Genetic structure and distribution of Gelidium elegans (Gelidiales,Rhodophyta) in Korea based on mitochondrial cox1 sequence data

Gelidium elegans Kützing is commonly found in Korea, China, and Japan, and is the economically most important agarophyte in the northwest Pacific. To assess the genetic structure of the Korean species, we analyzed 1200 base pairs of the mitochondrial cox1 gene from 272 individuals collected from 36 locations. A total of 34 haplotypes were found, most of which were unique, including 27 (79%) ‘private’ haplotypes. The nucleotide and haplotype diversities of cox1 within G. elegans were 0.711 ± 0.028 (H) and 0.00736 ± 0.00038 (π), respectively. The distribution of cox1 haplotypes, pairwise F_ST values, results of neutrality tests, AMOVA, and mismatch distribution revealed the existence of a deep genetic break between central Pacific Japan and all the other locations, corresponding to the surface seawater current patterns as well as the genetic signature of potential demographic expansion.     

Population structure, recruitment, and succession of the brown alga, Padina boryana Thivy (Dictyotales, Heterokontophyta), at an exposed shore of Sirinart National Park and a sheltered area of Tang Khen Bay, Phuket Province, Thailand

The brown algal genus, Padina, has a worldwide distribution in tropical and subtropical climate zones. Padina individuals are common and sometimes dominant in both the intertidal and shallow subtidal regions associated with coral reefs. We investigated the population structure and recruitment of two populations of Padina boryana at an exposed shore in Sirinart National Park (SNP) and a sheltered shore of Tang Khen Bay (TKB), Phuket Province, Thailand. From September 2005 to August 2006, the number of released spores and the height and radius of fronds were measured monthly. New recruits were recorded and monitored on manipulated permanent plots 0.25 m². The experiment was carried out monthly over a year. We found Padina recruits 1 month after the plots were cleared at both sites. There was a significant difference in percentage cover by new individuals between the two locations (P < 0.05) and also at the shore levels within the sites. At SNP, the highest recruitment was found in the uppermost zone, while at TKB high recruitment occurred at all shore levels except at 80-100 m from the shore. The factors influencing recruitment of P. boryana include wave motion, competition with other earlier successional species and sediment. The high availability of reproductive spores throughout the year, the successful recruitment and the Dictyerpa stage promote the successful establishment of Padina populations.     

Effects of depth, sediment and grazers on the degradation of drifting filamentous algae (Cladophora glomerata and Pilayella littoralis)

Eutrophication in the northern Baltic Sea promotes growth of annual filamentous algae. The algae detach, gather at the bottom and give rise to algal mats of varying size, density, composition and condition. Dense mats of filamentous algae induce anoxia, which in turn leads to faunal mortality. By a set of field experiments, we have studied the fate of the abundant Cladophora glomerata after detaching from the rocky substrate, and the effect of water depth and sediment on its decay. Further, we have studied the importance of common mesograzers (Gammarus and Idotea) on the rate of degradation of C. glomerata and Pilayella littoralis.Our results show that loose algae at shallow sites (8 m) decompose faster than algae in deeper (18 m) areas. Drifting C. glomerata on the sediment is more rapidly broken down and dissolved than algae floating in the water column, which depends on higher microbiological activity. Dominant amphipods (Gammarus spp) colonise near-shore drift algae quickly, and juvenile bivalves (Cerastoderma glaucum) utilise algae in the water column for settling. Moderate natural densities of grazers (Gammarus spp and Idotea baltica) in the drifting algae did not increase the degradation rates of C. glomerata and P. littoralis. C. glomerata was completely decomposed in 4 months.Our experiments demonstrate the effects of position (depth, water/sediment) and grazing on the degradation of drifting filamentous algae. Mass developments of opportunistic algae occur annually in the study area, and information on the destiny of detached drift algae may help us to predict their longevity and the damage they cause, and hence, to decide on long-term measures needed to improve environmental conditions.     

