JOURNAL OF MORPHOLOGY 255:131–145 (2003)
Further Structures in the Jaw Apparatus of
Limnognathia maerski (Micrognathozoa), With Notes
on the Phylogeny of the Gnathifera
Martin Vinther Sørensen*
Invertebrate Department, Zoological Museum, University of Copenhagen, Denmark
ABSTRACT The jaws of Limnognathia maerski, Micro-
gnathozoa, were investigated with light- and scanning
electron microscopy. The study yielded several new struc-
tures and sclerites, including the ventral part of main jaw,
the pharyngeal lamellae, the manus, the dorsal and ven-
tral fibularium teeth, and a reinterpretation of the fibu-
larium compartmentalization. Furthermore, it was shown
that several jaw elements are composed of densely packed
rods. Comparison with Rotifera and Gnathostomulida
suggested that the micrognathozoan main jaw is homolo-
gous with the rotifer incus and the gnathostomulid articu-
larium and that the pseudophalangids (the ventral jaws)
and their associated sclerites correspond to the rotifer
mallei. These results imply that Micrognathozoa is more
closely related to Rotifera than to Gnathostomulida. J.
Morphol. 255:131–145, 2003. © 2002 Wiley-Liss, Inc.
KEY WORDS: jaws; morphology; phylogeny; taxonomy;
Gnathifera
Limnognathia maerski was recently described
from a lake and a cold spring on Disko Island,
Greenland (Kristensen and Funch, 2000). Its unique
morphology, including the combination of cellular
intraskeletal lamina in the dorsal epidermis, multi-
ciliate ciliary bands in the head region, and cilia
arranged as ciliophores in the ventral parts of the
head and trunk, a temporarily functional anus,
monociliate protonephridia, and an extremely com-
plicated jaw apparatus demanded the establishment
of a new class, Micrognathozoa Kristensen and
Funch, 2000, placed in Gnathifera Ahlrichs, 1997
together with Acanthocephala, Rotifera, and Gna-
thostomulida. One of the most exciting features in L.
maerski is the combination of being among the
smallest known Metazoa and simultaneously pos-
sessing the most complex pharyngeal apparatus.
The ultrastructure of the jaw elements suggests that
the jaws are homologous with those in Rotifera and
Gnathostomulida (Rieger and Tyler, 1995; Ahlrichs,
1997; Kristensen and Funch, 2000; Sørensen et al.,
2000), but the jaws in L. maerski comprise far more
elements than present in any species of Rotifera or
Gnathostomulida. Even though Kristensen and
Funch (2000) presented a thorough description of
the jaws, several elements and structures were
© 2002 WILEY-LISS, INC.
poorly understood or totally undescribed. Thus, fur-
ther studies on the micrognathozoan jaws were most
desirable, both to increase our knowledge of the
complexity of the creature and to obtain phyloge-
netically significant information about possibly ho-
mologous elements within Gnathifera.
In two recent cladistic studies on the phylogeny of
Gnathostomulida and Rotifera (Sørensen, 2002a,b),
micrognathozoan jaw morphology was used to polar-
ize some of the characters. These studies presented
a possible phylogeny on lower taxonomic levels
within the two groups, but the relationships be-
tween the three gnathiferan taxa possessing jaws
still remain unsettled. Ahlrichs (Ahlrichs 1995,
1997) placed Micrognathozoa (referred to as New
Group A) as a sister-group to Syndermata (Rotif-
era ⫹ Acanthocephala) based on the presence of an
intraskeletal lamina and the absence of lateral and
dorsal ciliation of the trunk. Kristensen and Funch
(2000) also preferred this solution, but mentioned a
Micrognathozoa-Gnathostomulida sister-group rela-
tionship as an alternative possibility, based on other
jaw similarities and because both groups lack a syn-
cytial epidermis.
The present study presents new data on the jaw
morphology in Limnognathia maerski. The obtained
results are compared with the jaw morphology in
Rotifera and Gnathostomulida and discussed in a
phylogenetic context. The study focuses on jaw mor-
phology, hence the position of Acanthocephala will
not be treated further because the animals lack a
mouth, pharynx, and digestive system.
Contract grant sponsor: the Danish Research Agency; Contract
grant number: 51-00-0278.
*Correspondence to: M.V. Sørensen, Invertebrate Department, Zoo-
logical Museum, Universitetsparken 15, DK-2100 Copenhagen, Den-
mark. E-mail: mvsorensen@zmuc.ku.dk
DOI: 10.1002/jmor.10038
132 M.V. SØRENSEN
MATERIALS AND METHODS
The jaws in Limnognathia maerski were investigated using
scanning electron microscopy (SEM) and light microscopy (LM).
Jaws from seven specimens were prepared for SEM and one for
LM. Furthermore, several unpublished LM photos taken by R.M.
Kristensen were reinvestigated. All material was obtained from
the species’ type locality, a cold spring near Isunngua, Disko
Island, Greenland (see Kristensen and Funch, 2000). Parts of the
material were collected by the author in June 1997 and 1999.
Other parts were collected by R.M. Kristensen and P. Funch in
July 1994. The samples were taken from submerged mosses and
animals were extracted by squeezing the water from the mosses
through a 30-␮m mesh sieve. The concentrate was either fixed
immediately in a 4% borax-buffered formaldehyde solution or
transferred to the laboratory at the Danish Arctic Station or the
Zoological Museum, University of Copenhagen, where the live
animals were sorted out.
Jaws for both LM and SEM were prepared by dissolving the
specimens with dilute sodium hypochlorite (approximately 1:10
ratio water). Specimens for SEM were placed in a drop of hypo-
chlorite in the center of a 12-mm circular coverslip, which was
fixed with a drop of water on a glass slide. When the specimen is
placed in the hypochlorite the softer tissues start to dissolve. The
process is easily observed with a light microscope and after 2–10
min the animal will be fully dissolved, leaving only the pharyn-
geal hard parts on the coverslip. A micropipette prepared from a
50-␮l capillary tube was used to remove the hypochlorite by
repeatedly adding and removing distilled water to the drop. The
washing should be done extremely carefully and the jaws should
continuously be observed during the process. When the hypochlo-
rite has been replaced with distilled water, most of the water is
removed. The remaining water will evaporate, causing the jaws to
stick to the cover slip. If the washing has been insufficient, chlo-
rite crystals will appear after the evaporation and more water
should then be added immediately. If the preparation process is
performed carefully, without too long treatment with hypochlo-
rite, most of the ligaments that connect the sclerites will be intact
and the different jaw elements will be arranged regularly in their
normal position relative to each other. However, it may be an
advantage to damage the ligaments to isolate certain sclerites or
to see jaw parts that otherwise would be hidden. This can be done
by repeatedly adding water and then letting it evaporate totally.
When the preparation is dry the coverslip is transferred to an
SEM aluminum stub. In this study the preparations were
sputter-coated with gold and examined with a JEOL JSM-840
microscope.
