Phylactolaemate Bryozoans Research Articles

Genomic and transcriptomic survey of bryozoan Hox and ParaHox genes with emphasis on phylactolaemate bryozoans

BackgroundBryozoans are mostly sessile aquatic colonial invertebrates belonging to the clade Lophotrochozoa, which unites many protostome bilaterian phyla such as molluscs, annelids and brachiopods. While Hox and ParaHox genes have been extensively studied in various lophotrochozoan lineages, investigations on Hox and ParaHox gene complements in bryozoans are scarce.ResultsHerein, we present the most comprehensive survey of Hox and ParaHox gene complements in bryozoans using four genomes and 35 transcriptomes representing all bryozoan clades: Cheilostomata, Ctenostomata, Cyclostomata and Phylactolaemata. Using similarity searches, phylogenetic analyses and detailed manual curation, we have identified five Hox genes in bryozoans (pb, Dfd, Lox5, Lox4 and Post2) and one ParaHox gene (Cdx). Interestingly, we observed lineage-specific duplication of certain Hox and ParaHox genes (Dfd, Lox5 and Cdx) in some bryozoan lineages.ConclusionsThe bryozoan Hox cluster does not retain the ancestral lophotrochozoan condition but appears relatively simple (includes only five genes) and broken into two genomic regions, characterized by the loss and duplication of serval genes. Importantly, bryozoans share the lack of two Hox genes (Post1 and Scr) with their proposed sister-taxon, Phoronida, which suggests that those genes were missing in the most common ancestor of bryozoans and phoronids.

Phylogenomics reveals deep relationships and diversification within phylactolaemate bryozoans.

Bryozoans are mostly sessile colonial invertebrates that inhabit all kinds of aquatic ecosystems. Extant bryozoan species fall into two clades with one of them, Phylactolaemata, being the only exclusively freshwater clade. Phylogenetic relationships within the class Phylactolaemata have long been controversial owing to their limited distinguishable characteristics that reflect evolutionary relationships. Here, we present the first phylogenomic analysis of Phylactolaemata using transcriptomic data combined with dense taxon sampling of six families to better resolve the interrelationships and to estimate divergence time. Using maximum-likelihood and Bayesian inference approaches, we recovered a robust phylogeny for Phylactolaemata in which the interfamilial relationships are fully resolved. We show Stephanellidae is the sister taxon of all other phylactolaemates and confirm that Lophopodidae represents the second offshoot within the phylactolaemate tree. <i>Plumatella fruticosa</i> clearly falls outside Plumatellidae as previous investigations have suggested, and instead clusters with Pectinatellidae and Cristatellidae as the sister taxon of Fredericellidae. Our results demonstrate that cryptic speciation is very likely in <i>F. sultana</i> and in two species of <i>Plumatella</i> (<i>P. repens</i> and <i>P. casmiana</i>). Divergence time estimates show that Phylactolaemata appeared at the end of the Ediacaran and started to diverge in the Silurian, although confidence intervals were large for most nodes. The radiation of most extant phylactolaemate families occurred mainly in the Palaeogene and Neogene highlighting post-extinction diversification.

Reconstructing the neuromuscular ground pattern of phylactolaemate bryozoans: new data from the Lophopodidae

BackgroundThe solely freshwater inhabiting Phylactolaemata is a sister taxon to all other bryozoans. Among phylactolaemates, Lophopodidae represents an early branching clade that is therefore crucial for ground pattern reconstruction. While more recent morphological data of most phylactolaemate families are present, data of lophopodids are scarce. The genus Asajirella especially, which was previously assigned to the family Pectinatellidae, lacks any detailed analysis with more recent morphological methods.ResultsThis study provides the first morphological analyses of three lophopodid species using serial-sectioning histology and 3D reconstruction, but also immunocytochemical stainings and confocal laserscanning microscopy. There are several lophopodid-specific traits in the nervous system such as the large ganglion with extensive lumen and two prominent protrusions referred to as epistomial horns. The epistome in all lophopodids is rather small and dome-shaped. Contrary to previous reports, we can confirm that duplicature bands insert at the tentacle sheath rather than the diaphragmatic sphincter in all phylactolaemates. The morphology of the digestive tract of lophopodids is identical to other phylactolaemates and possesses exclusively circular muscles.ConclusionsAltogether, this study fills significant gaps in our knowledge on phylactolaemate neuromuscular systems and general morphology. It shows that the insertion of the duplicature bands at the tentacle sheath and the circular musculature of the digestive tract to be the ground pattern in phylactolaemates. In addition, we found apomorphic characters for lophopodids such as the dome-shaped epistome with its musculature and the voluminous ganglion with its epistomial horns, which aid in defining and delineating the family.

