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The British species of Bugula (Polyzoa)
Abstract
Eleven species of Bugula are known to occur in British waters, including three that have not previously been described. The characters of greatest value for separating these species are reviewed. Full descriptions are given of B. neritina, B. plumosa, B. purpurotincta, B. stolonifera sp. nov., B. avicularia, B. turbinata, B. flabellata, B. fulva sp. nov., B. calathus, B. aquilirostris sp. nov. and B. simplex. The account includes notes on habitat, breeding season and geographical distribution; and a key is provided to facilitate identification.
B. ditrupae, from Madeira, is redescribed and compared with B. fulva and B. aquilirostris, as the three species have hitherto been confused. Synonomies of each are given.
The B. simplex of most authors is not that of Hincks, but another species: possibly of form konwn as B. plumosa var. aperta. B. simplex is the B. flabellata of most American authors, and the B. sabatieri of those writing on Mediterranean Polyzoa. Synonomies of B. simplex and B. flabellata are given.
A full description of B. turrita and summary descriptions of B. californica, B. pedata and B. geermanae are given, and these species are compared with B. stolonifera. The characters of B. spicata are mentioned as it also has been confused with some of the above species.
... Autozooids fix by rhizoids which issue from frontal, lateral and basal surfaces. Colonies comprise two or more series of boat-shaped zooids (Ryland 1960). The type species, Bugula neritina, shows remarkable exception from other Bugulidae species in which bird's head avicularia are a common feature. ...
... It is a fouling species, but its distribution is not restricted to ports and harbours. Its distribution ranged from shallow water to depth of 70 m (Ryland 1960;Ryland and Hayward 1991). B. fulva was found on natural (attached on another bryozoan, Pentapora fascialis) (Novosel et al. 2004) and artificial substrates (Ramalhosa et al. 2017), and it was reported from brackish lagoons and location close to hydrothermal vent system (Occhipinti Ambrogi 1981;Morri et al. 1999). ...
... B. fulva is probably of exotic in origin (Hayward and McKinney 2002;Ryland et al. 2011) and it was reported as established for western Mediterranean and casual for Central, Eastern Mediterranean and Adriatic (Zenetos et al. 2010). Table 1) Bugula simplex Hincks, 1886: Ryland 1960Prenant and Bobin 1966: 505, Fig. 168;Zabala and Maluquer 1988: 104, Fig. 201-203;Ryland and Hayward 1991: 33, Fig.48;Hayward and Ryland 1998: 226, Fig. 71. Bugulina simplex (Hincks, 1886): Fehlauer-Ale et al. 2015; 6. Material Examined: Colonies collected from just one locality, fishing ports in Mordoğan (S4). ...
Along the Turkish coast, Bugulidae family represented with tree valid genera, Bugula, Bugulina and Crisularia, contains mostly common fouling species in shallow and deep water. Five species belonging to two genera (Bugula neritina, Bugulina flabellata, Bugulina simplex, Bugulina stolonifera, Bugulina fulva) were investigated during different sampling periods. This study provides descriptions, illustrations, and information about ecological characteristics and biogeographical distributions of the above mentioned species. Bugulina fulva was reported for the first time from the Eastern Mediterranean coast of Turkey (Marmaris).
... Recently, an extensive revision using molecular and morphological techniques was performed for Bugula sensu lato resulting in four valid genera, Bugula sensu stricto, Bugulina [44], Crisularia [44] and Virididentula [42]. Several of these species, such as Bugula neritina [45], Virididentula dentata [46] and Bugulina stolonifera [47], have a widespread distribution and are abundant among fouling communities [40,[48][49][50]. ...
... The present study aims to review a complete list of the Bugulidae species colonizing artificial substrates in the Madeira archipelago. In this paper, we include descriptions of seven Bugulidae species with Scanning Electron Microscopy (SEM) illustrations, including four new records: Bugulina fulva [47] and Bugulina simplex [53] for the Madeira Island and Bugula neritina and Crisularia gracilis for the neighbouring Porto Santo Island. Furthermore, we report here the correction of the previous identification of Bugulina calathus minor earlier reported from Funchal harbour in 1998, now as Bugulina flabellata [44] for the Madeira Island. ...
... It presents the beak of the avicularia hooked almost rectangularly, as it happens in our specimens (can also see in Alves and Cocito Fig. 1a, note that it is not possible to see in the Fig. 1b because the position of the avicularia), been the beak of the avicularia smoothly down-curve in B. calathus [e.g. 47,62]. Beside, in the avicularia of the specimens from Alves and Cocito [36] have the edge from the peduncle to the mandibular proximal border shorter and more curve than in the avicularia of Bugula calathus [e.g. ...
Until very recently the Madeira Archipelago registered a total of eight Bugulidae species. In the present study we include descriptions of seven Bugulidae species, now with Scanning Electron Microscopy images, with four new records for the Archipelago: Bugulina fulva and Bugulina simplex for Madeira Island and Bugula neritina and Crisularia gracilis for the neighbouring Island of Porto Santo. Furthermore, we report the correction of the previous identification of Bugulina calathus minor earlier reported from Funchal harbour in 1998, now as Bugulina flabellata. This study is part of an ongoing monitoring program for detecting non-indigenous species in all marinas from the Madeira archipelago, between 2007 and 2015. Specimens were collected in previously deployed PVC settling plates, marina pontoons, and also on recreational hull vessels while performing dry dock inspections at a local shipyard. Our study reveals that the Madeira archipelago now registers a total of ten Bugulidae species, contributing therefore to the total bryozoan fauna of the Archipelago, now with more than 100 records. These numbers could increase, as Madeira is considered to be a “hotspot” for bryozoan diversity when compared to other nearby regions. Finally, hull fouling is considered as the most likely vector of introduction for the non-indigenous species of Bugulidae detected in Madeira.
http://link.springer.com/article/10.1186/s10152-016-0465-8
... In Table 10 of the report, four species of bryozoans were listed as 'Doorknockers'; i.e., species that were expected to arrive but had not yet been recorded. These species were: Watersipora subtorquata (d'Orbigny,1852); Tricellaria inopinata d' Hondt and Occhipinti Ambrogi, 1985; Bugula neritina (Linnaeus, 1758); and Bugula stolonifera Ryland, 1960. From 29 June to 25 July 2014, scientists from Heriot Watt University and the Natural History Museum, London, undertook a research cruise aboard the vessel MV Halton. ...
... The marinas surveyed along the Norwegian coast fell into three categories: i) those with freshwater input and no bryozoans present; ii) fully saline with native or naturalized fouling species; or iii) fully saline with a mixture of native or naturalized fouling species and invasive nonnative species (Figure 1). The species is also present in the Eel Pond at Woods Hole, New England (Ryland 1960). Recently (September 2014), Bugulina simplex has been collected by Rachel Shucksmith, in Lerwick Harbour, Shetland (NHMUK 2014.11.1.12). ...
... Bugulina stolonifera (Ryland, 1960) (Figure 5) The distribution and historical records of B. stolonifera are reviewed in Ryland et al. (2011), and the status of the species in Western Europe is unclear. Its distribution appears to be very patchy and one of the more recent records is from the Outer Harbour, Harwich (Ashelby 2005). ...
From 29 June to 25 July 2014, scientists from Heriot Watt University and the Natural History Museum, London, aboard the vessel MV Halton, undertook a research cruise along the Norwegian coast. The cruise started in Bergen and over the two-week period surveys were conducted at intervals along the coastline, heading northwards, and including the major ports of Ålesund, Kristiansund, and Trondheim. When the vessel moored up in each harbour, surveys of the local pontoons were conducted to identify fouling species and to ascertain whether any non-native Bryozoa were present. Seven species of fouling Bryozoa were identified. Two of these were the non-native species Tricellaria inopinata d'Hondt and Occhipinti Ambrogi, 1985 and Schizoporella japonica Ortmann, 1890. This study represents the first confirmed records for these species from Norwegian coastal waters. It is likely, given the locations of occurrence, that small boat traffic was a vector for the introduction of these species. Recommendations are given for the amendment and addition of species to the Norwegian Black List.
