The ovary structure, previtellogenic and vitellogenic stages in parthenogenetic species Dactylobiotus dispar (Murray, 1907) (Tardigrada: Eutardigrada)

The reproductive system of Dactylobiotus dispar consists of the ovary and the oviduct that opens into the rectum. The sack-like ovary is filled with the developing oocytes, which are assisted by the trophocytes. In D. dispar, the mixed vitellogenesis takes place. One part of the yolk material is produced inside the oocyte (autosynthesis), the second part is absorbed by micropinocytosis while the third part is synthesized in the trophocytes and is transported to the oocytes through the cytoplasmatic bridges. Moreover, rRNA, lipids and mitochondria are transfered from the trophocytes to the oocytes. The histochemical researches show that the reserve material accumulated in the oocytes contains proteins, polysaccharides and lipids.     

Effect of temperature and thermal acclimation on locomotor performance of Macrobiotus harmsworthi Murray (Tardigrada, Macrobiotidae)

We investigated the effects of acute and acclimation temperature on the locomotor performance and behavior of the tardigrade Macrobiotus harmsworthi collected from Qinling Mountains in central China. Tardigrades were acclimated to either 10 or 25 °C for 2 weeks. Then we recorded their walking speed, percentage of time moving, and the maximum distance covered by continuous locomotion at either 10 or 25 °C as the rate parameters of locomotor performance. The walking speeds of M. harmsworthi varied from 1.98 to 4.8 mm min^–1. The locomotor performance rates were significantly influenced by both acclimation temperature and performance temperature and by the interaction of the performance temperature and acclimation temperature. The data from our studies support the Beneficial Acclimation Hypothesis (BAH) which predicts that animals acclimated to a particular temperature have enhanced performance or fitness at that temperature in comparison with animals acclimated to other temperatures. The data, at least potentially, also support the Warmer is Better Hypothesis which predicts that organisms raised at high temperatures have higher relative fitness across all temperatures than do those raised at intermediate or cool temperatures. Some of the results from our studies testify the inference from the BAH that performance temperature that deviates from the acclimation temperature could cause the reduction of the locomotor performance rate.     

Light and electron microscopic studies on the excretory system of Macrobiotus richtersi Murray, 1911 (Eutardigrada)

Summary The excretory system of Macrobiotus richtersi consists of one dorsal and two lateral components and shows a high degree of structural complexity. In each of these a tricellular external lobe and a column can be distinguished, the two parts being connected distally. The surface of the lobe cells is increased by deep basal infoldings and fingerlike processes which form a labyrinth next to the basal lamina. Their cytoplasm contains numerous mitochondria, a well developed rough endoplasmic reticulum, dictyosomes, and granules in amounts depending on the physiological state of the animal. Excretory crystals occur in caveolae located in the lobe: between the fingershaped processes of the cell and in the space enclosed by the basal lamina on one side and the column on the other.The column faces an extracellular channel meandering along its whole length which is surrounded on the outside by a basal lamina. Morphologically the column is similar to the protonephridial channel of Rotifera. At the ultrastructural level, the cytoplasm of the column shows numerous mitochondria, rough endoplasmic reticulum, lysosomes, and a well developed Golgi apparatus. The lumen of the channel is coated by glycocalyx. At the base of the column several small cells form the proximal part of a duct that communicates with the gut.The morphology and ultrastructure of the excretory system of M. richtersi have been compared with similar a system in Isohypsibius megalonyx (Greven, 1979), and on these grounds a proposal is put forward to call the excretory organs of Tardigrada nephridia instead of Malpighian tubules.     

Global diversity of tardigrades (Tardigrada) in freshwater

Tardigrada is a phylum closely allied with the arthropods. They are usually less than 0.5 mm in length, have four pairs of lobe-like legs and are either carnivorous or feed on plant material. Most of the 900+ described tardigrade species are limnoterrestrial and live in the thin film of water on the surface of moss, lichens, algae, and other plants and depend on water to remain active and complete their life cycle. In this review of 910 tardigrade species, only 62 species representing13 genera are truly aquatic and not found in limnoterrestrial habitats although many other genera contain limnoterrestrial species occasionally found in freshwater. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Cuticle structure and systematics of the Macrobiotidae (Tardigrada, Eutardigrada)