Wastewater Treatment Efficiency by a Freshwater Phylactolaemate Bryozoan and Experimental Feeding with Protozoa

The wastewater treatment ponds of the King’s Royally Initiated Laem Phak Bia Environmental Research and Development (LERD) Project in west-central Thailand provide habitats for freshwater bryozoans, which are colonial invertebrate animals. Bryozoans sieve food particles out of the water using a retractable lophophore and can play an important ecological role in wastewater treatment. In this unique environment, we: (1) investigated the efficiency of a phylactolaemate bryozoan (Plumatella casmiana) in wastewater treatment, measured by BOD5, chlorophyll a and turbidity; and (2) determined the role of protozoans in the diet of the bryozoan P. casmiana. Comparison of growth rate and fecal pellet characteristics between protozoan-fed bryozoans and phytoplankton-fed bryozoans was investigated. At the end of our wastewater treatment experiment, water quality parameters were markedly improved in the treatment with bryozoans compared to the control (without bryozoans). The treatment efficiency levels for BOD5, turbidity, and chlorophyll a were 24.04%, 59.21%, and 55.13%, respectively. The growth rates of bryozoans in the experimental treatment increased over time. Our study also revealed that this bryozoan can feed on a diet of protozoans under experimental conditions. However, the average daily growth rate of protozoan-fed bryozoans -20 zooids per day was lower than that of phytoplankton-fed bryozoans 19 zooids per day. This may have been due to incomplete digestion of protozoans or insufficient nutrition in the bryozoans. The results from this study provide better understanding of bryozoan ecology and their role in wastewater treatment systems.

Reproductive biology, embryonic development and matrotrophy in the phylactolaemate bryozoan Plumatella casmiana

Bryozoa is a phylum of aquatic, colonial suspension-feeders within the Lophotrochozoa. In the Phylactolaemata embryonic development occurs in an internal brood sac on the body wall accompanied by extraembryonic nutrition. Owing to previous contradictive descriptions, many aspects of their sexual reproduction require restudy. Consequently, this study analyses embryogenesis of the freshwater bryozoan Plumatella casmiana by serial sections, 3D reconstruction and transmission electron microscopy. Early embryos cleave and soon develop into blastulae with a small central cavity. The mesoderm forms by delamination starting from the distal side towards the proximal end. In later embryos two polypides form on the posterior side that ultimately will be covered by a ciliated mantle in the larva. Embryos increase in size during development and form temporary cell contacts to the embryo sac. Mesodermal cells of the embryo sac show signs of transcellular transport indicating that embryos are nourished by transferring nutrients from the maternal coelom towards the brood cavity. This study clarifies several details such as mesoderm formation and the onset of bud development. Embryos are connected to their respective embryo sacs by a variety of temporary cytoplasmic processes formed by both tissues during embryogenesis, including a ‘placental’ ring zone. Although ultrastructural data of these cell contacts are not entirely conclusive about their function, we suggest that embryos absorb nutrients via the entire surface. The close opposition of embryos to the embryo sac implies placentation as matrotrophic mode in phylactolaemate bryozoans, with embryo sacs acting as placental analogues.

