Phylogenomic analyses of lophophorates (brachiopods, phoronids and bryozoans) confirm the Lophotrochozoa concept

Based on embryological and morphological evidence, Lophophorata was long considered to be the sister or paraphyletic stem group of Deuterostomia. By contrast, molecular data have consistently indicated that the three lophophorate lineages, Ectoprocta, Brachiopoda and Phoronida, are more closely related to trochozoans (annelids, molluscs and related groups) than to deuterostomes. For this reason, the lophophorate groups and Trochozoa were united to Lophotrochozoa. However, the relationships of the lophophorate lineages within Lophotrochozoa are still largely unresolved. Maximum-likelihood and Bayesian analyses were performed based on a dataset comprising 11,445 amino acid positions derived from 79 ribosomal proteins of 39 metazoan taxa including new sequences obtained from a brachiopod and a phoronid. These analyses show that the three lophophorate lineages are affiliated with trochozoan rather than deuterostome phyla. All hypotheses claiming that they are more closely related to Deuterostomia than to Protostomia can be rejected by topology testing. Monophyly of lophophorates was not recovered but that of Bryozoa including Ectoprocta and Entoprocta and monophyly of Brachiozoa including Brachiopoda and Phoronida were strongly supported. Alternative hypotheses that are refuted include (i) Brachiozoa as the sister group of Mollusca, (ii) ectoprocts as sister to all other Lophotrochozoa including Platyzoa, and (iii) ectoprocts as sister or to all other protostomes except chaetognaths.     

苔藓动物18S rRNA基因的分子系统发生初探

本文对我国沿海较为常见的8种唇口目苔藓动物的18SrRNA基因进行了PCR扩增和序列测定。结合已知的其它苔藓动物（包括内肛动物和外肛动物）以及腕足动物和帚虫的相应序列,运用分子系统学方法,研究苔藓动物门的系统发生关系,结果表明,外肛动物和内肛动物构成苔藓动物分子系统树中的二大平行支;本文测定的大室膜孔苔虫与Giribet等测定的膜孔苔虫在系统树中的位置间隔较远。结果也支持外肛动物包含被唇纲和裸唇纲两大类群的形态划分,而关于裸唇纲特别是唇口目内部的系统发生关系。分子数据的分析结果和形态分类之间的分歧有待于进一步研究。     

Microarchitecture of Body Wall of Extant Cyclostome Ectoprocts

Scanning electron microscope and light microscope studies ofthe body wall in cyclostome stenolaemate ectoprocts show thatthis wall has a distinctive microarchitecture. The microstructuresconsist of a linear series of longitudinal canals in the centralregion. Adjoining regions on either side have laminae of overlappingcalcareous tablets. In some cyclostomes at certain stages ofgrowth, laminae may show a less orderly meshwork pattern nearthe canals. The laminate layers of calcareous tablets are enclosedin an organic matrix and are penetrated by tubuli which passfrom the inner part of the body wall to the canals and alsolink the canals. The canals with organic and mineralized matricesopen into pores in the body wall. This elaborate meshwork andcanal and pore system provide the framework for growth and resorptionof the body wall and facilitate transport of cellular materialsfrom one zooid to another in a colony. Calcification at thedistal part of a colony seems to proceed in a series of stagesaround sites of calcification. The pattern of the body wallof the cyclostome ectoproct does not parallel a molluscan patternof skeletal growth as previously proposed. The hypothesis ofa hypostegal coelom in some cyclostome ectoprocts is rejected.     

Trochophora larvae: cell-lineages, ciliary bands and body regions. 2. Other groups and general discussion

The embryology of sipunculans, entoprocts, nemertines, platyhelminths (excluding acoelomorphs), rotifers, ectoprocts, phoronids, brachiopods, echinoderms and enteropneusts is reviewed with special emphasis on cell-lineage and differentiation of ectodermal structures. A group Spiralia comprising the four first-mentioned phyla plus annelids and molluscs seems well defined through the presence of spiral cleavage with early blastomere specification, prototroch with characteristic cell-lineage, cerebral ganglia developing from cells of the first micromere quartet (i.e., the episphere) and a ventral nervous system developing from the hyposphere. The planktotrophic trochophore was probably the larval type of the ancestor of this group. Another group comprising phoronids, brachiopods, echinoderms and enteropneusts appears equally well delimited. It is characterized by radial cleavage with late blastomere specification, possibly by the presence of a neotroch consisting of monociliate cells, by the absence of cerebral ganglia and of a well-defined brain and paired longitudinal nerve cords developing in connection with the blastopore, and by coelomic organization. Its ancestral larval type was probably a dipleurula. Several characters link rotifers with the spiralians, although they do not show the spiral pattern in the cleavage. Ectoprocts are still a problematic group, but some characters indicate spiralian affinities.     