The cuticle of tardigrades is characterized by three main layers: epicuticle, intracuticle and procuticle. Pillars are present in the epicuticle of almost all heterotardigrades, but these structures are also known in a few species of eutardigrades. The apparent heterogeneity of the cuticular ultrastructure in several species of the Macrobiotidae (Eutardigrada) prompted us to analyse the structure of the cuticle in this family. Eleven species in several genera were investigated with light and/or electron microscopy. All the species of the genera Murrayon and Dactylobiotus showed pillars in the epicuticle, whereas the examined species of Macrobiotus , Richtersius and Xerobiotus completely lacked pillars. Therefore, in the Macrobiotidae, in contrast to what appears with light microscopy, the cuticle is homogeneous within each genus examined at the electron microscopic level. Considering the absence of pillars in the Macrobiotidae a synapomorphy, we propose the erection of two new subfamilies. Macrobiotinae subfam. n. is characterized by the absence of pillars in the epicuticular layer and includes, in addition to the genera Macrobiotus, Xerobiotus and Richtersius, the genera Pseudohexapodibius, Adorybiotus, and probably also Minibiotus, Calcarobiotus and Pseudodiphascon . Murrayinae subfam. n. is characterized by the presence of pillars in the epicuticular layer and includes the genera Murrayon, Dactylobiotus and, probably, Macroversum.     

Behavior of red king crab larvae: Phototaxis,geotaxis and rheotaxis

Zoeae of Paralithodes camtschatica were positively phototactic to white light intensities above 1 × 10¹³ q cm^?2 s^?1. Negative phototaxis occurred at low (1 × 10¹² q cm^?2 s^?1), but not high intensities (2.2 × 10¹⁶q cm^?2 s^?1). Phototactic response was directly related to light intensity. Zoeae also responded to red, green and blue light. Zoeae were negatively geotactic, but geotaxis was dominated by phototaxis. Horizontal swimming speed of stage 1 zoeae <4 d old was 2.4 ± 0.1 (SE) cms^?1 and decreased to 1.7 ± 0.1 cm s^?1 in older zoeae (P <0.01). Horizontal swimming speed of stage 2 zoeae was not significantly different from ≥4 d old stage 1 zoeae. Vertical swimming speed, 1.6 ± 0.1 cm s^?1, and sinking rate, 0.7 ± 0.1 cm s^?1, did not change with ontogeny. King crab zoeae were positively rheotactic and maintained position in horizontal currents less than 1.4 cm s^?1. Starvation reduced swimming and sinking rates and phototactic response.     

Graphical Presentation of the African Tardigrade FaunaUsing GIS with the Description of Isohypsibius malawiensis sp. n. (Eutardigrada: Hypsibiidae) from Lake Malawi

The tardigrade fauna of the African continent is reviewed and presented graphically using Worldmap, a Geographical Information System (GIS) developed for exploring geographical diversity patterns in large biological datasets. References to the African tardigrade fauna have been gathered from published literature and supplemented with unpublished species information from the collection of Prof. Reinhardt M. Kristensen (RMK), Zoological Museum, University of Copenhagen. 156 species belonging to 36 genera of tardigrades are present. They consist of 105 eutardigrade species and 51 heterotardigrade species, of which 42 species are semiterrestrial and 9 species are marine. The presence of tardigrades are reported from 20 countries, but of these 9 are represented by a single reference. Marine tardigrades in particular have been neglected, with only a single report from the shores of the African continent. Data from the RMK collection of samples from Egypt is included in the analysis.

The scattered and sparse knowledge of the African tardigrade distribution makes general conclusions difficult, but emphasises the large regions which require further investigation. The current distribution patterns corresponds with easy accessible or ‘tourist’ locations. Regions where GIS could be used to illustrate ecological preferences are also pointed out by the analysis.

Isohypsibius malawiensis sp. n. is described from Lake Malawi. It has a smooth body surface and lacks eyes. The heteronych claws are without lunulae, but below the claws a double arched cuticular bar is present. The dorsal and ventral apophyses of the buccal tube have a large drop-shaped swelling. The first macroplacoid is the largest followed by the second, which is only slightly larger than the third; microplacoid is absent. Males are smaller than females. Females lay only one to two very large eggs in the exuvium. The Malpighian tubules are clover-shaped. The new species is interstitial in coarse sediment and no other tardigrade species were present.     

Form and function of the feeding apparatus in Eutardigrada (Tardigrada)

Tardigrade feeding apparatus is a complex structure with considerable taxonomic significance that can be schematically divided into four parts: buccal ring, buccal tube, stylet system, and pharynx. We analyzed the fine morphology and the tridimensional organization of the tardigrade buccal?Cpharyngeal apparatus in order to clarify the relationships between form and function and to identify new characters for systematic and phylogenetic studies. We conducted a comparative analysis of the cuticular structures of the buccal?Cpharyngeal apparatuses of twelve eutardigrade species, integrating data obtained by SEM and LM observations. Morphological diversity was observed and new cuticular structures such as the stylet coat of the stylet system were identified. The synthesis of the buccal?Cpharyngeal apparatus during molting was also analyzed obtaining a clear developmental sequence of its resynthesis. These findings lead us to redefine the previous interpretations of the functioning mechanisms of the buccal?Cpharyngeal apparatus and provide a more specific relationship between tardigrade diet and the anatomy of their feeding apparatuses. In addition, the detection by energy-dispersive X-ray spectroscopy of calcium in the stylets, buccal tube, and placoids of eutardigrade species (i.e., Milnesium tardigradum, Paramacrobiotus richtersi) indicates that CaCO₃ incrustations are not an exclusive feature of heterotardigrades and lead to suppose that this trait was present in the ancestors of both classes.     