Population genetic structure of Penaeus merguiensis in Thailand based on nuclear DNA variation

Polymorphisms of nuclear DNA markers were used to study the genetic variation and differentiation among populations of Penaeus merguiensis in Thailand. The data consisted of three nuclear loci analysed for 163 individuals from five populations collected from the Gulf of Thailand (Trad, Surat Thani and Songkhla) and the Andaman Sea (Satun and Trang). The multilocus F_ST estimated at three nuclear loci revealed great and highly significant differentiation between the Gulf of Thailand and Andaman Sea (F_ST=0.203, P<0.001), mostly due to the polymorphism of locus PvAmy. In addition, significant population differentiation was also found within the Gulf of Thailand. These results were compared to that of a previous mitochondrial DNA survey spanning the same geographical range where two divergent mitochondrial clades were reported. The present study brings support to the fact that the existence of these two clades is not due to a mixture of cryptic species but reflects their phylogeographic origin. The strong genetic structure of P. merguiensis on each side of the Thailand peninsula for both mitochondrial and nuclear genes could thus be linked to the phylogeographic divide between Indian Ocean and Pacific forms on the west and east sides respectively.     

Hydrozoan and protozoan epibionts on two decapod species, Liocarcinus depurator (Linnaeus, 1758) and Pilumnus hirtellus (Linnaeus, 1761), from Scotland

Crabs of the species Liocarcinus depurator and Pilumnus hirtellus, collected on the west coast of Scotland, showed hydrozoan and protozoan epibionts. The epibionts of Liocarcinus depurator were the protozoan Ephelota plana and the hydrozoans Clytia gracilis and Leuckartiara sp. The epibionts of Pilumnus hirtellus were the protozoans E. plana and Zoothamnium sp. For Liocarcinus depurator the number of Ephelota per crab fluctuated between 0 and 47 and the greatest number of suctorians were located on the chelipeds, carapace and anterior pereiopods. The hydrozoans, for Liocarcinus depurator, showed densities of 0-20 (Clytia gracilis on the second pereiopod) and 0-507 individuals per crab (Leuckartiara sp., principally on chelipeds, carapace and the fourth right pereiopod). For Pilumnus hirtellus, Ephelota plana showed densities between 3 and 56 individuals per crab, the greatest number of suctorians being located on the same areas as on Liocarcinus depurator. There was a density of 10-69 individuals per crab of Zoothamnium sp. on Pilumnus hirtellus (located on the carapace). Ephelota plana has not been observed previously as an epibiont on crustacea, nor had Zoothamnium sp. as an epibiont on Pilumnus hirtellus. Both hydrozoans, Leukartiara and Clytia, have not been previously described as epibionts on Liocarcinus depurator. Data about the morphological characteristics and distribution of these epibionts are included.     

Effects of temperature on two Mediterranean populations of Dinophilus gyrociliatus (Polychaeta: Dinophilidae): I. Effects on life history and sex ratio

The effects of temperature on the life history characteristics of two populations of the polychaete Dinophilus gyrociliatus, one from Ravenna (northern Adriatic Sea) and the other from Genoa (Ligurian Sea), were investigated. The temperatures tested (6, 12, 18, 24 and 30 °C) cover a wider range than those prevailing in the natural environment. In the populations studied there are broad differences in timing of development and reproduction. At 6 °C, the adults of both populations survive for a long time but they are unable to reproduce. At 12 °C, only the animals from Ravenna manage to reproduce. At the higher temperatures (18, 24 and 30 °C), the development of the animals belonging to the Genoa strain is faster than that of the Ravenna strain. The duration of the various phases of the biological cycle is very similar in both populations, but that from Ravenna exhibits greater tolerance of low temperatures, slower development rate and lower development threshold temperature than does the Genoa population. Temperature and geographical origin also have strong effects on reproductive characteristics. The highest fecundity values were observed at 12 °C in the Ravenna strain, the lowest at 30 °C in both groups. At 18 °C, the Genoa population is more fecund than the Ravenna one, while the situation is reversed at 12 °C. The smallest ovigerous capsules are produced at 30 °C, the biggest at 12 °C, and the Genoa females produce larger capsules than do the females from Ravenna, except at 12 °C. The size of both male and female eggs varies in relation to temperature, the smallest female eggs generally being laid at the higher temperatures. At all the temperatures tested, the sex ratio of the Ravenna population is higher than that of the Genoa population. In the Ravenna strain, temperature has no effect on the sex ratio, while in the Genoa strain the sex ratio at 24 °C is lower than at 18 and 30 °C. Comparison of the two populations at the same temperature reveals considerable differences in the characteristics of their respective life histories and sex ratios. It is very likely that the extreme selectivity of the harbor environments has favored the fragmentation of the species into differentiated populations that have adapted to the conditions prevailing in the different localities.     

