Aplacophoran Mollusks—Still Obscure and Difficult?

Author(s): Christiane Todt

Source: American Malacological Bulletin, 31(1):181-187. 2013.
Published By: American Malacological Society
DOI: http://dx.doi.org/10.4003/006.031.0110
URL: http://www.bioone.org/doi/full/10.4003/006.031.0110

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Amer. Malac. Bull. 31(1): 181–187 (2013)

Aplacophoran mollusks–Still obscure and difficult?*

Christiane Todt

University Museum of Bergen, The Natural History Collections, University of Bergen, Allégaten 41, 5007 Bergen, Norway
Correspondence, Christiane Todt: Christiane.Todt@um.uib.no

Abstract: In numbers, the aplacophoran mollusks are of subordinate importance amongst the Mollusca and they remain a somewhat obscure
group as compared to gastropods, bivalves, or even chitons. The ongoing confusion when it comes to naming, categorizing, and classifying
what is known as either the class Aplacophora with subclasses Chaetodermomorpha and Neomeniomorpha, or the two classes Solenogastres
and Caudofoveata, is not helpful in this matter, either. Nevertheless the last decade has brought a number of important results when it
comes to aplacophoran biology, morphology, taxonomy, and currently also aplacophoran phylogeny. These results point to a sistergroup
relationship between Solenogastres and Caudofoveata, but also show that there are a number of crucial differences that support a classification
into two separate classes. Here, the current status of aplacophoran research is outlined and important open questions are discussed as focus
areas for future studies.

Key words: Solenogastres, Neomeniomorpha, Caudofoveata, Chaetodermomorpha, Mollusca

Amongst the Mollusca, the aplacophoran taxa are excep-
tional. Their body is worm-shaped and covered in cuticle and
calcareous sclerites (Fig. 1), the foot is narrow or lacking, the
radula has few teeth per row and is simple (or, as some would
say: highly reduced and derived), the digestive tract lacks a
midgut gland, the excretory system lacks a kidney, and the
gonad is connected to the pericardium. These morphological
features are unique as compared to other mollusks and put
the aplacophorans in a special place when discussing mollusk
relationships and evolution (for review see e.g., Todt et al.
2008). By some, they are considered to have the most plesio-
morphic morphology (Salvini-Plawen 1985a, Haszprunar
2000, Salvini-Plawen 2003); alternatively they are suggested
to represent a reduced sideline (Scheltema 1993, Ivanov 1996,
Vinther et al. 2011). By some, they are thought to represent
two class-level taxa, Solenogastres and Caudofoveata, of a
“primitive” mollusk grade *Aplacophora*, the asterisks indi-
cating a paraphyletic grouping (Salvini-Plawen 1985a, Salvini-
Plawen and Steiner 1996, Haszprunar 2000). Alternatively,
they have been classified as Neomeniomorpha and Chaeto-
dermomorpha in the class Aplacophora (Pelseneer 1906,
Scheltema 1978, 1988, 1993). To conclude, analyses based on
morphological characters clearly have not been sufficient to
clarify phylogenetic relationships of the aplacophoran taxa
(see also contribution by Kevin Kocot, this issue).
Not only because of their exceptional morphology, the
aplacophoran mollusks are considered to be somewhat obscure
animals. They are mostly known from fixed material origi-
nating from large-scale deep-sea sampling cruises. Observa-
tions of living specimens have up to now been rare and were
to a large extent restricted to a few known shallow-water taxa
(see below). The two aplacophoran taxa appear to be much
less species rich than most other molluscan class-level taxa.
Currently there are about 280 species of Solenogastres de-
scribed, and 130 species of Caudofoveata (but see below on
species diversity estimates).
It is easy to identify an aplacophoran mollusk as a
solenogaster or caudofoveate, but detailed taxonomic knowl-
edge has at all times been kept to an extremely small group of
people. Biodiversity inventories, for example, generally treat
all aplacophorans as a single category (Aplacophora; or some-
times: Caudofoveata) and even most malacologists shy away
from dealing with them. The most common explanations are:
(1) Aplacophorans are difficult to collect, (2) they are te-
dious to identify and describe (taxonomically difficult), (3)
their DNA is problematic in sequencing (contamination is-
sues), and (4) they are almost impossible to observe in life
and to keep under laboratory conditions.
In the following paragraphs I want to show that even if
some of these discouraging statements may have justification,
they do not necessarily have to be an impediment to studying
aplacophoran mollusks.
1) “Aplacophorans are difficult to collect.”—Most apla-
cophorans indeed live in deep subtidal to bathyal habitats