Mitochondrial gene order of the freshwater bryozoan Cristatella mucedo retains ancestral lophotrochozoan features

Bryozoans are aquatic colonial suspension-feeders abundant in many marine and freshwater benthic communities. At the same time, the phylum is under studied on both morphological and molecular levels, and its position on the metazoan tree of life is still disputed. Bryozoa include the exclusively marine Stenolaemata, predominantly marine Gymnolaemata and exclusively freshwater Phylactolaemata. Here we report the mitochondrial genome of the phylactolaemate bryozoan Cristatella mucedo. This species has the largest (21,008 bp) of all currently known bryozoan mitogenomes, containing a typical metazoan gene compendium as well as a number of non-coding regions, three of which are longer than 1500 bp. The trnS1/trnG/nad3 region is presumably duplicated in this species. Comparative analysis of the gene order in C. mucedo and another phylactolaemate bryozoan, Pectinatella magnifica, confirmed their close relationships, and revealed a stronger similarity to mitogenomes of phoronids and other lophotrochozoan species than to marine bryozoans, indicating the ancestral nature of their gene arrangement. We suggest that the ancestral gene order underwent substantial changes in different bryozoan cladesshowing mosaic distribution of conservative gene blocks regardless of their phylogenetic position. Altogether, our results support the early divergence of Phylactolaemata from the rest of Bryozoa.

Morphology and ontogeny of Lophopus crystallinus lophophore support the epistome as ancestral character of phylactolaemate bryozoans

Phylactolaemate bryozoans are the sister-group to all remaining bryozoan taxa. Consequently, their study is essential to reveal and analyze ancestral traits of Phylactolaemata and Bryozoa in general. They are the only bryozoans to possess an epistome which traditionally has been regarded as shared with phoronids and brachiopods. Contrary to older observations, an epistome was recently reported to be missing in the early branching phylactolaemate Lophopus crystallinus. In this study, the ontogeny of the lophophoral base and also its three-dimensional structure in adult specimens was reinvestigated to assess whether an epistome is never formed during ontogeny and absent in adult specimens. The results show that organogenesis during the budding process in this species is similar to other, previously investigated, species. The epistome anlage in L. crystallinus forms in early buds from the outer budding layer which penetrates the two shanks of the u-shaped gut. This ingression of the epithelium further proceeds distally and starts to wrap over the forming ganglion. The adult epistome is a rather short, but present bulge above the cerebral ganglion with prominent muscle bundles traversing its cavity. Distally it is arched by the forked canal that in L. crystallinus has a particularly thick and prominent epithelium in the three median tentacles. This study shows that neither during ontogeny nor in the adult stage an epistome is absent. The epistome is less pronounced than in other phylactolaemates, but otherwise similar in its general structure. Consequently, an epistome can be assumed to be present in the ground pattern of Phylactolaemata.

Reconstructing the muscular ground pattern of phylactolaemate bryozoans: first data from gelatinous representatives

BackgroundPhylactolaemata is commonly regarded the earliest branch within Bryozoa and thus the sister group to the other bryozoan taxa, Cyclostomata and Gymnolaemata. Therefore, the taxon is important for the reconstruction of the bryozoan morphological ground pattern. In this study the myoanatomy of Pectinatella magnifica, Cristatella mucedo and Hyalinella punctata was analysed by means of histology, f-actin staining and confocal laser-scanning microscopy in order to fill gaps in knowledge concerning the myoanatomy of Phylactolaemata.ResultsThe retractor muscles and muscles of the aperture, gut, body wall, tentacle sheath, lophophore constitute the most prominent muscular subsets in these species. The lophophore shows longitudinal muscle bands in the tentacles, lophophoral arm muscles, epistome musculature and hitherto undescribed muscles of the ring canal. In general the muscular system of the three species is very similar with differences mainly in the body wall, tentacle sheath and epistome. The body wall contains an orthogonal grid of musculature. The epistome exhibits either a muscular meshwork in the epistomal wall or muscle fibers traversing the epistomal cavity. The whole tentacle sheath possesses a regular mesh of muscles in Pectinatella and Cristatella, whereas circular muscles are limited to the tentacle sheath base in Hyalinella.ConclusionThis study is the first to describe muscles of the ring canal and contributes to reconstructing muscular features for the last common ancestor of all bryozoans. The data available suggest that two longitudinal muscle bands in the tentacles, as well as retractor muscles and longitudinal and circular muscles in the tentacle sheath, were present in the last common bryozoan ancestor. Comparisons among bryozoans shows that several apomorphies are present in the myoanatomy of each class- level taxon such as the epistomal musculature and musculature of the lophophoral arms in phylactolaemates, annular muscles in cyclostomes and parietal muscles in gymnolaemates.