Comparison of articulate brachiopod nuclear and mitochondrial gene trees leads to a clade-based redefinition of protostomes (Protostomozoa) and deuterostomes (Deuterostomozoa)

Nuclear and mtDNA sequences from selected short-looped terebratuloid (terebratulacean) articulate brachiopods yield congruent and genetically independent phylogenetic reconstructions by parsimony, neighbour-joining and maximum likelihood methods, suggesting that both sources of data are reliable guides to brachiopod species phylogeny. The present-day genealogical relationships and geographical distributions of the tested terebratuloid brachiopods are consistent with a tethyan dispersal and subsequent radiation. Concordance of nuclear and mitochondrial gene phylogenies reinforces previous indications that articulate brachiopods, inarticulate brachiopods, phoronids and ectoprocts cluster with other organisms generally regarded as protostomes. Since ontogeny and morphology in brachiopods, ectoprocts and phoronids depart in important respects from those features supposedly diagnostic of protostomes, this demonstrates that the operational definition of protostomy by the usual ontological characters must be misleading or unreliable. New, molecular, operational definitions are proposed to replace the traditional criteria for the recognition of protostomes and deuterostomes, and the clade-based terms ''Protostomoza'' and ''Deuterostomozoa'' are proposed to replace the existing term ''Protostomia'' and ''Deuterostomia''.     

Field studies of the diet of Rhabdosargus holubi (Pisces: Teleostei: Sparidae)

Rhabdosargus holubi is a marine teleost endemic to south east Africa. The juveniles occur mainly in estuaries but the adults are largely confined to the sea. Stomach contents of over 2000 juvenile R. holubi were examined from fish captured in the open Kowie and Msikaba estuaries, and from the closed West Kleinemond and Kasouga estuaries. Juvenile R. holubi feed mainly on aquatic vegetation. The fish cannot, however, digest the vegetation eaten due to the absence of a cellulase or a method of breaking up the plant tissue. Therefore, the plant material is passed out in an undigested state. Epiphytic diatoms and sessile ectoprocts (Bryozoa) are, however, removed from the macrophytes or multicellular algae in the stomach, and are subject to digestion. Diatoms may form up to 50% of the dry weight of the plants consumed. A variety of animals was also eaten. Maximum total food consumption was estimated at 1·9% of body weight per day although this is probably an underestimate. Feeding only took place during daylight hours. The teeth of juvenile R. holubi are specialized for grazing aquatic plants and differ from those of adult R. holubi. Adults consume mainly bivalves and large crustaceans. It is suggested that the different food requirements of the adults and juveniles determine at what stage this species enters and leaves estuaries.     

白垩纪缅甸琥珀双翅螳蛉科一新属新种(昆虫纲: 脉翅目)

基于三枚晚白垩世Cenomanian期早期缅甸琥珀化石标本,记述了脉翅目双翅螳蛉科Dipteromantispidae一新属新种:卓德曲双翅螳蛉Kurtodipteromantispa zhuodei gen.etsp.nov.。新种与格氏棒翅螳蛉Halteriomantispa grimaldii Liu et al., 2016相似,但以下特征区别于后者:1)前翅无色斑; 2)前翅ScP端部在scp-ra横脉处具明显弯折,不平滑; 3)前翅RP至少具1末端分叉的分支; 4)雌外生殖器肛上板与第9背板融合,第9生殖基节近半圆形,末端无近三角形突起。此外,描述并讨论了新属雌外生殖器的形态结构。     