... In this paper we update the known distributions in Britain and Ireland of non-native bryozoans of the genus Bugula Oken, 1815, especially the warm-water species B. neritina (Linnaeus, 1758). Two other species of Bugula that appear clearly to be non-native in Western Europe are B. simplex Hincks, 1886, and B. stolonifera Ryland, 1960, although no-one in Britain has been recording them as such. Neither species is difficult to identify and full descriptions are available (Ryland 1960;Prenant and Bobin 1966;Zabala 1986Zabala , 1988Ryland 1990, 1998;Ryland and Hayward 1991;De Blauwe 2009); nevertheless, for convenience with recognition, we include photographs and scanning electron micrographs (SEMs) of all three species (Figures 1, 6 and 8). ...
... Two other species of Bugula that appear clearly to be non-native in Western Europe are B. simplex Hincks, 1886, and B. stolonifera Ryland, 1960, although no-one in Britain has been recording them as such. Neither species is difficult to identify and full descriptions are available (Ryland 1960;Prenant and Bobin 1966;Zabala 1986Zabala , 1988Ryland 1990, 1998;Ryland and Hayward 1991;De Blauwe 2009); nevertheless, for convenience with recognition, we include photographs and scanning electron micrographs (SEMs) of all three species (Figures 1, 6 and 8). It should be noted, however, that several native species of Bugula also occur in the region, some known to occur in fouling communities, and a key including all candidate species (e.g. ...
... Good descriptions, based on Mediterranean material, are available in several languages (Gautier 1962;Ryland 1965;Prenant and Bobin 1966;Riedl 1970;Occhipinti Ambrogi 1981;Zabala 1986;De Blauwe 2009). British-sourced specimens were used for the descriptions appearing in Ryland (1960), Ryland and Hayward (1977) and Ryland (1990, 1998), while Winston and Woollacott (2008) have recently provided a description, illustrated with SEMs, of material from Florida. Particularly notable in most freshly collected material is the purple coloration (Villela 1948), though not always as vivid as in Figure 1A and adverse environmental conditions may cause its loss. ...
Three apparently non-native species of Bugula occur in marinas and harbours in Atlantic Europe. The most common, B. neritina, was known from a few sites in southern Britain and northern France during the 20(th) century, following its discovery at Plymouth by 1911. During the 1950-60s it was abundant in a dock heated by power station effluent at Swansea, south Wales, where it flourished until the late 1960s, while water temperatures were 7-10 degrees C above ambient. It disappeared after power generation ceased, when summer temperatures probably became insufficient to support breeding. Details of disappearances have not been recorded but B. neritina was not seen in Britain between c1970 and 1999. Since 2000, it has been recorded along the south coast of England, and subsequently in marinas in the southern North Sea, Ireland and southern Scotland, well to the north of its former range, as well as along the Atlantic coast from Spain to The Netherlands. It has also been introduced to outlying localities such as the Azores and Tristan da Cunha. We report that this rapidly spreading form has the same COI haplotype as B. neritina currently invasive elsewhere in the world. B. simplex has been reported less, with 1950s records from settlement panels in some Welsh docks. It has not been targeted in most recent marina surveys but has been observed in southwest England, Belgium and The Netherlands. There are almost no recent records of B. stolonifera, though it was probably introduced to a few British and Irish ports prior to the 1950s. Its current status in most of western Europe is unknown but it has been reported as expanding throughout most of the world during the last 60 years. Having poorly known distributions, B. simplex and B. stolonifera should be recorded during future monitoring of alien species in Atlantic Europe. Illustrations to aid identification are included for all three species.
... Species in the genus Bugula are globally distributed, inhabiting shallow water in tropical to cold water regions (Kluge 1962;Hayward 1995;Hayward & Ryland 1998;Winston & Woollacott 2008;Vieira et al. 2012), with some representatives reported from deeper waters of the continental shelf and beyond (Hayward 1981). Some species are known from fouling communities and also may form wellestablished introduced populations (Mackie et al. 2006;Johnson & Woollacott 2012;Karlson & Osman 2012;Fehlauer-Ale et al. 2014); others are apparently endemic to particular regions (Ryland 1960;Winston & Woollacott 2008;Vieira et al. 2012). ...
... The genus has been characterized by bird's head pedunculate avicularia, erect colonies and branches comprising two or more series of boat-shaped zooids (Ryland 1960). Almost all species have pedunculate bird's head avicularia, the type species Bugula neritina (Linnaeus, 1758) being a remarkable exception. ...
... C, type 3, found in Crisularia. D, type 5, found in some Bugulina species (A, C, and D redrawn from Ryland, 1960). ...
Species in the genus Bugula are globally distributed. They are most abundant in tropical and temperate shallow waters, but representatives are found in polar regions. Seven species occur in the Arctic and one in the Antarctic and species are represented in continental shelf or greater depths as well. The main characters used to define the genus include bird's head pedunculate avicularia, erect colonies, embryos brooded in globular ooecia and branches comprising two or more series of zooids. Skeletal morphology has been the primary source of taxonomic information for many calcified bryozoan groups, including the Buguloidea. Several morphological characters, however, have been suggested to be homoplastic at distinct taxonomic levels, in the light of molecular phylogenies. Our purpose was to investigate the phylogenetic interrelationships of the genus Bugula, based on molecular phylogenetics and morphology. A Bayesian molecular phylogeny was constructed using original and previously published sequences of the mitochondrial genes cytochrome c oxidase subunit 1 (COI) and the large ribosomal RNA subunit (16S). Morphological characteristics from scanning electron and light microscopy were used to confirm the clades detected by the molecular phylogeny. Our results suggest that the genus is composed of four clades, for which we provide diagnoses: Bugula sensu stricto (30 species), Bugulina (24 species), Crisularia (23 species) and the monotypic Virididentula gen. n. Ten species could not be assigned to any of those genera, so they remain as genus incertae sedis. Our findings highlight the importance of using molecular phylogenies in association with morphological characters in systematic revisions of bryozoan taxa.
... All measurements were made with the use of a Wild stereomicroscope with 20ϫ oculars at 100ϫ magnification. We also included the ratio of avicularian length to zooid width (Lav/WZ) suggested by Ryland (1960) as potentially useful in distinguishing Bugula taxa. For species with more than one size class of avicularia, the length of the large avicularia was used in determining this ratio. ...
... In contrast to the case in many cheilostomes, zooid length and width are not particularly good characters to use to distinguish between species because the species in the subset of the genus studied here are very similar in zooid size. Characters that have been found useful in the genus, such as coloration, colony size and growth form of colony, number of series of zooids along branches, branch bifurcation patterns, and distal spine patterns (e.g., Ryland, 1960) are not as useful for this limited group of taxa. Although the two species with very small mature colonies could be distinguished from the others, most species were represented only by branch fragments, the complete size of their colonies unknown. ...
... Avicularia position and shape, relative proportion of head to beak, and shape and width of each part, however, seem to be the most distinctive characters at the species level. Additionally, the ratio of avicularian length to zooid width originally discussed by Ryland (1960) as a potentially useful character in distinguishing Bugula species appears to have value for at least some of the species studied (see Table 1). ...
In this study, we describe or redescribe nine species of red-pigmented aviculiferous Bugula and compare them with the type species of the genus, Bugula neritina (L.). The names Bugula robusta MacGillivray and Bugula minima (Waters) have both been used (often interchangeably) for red-pigmented aviculiferous Bugula specimens collected from localities ranging from Tasmania to the Red Sea. Our analysis indicates, however, that Bugula robusta is a cool-water species whose distribution appears to be limited to the southern Australian region, whereas Bugula minima is a warm-water species distributed from the Red Sea to the Indo-Pacific. These, as well as some of the other species for which these names have been used by various authors, are described and illustrated here from museum material. Bugula providensis, Bugula miniatella, Bugula ceylonensis, Bugula robustoides, Bugula solorensis, and Bugula paternostrae are described as new. The descriptions of Bugula robusta, Bugula minima, and Bugula crosslandi are revised.