Comparative morphology of the nervous system in three phylactolaemate bryozoans.

BackgroundThough some elements of the bryozoan nervous system were discovered 180 years ago, few studies of their neuromorphology have been undertaken since that time. As a result the general picture of the bryozoan nervous system structure is incomplete in respect of details and fragmentary in respect of taxonomic coverage.ResultsThe nervous system of three common European freshwater bryozoans – Cristatella mucedo, Plumatella repens (both with a horseshoe-shaped lophophore) and Fredericella sultana (with a circular lophophore) had numerous differences in the details of the structure but the general neuroarchitecture is similar. The nervous system of the zooid consists of the cerebral ganglion, a circumpharyngeal ring and lophophoral nerve tracts (horns), both sending numerous nerves to the tentacles, and the nerve plexuses of the body wall and of the gut. A number of the important details (distal branching of the additional radial nerve, pattern of distribution of nerve cells and neurites in the ganglion, etc.) were described for the first time. The number and position of the tentacle nerves in Cristatella mucedo was ascertained and suggestions about their function were made. The revealed distribution of various neuromediators in the nervous system allowed us to suggest functional affinities of some major nerves.ConclusionsDespite the basic similarity, both the ganglion and the lophophore nervous system in Phylactolaemata have a more complex structure than in marine bryozoans (classes Gymnolaemata and Stenolaemata). First of all, their neuronal network has a denser and more complex branching pattern: most phylactolaemates have two large nerve tracts associated with lophophore arms, they have more nerves in the tentacles, additional and basal branches emitting from the main radial nerves, etc. This, in part, can be explained by the horseshoe shape of the lophophore and a larger size of the polypide in freshwater species. The structure of the nervous system in Fredericella sultana suggests that it underwent a secondary simplification following the reduction of the lophophore arms. Colony locomotion in Cristatella mucedo is based on co-ordinated activity of two perpendicular muscle layers of the sole and the plexus of motor neurons sandwiched between them. The trigger of this activity and the co-ordination mechanism remain enigmatic.Electronic supplementary materialThe online version of this article (doi:10.1186/s12983-015-0112-2) contains supplementary material, which is available to authorized users.

Freshwater Bryozoa of Okinawa, Japan, with descriptions of Rumarcanella gen. nov. (Phylactolaemata: Plumatellidae) and two new species

We identified nine species of phylactolaemate bryozoans collected from ponds and dams on Okinawa Island, Japan. This is the first report on freshwater bryozoans from Okinawa and includes the first record of Plumatella javanica from Japan. Phylactolaemate species were identified on the basis of colony and statoblast morphology and are illustrated by light microscopy and scanning electron microscopy (SEM). We detected two new plumatellid species that, like Plumatella minuta and Plumatella vorstmani, have a transparent, weakly chitinized colony and hypertubercles on the floatoblast fenestra. A previous molecular study showed these four species to form a clade separate from Plumatella and Hyalinella. We here establish the new genus Rumarcanella to accommodate these species and describe Rumarcanella gusuku n. sp. and R. yanbaruensis n. sp. We discuss the dispersal of bryozoans to and from Japan by birds carrying statoblasts and provide a key to the Phylactolaemata of Japan.