* From the “Mollusks: The Great Unanswered Questions. The James H. Lee Memorial Symposium” presented at 77th Annual Meeting of the
American Malacological Society on 24 July 2011 in Pittsburgh, Pennsylvania. All symposium manuscripts were reviewed and accepted by
the Symposium Organizer and Guest Editor, Dr. Timothy A. Pearce.
181
182 AMERICAN MALACOLOGICAL BULLETIN · ·
31 1 2013

Figure 1. Living specimens of aplacophoran mollusks from Norway. A–C, Caudofoveata. A, Chaetoderma nitidulum Lovén, 1844, scale bar:
1 mm; B, Scutopus ventrolineatus Salvini-Plawen, 1968, scale bar: 0.3 mm; C, Falcidens crossotus Salvini-Plawen, 1968, scale bar: 0.2 mm; D–H,
Solenogastres: D, Neomenia carinata Tullberg, 1875, scale bar: 3 mm; E, Stylomenia sulcodoryata Handl and Salvini-Plawen 2001, scale bar:
1 mm; F, Simrothiella margaritacea (Korén and Danielssen, 1877), scale bar: 1 mm; G, Helluoherpia aegiri Handl and Büchinger, 1996, scale
bar: 1mm; H, Kruppomenia borealis Odhner, 1921, scale bar: 1 mm.