The phylogenetic position of entoprocta, ectoprocta, phoronida, and brachiopoda

Ectoprocts, phoronids and brachiopods are often dealt with underthe heading Tentaculata or Lophophorata, sometimes with entoproctsdiscussed in the same chapter, for example in Ruppert and Barnes(1994). The Lophophorata is purported to be held together bythe presence of a "lophophore," a mesosomal tentacle crown withan upstream-collecting ciliary band. However, the mesosomaltentacle crown of pterobranchs has upstream-collecting ciliarybands with monociliate cells, similar to those of phoronidsand brachiopods, although its ontogeny is not well documented.On the contrary, the ectoproct tentacle crown carries a ciliarysieving system with multiciliate cells and the body does notshow archimery, neither during ontogeny nor during budding,so the tentacles cannot be characterized as mesosomal. The entoproctshave tentacles without coelomic canals and with a downstream-collectingciliary system like that of trochophore larvae and adult rotifersand serpulid and sabellid annelids. Planktotrophic phoronidand brachiopod larvae develop tentacles at an early stage, buttheir ciliary system resembles those of echinoderm and enteropneustlarvae. Ectoproct larvae are generally non-feeding, but theplanktotrophic cyphonautes larvae of certain gymnolaemates havea ciliary band resembling that of the adult tentacles. The entoproctshave typical trochophore larvae and many feed with downstream-collectingciliary bands. Phoronids and brachiopods are thus morphologicallyon the deuterostome line, probably as the sister group of the"Neorenalia" or Deuterostomia sensu stricto. The entoproctsare clearly spiralians, although their more precise positionhas not been determined. The position of the ectoprocts is uncertain,but nothing in their morphology indicates deuterostome affinities."Lophophorata" is thus a polyphyletic assemblage and the wordshould disappear from the zoological vocabulary, just as "Vermes"disappeared many years ago.     

The evolution of the serotonergic nervous system

The pattern of development of the serotonergic nervous system is described from the larvae of ctenophores, platyhelminths, nemerteans, entoprocts, ectoprocts (bryozoans), molluscs, polychaetes, brachiopods, phoronids, echinoderms, enteropneusts and lampreys. The larval brain (apical ganglion) of spiralian protostomes (except nermerteans) generally has three serotonergic neurons and the lateral pair always innervates the ciliary band of the prototroch. In contrast, brachiopods, phoronids, echinoderms and enteropneusts have numerous serotonergic neurons in the apical ganglion from which the ciliary band is innervated. This pattern of development is much like the pattern seen in lamprey embryos and larvae, which leads the author to conclude that the serotonergic raphe system found in vertebrates originated in the larval brain of deuterostome invertebrates. Further, the neural tube of chordates appears to be derived, at least in part, from the ciliary band of deuterostome invertebrate larvae. The evidence shows no sign of a shift in the dorsal ventral orientation within the line leading to the chordates.     

The nervous system of <Emphasis Type="Italic">Paludicella articulata</Emphasis> - first evidence of a neuroepithelium in a ctenostome ectoproct

Introduction

Comparatively few data are available concerning the structure of the adult nervous system in the Ectoprocta or Bryozoa. In contrast to all other ectoprocts, the cerebral ganglion of phylactolaemates contains a central fluid-filled lumen surrounded by a neuroepithelium. Preliminary observations have shown a small lumen within the cerebral ganglion of the ctenostome Paludicella articulata. Ctenostome-grade ectoprocts are of phylogenetic relevance since they are considered to have retained ancestral ectoproct features. Therefore, the ctenostome Paludicella articulata was analyzed in order to contribute to the basal neural bauplan of ctenostomes and the Ectoprocta in general.

Results

The presence of a lumen and a neuroepithelial organization of the nerve cells within the cerebral ganglion are confirmed. Four tentacle nerves project from the cerebral ganglion into each tentacle. Three of the tentacle nerves (one abfrontal and two latero-frontal nerves) have an intertentacular origin, whereas the medio-frontal nerve arises from the cerebral ganglion. Six to eight visceral nerves and four tentacle sheath nerves are found to emanate from the cerebral ganglion and innervate the digestive tract and the tentacle sheath, respectively.

Conclusions

The situation in P. articulata corresponds to the situation found in other ctenostomes and supports the notion that four tentacle nerves are the ancestral configuration in Ectoprocta and not six as proposed earlier. The presence of a lumen in the ganglion represents the ancestral state in Ectoprocta which disappears during ontogeny in all except in adult Phylactolaemata and P. articulata. It appears likely that it has been overlooked in earlier studies owing to its small size.