Effects of population density on the sediment mixing induced by the gallery-diffusor Hediste (Nereis) diversicolor O.F. Müller, 1776

The aim of this work was to quantify the intensity of sediment mixing induced by the gallery-diffusor (functional bioturbation group) Hediste diversicolor as a function of density, using particles tracers (luminophores). In order to assess the impact of density on sediment reworking, a 1-D model was used to obtain sediment reworking coefficients such as Db (biodiffusion-like) and r (biotransport). Densities used in this experiment corresponded to population densities observed in the sampling area (Saint-Antoine Canal, Gulf of Fos, France): 144, 288, 577, 1153 indiv/m². At first, results showed that neither luminophore maximum burying depth nor the more marked tracer accumulation areas were influenced by density. Thus density did not seem to have any influence on size of galleries or complexity of structure. Then, density-dependent relations with Db (biodiffusion-like mixing) and r (biotransport) were highlighted with an observed process intensity rate twice as high at highest worm density. On the other hand, Db and r per capita coefficients were negatively influenced by density. Db and r per capita at highest density were equal to ∼ 20% of individual Db and r obtained at the lowest density. Finally, this study showed the importance of density which appears to be a key parameter in the functioning of the sedimentary ecosystem.     

Effects of live coral,epilithic algal communities and substrate type on algal recruitment

The recruitment of algae is a critical process during algal colonization and invasions, including coral-to-algal phase shifts. Although algae are widely assumed to colonize and kill corals, there is very little known about the recruitment dynamics of coral reef algae. This study tested the ability of two dominant macroalgae (Fucales including Sargassum spp. and Lobophora variegata) to settle and grow on healthy coral tissue. The study also explored the effects of interactions with prior occupants, and of abiotic substrate properties (texture, and ceramic and carbonate material). The results indicate that healthy corals were able to prevent attachment or survival of recruits of these macroalgae. This is a significant point, since it suggests that the replacement of corals by algae may often require prior stress or death in the coral tissue. Pre-conditioning of plates at different sites had some effects, but these were relatively minor, whereas there was considerable variation within sites. Some of this variation appeared to be related to the amount of turf algae or crustose coralline algae on the plates. Recruitment was generally, but not always, higher on plates with rougher texture. Overall, this preliminary exploration indicates considerable potential for variability in outcomes of algal colonization, with implications for the dynamics of algal invasions. In particular, the results do not support suggestions that planktonic algal propagules can directly settle on and colonize healthy coral tissue.Communicated by Topic Editor D. Barnes     

Microsatellite diversity and population genetic structure of yellowcheek, Elopichthys bambusa (Cyprinidae) in the Yangtze River

Yellowcheek carp (Elopichthys bambusa) is the only species of genus Elopichthys. It is widely distributed in Chinese freshwaters but currently its populations have declined to threatening level. We examined the genetic diversity and population structure of E. bambusa in the Yangtze River basin. A total of nine polymorphic microsatellite markers were employed to study five populations occurring in middle and lower reaches of the river. The results revealed low-to-moderate genetic diversity. The number of alleles per locus varied between 3 and 8 with an average of 4.8. Observed heterozygosity ranged from 0.15 to a maximum of 1.00. Significant deviations (P < 0.01) from Hardy–Weinberg equilibrium were observed for all the tested locus-population combinations with clear heterozygosity deficits. AMOVA indicated that majority of the variance lies within populations (93.81%) than among the populations (7.05%). Pairwise F_ST and unbiased genetic distance pointed out significant differentiation among the samples from populations with different connections to the Yangtze River. In the UPGMA dendrogram, clustering pattern of populations indicated that most of the populations are reproductively isolated due to anthropogenic interventions. Clustering of PYL and DTL populations shows ongoing gene flow through the mainstream. The recent hydrological alterations and overfishing are major factors shaping the current genetic structure. These results can be helpful for effective management and sustainable conservation of E. bambusa populations.     