... 567), à la suite de recherches complémentaires qui nous ont permis d'accéder à une meilleure connaissance des limites du genre Dendrobeania et de la hiérarchie des carac¬ tères taxinomiques. Ryland, 1960Ryland, 1960Prenant et Bobin, 1966 : 541-544 ;Ryland et Hayward, 1977 : 170-171. ...
... 567), à la suite de recherches complémentaires qui nous ont permis d'accéder à une meilleure connaissance des limites du genre Dendrobeania et de la hiérarchie des carac¬ tères taxinomiques. Ryland, 1960Ryland, 1960Prenant et Bobin, 1966 : 541-544 ;Ryland et Hayward, 1977 : 170-171. ...
... Bugulina flabellata (Thompson in Gray, 1848) (Fig. 16) Bugula flabellata: Ryland, 1960 (Gordon & Mawatari 1992). Its presence in Brazil, however, has been regarded as doubtful (Vieira et al. 2008 ...
Non-indigenous bryozoans are frequent components of biofouling assemblages in harbour environments worldwide. We performed a survey of artificial hard substrates in six harbours spanning 16 degrees of latitude along the coast of Argentina, from Ingeniero White (38º47′ S) to Ushuaia (54º48′ S). Microeciella argentina n. sp., distributed in warm-temperate waters of northern Patagonia and Buenos Aires Province, is described. The non-indigenous species (NIS) Callopora dumerilii, Smittoidea spinigera and Stephanollona boreopacifica are recorded here for the first time in the Southwest Atlantic. Their occurrence in fouling assemblages of warm-temperate harbours and their previous absence in these areas suggest that they probably arrived in Argentina by international shipping traffic. Callopora dumerilii is native to Europe and the northeast Atlantic. Smittoidea spinigera and Stephanollona boreopacifica are native to China and Korea, respectively. The morphological differences between S. spinigera and the Californian species S. prolifera, which is invading European harbours in the North Sea, are discussed. Bugula neritina, Bugulina flabellata, Cryptosula pallasiana and Fenestrulina delicia, common in harbours of Buenos Aires Province, are here recorded in Patagonia, highlighting the importance of regional shipping traffic to secondary spread of NIS. Arachnidium fibrosum, Buskia socialis, Anguinella palmata, Aetea cultrata, Bicellariella edentata and Exochella moyanoi, previously known from Brazil, and Hippothoa divaricata, are here recorded in Argentina. This study shows that warm-temperate harbours in Buenos Aires Province and northern Patagonia are particularly prone to the introduction of non-indigenous bryozoans.
... In this study, we report on the first confirmed records of this species in the UK and discuss the biogeographical implications. (Linnaeus, 1758) (abundant) plus Bugulina stolonifera (Ryland, 1960) and Bugulina simplex (Hincks, 1886) (both occasional), that fouled the floating pontoons at shallow depth. The identity of the caprellids was not investigated at that time, but they were preserved for future examination. ...
Early detection of invasive species becomes particularly challenging in the case of small-sized, neglected or taxonomically challenging taxa. The invasive skeleton shrimp, Caprella scaura Templeton, 1836, stands out as a primary example of this and is a successful widespread invader in Europe. This study presents its first record in the UK, confirming that the species is now expanding towards higher latitudes by means of commercial shipping and recreational boating. Molecular analysis indicates that the UK population bears the most widespread haplotype across the northern hemisphere and suggests a potential introduction pathway shared with populations from the Iberian Peninsula, the Mediterranean Sea, Sea of Japan and Eastern US. We propose an Indo-Pacific origin, via the Suez Canal, with Mediterranean and Iberian populations acting as stepping stones. Considering the continuous development of European coastlines, the high molecular connectivity of C. scaura with transoceanic donors and its wide environmental tolerance, a further spread is expected in this continent. Lessons learned from its invasion in Southern Europe suggest the potential for competitive exclusion of native taxa. We outline the importance of promoting taxonomic expertise and effective dissemination of records, in order to reduce misidentifications, unresolved cases and lags in the detection of alien species.
... (Hincks) HINCKS, 1880, p. 464;IFMR, 1951, p. 30;PMF, 1957, p. 346; GUÉRIN-GANIVET, 1912, p. 21 BORG, 1926, p. 396;BUSK, 1875, p. 30;IFMR, 1951, p. 30;PMF, 1957, p. 346;GUÉRIN-GANIVET, 1912, p. 21. Forme mamillée très commune en dessous de pierres (zone des marées à 40 m), mais aussi sur les parois rocheuses : horizontales et parois inclinées à 45°, légèrement recouvertes de sédiment (6-13 m), planchers de grotte (7 m Ainsi, les Bugules, dont notre région offre un riche éventail, sont toutes présentes dans les eaux britanniques (RYLAND, 1960), si elles ne sont pour autant mentionnées dans les catalogues (PMF et IFMR). Par contre, il est intéressant de noter la grande pauvreté en Cellariidae : nous avons pu en rapporter 2 espèces, confinées à 30 m et plus : Cellaria salicornioides (30 m) et C. sinuosa (80 m). ...
... 60 Bryozoans are a fascinating phylum of aquatic metazoans that form 61 colonies with an exceptionally high degree of colony polymorphism 62 (Waeschenbach et al. 2012). All bryozoan colonies are composed of autozooids, 63 the basic feeding and reproductive zooid form, that constitutes a feeding unit 64 (polypide) that captures food particles with a ring of ciliated tentacles 8 this study, these questions are explored using the autozooids and two 103 heterozooids (the bird's beak-shaped avicularia and root-like substrate 104 attachment zooids called rhizoids) of the arborescent bryozoan Bugulina 105 stolonifera (Ryland 1960) (Fig. 1). 106 Cheilostomes such as B. stolonifera have an extensive and relatively well-107 preserved fossil record and about 4,800 described extant species that inhabit 108 almost every benthic marine habitat (Brood 1998 Approximately 20 hours later, colonies were exposed to bright light (from 151 a fiber optic microscope lamp) in order to trigger the release of larvae, which are 152 positively phototactic. ...
Bryozoans are a diverse phylum of marine and freshwater colonial invertebrates containing approximately 6,300 described living species. Bryozoans grow by budding new physiologically connected colony members (zooids) from a founding individual that forms from a metamorphosed larva. In some species these zooids come in different shapes and sizes and are specialized to serve different tasks within the. A complex interaction of genotype, environment, and developmental pathway shapes zooid fate, however, the specific mechanisms underlying the establishment of this division of labor remain unknown. Here, the first characterization of differential gene expression between polymorphic zooids of a bryozoan colony is presented. The development of different zooid types of lab-cultured Bugulina stolonifera colonies including feeding autozooids, avicularia (derived non-feeding zooids that are homologous to feeding autozooids but shaped like a bird's beak), and rhizoids (a branching network of non-feeding anchoring zooids) was explored using RNA sequencing, de novo transcriptome assembly, and differential gene expression analyses. High throughput sequencing of cDNA libraries yielded an average of 14.9 ± 1.3 (SE) million high-quality paired-end reads per sample. Data for the first de novo transcriptome assemblies of B. stolonifera and the first characterization of genes involved in the formation and maintenance of zooid types within a bryozoan colony are presented. In a comparison between autozooid and avicularium tissues, 1,097 significant differentially expressed genes were uncovered. This work provides a much-needed foundation for understanding the mechanisms involved in the development of polymorphic zooids and the establishment of division of labor in bryozoans.
... B. calathus Norman, 1968Gautier, 1962, p. 76;Prenant et Bobin, 1966, p. 498. Ryland, 1960. B. fulva Ryland, 1960, p. 86;Prenant et Bobin, 1966, p. 510. ...
... Although some differences have been recorded in the timing of preferential reproductive activity by bryozoans in Ponta Delgada Harbour, the continuous recruitment of larvae and the lack of die back in the winter months indicate a potential advantage of the three invasive species over native ones (for example, Bugula flabellata and Crisia denticulata) that are known from the literature to disappear in winter and then re-establish themselvescreating a yearly cycle, settling only during a short period [19,63,72]. In Eel Pond (Woods Hole, Massachusetts), observations on T. inopinata showed no sign of colony regrowth until May and no new colonies until June [36]. ...