Organogenesis during budding and lophophoral morphology of Hislopia malayensis Annandale, 1916 (Bryozoa, Ctenostomata)

BackgroundBryozoans represent a large lophotrochozoan phylum with controversially discussed phylogenetic position and in group relationships. Developmental processes during the budding of bryozoans are in need for revision. Just recently a study on a phylactolaemate bryozoan gave a comprehensive basis for further comparisons among bryozoans. The aim of this study is to gain more insight into developmental patterns during polypide formation in the budding process of bryozoans. Particular focus is laid upon the lophophore, also its condition in adults. For this purpose we studied organogenesis during budding and lophophoral morphology of the ctenostome bryozoan Hislopia malayensis.ResultsPolypide buds develop on the frontal side of the developing cystid as proliferation of the epidermal and peritoneal layer. Early buds develop a lumen bordered by the inner budding layer resulting in the shape of a two-layered sac or vesicle. The hind- and midgut anlagen are first to develop as outpocketing of the prospective anal area. These grow towards the prospective mouth area where a comparatively small invagination marks the formation of the foregut. In between the prospective mouth and anus the ganglion develops as an invagination protruding in between the developing gut loop. Lophophore development starts with two lateral ridges which form tentacles very early. At the lophophoral base, intertentacular pits, previously unknown for ctenostomes, develop. The ganglion develops a circum-oral nerve ring from which the tentacle nerves branch off in adult zooids. Tentacles are innervated by medio-frontal nerves arising directly from the nerve ring, and medio-frontal and abfrontal nerves which originate both from an intertentacular fork.ConclusionsWe are able to show distinct similarities among bryozoans in the formation of the different organ systems: a two-layered vesicle-like early bud, the ganglion forming as an invagination of the epidermal layer in between the prospective mouth and anal area, the digestive tract mainly forming as an outpocketing of the prospective anal area, and the lophophore forming from two lateral anlagen that first fuse on the oral and afterwards on the anal side. Future studies will concentrate on cyclostome budding to complement our knowledge on developmental patterns of bryozoans.

Particle capture in ciliary filter‐feeding gymnolaemate and phylactolaemate bryozoans – a comparative study

AbstractRiisgård, H.U., Okamura, B. and Funch, P. 2009. Particle capture in ciliary filter‐feeding gymnolaemate and phylactolaemate bryozoans – a comparative study. —Acta Zoologica (Stockholm) 91: 416–425.We studied particle capture using video‐microscopy in two gymnolaemates, the marine cheilostome Electra pilosa and the freshwater ctenostome Paludicella articulata, and three phylactolaemates, Fredericella sultana with a circular funnel‐shaped lophophore, and Cristatella mucedo and Lophophus crystallinus, both with a horseshoe‐shaped lophophore. The video‐microscope observations along with studies of lophophore morphology and ultrastructure indicated that phylactolaemate and gymnolaemate bryozoans with a diversity of lophophore shapes rely on the same basic structures and mechanisms for particle capture. Our study also demonstrates that essential features of the particle capture process resemble one another in bryozoans, brachiopods and phoronids.

Development of Tetracapsuloides bryosalmonae (Myxozoa: Malacosporea) in bryozoan hosts (as examined by light microscopy) and quantitation of infective dose to rainbow trout ( Oncorhynchus mykiss)

The myxozoan parasite Tetracapsuloides bryosalmonae is the causative agent of proliferative kidney disease (PKD), a highly damaging disease of cultured salmonid fish. Within this study, phylactolaemate bryozoans were collected from a river known to be endemic for PKD and subsequently cultured in the laboratory. Sequential developmental stages of T. bryosalmonae were studied by light microscopy within the living bryozoan colonies, allowing the identification of stages attached to host peritoneum, consistent with previous molecular evidence of cryptic stages. Infection resulted in the production of large numbers of spores, which were released from the bryozoans. Experimental exposure of rainbow trout ( Oncorhynchus mykiss) to medium in which infected bryozoans were cultured resulted in clinical PKD. Rainbow trout were exposed to known numbers of T. bryosalmonae spores collected by micromanipulation, which had been released from mature spore sacs within colonies of the bryozoan Fredericella sultana. Exposure to one spore was sufficient to lead to development of PKD. These findings indicate that small numbers of bryozoans are capable of releasing sufficient spores to infect large numbers of fish, having implications for future control methods for PKD in salmonid farming.

