Aplacophora - Eumollusca: - Autapomorphies (Fig. 6 2)

Aplacophora 11
Aplacophora - Eumollusca
The Aplacophora and the other taxa combined within the Eumollusca are
identified as the highest-ranking adelphotaxa of the Mollusca. Both are
characterized by a series of apomorphous features. They both also have a
number of plesiomorphies. They therefore fulfill the criteria for their re-
cognition as sister groups.
The most conspicuous differences are found in the periphery. Through
out-group comparison the Aplacophora can be interpreted as primary
shell-less Mollusca; their worm-shaped form is, however, an apomorphy.
The Eumollusca continue to have as a plesiomorphy the ground pattern di-
vision into a broad foot and flat visceral mass. In their stem lineage the
shell arises as an evolutionary novelty.
Aplacophora
For the following features an evolution within a common stem lineage of

the Solenogastres and Caudofoveata is hypothesized .
• Autapomorphies (Fig. 6 ---+ 2)

- Evolution of a vermiform body together with a change in habitat from
the sediment surface to within soft sediment.
- Reduction of the broad foot with creeping sole. In the ground pattern of
the Aplacophora only a ventral groove with a folded foot remains.
- Papilla cells of the epidermis with a presumed excretory function. Single
large cells are pushed with the cuticle into the periphery; they evacuate
the contents of apical vesicles to the exterior.
- Mantle cavity as an extension of the mantle groove at the posterior.
- Gonopericardioducts connect the gonads with the pericardium. Separate
gonoducts - as in the Eumollusca - are not seen. Instead the gametes
are led via the pericardium and the pericardioducts into the mantle cavity.
- Mucus channels (mucus grooves), differentiation products of the mantle
cavity, conduct gametes into the water.
Solenogastres - Caudofoveata
Solenogastres
Marine organisms which are either epibenthic or live within the sediment.
They exist partly on polyp colonies of Cnidaria and feed on them.
Body length can range from a few mm to 30 cm. Nematomenia, Doryme-
mao
P. Ax, Multicellular Animals

© Springer-Verlag Berlin Heidelberg 2000
12 ApJacophora
A prominent feature of the Solenogastres is the foot groove (ventral fur-

row, ventral fold) which comes from the ground pattern of the Aplaco-
phora. In the longitudinal slit on the ventral side, the foot is found in the
form of one or more folds. Mucus glands with openings into an anterior
ciliated pit and along the furrow produce a band of mucus on which the
Solenogastres glides.
Following separation from the Caudofoveata line, the features listed be-
low must have evolved in the stem lineage of the Solenogastres.
• Autapomorphies (Fig. 6 ~ 3)
- Reduction of the gills.
- Change to hermaphroditism.
- Opening of the gonoducts into voluminous spawn ducts formed from in-
vagination of mucus channels of the mantle cavity into the body.
Caudofoveata
Inhabitants of soft marine sediment out of which the posterior end with
gills may project. Body length can vary from between 3 mm to 14 cm.
Chaetoderma, Falcidens.
The Caudofoveata have a cylindrical body that is, with the exception of
the foot shield, fully covered with calcareous scales embedded in the cuti-
cle. The ventral groove is reduced, as is the foot along the ventral side. As
a presumed remnant of the foot a shield-like flexible structure behind the
oral opening is formed. This functions as a burrowing tool in the sedi-
ment. The oral shield is one of the basic autapomorphies of the Caudofo-
veata.
The Caudofoveata are plesiomorphous in being gonochoristic organisms
and in possessing two gill combs (ctenidia) in the mantle cavity.
- Oral shield (Fig. 9 E).
- Free hemoglobin in serum.
- Fusion of the pleurovisceral and pedal nerves at the posterior to form a
longitudinal stem on both sides.
Eumollusca (Testa ria)
- Solid calcareous shell in the form of eight plates on the dorsal surface.
Eumollusca 13
B c
o E
Fig. 9. Aplacophora. A-C Solenogastres. A Nematomenia banyulensis on a Cnidaria colony.

B Epimenia verrucosa. View of the ventral side with foot groove. C Dorymenia antarctica.
Transverse section through the groove and foot. D-G Caudofoveata. D Scutopus ventrolinea-
tus. E Limifossor talpoideus. Ventral view of the anterior with the partitioned foot shield in
various states. F Chaetoderma nitidulum. Section through the epidermis with calcareous
stalk and papilla cells (latter: autapomorphy of the Aplacophora). G Falcidens CTOssotuS. Pos-
terior ctenidia. cu Cuticle; ec epidermis cell; f foot; pc papilla cell; sp spiculum. (A,B,D-G
Salvini-Plawen 1971; C Kilias 1993a)
- 16 pairs of dorsoventral muscle bundles used as foot retractors, two

pairs per shell plate. Through contraction of the muscles the body is an-
chored firmly to the substrate.
- One pair of kidneys - differentiation products of the pericardioducts.
The sac-like kidneys are located at the sides of the body.
14 Polyplacophora
The number of gills in the stem species of the Eumollusca is disputed.

Most probably, a pair of bipectinate ctenidia was inherited as a plesiomor-
phy from the ground pattern of the Mollusca. Independent additions then
occurred in the stem lineages of the Neopilinida, Polyplacophora and Nau-
tiloidea (Cephalopoda).
Polyplacophora - Conchifera
Polyplacophora
The chi tons inhabit marine littoral hard substrates, especially the surf
zone of rocky coastlines. Their body length usually ranges from between a
few mm to a few cm; the maximum reached is seen in Cryptochiton stelleri
at more than 30 cm.
• Autapomorphies (Fig. 6 -+ 6)
- Tegmentum with aesthetes.
The outer calcareous layer of the shell plates contammg organic sub-
stances is traversed by cell processes which are components of the sen-
sory apparatus.
- Articulamentum.
Differentiation of the hypostracum to anchor the shell plates beneath
one another and into the girdle.
Their eight dorsal shell plates create the most noticeable feature of the
Polyplacophora - indeed, these plates gave rise to the name of the unity
and make them ideally suited for identification. Nevertheless, the eight cal-
careous plates are not in themselves a reason for establishing the Poly-
placophora as a monophylum. It can be hypothesized that a shell consist-
ing of eight plates is more likely to be a feature that was already present in
the ground pattern of the Eumollusca as an evolutionary novelty. In com-
parison to the single shell of its sister group Conchifera, this feature must
be viewed as a plesiomorphy of the Polyplacophora.
Two pairs of foot retractors are inserted at every plate. These must also
be considered as a plesiomorphy which originated together with the evolu-
tion of the eight plates in the stem lineage of the Eumollusca. The eight
shell plates overlap each other rather like roofing tiles from the anterior to
the posterior. They are framed by the perinotum, a further plesiomorphic
feature of the Polyplacophora. The soft girdle with cuticle and calcareous
spicules or spines is the remains of the original body wall of the Mollusca.
The most usual relationship between the broad plates and narrow girdle is
best seen in Lepidochiton cinereus or Tonicella lineata. Secondary enlarge-

Aplacophora - Eumollusca: - Autapomorphies (Fig. 6 2)

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Aplacophora 11

For the following features an evolution within a common stem lineage of

• Autapomorphies (Fig. 6 ---+ 2)

P. Ax, Multicellular Animals

A prominent feature of the Solenogastres is the foot groove (ventral fur-

Eumollusca (Testa ria)

Fig. 9. Aplacophora. A-C Solenogastres. A Nematomenia banyulensis on a Cnidaria colony.

- 16 pairs of dorsoventral muscle bundles used as foot retractors, two

The number of gills in the stem species of the Eumollusca is disputed.

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