In the marine environment, control of invasive species’ population levels, that is, keeping them at an abundance level which is below a density-dependent adverse effect, may be the most attainable goal for the management of introduced bryozoans. An improved understanding of reproductive strategies and life history traits is key in order to understand the spreading potential. The assessment of the magnitude and temporal dynamics of propagule pressure from the reproducing population important for the success of control actions and needs is to be determined prior to any field intervention. The reproductive cycle of three fouling bryozoans (Bugula neritina, Tricellaria inopinata and Virididentula dentata) in the waters of the Azores Archipelago was assessed. The study revealed that although the release of larvae can occur throughout the entire year, its intensity and developmental and the attachment success of the ancestrula are not even throughout the year and that each species’ reproductive development needs to be determine independently. In the light of these findings, it is possible to determine the best time to apply field actions aimed at controlling invasive population’s density levels, optimizing the always scarce financial resources for marine management.
... In the UK, an example of a species with this type of distribution is the bryozoan Bugula neritina, well-known for producing bryostatin, with anti-cancer effects (Davidson & Haygood, 1999). B. neritina had been introduced from warm temperate or tropical regions to southern Britain early in the 20th century (Ryland, 1960). It was apparently confined to the vicinity of power station cooling water discharges, but may have been eliminated by the cold winter of 1963 (Ryland et al., in press;Eno et al., 1997) regarded it as -no longer established [in Britain] in the wild‖. ...
Climate change can affect every stage of the introduction, colonization, establishment and invasion of alien species; this report examines the evidence for marine species in the UK and Ireland. The introduced Pacific oyster (Crassostrea gigas) spread from oyster farms in the early 1990s, becoming established in southern England. Similarly, new self-sustaining populations are now established in Northern Ireland with recruitment occurring in favourable years. Current sea temperature projections are thought likely to result in certain species such as Crassostrea gigas recruiting every year in Northern Ireland, Wales and south-west England by 2040. Rising water temperatures may have contributed to the expansion in range of a number of species such as the bryozoan Bugula neritina, previously restricted to warm water areas such as power station outlets, and the red seaweed Caulacanthus ustulatus which was introduced from Asia and spread rapidly to Devon in 2004, Cornwall in 2005 and Kent in 2009. The invasive gastropod, the slipper limpet Crepidula fornicata, which can build up enormous populations in southern England, seems to be expanding in range, possibly in association with rising temperatures, although its spread may be more related to aquaculture activities.
... In Portugal, B. flabellata was reported before only from Costa da Arrábida (Saldanha, 1974). Rosas (1944) and Ryland (1960) also reported this species from Portugal, but without further data. ...
Unpublished data on 37 bryozoan species collected in different parts of the Portuguese coast are provided. 21 of them are reported from Portugal for the first time. Among them, Sessibugula barrosoi, Smittina jordii and Schizomavella linearis profunda are reported for the first time since their original descriptions. Anguinella palmata was never reported before in Iberian waters, including both Atlantic and Mediterranean coasts. The record of Celleporella angusta extends the species' known southern limit of distribution. Another 12 species were previously reported only once in Portugal, sometimes with some doubts. In addition, the ovicell of Crepis longipes and the ancestrula of Herentia thalassae are here described for the first time. The presence in Smittina affinis of a denticulate distal border in the primary orifice and a well-developed peristome are reported. Two species of the genera Hincksina and Hemicyclopora, left in open nomenclature, are described.
... A third Bugula species, B. stolonifera also had very few records with the most recent from ports in Britain and Ireland prior to 1950 (Ryland et al. 2011). This species was reported from Cobh harbour, Cork by Ryland (1960). These three Bugula species have the potential to colonize other sites in Ireland and should be, if possible, targeted during invasive species surveys in marinas and harbours as this is the most likely place for first introduction. ...
Invasive bryozoans in Ireland: first record of Watersipora subtorquata (d'Orbigny, 1852) and an extension of the range of Tricellaria inopinata d'Hondt and Occhipinti Ambrogi, 1985 Abstract Two invasive bryozoan species were discovered at Dun Laoghaire Harbour, Dublin during a bryozoan survey. Watersipora subtorquata (d'Orbigny, 1852) and Tricellaria inopinata d'Hondt and Occhipinti Ambrogi, 1985 were discovered in Dun Laoghaire marina. Research shows that this is a first record for W. subtorquata in Ireland and a range extension for T. inopinata. In recent years the numbers of invasive marine species in Ireland has increased. The presence of these species within the marina and not in areas surrounding the marina suggests that they have been spread via shipping activities.
... Acamarchis neritina d' Orbigny (1841Orbigny ( , 1847. Bugula stolonifera Ryland, 1960 Distribution. Rio de Janeiro and São Paulo; Marcus (1937: p. 71), Ramalho et al. (2005: p. 235), Azevedo et al. (2006: p. 267). ...
We present here a checklist of recent marine bryozoans recorded in the literature from Brazil. The total number of species recorded is 346. The most diverse group is the order Cheilostomata with 271 species, followed by the order Ctenostomata, with 42 species, and the order Cyclostomata, with 33 species. Included in the checklist are records by state and citations for species with synonyms utilized in Brazilian works.
... Discussion. -Ryland (1960) has prepared an excellent review of the generic characters and a valuable discussion of useful features for the separation of species. Most of these will become obvious in the following key and descriptions; however, two characters need some explanation here. ...
The species of Bugula, Caulibugula, and Beania which occur from Cape Hatteras to Florida are described and figured. The genus Bugula includes B. neritina, B. turrita, B. fulva, B. stolonifera, B. microoecia, B. rylandi sp. n., B. marcusi sp. n., and B. grayi sp. n. Records of Bugula avicularia and B. californica from the western Atlantic are erroneous or dubious. Bugula fulva and B. stolonifera are recognized for the first time in the American fauna. The northern range limit of Bugula microoecia is extended from the Gulf of Mexico to North Carolina. The genus Caulibugula is represented by C. dendrograpta (Waters) and C. pearsei sp. n. Caulibugula armata Verrill is redefined. Both Beania hirtissima and B. intermedia are now known to occur as far north as Cape Lookout, N. C.
... Isles (Ryland, 1960 ). It settles between early summer to mid-autumn and is well known as a fouling species. ...
国立情報学研究所で電子化したコンテンツを使用している。 Samples taken in 1997 from the Port of Nagoya, Japan, revealed the first record of the cheilostome bryozoan, Bugula stolonifera Ryland in Japanese waters. The species, which has been reported within the last years from several localities over the world, has probably been imported by ships coming to the Port of Nagoya. The species record illustrates the changes to which the Japanese bryozoan communities are subjected due to neozoan immigration. 名古屋大学博物館報告. v.19, 2003, p.9-19
The Alboran platform, located in the Alboran Sea, is an area of high productivity due to the mixture of the Atlantic and Mediterranean waters, which produce an almost permanent upwelling in the region. It has been considered one of the main biodiversity hotspots in the Mediterranean Sea. Previous taxonomic studies on bryozoans carried out around the Alboran platform mentioned a total of 127 species collected at depths between 28 and 480 m. Here we report on four Indemares (Inventario y designación de la Red Natura 2000 en áreas marinas del Estado español) samples collected in 2011 and 2012 at depths between 95 and 120 m from a biogenic gravel bottom (i.e. small rocks and shells). Sixty-eight taxa belonging to 52 genera and 34 families were identified. Two species are proposed as new (Escharella similis n. sp. and Hemicyclopora admirabilis n. sp.), eight taxa were identified only to genus level although two of them (Chorizopora sp., Hippothoa sp.) are likely to be undescribed species, and one taxon only at family level. Among the species identified, 30 are new records for the Alboran platform, 13 are new records for the Alboran Sea, and Microporella funbio is recorded for the first time in the Mediterranean Sea. These results covered only a small part of the material sampled (4 out of 44 samples), which suggests that the number of bryozoan species in the Alboran platform is even higher, and also shows the importance of this area for “bryodiversity”.
