Ulrich Ehlers

The position of the Arthropoda in the phylogenetic system

Traditionally, Panarthropoda (Euarthropoda, Onychophora, Tardigrada) are regarded as being closely related to Annelida in a taxon Articulata, but this is not supported by molecular analyses. Comparisons of gene sequences suggest that all molting taxa (Panarthropoda, Nematoda, Nematomorpha, Priapulida, Kinorhyncha, Loricifera) are related in a monophyletic taxon Ecdysozoa. An examination of the characters supporting Articulata reveals that only segmentation with a teloblastic segment formation and the existence of segmental coelomic cavities with nephridia support the Articulata, whereas all other characters are modified or reduced in the panarthropod lineage. Another set of characters is presented that supports the monophyly of Ecdysozoa: molting under influence of ecdysteroid hormones, loss of locomotory cilia, trilayered cuticle and the formation of the epicuticle from the tips of epidermal microvilli. Comparative morphology suggests Gastrotricha as the sister group of Ecdysozoa with the synapomorphies: triradiate muscular sucking pharynx and terminal mouth opening. Thus there are morphological characters that support Articulata, but molecular as well as morphological data advocate Ecdysozoa. Comparison of both hypotheses should prompt further thorough and targeted investigations. J. Morphol. 238:263–285, 1998.

Comments on a phylogenetic system of the Platyhelminthes

Using the principles and methods of phylogenetic systematics, the phylogenetic relationships among the basic closed descendent communities (monophyla) of the taxon Platyhelminthes are discussed.

Monociliary receptors in interstitial Proseriata and Neorhabdocoela (Turbellaria Neoophora)

SummaryThe ultrastructure of monociliary receptors in 10 species of the Proseriata and Neorhabdocoela is described, with particular reference to the epidermal dendritic part.Sensory cells with a single kinocilium situated at the level of the distal epidermis membrane are considered as mechano- or chemoreceptors.There exist sensory cells with a dendrite penetrating one epidermis cell and bearing an embedded kinocilium and a collar of 8 stereocilia or ridges with a fribrillose substructure. These collared receptors probably function as mechanoreceptors.In comparison with collared sensory cells in species of other turbellarian orders, the embedded receptors in the Proseriata and Neorhabdocoela are more advanced and possess synapomorphous characteristics. With the embedded receptors a new evidence is given for the close phylogenetic relationship between the Proseriata and Neorhabdocoela.The distribution of collared cells in the animal system and their phylogenetic implication for a choanoflagellate origin of the Metazoa are briefly discussed.

Ultrastructure of the subepidermal musculature of Xenoturbella bocki, the adelphotaxon of the Bilateria

Abstract Xenoturbella bocki is the only species of the high-ranked taxon Xenoturbellida. The species lives on marine mud bottoms at a depth of 20–120 m and moves extremely slowly by ciliary gliding. Nevertheless it possesses a well-developed body wall musculature with outer circular muscles, a prominent layer of inner longitudinal muscles and radial muscles that extend from the outer circular myocytes to the musculature surrounding the gastrodermis. The longitudinal myocytes are not compact cells, but form fascicles of fibrils running parallel to each other. Fine cytoplasmic cords connect the fibres of a cell to each other and with its nuclear region. The muscles are embedded within a sometimes expansive extracellular matrix (ECM) that lacks any fibrillar components. All muscle cells display conspicuous and numerous cytoplasmic extensions that are intermingled with each other. Tight coupling between adjacent cell membranes is not found, but zonula adhaerens-like junctions exist. Fibrils belonging to different myocytes, but also fibrils of the same cell, are coupled by such cytoplasmic extensions. Circular, radial and at least the peripheral longitudinal myocytes display cell-matrix connections with the internal lamina, a component of the subepidermal ECM. This internal lamina projects down into the centres of the fascicles with longitudinal muscle fibrils and forms extensive attachment zones with the muscle cells, reminiscent of focal contacts. For the ingestion of food, X. bocki opens the simple mouth pore and protrudes the aciliated gastrodermis. The body wall musculature is responsible for this protrusion and also for the withdrawal of the gastrodermis. In the past, possible phylogenetic kinships with the Acoelomorpha (Plathelminthes) or the Enteropneusta and Holothuroidea were discussed, but, on the basis of all information available, X. bocki is hypothesized to be the sister taxon of the Bilateria.

Ultrastructure and function of the pharynx of Gnathostomula paradoxa (Gnathostomulida)

Abstract The pharynx of Gnathostomula paradoxa consists of the partly syncytial pharyngeal musculature, a pharyngeal epithelium, myoepitheliocytes, receptors, nerves, and three solid parts, called the jugum, the basal plate, and the jaw. Extended non-contractile regions of both pharyngeal and body wall musculature form the so-called parenchymatous tissue between the digestive tract and the body wall. The pharyngeal epithelium mediates the force from the pharyngeal musculature to the solid parts. The basal plate and jaw contain longitudinal cuticular rods which are elastic antagonists of the musculature. There is no buccal ganglion in G. paradoxa. The study supports the monophyly of the Gnathostomulida and Gnathifera.