Rapid identification of lymnaeid snails and their infection with Fasciola gigantica in Thailand

Freshwater snails of the family Lymnaeidae are the intermediate hosts of the liver fluke Fasciola worldwide. While distinct species have been identified at the molecular level in other parts of the world such data have not been published for Thailand. In this study we collected Lymnaeidae from different localities across Thailand and analyzed their 16S rDNA sequences as a molecular signature for classification. In addition to the ubiquitous Radix rubiginosa, we have confirmed the presence of Austropeplea viridis and Radix swinhoei, for the latter of which the ribosomal rDNA sequences are reported for the first time, in North-Thailand. Based on the obtained 16S rDNA data three primer pairs were designed that allowed rapid identification of these snail species by PCR. To determine their infection status, PCR primers for F.gigantica cathepsin L were used in parallel with the snail 16S rDNA species-specific primers in multiplex PCR analyses. Western blot analysis of total snail protein with a monoclonal anti-F.gigantica cathepsin L antibody confirmed positive cathepsin L PCR results. The developed diagnostic PCR will be of use in risk assessment for transmission of fascioliasis in Thailand.     

Phylogenetic reassessment of Hyaloscyphaceae sensu lato (Helotiales, Leotiomycetes) based on multigene analyses

Hyaloscyphaceae is the largest family in Helotiales, Leotiomycetes. It is mainly characterized by minute apothecia with well-differentiated hairs, but its taxonomic delimitation and infrafamilial classification remain ambiguous. This study performed molecular phylogenetic analyses using multiple genes including the ITS-5.8S rDNA, the D1–D2 region of large subunit of rDNA, RNA polymerase II subunit 2, and the mitochondrial small subunit. The primary objective was to evaluate the phylogenetic utility of morphological characters traditionally used in the taxonomy of Hyaloscyphaceae through reassessment of the monophyly of this family and its genera. The phylogenetic analyses inferred Hyaloscyphaceae as being a heterogeneous assemblage of a diverse group of fungi and not supported as monophyletic. Among the three tribes of Hyaloscyphaceae only Lachneae formed a monophyletic lineage. The presence of hairs is rejected as a synapomorphy, since morphologically diversified hairs have originated independently during the evolution of Helotiales. The true- and false-subiculum in Arachnopezizeae are hypothesized to have evolved through different evolutionary processes; the true-subiculum is likely the product of a single evolutionary origin, while the false-subiculum is hypothesized to have originated multiple times. Since Hyaloscyphaceae sensu lato was not resolved as monophyletic, Hyaloscyphaceae sensu stricto is redefined and only applied to the genus Hyaloscypha.     

Effects of temperature on two Mediterranean population of Dinophilus gyrociliatus (Polychaeta: Dinophilidae): II. Effects on demographic parameters

The effects of temperature on demographic characteristics of two populations from Ravenna and Genoa of the polychaete Dinophilus gyrociliatus were investigated. Temperature affects age-specific survival and fecundity and all the demographic parameters often to a different degree in the two populations. Individuals from Ravenna survive longer than those from Genoa. The most evident differences in the age-specific fecundity curves of the experimental groups are related to age at maturity and the duration of the reproductive period that are in inverse proportion to temperature. In both populations of D. gyrociliatus, the maximum daily fecundity is observed at intermediate temperatures. In all cases, the Genoa females mature earlier, attain their maximum fecundity more quickly and have a shorter reproductive period than their Ravenna counterparts.Age at maturity, fecundity during the first reproductive events and juvenile survival are by far the most important characteristics in determining the fitness of the two populations at the tested temperatures. Even though the greatest net growth rates and highest expectation of life were recorded at 12 °C in the Ravenna population, the delay in the attainment of sexual maturity means that, at this temperature, the population growth rate is lowest. The higher juvenile survivorship and the greater fecundity observed at 24 °C is counter-balanced by the early attainment of sexual maturity induced at 30 °C. The comparison of the population growth rate calculated in laboratory with field data suggests that temperature is one of the main environmental parameters determining the fitness of D. gyrociliatus.     