(Table 1), but the necessity of using ship-based equipment
for collecting marine invertebrates has not much hindered
extensive research on such elusive organisms as Xenoturbella
bocki Westblad, 1949 (phylum Xenoturbella), or—as an ex-
treme example—the hydrothermal vent giant tubeworm,
Riftia pachyptila Jones, 1981. And not all aplacophorans are
deep-sea dwellers. Some solenogasters live in shallow subtidal
sands of the Atlantic coasts of Spain and France (Biserramenia
psammobionta Salvini-Plawen, 1967) or in knee-deep (and
slightly deeper) waters off Bermuda, Florida, and Panama
(e.g., species of Meiomenia Salvini Plawen, Rieger and Sterrer,
1985 and Meioherpia Salvini Plawen, Rieger and Sterrer,
1985). Giant Epimenia Nierstrasz, 1908, up to 30 cm in body
length, and slightly smaller species of Anamenia Nierstrasz,
1908, (e.g., Anamenia amabilis Saito and Salvini-Plawen, 2010)
inhabit tropical coral reefs off New Guinea, Australia, Japan,
and in the Red Sea. Moreover, there are extensive museum
collections of aplacophoran material, collections with an ex-
pected number of yet undescribed species surmounting the
known diversity at least tenfold. For Australian waters, for ex-
ample, there are six species of solenogasters and four species
of caudofoveates listed in the Zoological Catalogue of Australia
(Scheltema 2001), while the same author mentions a di-
versity of about 65—to date largely unknown—aplacophoran
species from three deep-water surveys (200–1850m) in the
Bass Strait and the continental slope off South East Australia
(Scheltema 1990). One sled sample from the latter survey
yielded no less than 536 specimens within 22 morphologically
 APLACOPHORAN MOLLUSKS
Table 1. Global bathymentric distribution of aplacophoran mollusks. Aplacophoran mollusks occur from shallow subtidal areas of the
continental shelf down to hadal depths of 9000 m. The highest diversity can be found on the continental slope and rise.
Bathymetric zone Depth (m)
Continental shelf < 200
Continental slope 200–1000
Continental rise 1000–4000
Abyss 4000–6000
Hadal I 6000–9000
Hadal II > 9000
clearly distinct species (Scheltema 1990). Such high abun-
dances of aplacophorans (e.g., Scutopus ventrolineatus Salvini-
Plawen, 1968, Wirenia argentea Odhner, 1921, Simrothiella
margaritacea (Korén and Danielssen, 1877), Micromenia
fodiens (Schwabl, 1955), Genitoconia rosea Salvini-Plawen,
1967) can also be found in Europe, for example along the
continental slope of the Norwegian Sea, including the larger
fjord systems. The species diversity here usually is somewhat
lower (up to 7 species within one sled tow), but my own
count of Norwegian solenogaster species currently has
reached a number of more than 40 (about one-third of these
are new to science)—one seventh of the known described
global diversity. Many of these can be collected fairly close
to the coast, albeit in deep waters (150–800 m). Grassle and
Maciolek (1992) and Scheltema and Ivanov (2009) report a
hitherto unsurpassed abundance of two species of the Pro-
chaetodermatidae in the Northwest Atlantic off the coast of
New Jersey, in water depths of 1500–2500m. The caudofove-
ates made up 5–23% of the total fauna count, with an aston-
ishing maximum density of these micro-omnivorous animals
of 346 specimens per square meter.
2) “Taxonomic work on aplacophoran material is
tedious.”—This cannot be denied (but then it is true for most
taxonomic work). Taxonomic literature for aplacophorans of
a given geographical area is usually highly dispersed and often
published in somewhat obscure journals, many in German,
some in Russian or Japanese. For solenogasters, García-Álvarez
and Salvini-Plawen (2007) have taken an effort to compile
taxonomic information on the generic level, including a key to
the families. One of the major impediments of using this key
(and solenogaster taxonomic literature in general) is the need
to produce histological serial sections for identification. Much
of the old (and new) descriptive work is based on histology
but information about histologic characters are difficult to ac-
cess for a general taxonomist who routinely sorts and identi-
fies fauna—for example for environmental monitoring or for
curatorial purposes. Using epidermal sclerite, copulatory stylet
and radular morphology for species identification is possible
in many cases (Scheltema and Schander 2000) but not always—
there are species lacking both radula and/or copulatory
183
No. species Solenogastres No. species Caudofoveata
37 20
122 53
58 58
9 7
1 2
0 0
stylets. But even for species with distinctive hard parts, de-
tailed illustrations of these important taxonomic characters
are often lacking, especially in older species descriptions. As a
result of dispersed taxonomic literature and the need for rela-
tively time-consuming techniques, aplacophoran taxonomy
currently has the status of a “secret science”. Could modern
techniques such as micro-computer tomography for the de-
piction of internal hard parts (radula, copulatory stylets) and/
or molecular barcoding provide future solutions for the iden-
tification (not description!) of aplacophorans? In January
2011 we started a systematic inventory of aplacophoran mol-
lusks in the European North Atlantic with special focus on
Norwegian waters. The goals of this three-year project is to
map species distributions, but also to summarize taxonomic
knowledge in a monograph including high-quality photo-
graphic illustration of species (see Figure 1 for some exam-
ples), to establish a reference collection at the University
Museum of Bergen (Norway), and to generate a catalogue of
DNA barcodes. This endeavor will not only facilitate future
species identification in the geographical area, but also the
training of a future generation of aplacophoran taxonomists.