Preparation of jaws for LM was done slightly differently. The
dissolving and washing procedures are done directly on a glass
slide. When the jaws are isolated and clean they are mounted in
a drop of modified Faure’s medium to prevent the jaws from
moving. The modified Faure’s solution is mixed in following ratio:
water 50 ml, chloralhydrate 25 ml, glycerine 20 ml, and gum
Arabic 30 g. The amount of chloralhydrate is decreased with
respect to the recipe given by Adam and Czihak (1964) in order to
avoid too much bleaching. In this study the jaws were investi-
gated with an Olympus BX51 Nomarski microscope with a digital
camera.
The illustrations based on SEM were edited and set up in
Corel-PHOTO–PAINT 8 and CorelDRAW 8 and the 3D recon-
structions were made in Rhinoceros v. 2beta.
DESCRIPTION OF JAWS
The jaw apparatus in Limnognathia maerski is a
complex of numerous moveable or anchoring scle-
rites (Figs. 1– 6). It comprises six independent den-
ticulate paired structures. Two of these have only
been described superficially. Furthermore, it has a
pair of large, not previously described, pharyngeal
lamellae. The jaw sclerites can generally be divided
into four portions: the ventral jaw elements, the
main jaw, the fibularium, and the dorsal jaw ele-
ments. This division will be followed in the descrip-
tion below. The terminology generally follows Kris-
tensen and Funch (2000) unless otherwise noted.
Ventral Jaw Elements
A relatively thick cuticular membrane covers most
of the ventral side of the jaw apparatus (Fig. 1C,D).
The only sclerites without this membrane are the
rostral parts of the basal plate, the pseudophalan-
gium (the ventral jaws) with its associated sclerites,
and the pharyngeal lamellae (see description below).
The membrane attaches to the fibularium and the
caudal rods of the basal plates.
Three large, paired elements are located rostroven-
trally to the fibularium. These are the rostral parts of
the basal plate, the pseudophalangium, and the pha-
ryngeal lamellae. The pharyngeal lamellae have not
been described previously, even though they are a pair
of rather prominent structures (Figs. 1A–E, 2A,B,D,
4A,B, 5A,B, 6A). They resemble the oral lamellae (see
Kristensen and Funch, 2000) but are smaller (ca. 6 ␮m
vs. 9 ␮m) and, contrary to the oral lamellae, resistant
to hypochlorite treatment. Furthermore, they are lo-
cated posterior to the mouth opening. The pharyngeal
lamellae consist of a short shaft with 20–25 long, thin
lamellae. The proximal parts of the lamellae are fused
laterally but the distal halves are free. The lamellae
gradually thicken from the most ventral ones towards
the most dorsal ones. The three most dorsal lamellae
are strong and almost tooth-like. The shaft is short
and forms a ball-and-socket joint with a small cylin-
drical sclerite (Fig. 1C,D).
The pseudophalangia, or the ventral jaws, are large
isolated sclerites located close to the pharyngeal lamel-
lae (Figs. 1B–D, 2A,B,D, 4– 6). Each pseudophalan-
gium comprises a long shaft and five distal pseudodig-
its (Figs. 2B, 4A, 5A). The rostral surface of the shaft
has a distinct longitudinal ridge that bears numerous
transverse rods on its dorsal side (Fig. 2A). Caudally,
each pseudophalangium joins an accessory sclerite (by
Kristensen and Funch, 2000, referred to as as1). The
accessory sclerite is made up of numerous strands that
twist helically around the main axis of the sclerite
(Fig. 1A,D). In dorsolateral view the accessory sclerite
appears to be a rod with a rostral inward-projecting
extension and a smaller caudal, outward-projecting tip
(Figs. 1C,D, 4A), but in lateral view the sclerite is
flattened and triangular (Figs. 1A, 5B, 6B). The pseu-
dophalangia are solely connected to the jaw apparatus
through the accessory sclerites and in live animals
they are able to function independently of the other
jaw elements. Hence, the animal can move the pseu-
dophalangium from a rostrocaudal orientation to a
dorsoventral orientation without moving other jaw
parts.
The basal plate is formed by two paired, dentic-
ulate structures with long caudally projecting
 JAW APPARATUS OF L. MAERSKI 133

Fig. 1. Limnognathia maerski. Jaws. SEM. A: Dorsal view. B: Right lateral view. C: Ventral view. D: Ventral view. E: Rostral view.
F: Caudal view. as, accessory sclerite; ba, basal plate; ca, cauda; cs, cylindrical sclerite; dft, dorsal fibularium teeth; dj, dorsal jaw; dmj,
dorsal part of main jaw; dt, dorsal main jaw teeth; fi, fibularium; fro, fibula rostralis; ft, fibula transversalis; li, ligament; ma, manus;
pl, pharyngeal lamella; ps, pseudophalangium (ventral jaw); rlf, rostrolateral fenestra; sy, symphysis; vft, ventral fibularium teeth;
vmj, ventral part of main jaw; vt, ventral main jaw teeth.

shafts (Figs. 1C,D, 2B,D, 4A, 6A). The shafts are
inserted between the fibularium and the main
jaws. The denticulate parts are crescentic, located
rostrally to the fibularium and main jaws and
orientated obliquely to the main axis of the jaws.
Each plate has five teeth on the concave inner
margin. The teeth gradually increase in size from
the most caudal one towards the rostral one. A
pair of more or less isolated sclerites is somehow
associated with the denticulate parts of the basal
plates (Fig. 2B,D). These sclerites carry five well-
developed teeth. They resemble a pair of gloves or
hands and will therefore be referred to as the
manus (singular and plural ⫽ manus). The manus
attach proximally in the medial part of the lateral
margin of the basal plate. The connection, how-
ever, is rather weak and the manus are often
loosened and lost during preparation.
Main Jaws
The main jaw constitutes the central part of the
jaw apparatus in Limnognathia maerski (Figs. 1A,
134 M.V. SØRENSEN

Fig. 2. Limnognathia maerski. Different sclerites in the jaw apparatus. SEM. A: Pseudophalangium and pharyngeal lamellae.
B: Basal plate and manus, ventral view. C: Dorsal jaw, dorsal view. D: Basal plate and manus, frontal view. bp, basal plate; dft, dorsal
fibularium teeth; dj, dorsal jaw; dt, dorsal main jaw teeth; er, extrasclerital rods; ma, manus; pd, pseudodigits; pl, pharyngeal lamella;
ps, pseudophalangium.

4 – 6). It comprises an unpaired caudal part, a pair of
dorsal, denticulate, pincer-like main jaws, and a
pair of ventral elements densely covered with knobs
and rounded teeth.
The unpaired caudal part is composed of a sym-
physis and a paired cauda (Fig. 1A,B,D,F). The sym-
physis is apparently unpaired but a double ridge on
the ventral side implies that it could be formed by
secondarily fused structures. The symphysis is nar-
row in dorsoventral view but broad in lateral view
(Fig. 1B). The sides have no traces of a regular
surface ornamentation. However, in one of the in-
vestigated preparations the symphysis was broken
at its proximal end, which enabled examination of
the internal morphology of the element and it ap-
pears to be composed of densely packed rods similar
to those on the pseudophalangium (Fig. 3F). The
cauda attaches on the dorsal side of the symphysis.