Die Struktur des Nervengewebes von Macrobiotus hufelandi C.A.S. Schultze (Tardigrada)

Zusammenfassung Das Nervengewebe von Macrobiotus hufelandi zeichnet sich durch stark verzweigte pseudunipolare Nervenzellen und relativ wenige Gliazellen aus. Die Neurone besitzen rauhes ER, freie Ribosomen, zahlreiche Mitochondrien, einen wenig ausgeprägten Golgi-Apparat und einen elektronenlichten Kern. In ihren Axonen finden sich Vesikel und Einschlüsse unterschiedlicher Größe und Zusammensetzung. Die Gliazellen verzweigen sich stark und besitzen glattes ER, viele freie Ribosomen und einen elektronendichten Kern. Ganglien und Nerven werden nur durch eine dünne Neurallamelle vom extraganglionären Raum getrennt. Die morphologische Ausbildung des Nervengewebes wird im Hinblick auf die extreme Lebensweise der Tardigraden diskutiert.

---

The structure of the nervous tissue in Macrobiotus hufelandi C. A. S. schultze (tardigrada)

Summary The typical elements of the nervous tissue of Macrobiotus hufelandi are strongly ramified pseudunipolar neurons and a small amount of glial cells. In the perikarya of neurons there are rough ER, free ribosomes, many mitochondria, a poor Golgi-apparatus, and an electron-light nucleus. Nerve fibers contain masses of vesicles and inclusions of different size and composition. The ramifying glial cells have smooth ER, many free ribosomes, and an electron-dense nucleus. Ganglia and nerves are separated from the extraganglionic cavity by a thin acellular sheath (neural lamella). The organization of the nervous tissue is discussed with regard to the extreme conditions of environment of the tardigrades.

     

A new species of the genus Macrobiotus(Tardigrada:Macrobiotidae) from Southeastern China

A new species Macrobiotus wuyishanensis sp.nov.(Tardigrada:Macrobiotidae) is described from Mt.Wuyi,Southeastern China.It is most similar to Macmbiotus spectabilis Thulin,1928,but differs by lacking fine punctuation on the distal half of the projections on the egg,a ring of polygons that circle the base of each projection,and punctation between the projections on the egg shell.It is also similar to M.grandis Richters,1911,but differs by smaller body size,lacking polygons and punctations between the projections on egg shell.     

Biogeography of limno-terrestrial Tardigrada, with particular reference to the Antarctic fauna

The Tardigrada is a cosmopolitan phylum of pre-Pangaean origin, yet tardigrade families and genera show distinct biogeographic components isolated by two major geological events. Separate Laurasian and Gondwanan familial clusters correlate with the Triassic disintegration of Pangaea, while discrete Antarctic, Australian and New Zealand familial/generic clusters relate to the subsequent Jurassic/Cretaceous disintegration of Gondwana.     

Spermatozoa in the reproductive system of a hermaphroditic marine tardigrade,Orzeliscus belopus (Tardigrada: Arthrotardigrada)

Orzeliscus belopus has long been regarded as the only hermaphroditic marine tardigrade, yet there has been no published detailed information on its internal anatomy. Our study elucidates the ultrastructure of the ovotestis and the spermatozoa of Orzeliscus cf. belopus from Bermuda. The ovotestis had no septum to separate male and female germ cells and, while early stages of spermatogenesis were not observed, many spermatozoa were found at the periphery in both anterior and posterior areas of the gonad. The nucleus and the mitochondria of the spermatozoa in the ovotestis extend backward from the centriole region, forming a half-headed arrow-shape with the nucleus on the outer side of the ‘arrowhead’. The cross section of a long vesicular body (the paranuclear vesicle) is dumbbell- or horseshoe-shaped and attached to almost the entire length of the nucleus. In the seminal receptacle, we found both a complete spermatozoon and some which started to degrade. There is an indication that sperm is further modified after discharge as within the receptacle duct the sperm is no longer half-headed arrow-shaped but has a straight nucleus. This modification might be correlated with the degeneration of the paranuclear vesicle. Our observations clearly show that O. belopus is a simultaneous hermaphrodite, and suggests that the reproductive mode includes copulation and cross-fertilization.     