     

Structure and occurrence of cyphonautes larvae (bryozoa,ectoprocta)

We have studied larvae of the freshwater ctenostome Hislopia malayensis with scanning electron microscopy (SEM), confocal laser scanning microscopy (CLSM), and LM of serial sections. Some additional observations on larvae of M. membranacea using SEM and CLSM are also reported. The overall configuration of muscles, nerves, and cilia of the two larvae are identical. However, the larva of H. malayensis is much smaller than that of M. membranacea, which may explain most of the differences observed. Although all major nerves and muscle strands are present in H. malayensis, they are generally composed of fewer fibers. The H. malayensis larva lacks the anterior and posterior intervalve cilia. Its pyriform organ is unciliated with only a small central depression. The adhesive epithelium is not invaginated as an adhesive sac and lacks the large muscles interpreted as adhesive sac muscles in the M. membranacea larva. The velum carries two rows of ciliated cells, though the lower “row” consists of only one or two cells. Both rows of ciliated cells are innervated by nerves, which have not been detected in the M. membranacea larva. The ciliated ridge of H. malayensis lacks the frontal cilia. The planktotrophic cyphonautes larvae in a number of ctenostome clades and in the “basal” cheilostome clade Malacostega (and probably in the earliest cheilostomes) support the idea that the cyphonautes larva is the ancestral larval type of the Eurystomata. It may even represent the ancestral larval type of the bryozoans (= ectoprocts). J. Morphol. 271:1094‐1109, 2010. © 2010 Wiley‐Liss, Inc.     

NOR-Chromosome morphology and evidence for rDNA selection in phylactolaemates

Silver-stained chromosome spreads from the phylactolaemate ectoprocts Fredericella indica, Fredericella browni, Plumatella repens, Pectinatella magnifica and Asajirella gelatinosa show that each of these species has a single pair of NORs. In the first four species each NOR is immediately adjacent to the centromere of a large metacentric chromosome. Giemsa-stained preparations from Cristatella mucedo show centromeric gapping in a similar large metacentric, indicating the same type of NOR-chromosome. In Asajirella gelatinosa the two NORs are on a pair of smaller chromosomes, and are not immediately adjacent to a centromere. A close NOR-centromere association would inhibit meiotic crossing over in the NOR region. Since in phylactolaemates reproduction is primarily asexual, often by formation of statoblasts, there is potential intraclonal selection for favorable ribosomal DNA (rDNA) mutations, including from simultaneous germination of otherwise genetically identical statoblasts within a limited area and competition between cell lines within statoblasts. Inhibition of meiotic crossing over within the NOR would maintain its integrity during periods when intraclonal selection would not be occurring. This suggests that a more rapid rate of rDNA evolution is possible in phylactolaemates than in most other organisms, and is consistent with a derivation of this group from the morphologically similar phoronids, despite phylogenetic interpretations of rDNA sequence data that suggest the two groups are not closely related.     

Ellobiophrya conviva n. sp., a Commensal of Marine Ectoprocts,and a Revision of the Family Ellobiophryidae (Ciliophora,Peritricha)1

The sessiline peritrich Ellobiophrya conviva n. sp. is described from marine ectoprocts of the genus Bugula, the first report of an ellobiophryid on bryozoan hosts. The new species is distinguished from others of its genus by its different body proportions, size, host, and structure of the clasping holdfast (for which the new name cinctum is chosen). Ellobiophrya conviva has been found only on B. neritina and B. turrita and shows a marked seasonal cycle of abundance. The family Ellobiophryidae Chatton & Lwoff is revised on the basis of new information provided by E. conviva, with the single species of the genus Clausophrya removed to Ellobiophrya as E. oblida Naidenova & Zaika n. comb. The genus Caliperia Laird remains unchanged. The two genera of the revised family are distinguished from one another by differences in the structure of the cinctum. Hypotheses are advanced to explain the morphogenesis of the cinctum and the evolution of ellobiophryids from other peritrichs.     

Morphology and the classification of the so-called Cladocera

The validity as a monophyletic taxon of the group of branchiopod crustaceans long regarded as constituting the Cladocera is questioned. This seems in fact to be a heterogeneous assemblage whose members probably merit assignment to four separate orders. The Onychopoda and Haplopoda (the so-called Gymnomera) clearly stand apart from the Ctenopoda and Anomopoda (the so-called Calyptomera) and differ in important respects from each other. While sharing several characters, some doubtless indicative of distant ancestral similarities, others probably convergent, the Ctenopoda and Anomopoda differ in many respects, have clearly evolved along different lines, and are probably much less closely related than is generally assumed.     

