Quantitative cellular analysis of growth and reproduction in freshwater planarians (Turbellaria; Tricladida). I. A cellular description of the intact organism

Summary

Changes in egg shape and surface morphology during maturation may be related to the localization of cytoplasmic determinants in the embryos of organisms with spiral cleavage. The eggs of the polyclad turbellarian Hoploplana inquilino undergo pronounced shape changes during the meiotic divisions which have been examined with the scanning electron microscope. Unfertilized eggs have a smooth surface that becomes covered with microvilli and microblebs within 10 min of fertilization. First polar body extrusion is accompanied by the asymmetric appearance of large blebs (Blebbing Cycle I) primarily in the animal hemisphere with one quadrant characteristically smoother than the others and bearing fewer blebs. Blebbing Cycle II, which is less pronounced than the first but is still characterized by a relatively un blebbed quadrant of the zygote, coincides with second polar body formation. These asymmetric shape changes in the animal hemisphere during meiosis may possibly correlate with a primitive form of morphogenetic segregation and beginning quadrant specialization in polyclads, the most primitive spiralians with mosaic development.     

Paläogene Eischalenreste von Hainin (Belgien)

From Paleocene deposits from Belgium fossil egg shells of turtles are reported. A systematically not interpretable form(Testudinarum ovum) is described, together withHaininchelys curiosa n. g. n. sp. as a new genus organus for fossil eggs. Egg shells from Dinosauna, Aves, Testudines, Crocodylia and Squamata are discussed for comparison.     

Morphology of the Turbellaria at the ultrastructural level

The paper reviews the following systems: body wall, nervous system, protonephridia, parenchyma, digestive system, reproductive system, and includes a summary of the literature. New information is presented for the catenulid and neoophoran body wall-construction, the kalyptorhynch proboscis, the catenulid and haplopharyngid protonephridial construction, and the prolecithophoran spermatozoon and female germ cell. Examples of new features, as well as examples of how electron microscopy has clarified the relative position of structures and their substructures are given from the subcellular level to the organization of whole organs. Fine structural features linking different turbellarian orders are summarized. They apparently support Karling's (1974) latest assessment of the affinities between the turbellarian orders which is based primarily on light histological data, they add the recognition of a special link between the Macrostomida and Haplopharyngida and they suggest the existence of three main evolutionary lines within the Turbellaria.     

Light and Electron Microscope Study of Urosporidium cannoni N. Sp., a Haplosporidian Parasite of the Polyclad Turbellarian Stylochus sp.

ABSTRACT. Urosporidium cannoni n. sp. was found in most tissues of a polyclad turbellarian associated with commercial oyster farms from eastern Moreton Bay, Australia. Mature spores had 11 to 13 episporal tails at irregular intervals around the spore wall. The spherule, prominent in development, dispersed as the spores matured. Spores were found within cysts that contained up to a few hundred spores at the same stage of development. Mature cysts occurred in groups and their masses of dark, golden spores resulted in black patches on the otherwise red-brown turbellarian. The infection may have commercial importance.     

Pelvis morphology suggests that early Mesozoic birds were too heavy to contact incubate their eggs

Numerous new fossils have driven an interest in reproduction of early birds, but direct evidence remains elusive. No Mesozoic avian eggs can be unambiguously assigned to a species, which hampers our understanding of the evolution of contact incubation, which is a defining feature of extant birds. Compared to living species, eggs of Mesozoic birds are relatively small, but whether the eggs of Mesozoic birds could actually have borne the weight of a breeding adult has not yet been investigated. We estimated maximal egg breadth for a range of Mesozoic avian taxa from the width of the pelvic canal defined by the pubic symphysis. Known elongation ratios of Mesozoic bird eggs allowed us to predict egg mass and hence the load mass an egg could endure before cracking. These values were compared to the predicted body masses of the adult birds based on skeletal remains. Based on 21 fossil species, we show that for nonornithothoracine birds body mass was 187% of the load mass of the eggs. For Enantiornithes, body mass was 127% greater than the egg load mass, but some early Cretaceous ornithuromorphs were 179% heavier than their eggs could support. Our indirect approach provides the best evidence yet that early birds could not have sat on their eggs without running the risk of causing damage. We suggest that contact incubation evolved comparatively late in birds.     