The biology and ecology of lotic Tardigrada

• 1 Tardigrades comprise a micrometazoan phylum that is a sister group of the arthropods.
• 2 They are components of the meiobenthos in lotic habitats, and ≈ 50–70 species have been reported in such habitats world‐wide. Approximately 800 species have been identified from all marine, freshwater, and terrestrial habitats.
• 3 Taxonomy is based primarily on the morphology of the claws, buccal‐pharyngeal apparatus, cuticle and eggs.
• 4 Reproductive modes include sexual reproduction (amphimixis) and parthenogenesis. The sexual condition of individuals may be either gonochorism, unisexuality, or hermaphroditism. Moulting occurs throughout the life of the tardigrade.
• 5 Latent states (cryptobiosis, including encystment, anoxybiosis, cryobiosis, osmobiosis and anhydrobiosis) enable tardigrades to withstand unfavourable environmental conditions.
• 6 Population densities, life histories, dissemination and biogeography of freshwater species are poorly known.

     

GEOTAXIS/PHOTOTAXIS AND BIOCHEMICAL PATTERNS IN HETEROCAPSA (=CACHONINA) ILLDEFINA (DINOPHYCEAE) DURING DIEL VERTICAL MIGRATIONS

Two separate experiments with Heterocapsa (= Cachonina ) illdefina Herman et Sweeney, one with and the other without water volume replacement, were performed in a 250-L laboratory mesocosm (45-cm diameter × 150-cm height) to examine how diel vertical migration (DVM) relates to taxis sign and strength and to cellular biochemical state. Although only the cell population grown with water volume replacement maintained a division per day over the course of the experiment, periodic measurements during both experiments demonstrated that cells aggregating at the surface during the light period generally were deficient in all measured biochemical constituents compared to cells obtained from a midcolumn depth. More specifically, H. illdefina cells that aggregated at the surface during the light period in both experiments exhibited weakened positive geotaxis but strengthened positive phototaxis and were very deficient in lipid and free amino acid compared to midcolumn cells. Cells sampled at midcolumn during the light period exhibited similar but weaker taxes changes compared to surface samples, and geotaxis strength was inversely correlated with cell diameter, cellular DNA and protein content, and RNA/DNA ratio. In comparison, published data on Gymnodinium breve Davis, a harmful algal bloom species, showed that cells aggregating at the surface during the light period generally exhibited weakened negative geotaxis and strengthened positive phototaxis and were very deficient in lipid and chl a compared to midcolumn cells. Although the persistent tendency toward negative geotaxis was weaker in midcolumn subpopulations throughout the day, its strength was inversely correlated with cell diameter and cellular lipid content. The combined results for both species support a revised conceptual model of optimized DVM in autotrophic marine dinoflagellates incorporating generalized expressions of taxis and biochemical state of individual cells.     

Vertical Migration of the Rice White-Tip Nematode,Aphelenchoides besseyi

In the laboratory, the vertical migration of Aphelenchoides besseyi was favored by rough surfaces and an inverse water gradient. In relation th gravity, the nematode migrated down or up equally, a circumstance suggesting that a geotaxis was not involved. Stems of rice seedlings were effective surfaces for vertical migration of nematodes only when the stems were continuously supplied with moisture.     

A survey of the terrestrial Tardigrada of the Vestfold Hills,Antarctica

A survey of the terrestrial tardigrades inhabiting growths of algae, lichens and mosses in the Vestfold Hills, Antarctica, was carried out at 11 and 35 sites during the austral summers of 1980 and 1982, respectively. In all, 24 species of plants were collected from which four genera and four species of Tardigrada were recovered. A key to the tardigrades of the area is presented. The distribution and associational patterns of the tardigrades are discussed in the context of other studies of antarctic Tardigrada.     

Phenotypic Variations in the Life History of Two Clones of Macrobiotus richtersi (Eutardigrada, Macrobiotidae)

A comparative study of life history traits of two clones (CDMr01 and CDMr02) of a triploid thelytokous apomictic population of the eutardigrade Macrobiotus richtersi has been carried out. Both clones were reared under the same lab conditions: temperature of 14 °C, photoperiod of 12 h/12 h (L/D), and nematodes ad libitum as food. Statistical analysis of the life history traits considered has indicated interclonal variability. The two clones were significantly different in the number of eggs per clutch (fertility), number of eggs laid per female per life span (fecundity), hatching percentage of eggs and hatching time. Similarities between clones have been observed with regard to life span, total number of ovipositions per life span, and age at first oviposition. In addition, significant differences in hatching time, with eggs hatched over a long period, were found within each clone. Interclonal variability in life history traits indicated the effects of genetic factors, whereas intraclonal variability reflected the effects of environmental factors. The evolutionary and adaptive significance of the life history phenotypic variations is discussed.