No new information was obtained for this structure
(see Kristensen and Funch, 2000, for a detailed de-
scription).
The paired parts of the main jaw can be divided
into a dorsal and a ventral part. The dorsal part
carries the dentarium (Fig. 1A). The two parts are
normally held together and appear as one unit only
separated by a longitudinal cleft on the internal
surfaces of the pincers (Fig. 1E), but in one prepa-
ration the ventral and dorsal parts came out as
totally separated elements (Fig. 3).
The dorsal part of the main jaw is composed of an
adentate caudal shaft and the dentarium (Fig. 3C–
E). The outer surfaces of the shaft and the den-
tarium are smooth, whereas the internal surface is
densely covered with serrulae and knobs. A small
nodus is present laterally at the base of the den-
tarium (Fig. 3E) and most proximally, near the ar-
ticulation point, a small fenestra (fenestra dorsalis)
opens into a chamber (see fig. 16 in Kristensen and
Funch, 2000). The size of the fenestrae differs in the
different specimens, which implies that they are
covered by thin membranes that become more or
less dissolved during the hypochlorite treatment.
The dentarium has two sets of well-attached
teeth, the dorsal and ventral teeth (Fig. 3B–E). Dis-
tinctive medial teeth, as described by Kristensen
and Funch (2000), were not found. However, the
 JAW APPARATUS OF L. MAERSKI 135

Fig. 3. Limnognathia maerski. Different sclerites in the jaw apparatus. SEM. A: Left fibularium, dorsal view. B: Rostral parts of
fibularia and main jaw, lateral view. C: Collapsed parts of main jaw, dorsal view. D: Collapsed parts of main jaw, frontal view.
E: Collapsed parts of main jaw, laterocaudal view. F: Joint section between main jaw pincers and symphysis, lateral view. as, accessory
sclerite; ca, cauda; cd1– cd2, camera dorsalis 1–2; cl1– cl3, camera lateralis 1–3; dft, dorsal fibularium teeth; dj, dorsal jaw; dmj, dorsal
part of main jaw; dt, dorsal main jaw teeth; fc, fibula caudalis; fd, fibula dorsalis; fl, fibula lateralis; fr, fibula radialis; ir, intrasclerital
rods; no, nodus; pl, pharyngeal lamella; ps, pseudophalangium; rlf, rostrolateral fenestra; vft, ventral fibularium teeth; vmj, ventral
part of main jaw; vt, ventral main jaw teeth.
136 M.V. SØRENSEN
largest of the dorsal teeth tends to be more isolated
than the rest (see Fig. 3C,D) and it could correspond
to the medial teeth sensu Kristensen and Funch
(2000). It could also be this tooth that projects ros-
trally from the jaws shown in Figures 1C and 2B.
The ventral tooth row has approximately 11 short-
shafted teeth that increase in size from the most
caudal towards the most rostral one (Fig. 3B). The
five most caudal teeth are rather small and are only
visible in ventral view, which explains why they
were formerly overlooked. The dorsal teeth, approx-
imately 4 –5, differ considerably from the ventral
ones. They have long proximal shafts and attach
ventral to the nodus, medial to the outer margin of
the dorsal surface of the main jaw (Fig. 3E). All the
dorsal teeth are slightly bent and form a basket-like
structure (Fig. 3B-E). Unpublished SEM photos by
R.M. Kristensen suggest that a third set of loosely
attached teeth, the terminal teeth, might be present
on the small nodus at the base of the dentarium.
However, this could neither be confirmed nor re-
jected in this present study. The ventral part of the
main jaw is extremely difficult to analyze. In most
preparations it is either covered by the ventral cu-
ticle membrane or hidden under the dorsal jaw
parts. However, despite its inconspicuousness it ap-
pears to be a set of rather large elements. They are
elongate and somehow triangular, broadest caudally
(Fig. 3C–F). The broad caudal ends have a lateral
extension that projects under the dorsal part of the
main jaw (Fig. 1B). Rostrally, the elements narrow
toward the pointed tips. The internal surfaces are
totally covered with rounded knobs and serrulated
bands. LM observations reveal that the caudal part
in each sclerite contains a large chamber.
Dorsal Jaw Elements
One pair of well-developed, isolated jaws is
present on the dorsal side of the main jaw (Figs. 1A,
2C, 3A, C-E, 4, 5, 6B). Each jaw element is composed
of a proximal shaft with a caudal cleft and a distal
denticulate area (Fig. 2C). The surface of the dorsal
jaw is slightly rough and after a longer treatment
with hypochlorite the whole sclerite seems to be
composed of tubular rods.
The distal ends of the dorsal jaws have approxi-
mately 16 teeth (Fig. 3A,C–E) arranged in a conspi-
cuous pattern. The teeth are arranged in four rows
with four teeth in each row and increase in size from
the smallest dorsal teeth towards the largest ventral
ones. The dorsal jaws are connected to the fibu-
larium through ligaments at the proximal ends of
the shafts. According to Kristensen and Funch
(2000), a caudally projecting spinula is present at
the base of the denticulate areas. This spinula, how-
ever, is part of the fibularium and not in any way
connected with the dorsal jaws. Kristensen and
Funch (2000) also describe a pseudodens located
medially on the jaw. The presence of this structure
could not be confirmed and the pseudodens observed
by Kristensen and Funch (2000) probably corre-
sponds to the dorsal fibularium jaws (see description
below).
Fibularium
Together with the main jaw the fibularium consti-
tutes the most conspicuous element in the jaw ap-
paratus (Figs. 1A–D, 4 – 6). The fibularia are a pair
of large sclerites with complexes of fibulae. Each
sclerite is somewhat drop-shaped (Fig. 3A), but with
the rostral part twisted upwards a one-quarter turn
around its longitudinal axis (Figs. 4 – 6). The ventral
part of the fibularium is completely covered by a
cuticular membrane (Fig. 1C,D), whereas the dorsal
side is fully open, revealing the presence of five
chambers (Figs. 1A, 3A). The fibularium is delimited
laterally by the fibula lateralis, caudally by the fi-
bula caudalis, and towards the internal surface by
the strong fibula dorsalis (Fig. 3A). Rostrally, the
fibula rostralis extends from the fibula dorsalis to-
wards the tip of the sclerite (Fig. 1A). The fibula
rostralis is identical to the spinula that Kristensen
and Funch (2000) interpreted as a projection from
the dorsal jaw. Inside each fibularium two further
fibulae, fibula radialis and fibula transversalis, are
present (Figs. 1A, 3A). The different fibulae delimit
the five chambers. Camera dorsalis1 is the largest
chamber in the fibularium and is delimited by the
fibula dorsalis, fibula lateralis, and fibula radialis.
Camera dorsalis2 is delimited caudally and towards
the internal surface by the fibula caudalis and lat-
erally by the most caudal part of the fibula dorsalis
(Fig. 3A). The three lateral chambers are located
laterally in the rostral two-thirds of the fibularium.