Initial morphological diversity as a criterion in deciphering turbellarian phylogeny

The most profound structural variety in morphofunctional systems and morphogenetic mechanisms, i.e. the highest morphological diversity, is observed in those groups where these systems and mechanisms are evolutionarily most primitive. Here, such variety can involve the basic body plan of a given phylum and the types of morphogenesis characteristic of it. This correlation provides a new criterion of evolutionary primitiveness, namely, the criterion of initial morphological diversity.The highest morphological diversity among turbellarian groups is observed in the order Acoela. Acoel turbellarians are archaic in most of their features, apparently being a group near the base of the turbellarian phylogenetic tree. Among other turbellarians there are a few groups that also are archaic in some few features (above all, the Catenulida), although on the whole they are more advanced than the Acoela. The Turbellaria as a whole is notable for its morphological diversity in comparison with other classes of the Scolecida.     

浙江天台晚白垩世伤齿龙(troodontids)蛋化石的新发现

浙江天台上白垩统赤城山组中发掘出属伤齿龙类的棱柱形蛋化石。研究认为,棱柱形蛋从伤齿龙类恐龙中产出就具有较高刚度的蛋壳,可直接插入松软的泥沙里,明显区别于其他大多数恐龙及现代爬行类所产的韧性蛋壳蛋,韧性蛋壳的恐龙蛋往往因重力作用产生变形而呈扁球形和扁椭球形。伤齿龙类的棱柱形蛋的刚性蛋壳很接近现代鸟蛋的蛋壳,暗示伤齿龙类恐龙生殖系统在蛋壳形成方面与现代鸟类有一定相似性。同时,伤齿龙类的棱柱形蛋或多或少地以向外倾斜的形式排布,反映出伤齿龙类恐龙产卵时可能面向巢穴外,是一种对生存环境的适应行为。     

Reaction of the mussel Mytilus galloprovincialis (Bivalvia) to Eugymnanthea inquilina (Cnidaria) and Urastoma cyprinae (Turbellaria) concurrent infestation

In total 480 individuals of Mytilus galloprovincialis were sampled monthly from October 2009 to September 2010, at the shellfish farm in the Mali Ston Bay, south Adriatic Sea (Croatia) in order to assess the extent of pathology imposed by two parasites, Eugymnanthea inquilina (Cnidaria) and Urastoma cyprinae (Turbellaria). Although a deteriorating impact on host reproduction or condition index was lacking, we evidenced ultrastructural and functional alteration in host cells at the attachment site. Ultrastructural changes included hemocytic encapsulation of the turbellarian and cell desquamation in medusoid infestation. Caspase positive reaction inferred by immunohistochemistry (IHC) was triggered in cases of turbellarian infestation, in contrast with hydroids, suggesting that the former exhibits more complex host-parasite interaction, reflected in the persistent attempts of the parasite to survive bivalve reaction. We have evidenced that both organisms trigger specific host reaction that although not costly in terms of host reproductive cycle or growth, results in mild tissue destruction and hemocyte activation. A lower degree of tissue reaction was observed in cases of hydroid infestation, compared to turbellarian.     

Reproductive investment in moa: a K-selected life-history strategy?

The exaggerated K-selected life-history strategy of moa has been suggested as an important factor causing their rapid extinction. Classically, this strategy is characterized by few, large offspring and low fecundity rates. Assuming clutches with one or two eggs as derived from the fossil record, we tested if eggs of moa were larger than the average of similar-sized birds, and estimated their unknown annual breeding frequencies. Therefore, we established allometries on body mass and different reproductive traits (i.e. egg mass, clutch mass and annual clutch mass). These were derived for r-selected (r-model) and for K-selected (K-model) bird species. In agreement with our initial expectations, moa had egg to body mass relations seen in “average” extant K-selected birds. While the K-model pointed to a clutch size of one or two eggs for moa corroborating fossil data and a K-selected life-history, the r-model predicted two to three times larger sizes. Under clutch sizes between one and two eggs and an annual clutch mass as observed in other similar-sized flightless island birds (e.g. rails, ratites), the annual clutch mass allometry suggested one to three clutches per year for moa. Even when assuming less than one brood per year (K-model predicts 0.5 clutches per year); annual clutch masses were still consistent with the K-model. Further studies are needed to clarify whether or not the reproductive strategy of flightless island birds and/or of the birds underlying the K-model fits better to the moa strategy. The approach presented herein, illustrates that combining biological and paleontological data can assist in the reconstruction of species traits, which are insufficiently or not preserved in fossils, but are necessary to understand the evolution of traits.     