Similar efforts in other parts of the world are sorely needed.
3) “Aplacophorans are problematic in DNA sequencing”—
Specifically the solenogasters have been notorious for
sequencing problems related to contamination by prey or-
ganisms (Okusu and Giribet 2003). This is certainly true for
the nuclear 18S rRNA gene, where complicated secondary
structures and GC rich sequence areas hamper the amplifica-
tion process (Meyer et al. 2010), while mitochondrial genes
are in general less prone for such issues (H. Dreyer, N.
Mikkelsen, and D. Osca Ferriol, pers. comm.). Contamination
problems even led to a phylogenetic placement of Solenogas-
tres outside of Mollusca (Wilson et al. 2010). In addition,
the aberrant sequence composition of at least some genes in
both Solenogastres and Caudofoveata seems to be prone to
produce artificial groupings in phylogenetic reconstructions
(see Meyer et al. 2010, for discussion). In a number of pub-
lished analyses the aplacophoran taxa for example group with
Cephalopoda and/or Scaphopoda (Passamaneck et al. 2004,
Giribet et al. 2006, Hejnol et al. 2009; Meyer et al. 2010,
184 AMERICAN MALACOLOGICAL BULLETIN
Wilson et al. 2010, Vinther et al. 2011), thus annihilating the
morphologically well-supported clade Conchifera (shell-
bearing mollusks). A combination of a broad gene sampling
(several hundred genes) with a wide taxon representation (at
least 2–3 representatives per mollusk class-level taxon) ap-
pears to lead to more plausible results, as recently indicated
by first phylogenomics approaches to resolve molluscan deep
phylogeny (Kocot et al. 2011, Smith et al. 2011). Having
learned from the experiences of the last decade, the difficul-
ties of analyzing the phylogenetic relationships of aplacopho-
rans with molecular tools now largely have been boiled down
to the restricted availability of fresh tissue or material fixed
properly for DNA- and RNA-based molecular work, espe-
cially regarding analyses of internal relationships.
(4.) “Aplacophorans are almost impossible to observe in
life and to keep under laboratory conditions”— There are in
fact only a few scientists who have ever seen a living aplacoph-
oran (most of them appear to be meiofauna experts having
experienced solenogasters in their sand samples) and nobody
has yet succeeded in establishing permanent cultures. But no
matter if dealing with tropical or cold-water species: if there is
no physical damage afflicted during the collecting process and
if ambient water quality and temperature are provided, these
animals persist even without appropriate food, allowing for
extended observation periods. Many solenogasters, especially
species coming from water depths less than 300 m, can be kept
for more than a month in the laboratory, some up to four
months (Thompson 1960, Salvini-Plawen and Benayahu
1990, Todt and Wanninger 2010). In these cultures they digest
(Baba 1940; Todt and Salvini-Plawen 2004b), move (Salvini-
Plawen 1968), breathe, react to stimuli, they reproduce and
their larvae develop (e.g., Thompson 1960, Okusu 2002,
Nielsen et al. 2007). In a recent study, Todt and Wanninger
(2010) followed the development of Wirenia argentea, one of
the most common Norwegian solenogaster species. These
animals bear very few ripe oocytes at a time (4–6), but produc-
tion is constant and the reproduction period extends over sev-
eral months. The oocytes are internally fertilized by sperm
stored within the “mother” (solenogasters are hermaphro-
dites and copulate to transfer sperm) and uncleaved embryos
are released. The embryos develop into typical pericalymma
larvae with the growing juvenile body protected by a cap of
cleavage-arrested cells bearing ciliary trochi for locomotion.
Development and metamorphosis are very gradual, the apical
cap (calymma) is partially shed and partially absorbed, the ju-
venile trunk enlarges and the foot ciliation emerges while the
larva still is able to swim. Crawling postlarvae (juveniles) were
kept until an age of two months in the laboratory. At that
time, the yolk deposits were used up and the digestive tract
(including a small radula) was almost fully developed. As ear-
lier described for some solenogaster larvae (Thompson 1960,
Okusu 2002) but not all (see: Scheltema and Ivanov 2002), the
· ·
31 1 2013
epidermal sclerites in the developing larva and juvenile were
not serially arranged and thus not reminiscent of chiton shell
fields. But is the status in Wirenia argentea plesiomorphic or
derived? To date, cell-lineage and gene expression studies are
lacking and only few species are investigated, but work is in
progress and hopefully soon the new results will shed a light
on some of the open questions.
The “worm-mollusks”—How different are the
aplacophoran taxa?
Despite of striking similarities, especially the worm-
shape and the epidermal sclerites, there are a number of
profound differences between the two aplacophoran taxa
(Solenogastres versus Caudofoveata): presence or absence of a
foot, lack or presence of typical mollusk gills (ctenidia), and
uniform midgut or midgut in compartments (stomach, mid-
gut sack, intestine), respectively. Solenogastres are carnivo-
rous, feeding mostly on cnidarian polyps (Salvini-Plawen
1967) or polychaetes (Todt and Salvini-Plawen 2005) al-
though some appear to be scavengers on jellyplankton (C.
Todt, pers. obs.). Most caudofoveates, in contrast, stuff their
digestive tracts with detritus rich in organic contents (Scutopus
Salvini-Plawen, 1968, and species of Chaetoderma Lovén, 1844
C. Todt, pers. obs.) or they feed selectively on foraminiferans
or small particles (Scheltema and Ivanov 2009). While at least
a number of caudofoveates use a typical molluscan hemocya-