The most rostral part of the fibula dorsalis separates
the camera lateralis1 from the camera lateralis2 and
the fibula transversalis separates the camera late-
ralis2 from the camera lateralis3 (Fig. 3A). Towards
the dorsal side the chambers open through large
fenestrae (Fig. 3A) and their terminology follows the
names of the chambers, e.g., camera dorsalis1 opens
through fenestra dorsalis1. Kristensen and Funch
(2000) showed that each chamber contains at least
one cell and a total of nine cells appear to be present
in each fibularium.
The fibularium has two pairs of teeth, the dorsal
and ventral fibularium teeth (Figs. 2B, 3B,D). The
dorsal fibularium teeth are located rostrally on the
lateral fibula. Most of its shaft is fused with the
fibula, whereas the denticulate distal part is free.
There are 5– 6 teeth that increase in size from the
very small caudal ones towards the larger rostral
ones. The surface of the shaft is smooth, whereas the
surface of the distal part has the same characteristic
rough and knobbed structure found in several other
jaw elements.
The ventral fibularium teeth are located at the angle
between the rostral and internal margin of each fibu-
 JAW APPARATUS OF L. MAERSKI 137

Fig. 4. Limnognathia maerski. Three-dimensional reconstruction of jaw apparatus with ventral cuticle membrane removed.
A: Dorsal view. B: Rostral view. The main sclerites are symbolized by different colors: green, main jaw; blue, fibularium; yellow,
pseudophalangium and associated sclerites; red, pharyngeal lamellae; turquoise, basal plates and manus; orange, dorsal jaws.
138 M.V. SØRENSEN

Fig. 5. Limnognathia maerski. Three-dimensional reconstruction of jaw apparatus with ventral cuticle membrane removed.
A: Dorsorostral view. B: Ventrorostral view. The main sclerites are symbolized by different colors: green, main jaw; blue, fibularium;
yellow, pseudophalangium and associated sclerites; red, pharyngeal lamellae; turquoise, basal plates and manus; orange, dorsal jaws.

larium. There are 10–15 teeth irregularly arranged in 3A-B, 5). The fenestra does not open towards a specific
three rows (Fig. 3C). A large and yet undescribed chamber but is more like a hole in the fibularium. It
fenestra is located caudolateral to the teeth (Figs. will be referred to as the rostrolateral fenestra.
 JAW APPARATUS OF L. MAERSKI 139

Fig. 6. Limnognathia maerski. Three-dimensional reconstruction of jaw apparatus with ventral cuticle membrane removed.
A: Caudoventral view. B: Caudodorsal view. The main sclerites are symbolized by different colors: green, main jaw; blue, fibularium;
yellow, pseudophalangium and associated sclerites; red, pharyngeal lamellae; turquoise, basal plates and manus; orange, dorsal jaws.
140 M.V. SØRENSEN
DISCUSSION
The present study has revealed the presence of
several new structures in the jaw apparatus of Lim-
nognathia maerski and has shown that some of the
known structures must be reinterpreted. It appears
that most, if not all, sclerites are composed of tubu-
lar rods. In the pseudophalangium, the ventral part
of the main jaw, and in the dorsal jaws, the ends of
the rods are uncovered and appear as serrulae and
knobs, so they are easy to recognize (Figs. 2A,C,
3A–E). Other jaw elements, like the symphysis,
have a smooth surface, but inside they also contain
rods (Fig. 3F). The rods are certainly identical to the
rods shown with TEM by Kristensen and Funch
(2000) and are probably also homologous to similar
structures in Gnathostomulida and Rotifera, as sug-
gested by Rieger and Tyler (1995). Further attention
will be given to this point below.
New Structures in the Jaws of
Limnognathia maerski
The new observations reveal that the morphology
of the main jaw is much more complex than earlier
suggested. In LM and in SEM preparations with
intact ligaments, the pincers appear to be rather
simple denticulate elements like the rami in rotifers
and the rostral jaw parts in Gnathostomulida. How-
ever, the main jaw pincers have never been fully
investigated before because the ventral cuticle cover
prevented observations on the main jaw from a ven-
tral view. A ventral view is only possible in jaws
with broken ligaments and with the sclerites ar-
ranged irregularly. In one of the investigated prep-
arations the jaw apparatus was collapsed and the
presence of two relatively large ventral elements in
the main jaw was revealed (Fig. 3). It is difficult to
imagine how these elements are arranged in vivo,
but it is likely that they are closely associated with
the dorsal pincers, but yet partly isolated. In intact
jaws observed in lateral view the well-developed,
triangular caudal part of the ventral part of the
main jaw is visible as a lateral extension (Fig. 1B)
and a distinctive cleft that separates the more ros-
tral parts of the dorsal and ventral main jaw is
visible in dorsal and rostral views (Fig. 1A,E). Thus,
in all preparations, except the collapsed one, the two
parts appear to be closely connected. Conclusively,
the morphology of the main jaw is not yet fully
understood but the presence of a new element, the
ventral part of the main jaw, is certain.
Several new features were found in the fibu-
larium. The most conspicuous one is the ventral
cuticle cover that extends over most the ventral side
of the jaws, including the caudal parts of the basal
plate. The perforations in the layer, present on some
preparations (Fig. 1C,D), are artifacts made by the
hypochlorite treatment. In dorsal and dorsocaudal
view a total of five chambers, each with a large
fenestra, are visible. Kristensen and Funch (2000)
described and named nine fenestrae, but this was
probably based on the assumption that the ventral
side also had fenestrae. Based on a comparison be-
tween the results from this study and their sche-
matic drawings (see figs. 8, 9 in Kristensen and
Funch, 2000), the following fenestrae are identical:
fl3 and fl4 ⫽ fd1, fd3 ⫽ fv2, fd2 ⫽ fv1. Furthermore, I
found no fibula separating fl1 and fl2 (sensu Kris-
tensen and Funch, 2000) in the most rostral part of
the fibularium. Because of this confusion I suggest
that the terminology proposed herein be used in the
future (see Results and Fig. 3A).
Two other new features in the fibularium are the
dorsal and ventral fibularium teeth (Fig. 3A–C). The
presence of two sets of teeth in the rostral part of the
fibularium implies that these sclerites also play an
active role during feeding. The fibularium articu-
lates rostrally with the dentarium through a strong
ligament (Fig. 1A) and it is therefore difficult to
imagine how the fibularium teeth, which are located
quite close to the attachment point, are able to ope-
rate. However, unpublished LM pictures by R.M.
Kristensen show that the rostral parts of the fibu-
laria can also be moved quite far from the main jaws,
which implies that the ligamental connection is
rather flexible.
The only new structures on the pseudophalan-
gium are the longitudinal ridge and the serrulated
margin (Fig. 2A). The morphology of the associated
sclerites also agrees with the observations of Kris-
tensen and Funch (2000); however, with one impor-
tant difference. Kristensen and Funch (2000) de-
scribe the presence of two pairs of relatively similar
associated sclerites, connected to the pseudophalan-
gium and the dorsal jaws, respectively. However,
the second pair of associated sclerites was not found
in any of the investigated specimens in this study.