BIRD EGGSHELL IN DOMINICAN AMBER

Abstract:  Here we report an eggshell in Dominican amber, representing the first vertebrate egg in any amber deposit. The eggshell is compared with present-day eggs of lizards, snails and birds. Based on the surface structure and type of shell breakage, it appears that the most likely candidate is a bird, and with that consideration, an avian group that produces eggs similar to the fossil in shape, size and colouration is the Trochilidae (hummingbirds). Several possible explanations of how the fossil could be preserved in amber are provided. If indeed a hummingbird was involved, this discovery would represent the first New World record of a fossil trochilid.     

A gravid fossil turtle from the Early Cretaceous reveals a different egg development strategy to that of extant marine turtles

Extant sea turtles develop and lay pliable (flexible) eggs; however, it is unknown whether they inherited this reproductive strategy from their closer fossil relatives or if it represents an evolutionary novelty. Here, we describe the first undisputable gravid marine fossil turtle ever found, from the early Cretaceous of Colombia, belonging to Desmatochelys padillai Cadena & Parham, which constitutes a representative of the Protostegidae. Using thin sectioning of one of the eggs, as well as scanning electron microscopy coupled with elemental characterization, cathodoluminescence, and computer tomography, we established that Desmatochelys padillai produced rigid eggs similar to those associated with some extant and fossil freshwater and terrestrial turtles. At least 48 spherical eggs were preserved inside this gravid turtle. We suggest that the development of rigid eggs in the extinct marine turtle Desmatochelys padillai resulted as an adaptation for egg‐embryo requirements dictated by the physical attributes of the nesting site.     

Oceanic transport of calcified eggs: taphonomic and biogeographic implications

Estimates of the temporal and spatial resolution of fossil egg assemblages are required to constrain the inferences that can logically be drawn during assemblage analysis. Consequently, understanding egg transport mechanisms is required before conclusions are developed. Bird eggs are buoyant during part of development and can float from near shore nests during high tides, storm surges, or flooding. Complete and unbroken eggs have survived transoceanic transport. Evidence of these transport processes operating in the past is suggested by the recovery of fossil eggshell in marine deposits. Transport of eggs can potentially confound fossil assemblage spatial analysis since evidence of oceanic transport may not be present on an eggshell when collected. Bird embryos exposed to cold temperatures and immersed in both fresh and salt water can survive from hours to two days, suggesting that oceanic transport of viable eggs is possible, a conclusion further supported by field observations. Eggs of other taxa show similar environmental survival trends and may also be capable of surviving oceanic transport through floatation or rafting. In addition, if viable eggs are transported across seas and hatch, emigration or gene flow of terrestrial egg-laying vertebrates may be possible between landmasses.     

First fossil gravid turtle provides insight into the evolution of reproductive traits in turtles

Here we report on the first discovery of shelled eggs inside the body cavity of a fossil turtle and on an isolated egg clutch, both referable to the Cretaceous turtle Adocus. These discoveries provide a unique opportunity to gain insight into the reproductive traits of an extinct turtle and to understand the evolution of such traits among living turtles. The gravid adult and egg clutch indicate that Adocus laid large clutches of rigid-shelled spherical eggs and established their nests near rivers, traits that are shared by its closest living relatives, the soft-shelled turtles. Adocus eggshell, however, was probably more rigid than that of living turtles, based on its great thickness and structure, features that may represent unique adaptations to intense predation or to arid nest environments. In light of the reproductive traits observed in Adocus, the distribution of reproductive traits among turtles reveals that large clutches of rigid-shelled eggs are primitive for hidden-necked turtles (cryptodirans) and that spherical eggs may have evolved independently within this group.     

湖北安陆新的恐龙蛋类型的发现及其意义

本文记述的恐龙蛋化石标本,采自湖北省江汉盆地公安寨组下部.蛋化石保存完好.在普通光学显微镜和扫描电镜下观察,蛋壳的显微结构完整,清晰可见.根据钙质蛋壳基本结构单位的形态及排列特征,笔者建立了—新属,新种——Dendroolithus wangdianensis gen. et sp. nov.,代表恐龙蛋类一个新科——Dendroolithidae fam. nov.此外,残存的卵壳膜纤维化石的发现,将为进一步研究恐龙蛋卵壳膜的结构和氨基酸组成提供宝贵材料.     