Effects of algal canopy clearance on plant, fish and macroinvertebrate communities on eastern Tasmanian reefs

Changes in assemblages of plants, macroinvertebrates and fishes on three eastern Tasmanian reefs were monitored over 12 months in replicated control blocks and adjacent 10×12-m blocks cleared of fucoid, laminarian and dictyotalean algae. Removal of canopy-forming plants produced less change to biotic assemblages than reported in studies elsewhere, with the magnitude of change for fish and invertebrate taxa lower than variation between sites and comparable to variation between months.The introduced annual kelp Undaria pinnatifida exhibited the only pronounced response to canopy removal amongst algal taxa, with a fivefold increase in cleared blocks compared to control blocks. Marine reserves are suggested to assist reef communities resist invasion by U. pinnatifida, through an indirect mechanism involving increased predation pressure on sea urchins and reduced formation of urchin barrens that are amenable to U. pinnatifida propagation.Large invertebrates were more associated with turfing algae or the reef substratum than the macroalgal canopy. The herbivorous sea urchin Heliocidaris erythrogramma and abalone Haliotis ruber showed the strongest response to clearing amongst common macroinvertebrate species, with a halving of population numbers. Observed densities of the common monacanthid fish Acanthaluteres vittiger also declined by about 50%. The relatively high level of resistance shown by eastern Tasmanian reef biota to patch disturbance was attributed largely to high diversity and biomass of turfing macroalgae damping effects of canopy clearance.     

Systematics and evolution of the genus Torrubiella (Hypocreales, Ascomycota)

Torrubiella is a genus of arthropod-pathogenic fungi that primarily attacks spiders and scale insects. Based on the morphology of the perithecia, asci, and ascospores, it is classified in Clavicipitaceae s. lat. (Hypocreales), and is considered a close relative of Cordyceps s. 1., which was recently reclassified into three families (Clavicipitaceae s. str., Cordycipitaceae, Ophiocordycipitaceae) and four genera (Cordyceps s. str, Elaphocordyceps, Metacordyceps, and Ophiocordyceps). Torrubiella is distinguished morphologically from Cordyceps s. lat. mainly by the production of superficial perithecia and the absence of a well-developed stipitate stroma. To test and refine evolutionary hypotheses regarding the placement of Torrubiella and its relationship to Cordyceps s. lat., a multi-gene phylogeny was constructed by conducting ML and Bayesian analyses. The monophyly of Torrubiella was rejected by these analyses with species of the genus present in Clavicipitaceae, Cordycipitaceae, and Ophiocordycipitaceae, and often intermixed among species of Cordyceps s. lat. The morphological characters traditionally used to define the genus are, therefore, not phylogenetically informative, with the stipitate stromata being gained and/or lost several times among clavicipitaceous fungi. Two new genera (Conoideocrella, Orbiocrella) are proposed to accommodate two separate lineages of torrubielloid fungi in the Clavicipitaceae s. str. In addition, one species is reclassified in Cordyceps s. str. and three are reclassified in Ophiocordyceps. The phylogenetic importance of anamorphic genera, host affiliation, and stipitate stromata is discussed.     

Structure and development of the style base in Abildgaardia, Bulbostylis, and Fimbristylis (Cyperaceae,Cyperoideae, Abildgaardieae)

The Abildgaardieae tribe within the family Cyperaceae comprises six or seven genera, among which Abildgaardia, Bulbostylis and Fimbristylis pose a challenge regarding their morphological delimitation. Molecular phylogenetic analyses including species of Abildgaardieae are rare, but in most of those studies, Abildgaardia and Fimbristylis appear as more closely related to each other than to the Bulbostylis genus. Duration of the style base has been one of the most widely used characters for delimiting these three genera. The style base is a persistent structure in most species of Bulbostylis and deciduous in Abildgaardia and Fimbristylis. The reasons why the style base may persist or fall off have been scarcely discussed. The assumption that abscission layers are present in the style base of all three genera and the fact that tracheids have been observed in the style base of Bulbostylis suggest that this structure might have histological complexity. In view of this, a complete ontogenetic and anatomical study of the gynoecium has been carried out for all these three genera. It turned out that the style base is histologically simple in Abildgaardia, Bulbostylis and Fimbristylis and shows similar structure and development in all three genera. The fact that the style base has a shorter duration in Abildgaardia and Fimbristylis than in Bulbostylis might be related to the lower number of sclerotised cells that make up such structures in the mature fruit of the former two genera. Abscission of the style and style base may be the result of much simpler reasons than the differentiation of an abscission layer, resulting merely from mechanical shear force effects. Differences among genera have been observed in the shape of the style base and the development of the style. The histological simplicity of the style base is consistent with the homoplastic appearance of this structure in genera that are not closely related (e.g. Rhynchospora). Because of this, while the presence of the thickened style base seems to be a synapomorphy in species of Abildgaardieae, its persistence on or detachment from the fruit might have emerged repeatedly during this clade evolution and might not be a suitable character for genera delimitation.     