Ciliary feeding structures and particle capture mechanism in the freshwater bryozoan Plumatella repens (Phylactolaemata)

Abstract. In contrast to marine bryozoans, the lophophore structure and the ciliary filter‐feeding mechanism in freshwater bryozoans have so far been only poorly described. Specimens of the phylactolaemate bryozoan Plumatella repens were studied to clarify the tentacular ciliary structures and the particle capture mechanism. Scanning electron microscopy revealed that the tentacles of the lophophore have a frontal band of densely packed cilia, and on each side a zigzag row of laterofrontal cilia and a band of lateral cilia. Phalloidin‐linked fluorescent dye showed no sign of muscular tissue within the tentacles. Video microscopy was used to describe basic characteristics of particle capture. Suspended particles in the incoming water flow, set up by the lateral ‘pump’ cilia on the tentacles, approach the tentacles with a velocity of 1–2 mm s‐1. Near the tentacles, the particles are stopped by the stiff sensory laterofrontal cilia acting as a mechanical sieve, as previously seen in marine bryozoans. The particle capture mechanism suggested is based on the assumed ability of the sensory stiff laterofrontal cilia to be triggered by the deflection caused by the drag force of the through‐flowing water on a captured food particle. Thus, when a particle is stopped by the laterofrontal cilia, the otherwise stiff cilia are presumably triggered to make an inward flick which brings the restrained particle back into the downward directed main current, possibly to be captured again further down in the lophophore before being carried to the mouth via the food groove. No tentacle flicks and no transport of captured particles on the frontal side of the tentacles were observed. The velocity of the metachronal wave of the water‐pumping lateral cilia was measured to be ∼0.2 mm s‐1, the wavelength was ∼7 μm, and hence the ciliary beat frequency estimated to be ∼30 Hz (∼20 °C). The filter feeding process in P. repens reported here resembles the ciliary sieving process described for marine bryozoans in recent years, although no tentacle flicks were observed in P. repens. The phylogenetic position of the phylactolaemates is discussed in the light of these findings.

Plumatella geimermassardi, a newly recognized freshwater bryozoan from Britain, Ireland, and continental Europe (Bryozoa: Phylactolaemata)

Plumatella geimermassardi is a newly recognized species of phylactolaemate bryozoan. Its known range extends from Ireland east through southern Norway and south into Italy. Colonies grow close to the substrate with little free branching; the body wall is mostly transparent and without an obvious raphe. Floatoblasts are broadly oval and relatively small, with distinctively large dorsal fenestra and uniformly narrow ventral annulus. The sessoblast basal valve is low and dish-shaped; the annulus bears tubercles which vary in their prominence. This species brings to 14 the number of phylactolaemate bryozoans known in the region.

A New Class and Order of Myxozoans to Accommodate Parasites of Bryozoans with Ultrastructural Observations on Tetracapsula bryosalmonae (PKX Organism)

Tetracapsula bryosalmonae, formerly PKX organism, is a myxozoan parasite that causes proliferative kidney disease in salmonid fish. Its primary hosts, in which it undergoes a sexual phase, are phylactolaemate bryozoans. It develops in the bryozoan coelomic cavity as freely floating sacs which contain two types of cells, stellate cells and sporoplasmogenic cells, which become organised as spores. Eight stellate cells differentiate as four capsulogenic cells and four valve cells which surround a single sporoplasmogenic cell. The sporoplasmogenic cell undergoes meiosis and cytoplasmic fission to produce two sporoplasms with haploid nuclei. Sporoplasms contain secondary cells. The unusual development supports previously obtained data from 18S rDNA sequences, indicating that species of Tetracapsula form a clade. It diverged early in the evolution of the Myxozoa, before the radiation that gave rise to the better known genera belonging to the two orders in the single class Myxosporea. The genus Tetracapsula as seen in bryozoans shares some of the characters unique to the myxosporean phase and others typical of the actinosporean phase of genera belonging to the class Myxosporea. However, it exhibits other features which are not found in either phase. A new class Malacosporea and order Malacovalvulida are proposed to accommodate the family Saccosporidae and genus Tetracapsula. Special features of the new class are the sac-like proliferative body, valve cells not covering the exit point of the polar filament, lack of a stopper-like structure sealing the exit, maintenance of valve cell integrity even at spore maturity, absence of hardened spore walls and unique structure of sporoplasmosomes in the sporoplasms.