Embryonic development of the nervous system of the rhabdocoel flatwormMesostoma lingua (Abildgaard, 1789)

We have analyzed the embryonic development of the Mesostoma nervous system, using a combination of histology, transmission electron microscopy, and wholemount immunohistochemistry. Neural progenitors are formed at an early stage when the Mesostoma embryo constitutes a multilayered mesenchymal mass of cells. A neurectoderm as in vertebrates or arthropods is absent. Only after neurons in the deep layers of the embryo have started differentiating do superficial cells reorganize into an epithelium that will give rise to the epidermis. Neurons are clustered in two anterior, bilaterally symmetric brain hemispheres. An antibody against acetylated beta‐tubulin (anti‐acTub) that labels neurotubules reveals an invariant pattern of pioneer neurons in the brain of midstage embryos. Pioneer neurons are grouped in several small clusters at characteristic positions. They pioneer several commissural tracts of the brain and two pairs of ventral and dorsal connectives, respectively. J. Comp. Neurol. 416:461–474, 2000.

Comparative morphology of statocysts in the Plathelminthes and the Xenoturbellida

The general fine-structural organization of statocysts in Catenulida, Nemertodermatida, Acoela, Proseriata, Lurus (Dalyellioida), and Xenoturbella are summarized. In lithophorous (statocyst-bearing) members of the Catenulida, the statocysts exhibit a few parietal cells and one or several movable statoliths within a spacious intracapsular cavity. Statocysts in the Nemertodermatida have several parietal cells and two lithocytes, each equipped with one statolith, whereas those of the other acoelomorphan taxon, the Acoela, always have two parietal cells and one movable lithocyte. The statocysts of lithophorous members of the Proseriata represent more sophisticated systems: each has two clusters of accessory cells in addition to several parietal cells and a voluminous lithocyte in which the statolith is movable. In catenulids and proseriates, processes of outer neurons penetrate the capsule of the statocyst, whereas such innervations have not been found in the Nemertodermatida and Acoela. I conclude that the different types of statocysts have evolved independently within the Plathelminthes. Xenoturbella displays an intraepidermal statocyst with many monociliary parietal cells and several mobile cells (‘lithocytes’) within the central cavity of the statocyst. Each of these mobile cells carries a statolith-like structure and one prominent cilium. The statocyst of Xenoturbella does not correspond to any type of plathelminth statocyst.

Paddle cilia and discocilia — Genuine structures?

SummaryKinocilia of epidermal sensory cells in fixed marine Turbellaria often terminate as flattened biconcave discs. The distal part of the ciliary axoneme curves back upon itself forming a 360° loop which is enveloped by the plasmalemma. In living animals this structure can be induced by the addition of sodium cacodylate, monobasic sodium phosphate, dibasic sodium phosphate, sucrose, calcium chloride, or formaldehyde to the sea water. Specimens treated with sodium chloride, glutaraldehyde, or osmium tetroxide do not show modified cilia. In animals prepared for EM at low temperature and with a buffered hypotonic fixative less kinocilia are modified than in animals treated with a buffered iso- or hypertonic fixative and at a higher temperature. It is assumed that the unusually shaped cilia, described as “paddle cilia” or “discocilia” in other invertebrates, do not represent a genuine but an artificial structure.

Die Feinstruktur eines ciliären Lamellarkörpers beiParotoplanina geminoducta Ax (Turbellaria, Proseriata)

SummaryThe fine structure of a lamellate body in the head region ofParotoplanina geminoducta Ax, 1956 is described. It is constructed by some modified cilia, which protrude into an intracellular cavity. The plasmalemma of these cilia are prolonged and rolled up, thus forming a whorl. The lamellate body is supposed to have a photoreceptive function.ZusammenfassungAus der Kopfregion vonParotoplanina geminoducta Ax, 1956 wird die Feinstruktur eines Lamellarkörpers beschrieben. Er besteht aus mehreren modifizierten Cilien, die in ein intracelluläres Lumen hineinragen und deren verlängertes Plasmalemm spiralig aufgerollt ist. Dem Lamellarkörper wird eine vermutlich photoreceptorische Funktion zugeschrieben.

Frontal glandular and sensory structures inNemertoderma (Nemertodermatida) andParatomella (Acoela): ultrastructure and phylogenetic implications for the monophyly of the Euplathelminthes (Plathelminthes)

SummaryThe ultrastructural organization of the frontal glandular and sensory structures in aNemertoderma species and in the acoelParatomella rubra is described. In both species, a frontal gland complex with an accumulation of different body wall glands and of ciliated sensory receptors exists at the anterior tip of a specimen. Such an organization can be hypothesized as a ground pattern characteristic of the Acoelomorpha. A “frontal organ” with a common apical pore of many necks of mucoid glands is known for most species of the Acoela; however, such an organ is not a basic feature either of this taxon or of the Acoelomorpha. This substantiates the hypotheses of a homology of the frontal glandular structures in the Acoelomorpha and the Rhabditophora and of the monophyly of the taxon Euplathelminthes. Within the Euplathelminthes, positional, morphological and histochemical differences of the frontal glands and the secretions exist between distinct taxa. These different organizations contribute to the phylogenetic systematization of the Plathelminthes, as shown in a diagram of the phylogenetic relationships among the basic closed descendent communities of this taxon.