Metabolic consequences of living in a wave-swept environment: Effects of simulated wave forces on oxygen consumption, heart rate, and activity of the shell adductor muscle of the abalone Haliotis iris

Animals in wave-exposed habitats must constantly contend with the hydrodynamic forces of lift and drag. In this study, we investigated aspects of the metabolic response of Haliotis iris to simulated wave forces varying in magnitude up to 9.6 N applied to the shell at 69° to horizontal, alternately from anterior and posterior directions, with a period of 10s. Shell adductor muscle activity (electromyogram, EMG), heart rate, and oxygen consumption were monitored during force application and during extended recovery. EMG spiking was absent at zero force, but increased markedly with increasing force, in synchrony with the wave cycle. In contrast, heart rate was unaffected by wave forces and varied by only 5% over the whole range of applied forces. During force application, oxygen consumption increased by 10-25% above resting rates and remained elevated throughout a 5-hour recovery period, indicating a switch to anaerobic metabolism. It is concluded that living in a wave-swept environment is metabolically costly for abalone although this may be compensated by improved food availability and more efficient ventilation induced by external flow.     

Carrageenan from Sarconema scinaioides (Gigartinales, Rhodophyta) of Indian waters

Carrageenan was extracted from the red seaweed Sarconema scinaioides of Indian waters and was characterized. The crude carrageenan as well as its alkali modified derivative was composed of 3,6-anhydro galactose, 6-O-methyl galactose as well as galactose moieties in various proportions. Linkage analysis exhibited that these two carrageenan samples consisted of 4-linked 3,6-anhydrogalactose residue sulphated at position 2, and 3-linked galactose residue sulphated at position 4. The physicochemical and rheological data along with molecular weight data, FT-IR, 1D and 2D NMR (¹H, ¹³C, COSY and HSQC) spectrometry suggested that the polysaccharide was composed predominantly of iota- along with a small amount of its precursor nu (ν)-carrageenan, unlike the hybrid carrageenans (iota-, pyruvated- and kappa-carrageenans) from this seaweed reported in the literature. This Indian seaweed species would be a potentially important source of iota-carrageenan.     

Female biased sex-ratio in Schistosoma mansoni after exposure to an allopatric intermediate host strain of Biomphalaria glabrata

For parasites that require multiple hosts to complete their development, the interaction with the intermediate host may have an impact on parasite transmission and development in the definitive host. The human parasite Schistosoma mansoni needs two different hosts to complete its life cycle: the freshwater snail Biomphalaria glabrata (in South America) as intermediate host and a human or rodents as final host. To investigate the influence of the host environment on life history traits in the absence of selection, we performed experimental infections of two B. glabrata strains of different geographic origin with the same clonal population of S. mansoni. One B. glabrata strain is the sympatric host and the other one the allopatric host. We measured prevalence in the snail, the cercarial infectivity, sex-ratio, immunopathology in the final host and microsatellite frequencies of individual larvae in three successive generations.     

Septal pore complex morphology in the Agaricomycotina (Basidiomycota) with emphasis on the Cantharellales and Hymenochaetales

The ultrastructure of septa and septum-associated septal pore caps are important taxonomic markers in the Agaricomycotina (Basidiomycota, Fungi). The septal pore caps covering the typical basidiomycetous dolipore septum are divided into three main phenotypically recognized morphotypes: vesicular-tubular (including the vesicular, sacculate, tubular, ampulliform, and globular morphotypes), imperforate, and perforate. Until recently, the septal pore cap-type reflected the higher-order relationships within the Agaricomycotina. However, the new classification of Fungi resulted in many changes including revision of existing and addition of new orders. Therefore, the septal pore cap ultrastructure of more than 325 species as reported in literature was related to this new classification. In addition, the septal pore cap ultrastructures of Rickenella fibula and Cantharellus formosus were examined by transmission electron microscopy. Both fungi have dolipore septa associated with perforate septal pore caps. These results combined with data from the literature show that the septal pore cap-type within orders of the Agaricomycotina is generally monomorphic, except for the Cantharellales and Hymenochaetales.It appears from the fungal phylogeny combined with the septal pore cap ultrastructure that the vesicular-tubular and the imperforate type both may have arisen from endoplasmic reticulum. Thereafter, the imperforate type eventually gave rise to the perforate septal pore cap-type.