New insights into the evolution of portunoid swimming crabs (Portunoidea, Heterotremata, Brachyura) and the brachyuran axial skeleton.

Portunoidea (Heterotremata) is a morphologically disparate taxon of true crabs (Brachyura) best-known for many of its representatives being considered "swimming crabs". The term "swimming crab", however, sometimes refers to a distinct taxon (traditionally to Portunidae within Portunoidea), and sometimes to a certain morphotype in which the 5th pereiopod (P5) has a specific shape that facilitates swimming. We use the term "P5-swimming crab" or "P5-swimmer" herein, not only to restrict it to the morphotype, but also to distinguish the swimming in question from other kinds of swimming in Brachyura. The evolution of P5-swimming crabs has not yet been satisfactorily investigated. In particular, it is not known whether the morphotype evolved several times independently in different lineages of Portunoidea or whether it evolved only once and was lost in several lineages. Ours is the first approach combining molecular with morphological data to result in a new phylogenetic positioning of some members of Portunoidea. For the first time, data from the axial skeleton and extrinsic musculature are used. Morphological examinations reveal that the axial skeleton and extrinsic musculature in P5-swimming crabs are more diverse than previously thought, with the exception of the P5 anterior coxa muscle, which originates at the median plate in all P5-swimmers. Ancestral state reconstructions based on parsimony reveal that the stem species of Portunoidea already showed the morphotype of a P5-swimming crab, but with a long merus which probably resulted in less effective P5-swimming than in extant P5-swimming crab species with a short merus. Several other extant taxa represent a reversal of the P5-swimmer morphotype to varying degrees, with some extant species showing a complete reversal of unambiguous P5-swimming crab character states-one example being the well-known common shore crab Carcinus maenas. The absence of a connection between interosternite 7/8 and the sella turcica (the secondary loss of the "brachyuran sella turcica") in the stem species of Heterotremata, resulting in a junction plate which forms a cavity that offers room and attachment sites for the P5 extrinsic musculature is uncovered as preadaptation to the P5-swimmer morphotype in Heterotremata. This preadaptation is missing in Podotremata and Thoracotremata, the other two traditional main taxa of Brachyura.

Kinematics and morphology: A comparison of 3D-patterns in the fifth pereiopod of swimming and non-swimming crab species (Malacostraca, Decapoda, Brachyura).

Swimming crabs of the taxon Portunoidea show specialized, paddle-shaped fifth pereiopods (P5), which play a role in these crabs' ability to swim. In this study, the morphology of the fifth pereiopod in swimming and non-swimming crabs was studied in detail and the mobility in the articulations between podomeres was calculated from reconstructed three-dimensional (3D)-models. This way, we aimed to provide new estimates of kinematic parameters, and to answer the question on a possible homology of the P5 within several portunoid clades. We measured and compared podomere length ratios, orientations of the joint axes, and modeled single range of motion (sROM) of each joint as well as the total range of motion (tROM) of all joints of the P5 as a whole. Seven Portunoidea species, four of them belonging to the P5-swimming crab morphotype (Liocarcinus depurator, Polybius henslowii, Callinectes sapidus, Portunus pelagicus) and three not belonging to this morphotype (Carcinus aestuarii, Portumnus latipes, and with uncertain status Carupa tenuipes) were compared with the non-portunoids Sternodromia monodi, Ranina ranina, Raninoides bouvieri, Eriocheir sinensis, Varuna litterata, Ashtoret lunaris, and Cancer pagurus. The study was carried out using a combination of microcomputer tomography (µCT)-techniques and 3D-reconstructions. The µCT-data were used to create surface models of the P5 in Amira, which were then 3D-animated and manipulated in Maya to qualitatively compare modeled kinematic parameters. Results show that the merus and carpus in swimming crabs are shorter than in non-swimming crabs, while sROM angles are generally larger. The tROM of all joints expressed as Euclidean distances is generally higher in the portunoids (except for Carcinus). Our comparison of the complete trajectory of the dactylus tip regarding all maximum joint positions of the studied species suggests that the P5-swimming leg might have evolved once in the Portunoidea and got lost afterward in certain clades.

Why "swimming crabs" are able to swim - The importance of the axial skeleton: A comparison between the "swimming crab" Liocarcinus depurator and two other brachyuran crabs (Cancer pagurus, Carcinus maenas) using µCT and 3D-reconstruction.

Most brachyuran crabs use their pereiopods as walking legs, but there are also a number of species, in which the last (5th) pair of pereiopods (P5) are specialized to permit a unique mode of swimming. One of these P5-swimming crabs is Liocarcinus depurator, commonly found on European shores. We present 3-dimensional µCT-based reconstructions of the axial skeletons and 5th pereiopods (P5), including the intrinsic and extrinsic musculature of the P5, of L. depurator and of two other non-swimming brachyuran crabs, Cancer pagurus and Carcinus maenas. In Liocarcinus, we also present a reconstruction of the 4th pereiopod (P4) together with its intrinsic musculature. We further use 3-dimensional prints of the reconstructed P5 coxa and basi-ischium of L. depurator as well as thoracal parts near the thoracal-coxal arthrodial cavity to build a model which can simulate the effects of muscular activity based on muscle insertions in our 3D reconstruction and under various simulated tensile vectors. This enables us to test muscular functions that had previously been speculated upon. Reconstructed morphological structures are compared to find differences that may explain why of the three species, only L. depurator is able to swim. Significant differences between Liocarcinus and the non-swimmers Cancer and Carcinus were found in the shape of the axial skeleton, external P5 morphology and the dimensions and origin sites of the extrinsic P5 musculature, but not in the intrinsic musculature of the P5. Inclination angle measurements of P1 thoracal-coxal articulation axes against P5 axes showed that in Cancer and Carcinus, angles in the longitudinal plane were smaller than in the lateral one, whereas in Liocarcinus, they were greater. Inclination angles in the longitudinal plane were also much greater in Liocarcinus than in Cancer and Carcinus. 3D print muscular activity simulation showed that muscles inserting at the basi-ischium, which are often referred to as "levator" or "depressor" muscles, may actually also function as promotors or remotors, depending on the tensile vector within which the muscle is acting.

Viral infections in common carp lead to a disturbance of mucin expression in mucosal tissues.

In response to the constant challenge by potential pathogens, external surfaces of fish, their skin, gills and intestinal tract, are coated with mucus, a gel like substance which largely prevents the entry of pathogens. This mucus gel consists mainly of water and mucins, large O-glycosylated proteins, which are responsible for forming a gel like mixture. A modulation of the mRNA expression of mucins, was described in viral diseases in mammals however there is a knowledge gap about the regulation of mucins during viral infection in fish. Therefore, novel sequences for common carp mucins were located in an early version of the common carp genome and their mRNA expression measured in carp under infection with three different viral pathogens: (i) the alloherpesvirus cyprinid herpesvirus 3, (ii) the rhabdovirus spring viremia of carp virus and (iii) the poxvirus carp edema virus. The results showed a downregulation of mucin mRNA expression in gills and gut of common carp under infection with these pathogenic viruses. This could be a sign of a severe distress to the mucosal tissues in carp which occurs under viral infection. The reduced expression of mucins could help explaining the increased susceptibility of virus-infected carp to secondary bacterial infection.