Furthermore, the dorsal jaw attaches on the inter-
nal side of the lateral fibula in the fibularium (Fig.
1A) and it is difficult to imagine where the second
pair of associated sclerites should be inserted. Thus,
the second pair of associated sclerites is probably a
misinterpretation or, alternatively, they correspond
to the small, cylindrical sclerites that articulate with
the pharyngeal lamellae (Fig. 1C).
The jaw apparatus of Limnognathia maerski con-
tains two sets of lamellar structures—the oral la-
mellae (see Kristensen and Funch, 2000) and the
pharyngeal lamellae. The oral lamellae are not
shown here because they very easily are dissolved
during the hypochlorite treatment. The pharyngeal
lamellae (Fig. 2A) are a new pair of sclerites. The
sclerites resemble the oral lamellae, but are smaller,
have a more well-developed proximal shaft, and are
located more caudally in the jaws, ventrolateral to
the fibularia. In figure 14 in Kristensen and Funch
(2000) both the oral and the pharyngeal lamellae are
visible. The remains of the oral lamellae are located
in the upper left corner of the picture, while the
 JAW APPARATUS OF L. MAERSKI
pharyngeal lamellae are a pair of rather strong scle-
rites laterorostral to the fibularia. In SEM the pha-
ryngeal lamellae often appear to be connected with
the pseudophalangium through a very flexible liga-
ment between the proximal part of the pseudopha-
langium and the small cylindrical sclerite that ar-
ticulates with the pharyngeal lamella in a ball-and-
socket joint. However, it is noteworthy that
unpublished LM photos by R.M. Kristensen show
that the two pairs of elements can be moved totally
independent of one another.
No new data were accumulated on the basal plate
because most of it was always hidden behind the
ventral cuticle of the fibularium. However, a new
pair of sclerites associated with the basal plate was
found. The sclerites articulate with the outer ros-
troventral margins of the basal plate and are re-
ferred to as the manus because they resemble a pair
of hands or gloves (Fig. 2B,D). The teeth of the
manus are easily distinguished from those of the
basal plate teeth because the latter have smaller
and more pointed teeth.
The dorsal jaws have been described by Kris-
tensen and Funch (2000) but more details are added
here. The peculiar arrangement of the teeth in four
rows (Fig. 2C) has not been described before and is,
to my knowledge, not found in jaws from other
groups. The elements attach caudally on the inter-
nal side of the lateral fibula in the fibularium and in
relaxed jaws they appear to be closely associated
with the fibularium and the dorsal fibularium teeth.
However, LM observations show that the dorsal
jaws are highly moveable and able to stand at a
right angle to the axis of the main jaw.
Comparison With Gnathostomulida
and Rotifera
The most consistent link between the jaws in the
gnathiferan taxa is probably the rods that appear to
be a basic brick in most of the jaw elements (Ahl-
richs, 1995, 1997; Rieger and Tyler, 1995; Kris-
tensen and Funch, 2000; Sørensen et al., 2000). In
Rotifera the rods were first shown with TEM in the
rami of Asplanchna sieboldi and Philodina acuticor-
nis by Koehler and Hayes (1969a,b) and later also in
the rami of A. brightwelli and Notommata copeus
(Clément and Wurdak, 1991) and in the rami, ful-
crum, and epipharynx of Seison annulatus (Ahl-
richs, 1995). In the fulcrum the rods are also visible
on the lateral surfaces and Markevich (1989) refers
to the rods as sclerofibrillae and suggests that they
are homologous with the small hair-like scleropili
present in the rami and unci of several rotifer spe-
cies (see Markevich, 1989; Melone et al., 1998; Sø-
rensen, 2002a). Thus, the rods appear to be present
widespread in the rotiferan trophi. Furthermore,
studies on trophi from embryonic Floscularia meli-
certa show that at least the rami, fulcrum, and unci
are formed by fusion of numerous scleropili or
141
sclerofibrillae (Sørensen, 2002a). The presence of
rods or sclerofibrillae in mostly all sclerites in Mi-
crognathozoa and Rotifera is therefore considered
an important link between the two groups.
The rods are also present in the jaws of Gnathos-
tomulida. The rods were first shown with TEM in
the lamellae symphysis of Rastrognathia macros-
toma by Kristensen and Nørrevang (1977) and later
also in Haplognathia rosea, Pterognathia meixneri,
and Gnathostomula paradoxa (see Lammert, 1986),
which imply that they are a common trait in Gna-
thostomulida. This was confirmed by Sørensen and
Sterrer (2002), who showed with SEM that the rods
were present in the lamellae symphysis of at least
10 different genera. Herlyn and Ehlers (1997) also
demonstrated the presence of rods in the gnathosto-
mulid basal plate using TEM, but Sørensen and
Sterrer (2002) doubted this observation and noted
that the rods could be transverse sections of the
basal plate dentes. The rods have never been found
in the suspensorium or in the dentaria in Bursovagi-
noidea, but the small needle-like denticles present
in the dentaria of filospermoids are probably homol-
ogous with the rotiferan scleropili (Sørensen and
Sterrer, 2002).
Some of the sclerites in the jaw apparatus in Lim-
nognathia maerski are definitely autapomorphic,
but others may be present in Rotifera and/or Gna-
thostomulida. Kristensen and Funch (2000) suggest
that the main jaw is homologous with the rotifer
incus (rami ⫹ fulcrum) and the gnathostomulid ar-
ticularium ⫹ dentarium, but with the greatest sim-
ilarity with the gnathostomulids. They especially
interpret the following similarities as homologous:
basket-like arrangement of the teeth, teeth in three
rows, paired pincers that join in a symphysis, and
the presence of a cauda. Some of these traits are
truly very similar in the two groups, but other data
imply that they might be either plesiomorphic or
convergently evolved. In L. maerski the dorsal teeth
in the main jaw form a basket but the ventral teeth
are arranged in a straight row, which is in contrast
to the onychognathiid gnathostomulids, where both
the dorsal and ventral teeth form the basket (Ster-
rer, 1972; Sørensen and Sterrer, 2002). Further-
more, a cladistic study of the gnathostomulid phy-
logeny shows that the basket-like arrangement of
the teeth probably evolved convergently in L. maer-
ski and Onychognathiidae (Sørensen, 2002b). Teeth
arranged in three rows, as in Gnathostomula, are
not present in L. maerski. Even if a medial set of
teeth is present, it is definitely not located on the
internal side of the pincer, so the similarity between
the jaws of L. maerski and Gnathostomula can be
ruled out. A cauda-like structure, as in some gna-
thostomulids, is present in L. maerski, but according
to the cladistic analysis of Sørensen (2002b), the
cauda evolved convergently three times in Gnatho-
stomulida, and none of them are homologous with
the cauda in L. maerski. Furthermore, it is notewor-
142 M.V. SØRENSEN
thy that a paired cauda, as in L. maerski, is only
present in Gnathostomulidae, and this family is tra-
ditionally considered among the most advanced con-
cerning jaw morphology (Sterrer, 1972; Sørensen
and Sterrer, 2002; Sørensen, 2002b). The arrange-
ment of pincer-like jaws that articulate in a symphy-
sis is definitely shared between Micrognathozoa and
Gnathostomulida, but a similar trait is present in
the incus of most rotifers. Sørensen (2000) showed
the great similarities between the rotifer fulcrum
and the caudally extended symphysis found in some
gnathostomulids. Markevich (1989) suggested that
the rotifer fulcrum was formed as fused caudal ex-
tensions from the rami and SEM observations from
different species show the presence of a thin suture
on the fulcrum, which supports that it is formed
by secondarily fused structures (Sørensen, 2000,
2002a). This suggests that the micrognathozoan
main jaw, the gnathostomulid articularium, and the
rotifer incus probably are homologous, and hence
apomorphic for Gnathifera.