嵩草属植物硅酸体系统的研究

对国产狭义嵩草属 (K obresia,Cyperaceae) 3个组 2 7种植物进行了硅酸体系统分析研究 ,同时结合近缘的苔草属 (Carex) 2个组的植物硅酸体特征 ,深入探讨了植物硅酸体系统分析研究在莎草科苔草族 (Cariceae)中的系统分类学意义。认为 :(1 )嵩草属及其临近类群在属间、种间及种下水平的系统分类学比较研究都可以借助植物硅酸体系统研究的证据。 (2 )穗状嵩草组 (sect. Elyna)的硅酸体系统特征具有较高的一致性 ,除矮嵩草 (K.humilis)及其近缘种同其他穗状嵩草组的种类有所不同 ,它很可能有一个不同于穗状嵩草组其他类群的嵩草组(sect. K obresia)的亲缘。 (3 )从植物硅酸体系统特征的研究来看 ,拟苔草组 (sect. H emicarex)的一些种接近于穗状嵩草组 ,如高山嵩草 (K . pygmeae) ;而另一些种接近于嵩草组 ,如禾叶嵩草 (K . graminifolia) ;也有一些种类的硅酸体系统特征更接近苔草属 ,说明这个组很可能是多源的 ,即本组的单一花穗和果囊形态特征由不同类群的复合花序种类趋同演化而出。(4 )嵩草组的硅酸体系统特征具有较高的一致性 ,说明嵩草属原始类群在硅酸体系统特征方面与外部形态特征演化的某些相关性。     

Effect of age on water loss of leaves

The water loss of leaves of the same level of insertion but of various ages was examined. The results show that the transpiration of youngest leaves is higher than that of the older ones. The oldest leaves which have started yellowing show an increase of transpiration. This is probably related to the premortal changes of the leaf. The differences are probably caused by various physiological states of leaf because the leaf area was not different.     

Hereditary resistance to anthrax and sensitivity to the anthrax toxin in mice

The degree of the hereditary susceptibility of mice to anthrax caused by noncapsular and capsule-forming Bacillus anthracis strains has been found to be directly related to the sensitivity of the animals to the edematogenic and immunosuppressing action of anthrax toxin. The genetic analysis indicates that resistance to anthrax is probably controlled by a dominant gene, not linked with histocompatibility complex H-2 and, probably, unrelated to the presence of hemolytic activity in mouse sera, determined by component C5 of the complement.     

Relationships and origin of endemic Lake Baikal gastropods (Caenogastropoda: Rissooidea) based on mitochondrial DNA sequences

The phylogenetic relationships and the origin of two groups of rissooid freshwater snails endemic to Lake Baikal were investigated using partial mitochondrial COI, 12S rDNA, and 16S rDNA sequences. The Baikalian Benedictiinae proved to be closely related to the Lithoglyphinae. According to a molecular clock estimate the two groups diverged in the Paleogene. The Benedictiinae might have evolved autochthonously in precursors of Lake Baikal. The Baikalian Baicaliidae are probably most closely related to the Amnicolidae and the Bithyniidae. These groups diverged at the latest during the Cretaceous. Thus the origin of the Baicaliidae predates the origin of the Baikal rift zone. The Baicaliidae evolved probably in other Central Asian freshwater reservoirs. However, the radiation of the extant Baicaliidae only started in the Neogene and might have occurred autochthonously in Lake Baikal. The conchological similarity of the Baicaliidae and the Pyrgulidae is due to convergence. The Pyrgulidae diverged from the common stem lineage of the other hydrobiid families at the latest in the Jurassic. The Bithyniidae is derived from hydrobiids and is related to the Amnicolidae.     

Phytohormones in soil after atrazine application

The herbicide atrazine influences the content of phytohormones (auxins, gibberellins, cytokinins) in maize-cultivated soils. The effect is probably related to variations in the metabolism of hormones in soil microorganisms.     

Circulating Rheumatoid and Antinuclear Factors in Asbestos Workers

In an unselected series of 80 patients with asbestos exposure referred to the London Pneumoconiosis Medical Panel, antinuclear and rheumatoid factors were found in 28 and 27% respectively. This probably represents at least a fourfold increase over their incidence in random populations. The presence of these autoantibodies was not related to the duration of exposure, but was related to the extent of radiographic abnormality.