Ports have long been considered ‘high-risk’ areas for the introduction of non-native species (NNS) and should therefore be a focus of NNS monitoring. The industrial nature of active ports can, however, provide various problems when attempting to carry out monitoring programmes. Current methodologies designed to identify NNS and to describe fouling communities have not been developed specifically for use in active ports and can encounter a number of issues when used in these environments. Here, two surveys were developed and trialled within an active port in South Wales, UK, designed to describe fouling communities, identify NNS and overcome some of the major limitations to conducting surveys within ports. Over a 6-month period, fouling communities dominated by solitary ascidians developed in each survey. Seven NNS were identified, mostly species already recorded in the 1950s, including the Mediterranean crab Brachynotus sexdentatus , and the more recently introduced Japanese skeleton shrimp Caprella mutica . Each survey was evaluated independently with respect to key factors, including the ability to detect NNS and practical aspects of using these survey methods in an applied context. We conclude that whilst each survey can function independently, the use of both survey types in conjunction offers the most robust solution to identifying NNS and describing wider fouling communities within active ports. This research has implications for the future monitoring and management of NNS within UK ports.
We report a first record of the widely invasive bryozoan Bugula neritina in Ketchikan, Alaska (USA), on Revillagigedo Island (southeast Alaska). This represents the northernmost record of this fouling organism in the northeast Pacific Ocean. We also report a new occurrence of the solitary ascidian Ciona savignyi not found in Alaska since 1903, along with recent occurrences of the invasive colonial ascidians Botryllus schlosseri and Botrylloides violaceus in new localities. The high level of vessel traffic in this region and the precedent for historical ship-borne invasions worldwide suggest that future population growth and establishment of these taxa in the Ketchikan area could set the stage for further poleward range expansion, highlighting the need for continued monitoring.
There are few genuine examples of the conscious introduction of marine animals to new areas. Immigrant species have usually been accidentally introduced; ships arriving from foreign ports are regarded as the most likely source of immigrants either as ballast or on hulls. The history is usually unknown and the spread of the immigrant species is largely conjectural, one of the major difficulties, of course, being that even bottom-living forms frequently have planktonic larval stages and can spread relatively rapidly with tides and currents.
The plan for this chapter is to present, for the generalist, an overview of those aspects of the biology of bryozoans that are related directly to their procurement for and maintenance in the laboratory and to draw attention to the pertinent literature, particularly that of the 15 years since the Hyman (l959) volume. The data and opinions presented are derived from the author’s field and laboratory research and experience with the bryozoa of the northeastern United States coastal waters, from word of mouth information from colleagues, and from the literature.
Seven species of the cheilostome bryozoan genus Bugula Oken, 1815 have been recorded to date from Brazil. However, this genus is almost unknown in Rio de Janeiro State where only one species has hitherto been recorded, Bugula neritina (Linnaeus, 1758). In this study we investigate the taxonomy and distribution of Bugula in three areas of Rio de Janeiro State: Arraial do Cabo, Guanabara Bay and Sepetiba Bay. Five species of Bugula were identified, including the previously recorded Bugula neritina and four new to the state: B. dentata (Lamouroux, 1816), B. uniserialis Hincks, 1884, B. carvalhoi Marcus, 1949, and B. stolonifera Ryland, 1960. Bugula dentata is also a new record for Brazil. Four of the five species are cosmopolitan or have disjunct distributions. These are considered to be cryptogenic for Rio de Janeiro State. The fifth species, B. carvalhoi, is endemic to southeastern Brazil.
One hundred six species of Bryozoa collected from the northern Adriatic in the vicinity of Rovinj, Croatia, are distributed among the orders Ctenostomata (8 species), Cheilostomata (79 species), and Cyclostomata (19 species). Ctenostomes are underrepresented in the collections relative to the two orders with calcified colonies. Five of the cheilostome species are new: Hagiosynodos hadros n. sp., Schizomavella subsolana n. sp., Cellepora adriatica n. sp., Celleporina siphuncula n. sp., and Rhynchozoon revelatus n. sp. (previously referred to as Rhynchozoon sp. II Hayward). Seven species named by Heller (1867) are stabilized by selection of lectotypes (Beania hirtissima, Adeonella pallasii, Hagiosynodos kirchenpaueri, Exidmonea triforis, Crisia recurva) and neotypes (Mollia circumcincta, Schizomavella cornuta) from Heller's collection in the University of Innsbruck Institute of Zoology. Lectotypes are designated for the Adriatic species Hippoporina lineolifera (Hincks, 1886) and for Schizomavella mamillata (Hincks, 1880). Beania cylindrica (Hincks, 1886) and Schizoporella asymetrica (Calvet, 1927) are recognized as species rather than as subspecific units. The species-rich cheilostome genus Schizoporella Hincks, 1877, which contains some of the most widely known fouling bryozoans, is designated a nomen protectum. The species name Smittina cheilostoma (Manzoni, 1869) is preserved as established usage.
Biofouling of international marine vessels is one of the most important mechanisms for the transfer of nonnative- invasive species around the world. Bryozoan species are some of the commonest of these marine biofouling organisms found worldwide. Whilst some efforts have been made to document the bryozoan species in Australian ports, these surveys are very limited in number, poorly resolved and lack repetition. This paper records two invasive bryozoan species new to Australian waters (Hippoporina indica and Biflustra grandicella), and a northerly range extension of a known invasive bryozoan (Zoobotryon verticillatum).
This chapter highlights the effects of the heated effluents upon marine and estuarine organisms. Biological effects that observed in areas affected by heated discharges include the elimination of some species, such as cool water stenothermal forms, together with possible increase in the abundance of remaining eurythermal species, perhaps also accompanied by the introduction of immigrant warm water stenothermal species. The effects of heated discharges clearly vary according to whether the hot water spills into enclosed areas, such as docks or estuaries, or into open areas on the coast. The siting of heated effluents in marine and estuarine localities should therefore be based on a sound ecological approach and future studies would clearly benefit by the investigation of faunal and hydrographical characteristics of a region before a heated effluent is discharged. The precise biological effects of heated effluents in marine localities vary according to the extent to which the discharge area is enclosed as a cooling pond, or is flushed by tidal streams or other water movements. The effect of heated effluents is very severe in tropical areas, particularly also since there are few species normally occurring elsewhere that would be adapt to extreme heating and which could serve as replacement organisms.
Ninety alien species have been identified from British marine and brackish environments; of which 58 are established. Their arrival has been principally due to shipping and imported consignments of cultured species. The majority of alien species were initially reported from the English Channel, with many subsequently spreading northwards to the North or Celtic Seas. The majority of aliens in Britain originate from the North Pacific (N=35), followed by the North-west Atlantic (N=22). Additional alien species may be expected as a result of continued trade, port, and marina developments. Alterations in climate and extreme weather events are likely to result in future changes to the distribution of marine and brackish water alien species around the British coast.
Oil was extracted from freeze-dried sardine (Sardinella lemur) fillets using supercritical carbon dioxide (SC-CO2) and a few milliliters of ethanol were optimized with response surface methodology (RSM). The impact of extraction pressure (200–400 bars) and temperature (40–70 °C) were studied on the total extraction yields, ratios of Eicosapentaenoic acid (EPA) and Docosahexaenoic acid (DHA). The results were compared with those of Soxhlet and modified Kinsella methods (MKM). The oils obtained using the SC-CO2 and MKM methods were significantly (P < 0.05) higher in oil yield (8.04% and 6.83%), EPA (5.43% and 5.45%) and DHA (18.76% and 18.54%), respectively, compared to the Soxhlet yield (5.10%), EPA (2.17%) and DHA (06.46%). Of the two independent variables, pressure had a critical effect on yield while EPA and DHA ratios were notably influenced by temperature. The combined optimal values were pressure at 328 bars and temperature at 40 °C, with corresponding responses of 7.20%, 5.68% and 20.09% for yield, EPA and DHA, respectively. The experimental values in this study were reasonably comparable to their predicted counterparts.