A synapomorphy for Rotifera and Micrognathozoa
could be the formation of two more or less closed
chambers that each contain a cell. Rotifers basally
have two or three chambers in each ramus that may
open through fenestrae (Markevich, 1989; Nogrady
et al., 1993; Sørensen, 2002a) and the chambers in
the main jaw of Limnognathia maerski may very
well be homologous with these. Chambers are also
found in the jaws of some gnathostomulids, i.e.,
some Onychognathiidae and Gnathostomulidae, but
in the latter the chambers apparently do not contain
cells and serve instead as muscle attachment sites
(Lammert, 1986; Herlyn and Ehlers, 1997), so these
are definitely convergently evolved. Sørensen (2000)
showed that each jaw half in Rastrognathia macro-
stoma had two chambers with a cell in each, and
later Sørensen and Sterrer (2002) showed that sim-
ilar chambers are present in Vampyrognathia mi-
nor. These chambers might be homologous with
those in Micrognathozoa and Rotifera, but they dif-
fer in being fully open and only delimited by fibulae.
The fibularium is extremely difficult to homolo-
gize with structures in any other group. Kristensen
and Funch (2000) suggest a homology between the
fibularium, the rotifer rami, and the gnathostomulid
suspensorium and involucrum in the family Gnatho-
stomulidae. However, I consider this rather doubt-
ful. First of all, as noted above, the suspensorium
chambers in Gnathostomulidae do not contain cells.
Furthermore, the jaw elements that form the hollow
parts in Gnathostomulidae have a special porous
substructure, which is only found in this and the
most closely related families. Sørensen and Sterrer
(2002) suggested that the presence of suspensoria
with this substructure is an advanced trait that has
evolved within Bursovaginoidea and this assump-
tion is supported by the cladistic analysis of Sø-
rensen (2002b). The compartmentalization with
cells inside the fibularium resembles the arrange-
ment in the rotifer rami, but I find it very unlikely
that the fibularium should be a part of the rami. It is
easy to imagine the transformation from a forceps-
like ancestral element into the micrognathozoan
main jaw and the rotifer incus, but nothing indicates
that the fibularium has evolved by separation from
another structure, or that the incus is formed by
fusion of forceps and fibularia-like structures. Thus,
I consider the fibularium as an autapomorphy for
Micrognathozoa.
Kristensen and Funch (2000) note that W.H. Ahl-
richs in a personal communication suggested that
the fibularium could be homologous with the rotifer
manubria. In an article by Bedwell (1878) concern-
ing the trophi morphology in the rotifer Floscularia
ringens, the manubria actually look quite similar to
the fibularium, but besides this superficial simila-
rity with a rather imprecise illustration there is
nothing in common between the fibularium and the
manubria. The rotifer manubrium consists basically
of three chambers and an elongate cauda1 formed by
elongation of one or more of the chambers (Marke-
vich, 1989; Nogrady et al., 1993) and this arrange-
ment is considered by some authors a basal trait in
the rotifer trophi (Melone et al., 1998; Sørensen,
2002a). Kristensen and Funch (2000) suggest that
the accessory sclerites and the pseudophalangids in
Limnognathia maerski are homologous with the ro-
tifer manubria and unci (together referred to as the
mallei) and I fully agree with this view. Three argu-
ments support this: the location of the sclerites, the
ligamental connection between the two sclerites so
they form one operational unit, and the morphology
of the sclerites. The unci are, in most rotifer species,
positioned ventral to the rami and their proximal
ends point outward, towards the lateral sides, where
they articulate with the manubria. The manubria
are located lateral to the rami and their distal ends,
the caudae, point in a caudodorsal direction
(Markevich, 1989; Nogrady et al., 1993; Sørensen,
2002a). These positions correspond well to the posi-
tions of the pseudophalangids and accessory scle-
rites in L. maerski. The pseudophalangids are ex-
tremely moveable and their distal ends can be
stretched beyond the mouth opening or protruded
out through the mouth (Kristensen and Funch,
2000), but the rotifer unci can also be highly move-
able, as, for instance, in Synchaetidae. The pseu-
dophalangids and accessory sclerites articulate in a
flexible joint like the two sclerites in the rotifer
malleus, but the associated arrangement of muscu-
lature differs slightly because the pseudophalangids
appear to be moved by a large muscle that attaches
in the fibularium (Kristensen and Funch, 2000). In
Rotifera the uncus is primarily moved by a large
1
Please note that the rotifer manubrial cauda is in no way homol-
ogous with the gnathostomulid and micrognathozoan cauda.
 JAW APPARATUS OF L. MAERSKI
extensor muscle that attaches distally on the manu-
brium (Remane, 1929 –1933). The large rod-shaped
pseudophalangium mostly resembles the uncus in
some rotifers, i.e., Seison and different Dicranopho-
ridae, but it is difficult to say whether this is due to
convergence. However, the accessory sclerite has a
special substructure, with helically twisted strands
that are also present in the manubrium of bdelloid
rotifers (Melone et al., 1998) and various different
monogononts such as Ascomorpha, Encentrum,
Itura, Lindia, and Trichotria (see Sørensen, 2002a).
Kristensen and Funch (2000) suggested that the
oral lamellae are homologous with some of the
epipharyngeal elements found in different rotifers,
and this may very well be true for the pharyngeal
lamellae also. The sclerites especially resemble the
pleural rods present in genera like Cephalodella and
Resticula. However, the morphology of the rotifer
epipharyngeal elements is extremely variable and
the ground pattern is difficult to establish.
The rest of the micrognathozoan jaw elements,
including the dorsal jaws, the basal plate, and the
manus, are considered autapomorphic. Kristensen
and Funch (2000) considered the basal plate as ho-
mologous with the gnathostomulid basal plate, but
as noted by Sørensen (2002b) and Sørensen and
Sterrer (2002), there is only very weak support for
this assumption. The gnathostomulid basal plate is
clearly an unpaired structure composed of five
plates (Riedl and Rieger, 1972), while the microgna-
thozoan basal plate is two articulating elements.