nin as respiratory protein, this was not found for solenogas-
ters collected from the same habitats (Lieb and Todt 2008).
Significant differences are also to be found when it comes to
reproductive biology: solenogasters are internally-fertilizing
hermaphrodites with introsperm and caudofoveates have
separate sexes and are free-spawning with ectaquasperm
(Buckland-Nicks 1985). Interestingly, the central nervous sys-
tem of the burrowing, micro-omnivorous, and free-spawning
caudofoveates is more complex than that of the epibenthic,
“carnivorous” solenogasters, most of which ingest dangerous
cnidarian prey in addition to having to coordinate courtship
and copulation (Faller et al. 2012). In summary, I propose that
the differences between Solenogastres and Caudofoveata can
be rated as so profound that, no matter whether the taxa are
sistergroups or not, they should be treated as separate classes.
Aplacophoran relationships
Are the Solenogastres and Caudofoveata derived from
chiton-like, shell plate bearing ancestors (Polyplacophora),
eventually via pedomorphosis (Scheltema 1993, Vinther et al.
2011)? Or can the recent morphology of aplacophorans, espe-
cially solenogasters, serve as a model for the ancestral state
within mollusks (see Haszprunar 2000, Salvini-Plawen 2003)?
To date, the internal relationships of Solenogastres and
Caudofoveata are not yet resolved but recently phylogenomics
(Kocot et al. 2011, Smith et al. 2011) and other molecular
 APLACOPHORAN MOLLUSKS
phylogenetic approaches (Vinther et al. 2011) begin to render
results that point to a sistergroup relationship of the apla-
cophoran taxa (see also Kocot 2013 in this volume). These re-
sults are in accordance with the Aplacophora hypothesis
(Scheltema 1988), as opposed to the Testaria hypothesis
(Salvini-Plawen and Steiner 1996), the latter seeing Soleno-
gastres and Caudofoveata as two independent early branches
of the molluscan tree of life. Some say the last word is not
spoken, while others think they found the answer. But despite
of—or even driven by—the prevailing disputes about phylo-
genetic relationships, current morphological research in-
cludes studies on the excretory system, nervous system,
radula morphology and development, sperm ultrastructure,
as well as neurogenesis, myogenesis, and developmental gene
expression of aplacophoran molluscs. Placed into the right
context, these analyses will finally help us to learn about evo-
lutionary mechanisms leading to the recent aplacophoran
morphologies.
Future challenges and open questions
Aplacophoran mollusks are fascinating study organisms
much more diverse than commonly appreciated. In marine
sediments we can find caudofoveates up to 40 cm long, such as
Chaetoderma felderi Ivanov and Scheltema, 2007, or tiny inter-
stitial solenogasters, not longer than half a millimeter (Salvini-
Plawen 1985b, García-Alvarez et al. 2000); many solenogaster
species are colorful (Fig. 1), some are reef dwellers (Salvini-
Plawen and Benayahu 1990, Scheltema and Jebb 1994, Saito
and Salvini-Plawen 2010), and some are part of the dominat-
ing fauna at hydrothermal deep-sea vents (Scheltema 2000,
Todt and Salvini-Plawen 2005, Salvini-Plawen 2008). Apla-
cophoran molluscs may occur in high densities in continental
slope soft-bottom habitats (see above) and members of the
caudofoveate family Prochaetodermatidae can even be called
true opportunistic organisms (Scheltema and Ivanov 2009).
But what is the ecological role of aplacophorans in habitats
where they are highly abundant? Could they be used as indica-
tor organisms for certain marine soft-bottom communities?
Many solenogasters feed on cnidarian polyps without being
harmed by the cnidocysts of their prey (Salvini-Plawen 1967,
Todt and Salvini-Plawen 2004a). Do they produce chemical
compounds to protect themselves? Adult solenogasters do not
possess an obvious system to modify their primary urine (no
“kidney”) and in general they are not very effective in regulat-
ing their inner osmotic pressure (C. Todt, pers. obs.). Thus,
how do the hydrothermal vent solenogasters cope with such
an extreme habitat? Nobody knows about the function of
the complex copulatory apparatus of the hermaphrodidic
solenogasters or about their reproduction behavior in general.
How do aplacophoran mates attract each other, what are the
sensory systems and/or the pheromones involved? Are they
able to self-fertilize? Could solenogasters be good model
185
organisms for investigating speciation and the underlying
genetics in internally fertilizing hemaphrodites?
Even if there are still many open questions—or maybe
because of them—aplacophoran research currently is alive
and kicking, working on different fronts, with different tech-
niques and in a number of laboratories in Austria, Germany,
Spain, Russia, Japan, Norway, Canada, and the United States.
In fact this is quite a large effort for studying taxa as “small”
as the aplacophoran mollusks. Hopefully all these effort will
soon render the solenogasters and caudofoveates less ob-
scure, less difficult, but all the more fascinating.
ACKNOWLEDGMENTS
I am grateful to all my colleagues dedicated to aplacopho-
ran research for exchange of material, fruitful discussions, and
inspiring ideas. The University of Bergen is acknowledged for
funding via a series of three free researcher initiated projects
(# 226270 Friforsk) in the years 2008–2010; this is a contribution
to the Norwegian Taxonomy Initiative project # 70184222.
Thanks to the American Malacological Society for inviting me
to give a talk at the 2011 symposium “Great Unanswered
Questions in Malacology” and to write this contribution.
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Submitted: 31 January 2012; accepted: 15 September 2012;

final revisions received: 12 November 2012