The invasive bryozoan, Tricellaria inopinata d'Hondt & Occhipinti Ambrogi, 1985 was first recorded in European waters in the early 1980's and has since been reported from 166 locations from the Mediterranean Sea to the north-east Atlantic coastline. This species is typically associated with human activity, including commercial and recreational vessels and aquaculture, where it has been found in abundance on boat hulls and propellers, floating pontoons and structures associated with shellfish cultivation. Tricellaria inopinata has a high tolerance to a wide range of salinities and temperatures, although following the harsh winter of 2010, populations disappeared in Germany suggesting that this species is near the northern limit of its range under current climatic conditions. It is expected to continue to disperse though, throughout the Mediterranean, English Channel, North and Celtic Seas.
Polystyrene test panels immersed half a meter deep in the water were used to collect the fouling, encrusting Bryozoa at three harbours in Alexandria city. These were Abu Qir Harbour, the Eastern Harbour and El-Dekheila Harbour. Four species of encrusting bryozoa are recorded in the present study. These are Conopeum reticulum, Watersipora subtorquata, Cryptosula pallasiana and Schizoporella errata. The first species is affiliated to the Membraniporidae family , belonging to the suborder Anasca while the other three species are affiliated to the Watersiporidae, Cryptosulidae and Schizoporellidae families respectively. These families belong to the suborder Ascophora. A re-description, supplied with full structural illustrations of the recorded species is given. Moreover, the temporal and spatial distributions of the species recorded are provided.
1. An apparatus is described for investigating the photic responses of marine larvae. A parallel beam of light is projected along horizontal Perspex troughs containing the larvae, while the use of a water bath prevents their being subjected to a rise in temperature.
2. Using a light beam of 5000 lux at normal summer sea temperatures, plotting the positions of the larvae in the troughs at regular intervals has revealed four types of behaviour:
I. Larvae at first all photopositive, later all becoming photonegative; for example, Celleporella hyalina.
II. Larvae at first all photopositive, only a small proportion of them later becoming photonegative; for example, Bugula neritina.
III. Larvae photopositive throughout the length of their natant phase; for example, Flustrellidra hispida.
IV. Larvae showing no response at all to directional illumination; for example, Alcyonidium polyoum.
3. It has been shown that the reversal of the sign of reaction to light in Celleporella hyalina larvae is dependent on the passage of time; it does not require their exposure to a certain amount of light for its inception.
4. When disks cut from Fucus serratus fronds were placed in the troughs as a substrate for settlement, the majority of Celleporella hyalina larvae developed a photonegative reaction before they metamorphosed.
5. Horizontally and vertically placed panels in the Menai Straits received settlement of polyzoans on all surfaces free from sediment and dense algal cover. These adverse factors would have masked any selection based on the behavioural responses of the larvae.
6. Laboratory experiments with opaque panels suspended vertically, obliquely and horizontally have shown that Celleporella hyalina larvae settle on the side away from the light. Thus they normally settle on the undersurfaces of horizontal and oblique panels, but this pattern is reversed if the panels are lit from below. Settlement on panels in the sea would therefore seem to depend on the reactions of larvae to light as well as on the unfavourability of upward-facing surfaces.
7. The nature of the larval responses is discussed.
The ultrastructure of potential photoreceptors in larvae of Tricellaria occidentalis and four species of Bugula is described and compared with previously reported photoreceptors in larvae of Bugula neritina and Scrupocellaria bertholetti. A single sensory cell forms the functional unit of each photoreceptor. This cell is distinguished by a concentration of pigment vesicles in its apical part, a direct connection with the nervous system, and a large number of cilia that form the photoreceptoral organelle. These cilia have axonemes morphologically identical to those of motile cilia. The membranes of sensory cilia are unbeaded and qualitatively less osmophilic than those of the motile cilia of adjacent accessory and coronal cells. Three photoreceptor types are designated based on topological complexity: Type I, in which the sensory cell is flush with adjacent coronal cells and the photoreceptoral organelle is unprotected; Type II, in which the apical surface of the sensory cell is invaginated, forming a lumen containing the photoreceptoral organelle; and Type III, in which the sensory cell is at the base of an epidermal invagination and the photoreceptoral organelle is protected in a lumen formed by the sensory cell and accessory cells. There is a greater range of morphological variation among photoreceptors in larvae of Bugula spp. than between those of two species of the related genera Scrupocellaria and Tricellaria.
Since the species was originally described in 1960, the cheilostomate bryozoan Bugula stolonifera is known to be widely distributed in temperate and sub‐tropical regions. Evidence also exists that it is an invasive species that can spread rapidly. Yet there have been no studies of its variability at the level of individual populations. One of the taxonomic characters cited in the diagnoses of Bugula spp. is the spination pattern surrounding the frontal membrane of the ancestrula, the founding member of the colony that arises from the metamorphosis of a larva. Although some authorities have recognized that the number and distribution of ancestrular spines can vary, there has been no systematic study of the extent of this variation. We examined the spination patterns of 11,162 ancestrulae derived from larvae released from 54 colonies collected at Woods Hole, Massachusetts, USA. The formula cited in descriptions of the species was present in only 53% of the ancestrulae. In total, 34 distinct patterns were recorded. Two trends in spine appearance were statistically significant. First, those individuals possessing a spine in the proximal position were more likely to gain an additional spine. Second, those individuals lacking a proximal spine were more likely to lose an additional spine. The significance of these trends remains to be explored. Fourteen second‐generation ancestrulae possessing five variant (other than the most common) spination patterns were placed in the field for 29 days. Colonies grown from these ancestrualae were returned to the laboratory and 225 larvae produced by these colonies were collected. The spine patterns of ancestrulae resulting from metamorphosis of these larvae were recorded. Twelve patterns not present among those of the second‐generation parents were observed in addition to the five original ones. Although all colonies grown in the field were from variant ancestrulae, 51% of the second‐generation ancestrulae possessed the basic pattern. Finally, there were no significant differences between left and right side asymmetries in either the first‐ or second‐generation ancestrulae. From these studies we conclude that spine pattern of ancestrulae is not a reliable taxonomic character, at least in the Woods Hole population of B. stolonifera. Second, there exists considerable variation in spine patterns yet there is no bias to left or right asymmetry. Third, possession of a particular spine pattern is not a heritable feature in one‐generation studies. The production of ancestrular spines appears to be strongly influenced by random events occurring in the developmental process.Present address: Paula J. Rodgers, Department of Biology, University of Maryland, College Park, MD 20742, USA.
The diversity of Bryozoa at the Pacific entrance to the Panama Canal is threefold that at the Atlantic. Three ecological components are recognized on the Pacific side, two on the Atlantic. Since its inception 56 years ago, successful transmigrations of Bryozoa through the Canal have been unidirectional, as demonstrated by the occurrence of three Atlantic fouling species at the Pacific entrance, i.e., Membranipora annae Osburn, Electra monostachys (Busk) and Bugula stolonifera Ryland. Gemelliporidra multilamellosa (Canu & Bassler) and Trematooecia aviculifera (Canu & Bassler), both previously known as fossils from the Upper Tertiary of Panama, have persisted into the Recent Panamic-Atlantic intertidal environment.
One hundred six species of Bryozoa collected from the northern Adriatic in the vicinity of Rovinj, Croatia, are distributed among the orders Ctenostomata (8 species), Cheilostomata (79 species), and Cyclostomata (19 species). Ctenostomes are underrepresented in the collections relative to the two orders with calcified colonies. Five of the cheilostome species are new: Hagiosynodos hadros n. sp., Schizomavella subsolana n. sp., Cellepora adriatica n. sp., Celleporina siphuncula n. sp., and Rhynchozoon revelatus n. sp. (previously referred to as Rhynchozoon sp. II Hayward). Seven species named by Heller (1867) are stabilized by selection of lectotypes (Beania hirtissima, Adeonella pallasii, Hagiosynodos kirchenpaueri, Exidmonea triforis, Crisia recurva) and neotypes (Mollia circumcincta, Schizomavella cornuta) from Heller's collection in the University of Innsbruck Institute of Zoology. Lectotypes are designated for the Adriatic species Hippoporina lineolifera (Hincks, 1886) and for Schizomavella mamillata (Hincks, 1880). Beania cylindrica (Hincks, 1886) and Schizoporella asymetrica (Calvet, 1927) are recognized as species rather than as subspecific units. The species-rich cheilostome genus Schizoporella Hincks, 1877, which contains some of the most widely known fouling bryozoans, is designated a nomen protectum. The species name Smittina cheilostoma (Manzoni, 1869) is preserved as established usage.