The rostral plates may show superficial resemblance
with the gnathostomulid basal plate, but the long
caudally extending shafts are not present in any
gnathostomulid. Also, the well-developed teeth and
the associated manus are unique for the microgna-
thozoan basal plate. The basal plate in Gnathosto-
mulida is either adentate or has smaller denticles
(Sterrer, 1972; Riedl and Rieger, 1972; Sørensen and
Sterrer, 2002) and it has no associated elements.
Finally, the basal plate in Micrognathozoa is a much
more integrated part of the jaw apparatus, while the
gnathostomulid basal plate is totally isolated. Thus,
I consider the micrognathozoan basal plate as auta-
pomorphic.
Phylogeny of Gnathifera and the
Taxonomic Status of Micrognathozoa
The jaw similarities discussed above support
Gnathifera as a monophyletic group and with the
inclusion of further characters, concerning other
morphological traits, a phylogeny of the Gnathifera
can be proposed (Fig. 7). Gnathifera is basically sup-
ported by the presence of jaws composed of translu-
cent rods with an electron-dense core. A set of rather
simple pincer-like jaws fused caudally and forming
an unpaired symphysis or fulcrum was also present
in the ground pattern. This kind of jaws is homolo-
gous to the micrognathozoan main jaw, the gnatho-
143
stomulid lamellae symphysis, and the rotifer rami.
Multiciliate epidermis and dioecious sexes are ple-
siomorphic for Gnathifera, which mean that the
gnathostomulid monociliate epidermis and her-
maphroditism are advanced traits. Gnathostomu-
lida is the first group to branch off in Gnathifera and
it is supported by the reversal of epidermal cells
from a multiciliate to a monociliate condition, her-
maphroditism, the presence of a basal plate, and
jaws belonging to either the compact type or the
open lamellar type. The presence of a temporarily
functioning anal pore and monociliate proto-
nephridia could be considered synapomorphic for
Gnathostomulida and Micrognathozoa. However,
the presence of a temporarily functioning anal pore
is not only found in Gnathostomulida and Micro-
gnathozoa, but also in several flatworms and gastro-
trichs, and even in Arthrotardigrada. Furthermore,
a true anus might be present in the gnathostomulid
genus Agnathiella (see Sterrer, 1971; Knauss, 1979),
which implies that such a character should be used
with caution. Apart from the Gastrotricha, Micro-
gnathozoa and Gnathostomulida are the only proto-
stomes with monociliated terminal cells in their pro-
tonephridia (Lammert, 1985; Kristensen and Funch,
2000) and this character is probably the most con-
sistent of the possible synapomorphies for the taxa.
However, their protonephridia differ in other ways,
such as the number of microvilli and terminal and
canal cells.
The sister-group relationship between the Micro-
gnathozoa and the Rotifera-Acanthocephala clade is
supported both by the above-mentioned characters
connected to the pharyngeal apparatus and to other
organs. In both groups the epidermis is covered by a
thin extracellular glycocalyx and inside it contains
an intracellular protein lamina (Storch and Welch,
1969, 1970; Kristensen and Funch, 2000). Although
the integument in Micrognathozoa is cellular and
the intraskeletal lamina forms a matrix of epider-
mal plates that cover the dorsal and lateral sides,
whereas the Rotifera-Acanthocephala integument is
modified from a cellular to a syncytial epidermis, the
presence of an intracellular lamina still provides a
strong synapomorphy for the groups. Another possi-
ble synapomorphy is the occurrence of eutely in
some organs. Eutely is present in most organs in
Acanthocephala and Rotifera (Hyman, 1951) and it
is probably also present in the integument of Mi-
crognathozoa (Kristensen and Funch, 2000). How-
ever, the latter needs to be confirmed in further
studies.
Even though Rotifera and Micrognathozoa share
some similarities, the monophyly of Rotifera with
respect to Micrognathozoa still remains unquestion-
able. Besides the syncytial epidermis with intracel-
lular lamina, shared by Acanthocephala and Roti-
fera, the Rotifera are supported by characters such
as epidermal ciliation restricted to the head region,
ciliary bands forming a wheel-organ, and division of
144 M.V. SØRENSEN

Fig. 7. Cladogram showing the phylogeny of Gnathifera including the supporting synapomorphies. 1. Gnathifera: jaws composed
of rods with a lucent material and an electron-dense core; jaws with pincers caudally articulating into unpaired pedicle. 2. Gnatho-
stomulida: epidermis cells reverse from multiciliate to monociliate condition; hermaphroditism develops; basal plate develops; jaws
belonging to either closed or open lamellar type. 3: Development of pseudophalangium/unci and associate sclerites/manubria;
development of oral and pharyngeal lamellae/epipharynx; cellular epidermis with intracellular lamina; eutely(?). 4. Micrognathozoa:
intracellular lamina forms epidermal plates restricted to dorsal and lateral sides; head and trunk with sensoria formed by multiciliate
cells; cilia on trunk and parts of head arranged as ciliophores; jaws with fibularium, paired basal plates, and dorsal jaws. 5.
Rotifera-Acanthocephala: syncytial epidermis; trunk ciliation reduced; cilia in head arranged in bands forming a wheel-organ; body
divided into head, trunk and foot. 6. Acanthocephala: obligatory endoparasite, body ciliation lost, mouth, pharynx and gut lost.

body into head, trunk, and foot regions. Some au-
thors have questioned the monophyly of Rotifera
with respect to the Acanthocephala (Ahlrichs, 1995,
1997; Sørensen et al., 2000) and this problem has
not been solved yet.
In the original description of Micrognathozoa
(Kristensen and Funch, 2000), the taxon was as-
signed class rank, based on the condition that
Gnathifera has phylum rank. The use of specific
taxonomic ranks is by many scientists considered an
obsolete relic from the Linnean system, but it still
enjoys widespread use, at least for historical rea-
sons. Gnathostomulida has for more than 30 years
been assigned phylum rank (Riedl, 1969; Sterrer,
1972) and in most recent classifications Rotifera is
also considered a phylum (Nogrady et al. 1993).
Based on our increased knowledge of the microgna-
thozoan phylogeny and its well-established sister-
group relationship to the Rotifera-Acanthocephala
clade, I suggest that future classification follow
these conventions and raise Micrognathozoa to phy-
lum rank.
CONCLUSIONS
The SEM and LM studies on the jaws of Limnog-
nathia maerski revealed the presence of several new
structures and elements. The pharyngeal lamellae
are a pair of sclerites that resemble the oral lamel-
lae, but they are smaller, located more laterocau-
dally, and are more resistant to hypochlorite treat-
ment. The pharyngeal lamellae may be homologous
with the rotifer epipharynx. Another new pair of
elements is the manus, which is a set of hand-
shaped sclerites associated with the basal plate. The
manus are considered autapomorphic for Microgna-
thozoa.
