PART V, REVISED, CHAPTER 12:
GLOSSARY OF THE HEMICHORDATA
JÖRG MALETZ,
1
DENIS E. B. BATES,
2
EDSEL D. BRUSSA,
3
ROGER A. COOPER,
4
ALFRED C. LENZ,
5
JOHN F. RIVA,
6
BLANCA A. TORO,
7
and YUANDONG ZHANG
8
[
1
Institut für Geologische Wissenschaften, Berlin, Germany, Yorge@zedat.fu-berlin.de;
2
Institute of Geography and Earth Sciences, Aberystwyth
University, UK, deb@aber.ac.uk;
3
Deceased;
4
DSIR GEO, New Zeandand Geological Survey, R.Cooper@gnx.cri.nz;
5
University of Western Ontario,
Canada, aclenz@uwo.ca;
6
INRS-ETA, Québec, Canaba, john_riva@ete.enrs.ca;
7
CICTERRA, Ciudad Universitaria, Cordoba, Argentina, btorogr@
mendoza-conicet.gob.ar;
8
Nanjing Institute of Geology and Palaeontology; Academica Sinica, Nanjing, China, ydzhang@nigpas.ac.cn]
PREFACE
The terminology of the Hemichordata
used herein differs considerably from the
terminology used in the previous versions
of the “Graptolite Treatise” (BULMAN, 1955,
1970), which were focused entirely on the
fossil members of the Graptolithina and the
few known tube-bearing extant and extinct
Pterobranchia. Both classes were regarded as
independent groups of the Hemichordata,
but they are now included as a single taxon,
the Pterobranchia, with the Graptolithina
and the Cephalodiscida as two subclades
(MITCHELL & others, 2013). A number of
terms had to be changed due to a better
understanding of the construction of the
graptolite tubarium and the phylogenetic
relationships within the Pterobranchia, the
main group of fossilized hemichordates, in
an attempt to homogenize the terminology
of these previously separated groups.
The term tubarium, for example, is rein-
troduced to the pterobranch terminology
to describe the organic housing or domi-
cile of all Pterobranchia, instead of using
the term rhabdosome for the fossil Ptero-
branchia (Graptolithina in BULMAN, 1955,
1970) and the coenecium for the extant
members (Pterobranchia sensu BULMAN,
1955, 1970). LANKESTER (1884) initially
introduced the term tubarium to describe
the housing of the extant Pterobranchia, as
he considered the terms coenecium or zooe-
cium inappropriate since they described the
housing construction of bryozoans, to which
the Pterobranchia were referred initially.
Tubarium describes the housing construc-
tion of extant and extinct Pterobranchia
more independently; it also describes it more
precisely as being formed by glandular secre-
tions and can easily be used for all members.
It is therefore extended herein to include the
homologous glandular constructions of the
domiciles of fossil graptolite taxa.
Other terms, such as the periderm, have
been eliminated due to constructional
considerations. The term periderm, intro-
duced by WIMAN (1895), suggests a dermal
construction of the tubarium, which is now
known to be incorrect. The tubarium is
formed from glands on the head shield of the
zooids and is not a dermal construction as is
the coenecium of a bryozoan. The tubarium
can perhaps be compared with the formation
of a hornet’s nest, though a hornet’s nest is
largely constructed with foreign material. A
comparison with hydrozoan colonies also
falls short, as these are often covered by an
organic exoskeleton, the perisarc (also iden-
tified as the periderm in HYMAN, 1940, p.
400), which is secreted by the epidermis of
the advancing stolon (e.g., BERKING, 2006).
Early graptolite literature is written in a
number of languages, including Chinese,
Czech, English, French, German, Latin,
Norwegian, Spanish, and Swedish. Thus,
paleontological terms describing grapto-
lite features were created in a variety of
languages. Many terms originally proposed
in other languages were translated into
English before they became the standard,
© 2014, The University of Kansas, Paleontological Institute, ISSN 2153-4012
Jörg Maletz, Denis E. B. Bates, Edsel D. Brussa, Roger A. Cooper, Alfred C. Lenz, John F.
Riva, Blanca A. Toro, & Yuandong Zhang. 2014. Part V, Revised, Chapter 12: Glossary of the
Hemichordata. Treatise Online 62:1–23, 6 fig.