The importance of turbellarians in the marine meiobenthos: a review

Recent data and opinions on meiofaunal ecology are briefly reviewed; and from scattered data, the place of turbellarians in the meiobenthic community is discussed. Turbellarian diversity, density, and biomass are higher in sandy habitats than in muddy bottoms. In sand, turbellarian diversity is of the same magnitude as that of other important meiofaunal taxa, while densities range between 7–25% of the total meiofauna. Mean individual turbellarian dry weight seems to be four times that of nematodes and in sandy habitats turbellarian biomass may be equal to or excede that of nematodes. Most turbellarian species may be considered as predators and in this respect may take the place occupied by macrofaunal species in muddy sediments. mens.

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Abstract

Described in 1930–31 by Steinböck who considered it the most primitive bilaterian, the turbellarian genus Nemertoderma is known for its role in platyhelminth phylogeny as much as for its muddled taxonomy. On the basis of material collected in the Mediterranean, Atlantic and Pacific Ocean since 1964 the known four genera and seven species (Nemertoderma bathycola Steinböck, 1930–31; N. westbladi Steinböck, 1938; N. psammicola Sterrer, 1970 (syn. N. rubra Faubel, 1976); Meara stichopi Westblad, 1949; Meara sp. Smith et al. 1994; Nemertinoides elongatus Riser, 1987; and Flagellophora apelti Faubel &; Dörjes, 1978) are redescribed and one new genus with two new species (Ascoparia neglecta n. g. n. sp. and A. secunda n. sp.). Live observations on morphological variability, body size vs. reproductive state, statocyst structure and statolith variability, and sperm morphology and dimorphism are provided. The paper concludes with a discussion and diagnoses of the known taxa of Nemertodermatida, including the new family Ascopariidae.     

Three Trematodes Including Two New Species from Fishes of the Karachi Coast

Three trematodes are described from three different species of Karachi coast fishes. Complexo-bursa magna sp.n., from Lates calcarifer , is characterized by its large body size, unlotxjd rounded ovary, large operculate eggs, and numerous vitelline follicles extending from the preacetabular region to the posterior end of the body. Monorchis heterorchis sp.n., from the fish Muraenesox cinereus , is also characterized by its relatively large body, median transversely elongated testes of variable shapes, ovary consisting of four distinct follicles and relatively large operculate eggs. Waretrema piscicola Srivastava, 1937 is reported here from the Karachi coast 17 years after its last report from the Philippines, in the same host ( Scato-phaugs argus ) but from a new locality.     

Tapeworm Eggs in a 270 Million-Year-Old Shark Coprolite

Remains of parasites in vertebrates are rare from the Mesozoic and Paleozoic. Once most parasites that live in – or pass through – the gastrointestinal tract of vertebrates, fossil feces (coprolites) or even intestinal contents (enterolites) can eventually preserve their remains. Here we announce the discovery of a spiral shark coprolite from the Paleozoic bearing a cluster of 93 small oval-elliptical smooth-shelled structures, interpreted as eggs of a tapeworm.The eggs were found in a thin section of an elasmobranch coprolite. Most of the eggs are filled by pyrite and some have a special polar swelling (operculum), suggesting they are non-erupted eggs. One of the eggs contains a probable developing larva. The eggs are approximately 145–155 µm in length and 88–100 µm in width and vary little in size within the cluster. The depositional and morphological features of the eggs closely resemble those of cestodes. Not only do the individual eggs have features of extant tapeworms, but their deposition all together in an elongate segment is typical to modern tapeworm eggs deposited in mature segments (proglottids). This is the earliest fossil record of tapeworm parasitism of vertebrates and establishes a timeline for the evolution of cestodes. This discovery shows that the fossil record of vertebrate intestinal parasites is much older than was hitherto known and that the interaction between tapeworms and vertebrates occurred at least since the Middle-Late Permian.     

Observations on lesser-known flatworms: Udonella

Udonella caligorum is acknowledged as a monogenean by some (Baer & Euzet, 1961; Yamaguti, 1963), but not by others (Ivanov, 1952; Bychowsky, 1957). It also has many similarities to Temnocephala, a rhabdocoel turbellarian. Light and electron microscope studies showed that the posterior sucker is glandular and that neither the adult nor eggs possess hooks; the head-gland secretion is eosinophilic and granular, and the tegument composed primarily of secretory granules and mitochondria. Ten pair of cells appearing to be paranephrocytes are present. Udonella larvae were shown to be larger than the host copepod nauplii, so transmission of worms between hosts must occur either during host mating, casual contact of hosts, or through free worms on the surface of the fish which the copepod infests.