More details were obtained on the morphology of
the main jaw and fibularium. The main jaw has a
 JAW APPARATUS OF L. MAERSKI
pair of partly isolated elements, the ventral part of
the main jaw, that are characterized by their rough
surfaces ornamented with knobs and serrulae. Two
sets of teeth were found on the fibularium. These are
referred to as the dorsal and ventral fibularium
teeth. Furthermore, it was clarified that each fibu-
larium contains only five chambers and that its
ventral side is totally covered by a dense cuticle
membrane. Generally, the micrognathozoan jaw ele-
ments appeared to consist of rod-like structures that
are homologous with the sclerofibrillae and sclero-
pili in the rotifer trophi and with the rods in the
lamellae symphysis in Gnathostomulida.
Comparison with the jaws in Rotifera and Gna-
thostomulida suggests that the main jaw, the rotifer
incus, and the gnathostomulid articularium are ho-
mologous and synapomorphic for Gnathifera. Fur-
thermore, the pseudophalangium and associated
sclerites are homologous with the rotifer malleus.
The fibularium, the basal plate, and the dorsal jaws
are autapomorphic for Micrognathozoa. The proba-
ble synapomorphies suggests a sister-group rela-
tionship between Micrognathozoa and the Rotifera-
Acanthocephala clade.
ACKNOWLEDGMENTS
I thank Reinhardt M. Kristensen and Peter Funch
for giving me valuable specimens from their samples
from Disko Island. I also thank R.M. Kristensen for
numerous interesting discussions on the gnathi-
feran jaw morphology and for his comments on the
manuscript. I thank Ditte Bruun for her great work
on the 3D reconstructions of the jaws, the board and
the staff on the Danish Arctic Station for their help
and support, and Mary E. Petersen for correcting
the language.
LITERATURE CITED
Adam H, Czihak, G. 1964. Arbeitsmethoden der makroskopis-
chen und mikroskopischen Anatomie. Stuttgart: Gustav Fi-
scher Verlag.
Ahlrichs WH. 1995. Ultrastruktur und Phylogenie von Seison
nebaliae (Grube 1859) und Seison annulatus (Claus 1876). Göt-
tingen: Cuvillier Verlag.
Ahlrichs WH. 1997. Epidermal ultrastructure of Seison nebaliae
and Seison annulatus, and a comparison of epidermal struc-
tures within Gnathifera. Zoomorphology 117:41– 48.
Bedwell FA. 1878. The mastax-framework in Melicerta ringens
and Conochilus. Trans R Microsc Soc 1:176 –185.
Clément P, Wurdak E. 1991. Rotifera. In: Harrison FW, Ruppert
EE, editors. Microscopic anatomy of invertebrates, vol. 4. As-
chelminthes. New York: Wiley-Liss. p 219 –298.
Herlyn H, Ehlers U. 1997. Ultrastructure and function of the
pharynx of Gnathostomula paradoxa (Gnathostomulida).
Zoomorphology 117:135–145.
Hyman LH. 1951. The invertebrates: Acanthocephala, Aschel-
minthes, and Entoprocta. New York: McGraw-Hill.
145
Knauss EB. 1979. Indication of an anal pore in Gnathostomulida.
Zool Scr 8:181–186.
Koehler JK, Hayes TL. 1969a. The rotifer jaw: a scanning and
transmission microscope study. I. The trophi of Philodina acu-
ticornis odiosa. J Ultrastruct Res 27:402– 418.
Koehler JK, Hayes TL. 1969b. The rotifer jaw: a scanning and
transmission microscope study. II. The trophi of Asplanchna
sieboldi. J Ultrastruct Res 27:419 – 434.
Kristensen RM, Funch P. 2000. Micrognathozoa: a new class with
complicated jaws like those of Rotifera and Gnathostomulida. J
Morphol 246:1– 49.
Kristensen RM, Nørrevang A. 1977. On the fine structure of
Rastrognathia macrostoma gen. et sp.n. placed in Rastrognathi-
idae fam.n. (Gnathostomulida). Zool Scr 6:27– 41.
Lammert V. 1985. The fine structure of protonephridia in Gna-
thostomulida and their comparison within Bilateria. Zoomor-
phology 105:308 –316.
Lammert V. 1986. Vergleichende Ultrastruktur-Untersuchungen
an Gnathostomuliden und die phylogenetische Bewertung ihrer
Merkmale. Dissertation, Georg-August-Universität zu Göttin-
gen, Göttingen, Germany.
Markevich GI. 1989. Morphology and the principal organisation
of sclerite system of the mastax in rotifers. In: Shilova IA,
editor. Biological and functional morphology of freshwater an-
imals. Leningrad: Proceedings of the Institute of the Biology of
Inland Waters 56:27– 82 [in Russian].
Melone G, Ricci C, Segers H. 1998. The trophi of Bdelloidea
(Rotifera): a comparative study across the class. Can J Zool
76:1755–1765.
Nogrady T, Wallace RL, Snell TW. 1993. Rotifera. In: Dumont
HJF, editor. Guides to the identification of the microinverte-
brates of the continental waters of the world, vol. 4. Rotifera,
vol. 1. Amsterdam: SPB Academic. p 1–142.
Remane A. 1929-33. Rotatoria. In: Bronn HG, editor. Klassen und
Ordnungen des Tier-Reichs, vol. 4, Vermes. Leipzig: Akade-
mische Verlagsgesellschaft.
Riedl RJ. 1969. Gnathostomulida from America. Science 163:445–
452.
Riedl R, Rieger RM. 1972. New characters observed on isolated
jaws and basal plates of the family Gnathostomulida (Gnatho-
stomulida). Z Morph Tiere 72:131–172.
Rieger RM, Tyler S. 1995. Sister-group relationship of Gnathosto-
mulida and Rotifera-Acanthocephala. Invertebr Biol 114:186–188.
Sørensen MV. 2000. An SEM study of the jaws of Haplognathia
rosea and Rastrognathia macrostoma (Gnathostomulida), with
a preliminary comparison with the rotiferan trophi. Acta Zool
(Stockh) 81:9 –16.
Sørensen MV. 2002a. On the evolution and morphology of the rotiferan
trophi, with a cladistic analysis of Rotifera. J Zool Syst Evol Res.
Sørensen MV. 2002b. Phylogeny and jaw evolution in Gnathos-
tomulida, with a cladistic analysis of the genera. Zool Scr.
Sørensen MV, Funch P, Willerslev E, Hansen AJ, Olesen J. 2000.
On the phylogeny of the Metazoa in light of Cycliophora and
Micrognathozoa. Zool Anz 239:297–318.
Sørensen MV, Sterrer W. 2002. New characters in the gnathos-
tomulid mouth parts revealed by scanning electron microscopy.
J Morphol 253:310 –334.
Sterrer W. 1971. Agnathiella beckeri nov. gen. nov. spec. from
southern Florida: the first gnathostomulid without jaws. Int
Rev Ges Hydrobiol 56:215–225.
Sterrer W. 1972. Systematics and evolution within the Gnathos-
tomulida. Syst Zool 21:151–173.
Storch V, Welch U. 1969. Über den Aufbau des Rotatorienintegu-
mentes. Z Zellforsch 95:405– 414.
Storch V, Welch U.1970. Über den Aufbau resorbierender Epi-
thelen darmloser Endoparasiten. Zool Anz Suppl 33:617–
621.