Erect helical colony forms have evolved at least six separate times within the Bryozoa, but only among those in which branches are composed of a single layer of feeding zooids. The four best known genera with helical colony forms evolved independently, and each occupied different benthic marine environments, achieved different growth habits, and utilized different aspects of an array of functional potentials resulting from the radially symmetrical colonies. Examination of the distribution of these four genera (Archimedes, Bugula, Crisidmonea, and Retiflustra) within a theoretical morphospace of hypothetical helical colony form reveals that each occupies its own characteristic region of morphospace, broadly overlapping in some dimensions but separated in others. The genera differ markedly in the branching densities within their filtration-sheet whorls, reflecting their phylogenetic legacies rather than constructional or functional constraints associated with helical growth. In contrast, all tend toward helices in which the radiating whorls of the unilaminate branches are held at an average of 50–60° to the central axis of the colony, and this may reflect a common functional optimum associated with the cilia-driven, auto-generated currents within the helix. The region of morphospace characterized by high values of surface area – i.e. helical geometries with branches orientated at very low angles to the central axis, and with very closely spaced whorls along the axis – is entirely empty of bryozoans, and these geometries apparently represent functionally unrealistic colony forms. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 235–260.
The coronate larva of the ascophoran bryozoan Watersipora arcuata has a ring of 32 large, multiciliated coronal cells that are used for swimming. Fourteen pairs of small cells are intercalated between the lateral margins of adjacent coronal cells. These intercoronal cells are arranged in a precise pattern and are polymorphic: seven pairs have multiple cilia and seven pairs are mono- or oligociliated. Three pairs of multiciliated intercoronal cells have their cilia arranged as a whorl that is recessed in a pocket formed between the adjacent coronal cells, and they are thought to be photoreceptors that sense general light intensity. Two other pairs of multiciliated cells with cohesive tufts of cilia may be chemo- or mechanoreceptors. Roles of the other intercoronal cells in this species are not evident, but it is proposed that the majority, if not all, of them are sensory. The close proximity of all the intercoronal cells to the equatorial nerve ring is compatible with this interpretation. Analyses of the literature on cleavage patterns, pigment cup ocelli, and flagellar tufts that serve as balancers in coronate larvae lead us to propose that (1) an intercoronal cell is the sensory element of most, if not all, pigment cup ocelli of bryozoan larvae; and (2) intercoronal cells are not modified coronal cells but probably are specialized supra- and/or infracoronal ones that have migrated to an intercoronal position.
Induction of metamorphosis in larvae of the marine bryozoan Bugula stolonifera was examined for potential roles of G protein-linked cell-surface receptors. Two compounds that modulate components of the G protein system were tested in a bath-applied experimental design over concentrations of 10−8 to 10−3 M, plus a seawater control, and 2 controls to assess metamorphic competence (10 mM excess KCl and a biofilmed surface). The G protein activator Gpp[NH]p and the G protein inhibitor GDP-β-s did not affect rates of metamorphosis, under the conditions tested. We found no evidence that G protein-linked cell-surface receptors are the receptors of the metamorphic cue in this species. This negative finding parallels reports on a gastropod and a polychaete; by contrast, G proteins are involved in metamorphic induction in several cnidarians, a nematode, and barnacles. Thus, as no single model can account for induction across phyla, our results may serve to focus attention on identifying possible roles of the other 2 major classes of cell-surface receptors in triggering metamorphosis in B. stolonifera and other invertebrates.
Observations on the fauna of a heated, fully saline dock have been made by diving and by using suspended test panels. The seasonal abundance and breeding of native and immigrant species are described. The effects of subsequent cooling of the dock and the appearance of additional native species in the fauna are described and discussed.
The upper reaches of Milford Haven and the Daucleddau have an unusually rich fauna because of their depth and the abundance of hard substrata. The estuary is of commercial importance both in its potential for shell-fisheries and its development as a major oil port, bringing risks of serious pollution. Recent extensive sublittoral studies have added numerous faunistic records and have also cast further light upon changes in the zonation of sedentary marine animals as they penetrate brackish regions. Re-survey of a transit across the channel near Lawrenny has shown changes largely related to the spread of a vigorous immigrant competitor of the native oyster.
Bugula is a speciose genus of marine bryozoans, represented by both endemic and cosmopolitan species distributed in tropical and temperate waters and important to marine biologists because of the occurrence of many species in harbor and fouling communities, therefore as potential invaders. The southeastern Brazilian coast in the southern Atlantic hosts the highest known diversity of the genus, a status intimately associated with the intensity of collecting efforts.
Morphological data based on the examination of living specimens, scanning electron and light microscopic images, and morphometric analyses were used to assess the diversity of Bugula along the coastal areas of southern, northeastern, and southeastern Brazil. In this study, morphological species boundaries were based mainly on avicularian characters. For two morphologically very similar species, boundaries are partially supported by 16 S rDNA molecular data.
Nine species are newly described from Brazil, as follows: Bugula bowiei n. sp. ( = Bugula turrita sensu Marcus, 1937) from the southern, northeastern, and southeastern coasts; Bugula foliolata n. sp. ( = Bugula flabellata sensu Marcus, 1938), Bugula guara n. sp., Bugula biota n. sp. and Bugula ingens n. sp from the southeastern coast; Bugula gnoma n. sp. and Bugula alba n. sp. from the northeastern coast; Bugula rochae n. sp. ( = Bugula uniserialis sensu Marcus, 1937) from the southern coast; and Bugula migottoi n. sp., from the southeastern and southern coasts.
The results contribute to the morphological characterization and the knowledge of the species richness of the genus in the southwestern Atlantic (i.e., Brazil), through the description of new species in poorly sampled areas and also on the southeastern coast of that country. Additionally, the taxonomic status of the Brazilian specimens attributed to B. flabellata, B. turrita and B. uniserialis are clarified by detailed studies on zooidal and avicularia morphology.
An overview is presented of non-indigenous marine and estuarine plant and animal species recorded from The Netherlands. In this list both exotic species from outside NW Europe and non-indigenous species from elsewhere in NW Europe are enumerated. Species that have been suggested to be non-indig-enous in The Netherlands but for which insufficient evidence could be found are discussed shortly as well. The list is based mainly on literature data supplemented by own observations of the author. At least 99 plant and animal species have been introduced from elsewhere in the world. Another 13 species have been introduced from other parts of NW Europe. The third category of dubious non-indigenous species enu-merates 37 species. The list is preceded by an introduction describing the history of Dutch research on in-troduced species, the origin of the marine and estuarine flora and fauna of The Netherlands, natural and human-induced dispersal processes, and a summary of the geographic patterns of introduced species.
Seasonal life history phenomena were monitored through 1978 for two sublittoral cheilostome bryozoans from souther Britain: Chartella papyracea (Ellis and Solander), a non-placental brooder, and Bugula flabellata (Thompson in Gray), which is placental. The intracolonial relationships between growth by zooid budding, polypide recycling, and sexual reproduction, were analysed for both species — quantitatively in the case of C. papyracea. Although both species grow only in summer, their life histories differ. C. papyracea is more of a K-strategist, producing perennial fronds that release larvae in several successive years, most being liberated in winter months. The reduced need for maximization of the larval productivity rate is reflected in the intracolonial configuration of polypide and sexual recycling. B. flabellata is more of an r-strategist. The larvae are produced by the ephemeral fronds of two successive colony generations each summer. Colonies are dormant in winter. The need to maximize the larval productivity rate in this species is reflected in the polypide/sexual recycling configuration. It is believed that embryonic placentation contributes to the maximization of the larval productivity rate in this and other placental brooders.
