© The Norwegian Academy of Science and Letters 2005 • Zoologica Scripta, 34, 3, May 2005, pp313– 328 313
Worsaae, K., Nygren, A., Rouse, G. W., Giribet, G., Persson, J., Sundberg, P. & Pleijel, F.
(2005). Phylogenetic position of Nerillidae and Aberranta (Polychaeta, Annelida), analysed by
direct optimization of combined molecular and morphological data. — Zoologica Scripta, 34,
313- 328.
The phylogenetic position of the most speciose meiofaunal polychaete family, Nerillidae, has
remained contentious. Recent hypotheses have generally focused on the fact that Nerillidae
shares with Aciculata (a major polychaete subgroup) features such as compound chaetae, ven-
tral buccal organ and short ventrolateral palps. Here we present the first phylogenetic analysis
of Aciculata, together with Nerillidae, combining morphological and molecular data. We also
include Aberrantidae, previously referred to or placed near to spiomorph polychaetes, but
recently referred to Aciculata, possibly close to Nerillidae. The data sets of 24 terminals con-
tain 53 morphological characters and nearly complete sequences of 18S rRNA. The sequences
were analysed simultaneously with the morphological data by direct optimization in the
program POY with a variety of parameter settings (costs of gaps: transversions: transitions).
The various settings resulted in markedly different phylogenetic hypotheses, but on the basis
of congruence (ILD) the results of two parameter settings were chosen. In all analyses, the
three included nerillid species constituted a monophyletic group. Only two analyses provided
fully resolved cladograms. The morphological analysis gave poor resolution and the position
of the nerillids was equivocal. The two molecular-based cladograms (minimizing ILD) were
also poorly resolved, but one provided a position for nerillids next to Eunice pennata and
Nothria conchilega, from the subgroup Eunicida within Aciculata. The two cladograms of the
combined analyses (minimizing ILD) were fully resolved and placed nerillids in a terminal
position next to Aberranta sp., within a clade of eunicidan species. The study showed that
the analytical conditions for the homology assignment of 18S rRNA strongly influenced
the phylogenetic results. The various previous proposals on the phylogenetic position of the
Nerillidae are reviewed, some of which are in accordance with the results of the present study.
Katrine Worsaae, Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100
Copenhagen, Denmark. E-mail: kworsaae@zmuc.ku.dk
Arne Nygren, Jenny Persson, Per Sundberg, Zoological Institution, Göteborg University, Sweden
Greg W. Rouse, South Australian Museum, and Earth and Environmental Sciences, University of
Adelaide, Australia
Gonzalo Giribet, Department of Organismic & Evolutionary Biology and Museum of Comparative
Zoology, Harvard University, Cambridge, MA, USA
Fredrik Pleijel, National Museum of Natural History, Paris, France
Blackwell Publishing, Ltd.
Phylogenetic position of Nerillidae and Aberranta (Polychaeta,
Annelida), analysed by direct optimization of combined
molecular and morphological data
KATRINE WORSAAE, ARNE NYGREN, GREG W. ROUSE, GONZALO GIRIBET, JENNY PERSSON, PER SUNDBERG
& FREDRIK PLEIJEL
Accepted: 13 December 2004
doi:10.1111/j.1463-6409.2005.00190.x
Introduction
Nerillidae Levinsen, 1883 is, with 49 species in 18 genera, the
most speciose meiofaunal family in Polychaeta. Nerillids
possess seven to nine chaetigerous segments and are all
minute, with most species ranging from only 300 μm to
2 mm in length. They are found in marine and brackish
habitats, from the intertidal to abyssal depths, and on all con-
tinents except Antarctica. The single freshwater species is
reported from groundwater and mountain rivers in North
America and continental Europe. Nerillids are mainly known
from the interstitial environment, although several species
are found in muddy habitats or caves with silt. The wide