Biofouling of wastewater treatment plants by the freshwater bryozoan, Plumatella vaihiriae (Hastings, 1929)

We report large growths of a phylactolaemate bryozoan, Plumatella vaihiriae [Hastings, A. (1929) Notes on some little-known phylactolaematous polyzoa and description of a new species from Tahiti. Ann. Mag. Nat. Hist., 10(3), 300–311], from three widely separated domestic wastewater treatment plants in the United States. This relatively uncommon species is previously known only from Tahiti, Hawaii, Utah, and Argentina. The bryozoan colonies are capable of plugging pipes and filters. Free statoblasts are suspected of interfering with ultraviolet disinfection of wastewater. Laboratory reared specimens exhibit rapid, aggressive growth and produce vast quantities of free statoblasts which either float or sink. Devising effective control methods is impeded by limited knowledge of bryozoan physiology. Given the widespread distribution of known wastewater treatment facilities affected by the bryozoan, the problem is probably a common one.

Plumatella nitens, a new species of freshwater bryozoan from North America (Ectoprocta: Phylactolaemata), previously misidentified

Plumatella nitens is a newly recognized species of phylactolaemate bryozoan in North America. It has previously been combined with either P. repens or P. fungosa, from which it differs primarily by the floatoblast. In P. nitens the ventral floatoblast annulus is uniformly narrow with no significant widening at the poles. The dorsal floatoblast fenestra is conspicuously larger than in either P. repens or P. fungosa. Reticulated ridges on the dorsal fenestra are weak along the margins, becoming unusually prominent in the center. The sessoblast is densely covered with uneven papillae. Spotty sampling so far shows a distribution only in the northern half of North America, including Massachusetts, Michigan, Minnesota, Wisconsin, Ontario, and the northern regions of Illinois, Indiana, and Ohio.

Asynchronous and density-dependent germination: the spreading of risk in Plumatella emarginata.

I examined both temporal and spatial aspects of the germination of asexually produced sessoblasts of the phylactolaemate bryozoan Plumatella emarginata. The majority of sessoblasts germinated over a 5-week period from April to May; low frequencies of germination were observed from June to September. During May and June high sessoblast density further enhanced asynchrony. Risk spreading in response to both unpredictable disturbances and high sessoblast density may explain asynchronous germination in this system.

Chromosomal heteromorphism in a Japanese population of Pectinatella gelatinosa and karyotypic comparison with some other phylactolaemate bryozoans

Five different karyotypes were present in germinating statoblasts obtained from a sampling of a predominately or exclusively asexually reproducing population of Pectinatella gelatinosa from Lake Tatara-numa, Japan. Karyotypes were consistent in having 18 chromosomes, a unique telocentric, a pair of minute metacentrics, and at least one prominently secondarily constricted element. In a number of cases, a sampling of statoblasts from a single large colony yielded more than one karyotype, perhaps because there had been fusion of smaller colonies with different karyotypes. Some individual statoblasts may have been chimeric, but this was not conclusively demonstrated. The possibility of asexually-maintained permanent chimerism in phylactolacmates involving karyotypically distinct ectodermal and peritoneal cell lines is discussed. Karyotypically, P. gelatinosa is more similar to Lophopodella carteri than to Pectinatella magnifica.