The pigmented spots of bryozoan larvae have often been implicated in photoreception due to their preferential occurrence in larvae with positive phototactic behavior. Results of light and electron microscopic studies of Bugula neritina show that the larvae possess two spots, each with a basal sensory cell situated at the base of a pit-like depression. The embedment of the pit and its basal cell in a pad of subepidermal pigment cells allows for directional illumination. The basal sensory cell produces a ball-like mass of non-motile cilia. The configuration of the axoneme is typical of kinocilia and unlike the arrangement previously described for ciliary photoreceptors. Elaboration of receptor organelle membrane surface area is accomplished uniquely by multiple cilia oriented so that large portions of each cilium lie perpendicular to the direction of incident light. The pigmented spot directly contacts the underlying equatorial nerve ring which also connects with the major larval locomotor organ. The pigmented spots of B. neritina are the only potential photoreceptor structures which have been studied by electron microscopy in the three lophophorate phyla. The use of ciliary membranes as the potential photoreceptor organelle allies the bryozoan pigmented spot with the ciliary type photoreceptor which occurs most prevalently in deuterostome animals.
Due to the strong ecotoxicity of the biocide tributyltin (TBT), which is in use in anti-fouling paints on ships and the negative
ecological effects observed worldwide, it is planned to impose a global prohibition on the application of TBT on ships in
2003. Although substitutes are already on the market, research-efforts for novel anti-fouling technologies are intensified.
However, an essential requirement of any TBT-free anti-fouling product must be the environmental compatibility in order to
avoid pseudo-alternatives. Many of these new paints require supplements of toxic biocides and/or high loadings of copper compounds
to function effectively. Due to the lack of data the potential ecotoxicological risks to the marine environment from the use
of these alternative coatings can hardly be estimated in their range at present. Also first results of German pilot projects
on alternative ship coatings showed that in general macro-fouling is more pronounced on these materials than on TBT-containing
paints. It has to be assumed that in the future an increase in bioinvasion by shipping will occur if alternative anti-fouling
paints lack the effectiveness of TBT anti-foulants. Especially the biocide-free silicone coatings, from which fouling growth
is easily removable, have an extraordinarily high potential for introduction of alien species.
The larvae of several entoproct species and their metamorphosis or budding are described and the literature on entoproct development reviewed. Most species probably have spiral cleavage; some larvae, such as those of Loxosomella elegans and Loxosoma pectinaricola, are rather schematic trochophores, whereas most other larvae are highly characteristic, having a frontal organ and a large ciliated foot. The larvae of species such as Loxosomella harmeri undergo a metamorphosis; they attach by means of the contracted frontal organ and the alimentary canal rotates 90° in the median plane with the mouth first. Other loxosomatids, such as Loxosomella leptoclini and L. vivipara, have no metamorphosis; buds develop from the episphere of their larvae and the larvae die after the liberation of the buds. Loxosomella polita, in which the metamorphosed larvae never grow to a normal size or develop the differentiated “foot” characteristic of the adult specimens which have originated from buds, represents an inter...
IN September 1957, a species of Bugula was discovered at Holyhead, North Wales, which did not conform to any of the species described by Hincks1. Reference to the literature showed that it resembled the B. flabellata of American authors, although it was, in fact, clearly not B. flabellata (Thompson). Osburn2 identified the American form as B. flabellata and noted that Woods Hole material did not conform completely to the description given by Hincks1, although he evidently did not appreciate the full extent of the differences. Since the species is common in the Woods Hole area on submerged piles2 and has been the subject of extensive experiments, notably by Lynch3, its proper identification is of importance.
This species was originally found by Ehlers at Spiekeroog, an island off the coast of E. Friesland, in the North Sea. It has more recently been found by Joyeux-Laffuie at Luc-sur-Mer (Normandy), and by Prouho at Roscoff and at Banyuls. Its wide distribution, from the North Sea to the Mediterranean, made it almost certain that it would be found in British waters when looked for in the right place. Ehlers discovered it in the substance of the tubes of Terebella (Lanice) conchilega , while Joyeux-Laffuie and Prouho found it in the tubes of Chætopterus .
1. In calcium-free van't Hoff's solution and both Na-free and K-free mixtures having the other ions in the same proportion as in sea water at a pH of 7.1-8.0, the larvae of Bugula flabellata behaved exactly like those in sea water containing an excess of MgCl2. Phototropic reactions were lost, ciliated tissue was shed as material migrated basally from the pallial furrow and metamorphosis did not occur.2. In isotonic Mg-free solutions (pH = 8.0) metamorphosis was greatly accelerated, fixation occurring within 30 minutes to 3 hours in the majority of larvae; these organisms developed well-formed zooids in 10 hours. (Normally only about 77% of the larvae metamorphose by 12 hours.)3. Neutral red, methylene blue and, to a lesser extent, eosin accelerated metamorphosis. Neutral red, the most potent of the three, was effective in concentrations of 1:1,000,000 parts of sea water; methylene blue caused acceleration in concentrations of 1:500,000 parts. At a concentration of 1:100,000 parts of neutral red the avera...
Few observations concerning the effects of ionizing radiations have been made on bryozoan tissues. Oka ( 1954) x-rayed various regions of the fresh-water bryozoan, Lophopodella carteri, and noted that the more active embryonic tissues had greater radiosensitivity than other parts. But no observations known to the writer have been reported concerning the effects of x-radiation on the attachment and metamorphosis of bryozoan larvae. It is well known that some of the biological effects of ionizing radiations have been attributed to the production of H2O2 or organic peroxides when aqueous solu tions are x-rayed (Evans, 1947 ; Barron et al., 1949 ; Kimball and Gaither, 1952). Barron and his co-workers found that both irradiated sea water and hydrogen peroxide inhibited oxygen uptake in sea urchin sperm ; but the latter had two op posite effects, increasing respiration at great dilution and inhibiting oxygen uptake in higher concentration. Furthermore, Blum (1941, p. 96) has emphasized the probability that the effects of certain photodynamic dyes may be mediated through the production of H202. Since some of the basic dyes had been found to be potent inductors of attachment and metamorphosis of Bugula larvae when sea water solu tions of these dyes were exposed to light (Lynch, 1955a), it was of interest to de termine whether x-rays and hydrogen peroxide would have any effect on setting. If ionizing radiations were found to affect the rate of attachment of the larvae, the problem of determining whether the action of these rays was a direct or an indirect one would naturally arise. After x-raying the larvae proved to have a positive effect on the rate of setting, it was decided to employ sea water seeded immediately after being x-rayed with non-irradiated larvae. The results of these experiments led to an investigation of the possible effects of adding H2O2 to sea water and finally to observations on the action of two other oxidizing agents, sodium 2,6-dichloro benzenoneindophenol and 2,3,5-triphenyltetrazolium chloride, on the attachment and metamorphosis of Bugula larvae.
The breeding season of Bugula flabellata extends from June 10th to November 15th. The young embryos develop in brood pouches (ovicells) and are finally expelled from the colony as swimming embryos. They come from the colonies at dawn or early morning. After a free-swimming period of four to six hours, each larva becomes attached and after a profound metamorphosis which involves the loss of larval organs, it develops into the bryozoan colony by budding.The larvae at first are positive to light, but become negative before attachment. Their behavior is described in detail and the mode of attachment is explained.After the larva has become attached a period of rapid growth by budding ensues. The rate of growth is given in a table in which it is shown that the first individual of the colony is completed in two days and that a new series of buds is formed every two days. There are eight or ten individuals after one week and over a hundred in two weeks. In one month the colony is half-grown and becomes sexually mature. A colony becomes senescent in three months, when it measures 112 to 134 inches in diameter.Younger colonies hibernate successfully and resume growth in early May, when new polypides are formed.
Since a description of the larva of B. flabellata and its reactions to light and gravity has been given by Grave (1930), only contrasting features of the two larvae or additional details of their behavior will be presented here. B. flabellata, the smaller of the two (average, 0.17 by 0.19 mm.), has ten or twelve flagella in its pyriform region and is devoid of light-reactive organs. B. turrita is larger (average, 0.19 by 0.20 mm.), has four or five long slender flagella and four brilliant red, spherical eye-spots, two very close to the pyriform organ and two slightly larger ones located in the opposite hemisphere.' The whole body of the larva, ex cept the eye-spots fluoresces faintly in ultra-violet light of 3600 A. Ejected hold fast material and disintegrated larvae, however, do not fluoresce. This response to
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