with
the appalling difficulties associated with sponge systematics would
already know that keys usually do not work, except perhaps for the
very few well-known small regional faunas, or for small, well-
characterised taxonomic groups (such as some genera and families).
The message is: beware! These difficulties are mostly due to the fre-
quent losses of ‘pivotal characters’, or the modification of these char-
acters beyond recognition as being ‘typical’ for a particular taxonomic
group, or the difficulty in interpreting whether characters are the same
or different between groups of species. This latter problem concerns
the interpretation of ‘homology’, whereby some prominent characters
that may appear to be the same across several taxonomic groups actu-
ally represent analogous features that do not necessarily reflect
phylogeny, or translate into a convenient morphological classification.
Consequently, the secondary structure of this book – and one
that is more important to its future as a sound platform to develop
the sponge systematics – concerns extensive analyses of sponge
characters and their relative importance (homology vs. analogy), to
arrive at both a practical classification and a theoretical reconstruc-
tion of sponge phylogenies. For those who are mostly interested in
simply identifying sponges these phylogenetic analyses and
lengthy discussions may appear as ‘noise’, although we contend
that this ‘noise’ is an essential quality control. Accurate interpreta-
tion of characters is essential for the accurate identification of taxa.
Illustrated glossaries are useful to define characters and to assist in
recognising homologous from analogous characters. However, we
only provide a few of these here because many have already been
published in the contemporary literature (see below).
Thirdly, the Systema represents a concerted attempt to both
stabilize the sponge nomenclature (through our collective compre-
hensive re-evaluations of the entire literature), and to revise the
Poriferan classification (incorporating recent evidence held by
experts in the field – who have an estimated combined taxonomic
expertise of more than 600 person-years!). Thus, many of these
discussions and analyses are necessary to justify the new
taxonomic hypotheses presented here.
References to other works that may assist using this book
There are several illustrated glossaries and other useful publi-
cations that define sponge morphology. Ideally, these should have
been reproduced here, to make this book complete in itself, but this
was seen as a luxury and a redundancy. Already the size of this vol-
ume had doubled from our original assessment of the Phylum at the
outset of this project, as we continued to uncover numerous forgot-
ten taxa. Consequently, there are two publications in particular that
should be used in conjunction with this volume to identify
sponges: the Thesaurus of sponge morphology (Boury-Esnault &
Rützler, 1997) and Atlas of sponge morphology (De Vos et al.,
1991). Several other works may also be useful, providing addi-
tional glossaries, relevant illustrations and interpretations sponge
characters (Brien et al., 1973; Bergquist, 1978; Hooper &
Wiedenmayer, 1994 [available through bookshop@isb.csiro.au,
with an online version at http://www.environment.gov.au/abrs/
work/zoocat/]; Van Soest et al., 2000 [available through orders@
springer.de; http://www.eti.uva.nl/Products/CD-catalogue.html];
Lévi et al., 1998 [available through diffusion@bondy.orstom.fr].