© The Norwegian Academy of Science and Letters • Zoologica Scripta, 29, 3, July 2000, pp247–264 247
Grischenko, A. V., Mawatari, S. F. & Taylor, P. D. (2000). Systematics and phylogeny of the
cheilostome bryozoan Doryporella. — Zoologica Scripta, 29, 247–264.
The four nominal species of the Arctic-Boreal anascan bryozoan Doryporella Norman are
revised. Colonies of Doryporella are small, typically encrust shells and pebbles, and have
autozooids with frontal walls bearing a distinctive reticulate pattern of polygonal ridges.
High resolution SEM shows the frontal wall to be an interior wall (cryptocyst), contrary to
previous studies which have regarded it as an exterior wall (gymnocyst). The inter-relationships
between the species assigned to Doryporella are assessed cladistically using skeletal characters.
The type species (D. spathulifera) and two other species (D. alcicornis and D. armata) form a
well-supported clade, but the fourth species (D. reticulata) is more closely-related to certain
other calloporid genera. Therefore, the reticulate frontal wall is inferred to have evolved
twice and D. reticulata is removed to Doryporellina gen. n. Biogeographical distributions of the
four species are reviewed.
Andrei V. Grischenko, Department of Hydrobiology and Ichthyology, Faculty of Biology and Soil
Sciences, St Petersburg State University, 16 Liniya 29, St Petersburg, 199178, Russia (address for
correspondence: Phontannaya 4-92, Perm 614002, Russia). E-mail: gat@mail.perm.ru
Shunsuke F. Mawatari, Division of Biological Sciences, Graduate School of Science, Hokkaido
University, Sapporo 060, Japan. E-mail: shunfm@sci.hokudai.ac.jp
Paul D. Taylor, Department of Palaeontology, The Natural History Museum, Cromwell Road,
London SW7 5BD, UK. E-mail: pdt@nhm.ac.uk
lackwell Science, Ltd
Systematics and phylogeny of the cheilostome bryozoan
Doryporella
ANDREI V. GRISCHENKO, SHUNSUKE F. MAWATARI & PAUL D. TAYLOR
Accepted: 25 October 1999
Introduction
Doryporella is a distinctive and yet poorly known cheilostome
bryozoan with an Arctic-Boreal distribution and no fossil
record. Species of Doryporella are recognizable by the unusual
polygonal, reticulate ornamentation of their frontal walls, a
feature not found in the other genera of calloporid anascans
with which Doryporella has usually been grouped. In some
species, including the type species D. spathulifera (Smitt 1868),
the frontal wall is so extensive and ascophoran-like that the
identity of the genus as an anascan cheilostome has been called
into question. Levinsen (1916), for example, when placing
D. spathulifera into the anascan genus and subgenus Mem-
branipora (Callopora), noted that it had been referred by
previous workers to three different genera of ascophorans.
The origin and type of frontal wall in Doryporella has also been
a matter of contention (Ryland 1963: 10): is it a gymnocyst
(i.e. an exterior wall with epithelium on the inner side and
cuticle on the outer side) or a cryptocyst (i.e. an interior wall
with epithelium on both inner and outer sides)?
Given the problems over the affinities of Doryporella and the
lack of any modern descriptions utilizing scanning electron
microscopy (SEM), restudy of this distinctive genus is clearly
necessary. This is especially so in view of the need for data
for the forthcoming revision of the cheilostome bryozoan
volume of the ‘Treatise on Invertebrate Palaeontology’. Here
we take advantage of newly collected material from the north-
west Pacific, supplemented by existing material in The Natural
History Museum, London (NHM), to undertake a systematic
revision of Doryporella and to review what little is known about
the biogeographical distributions and ecologies of the four
nominal species. A further objective is to present an inter-
pretation of the phylogenetic relationships between the
species of Doryporella using cladistic methodology, and to ask
whether the genus as currently understood is monophyletic.
Materials and methods
New specimens were obtained during cruises of the ‘ R.V.
Nazarovsk’ (September 1988), the ‘R.V. Akademic Oparin’
(14th Cruise, August-September 1991) and the Middle Fishery
Refrigerator Trawler ‘ Rodino’ (September 1992) to the northern
Pacific. Additional material was furnished from the existing
reference collections of bryozoans held in the Department
of Zoology at The Natural History Museum, London. Most
of the SEM was undertaken on dried, uncoated colonies,