Skip to main content

Bangiophytes: From one Class to Six; Where Do We Go from Here?

Moving the Bangiophytes into the Genomic Age

  • Chapter
  • First Online:
Red Algae in the Genomic Age

Abstract

Traditionally, the red algae have been divided into two classes or subclasses: the Florideophyceae or Florideophycidae and the Bangiophyceae or Bangiophycidae, respectively; the latter taxon contains 46 genera and <400 species. Numerous studies have identified considerable variation in this group with respect to life-history, morphology, and habitat. In addition, based on biochemical, morphological, ultrastructural, and molecular analyses, the Bangiophycidae as an entity is clearly paraphyletic. A recent study proposed that the Rhodophyta be divided into seven classes within two subphyla: the Cyanidiophytina, which contains the enigmatic members of the Cyanidiophyceae and the Rhodophytina, which contains the remaining six classes, including the morphologically complex Florideophyceae. These taxonomic revisions have made considerable progress in unraveling the evolutionary history of this phylum. Despite that five of these classes comprise ∼1% of recognized species in the red algae, they are of considerable evolutionary importance and may have played a critical role in the origin of plastids in numerous other photosynthetic phyla. The first complete red algal genome was Cyanidioschyzon merloae (Cyanidiophyceae) and there are currently two other “bangiophyte” genome sequencing projects: Galderia sulphuraria (Cyanidiophyceae) and Porphyra umbilicalis (Bangiophyceae). There has been considerable research on the Cyanidiophyceae and Bangiophyceae but the remaining “bangiophyte” classes, the Compsopgonophyceae, Porphyrdiophyceae, Rhodellophyceae, and Stylonematophyceae are not as well understood. We would discuss basic aspects of each class, but additional research on these interesting groups in terms of evolutionary history, ecology, and other characteristics can only enhance the interpretation of future genomic information.

This is a preview of subscription content, log in via an institution to check access.

Access this chapter

Chapter
USD 29.95
Price excludes VAT (USA)
  • Available as PDF
  • Read on any device
  • Instant download
  • Own it forever
eBook
USD 169.00
Price excludes VAT (USA)
  • Available as EPUB and PDF
  • Read on any device
  • Instant download
  • Own it forever
Softcover Book
USD 219.00
Price excludes VAT (USA)
  • Compact, lightweight edition
  • Dispatched in 3 to 5 business days
  • Free shipping worldwide - see info
Hardcover Book
USD 219.99
Price excludes VAT (USA)
  • Durable hardcover edition
  • Dispatched in 3 to 5 business days
  • Free shipping worldwide - see info

Tax calculation will be finalised at checkout

Purchases are for personal use only

Institutional subscriptions

References

  • Baldauf, S.L., Roger, A.J., Wenk-Siefert, I. and Doolittle, W.F. (2000) A kingdom-level phylogeny of eukaryotes based on combined protein data. Science 290: 972–977.

    Article  PubMed  CAS  Google Scholar 

  • Barbier, G., Oesterholt, C., Larson, M.D., Halgren, R.G., Wilkerson, C., Garavito, R.M., Benning, C. and Weber, A.P.M. (2005) Comparative genomics of two closely related unicellular thermo-acidophilic red algae, Galdieria sulphuraria and Cyanidioschyzon merolae, reveals the molecular basis of the metabolic flexibility of Galdieria sulphuraria and significant differences in carbohydrate metabolism of both algae. Plant Physiol. 137: 460–474.

    Article  PubMed  CAS  Google Scholar 

  • Bhattacharya, D. and Medlin, L. (1995) The phylogeny of plastids: a review based on comparisons of small-subunit ribosomal RNA coding regions. J. Phycol. 31: 489–498.

    Article  CAS  Google Scholar 

  • Bold, H.C. and Wynne, M.J. (eds.) (1985) Introduction to the Algae. Structure and Reproduction. Prentice Hall, Englewood Cliffs, NJ.

    Google Scholar 

  • Broadwater, S.T. and Scott, J.L. (1994) Ultrastructure of unicellular red algae, In: J. Seckbach (ed.) Evolutionary Pathways and Enigmatic Algae: Cyanidium caldarium (Rhodophyta) and Related Cells. Kluwer, Dordrecht, The Netherlands, pp. 215–230.

    Google Scholar 

  • Broadwater, S.T., Scott, J.L., Goss, S.P.A. and Saunders, B.D. (1995) Ultrastructure of vegetative organization and cell division in Glaucoshphaeara vacuolata Korshikov (Porphyridiales, Rhodophyta). Phycologia 34: 352–361.

    Article  Google Scholar 

  • Broom, J.E., Jones, W.A., Hill, D.F., Knight, G.A. and Nelson, W.A. (1999) Species recognition in New Zealand Porphyra using 18S rDNA sequencing. J. Appl. Phycol. 11: 421–428.

    Article  CAS  Google Scholar 

  • Broom, J.E.S., Farr, T.J. and Nelson, W.A. (2004) Phylogeny of the Bangia flora of New Zealand suggests a southern origin for Porphyra and Bangia (Bangiales, Rhodophyta). Mol. Phy. Evol. 31: 1197–1207.

    Article  CAS  Google Scholar 

  • Burger, G., Saint-Louis, D., Gray, M.W. and Lang, B.F. (1999) Complete sequence of the mitochondrial DNA of the red alga Porphyra purpurea: cyanobacterial introns and shared ancestry of red and green algae. Plant Cell 11: 1675–1694.

    PubMed  CAS  Google Scholar 

  • Brodie, J.A. and Irvine, L.M. (2003) Seaweeds of the British Isles. Volume 1. Rhodophyta. Part 3B. Bangiophycidae. Intercept, Hampshire, United Kingdom.

    Google Scholar 

  • Cavalier-Smith, T. (1998) A revised six-kingdom system of life. Biol. Rev. 73: 203–266.

    Article  PubMed  CAS  Google Scholar 

  • Ciniglia C., Yoon, H.S., Pollio, A., Pinto, G. and Bhattacharya, D. (2004) Hidden biodiversity of the extremophilic Cyanidiales red algae. Mol. Ecol. 13: 1827–1838.

    Article  PubMed  CAS  Google Scholar 

  • Dixon, P.S. (1963) The taxonomic implications of the “opit connexions” reported in the Bangiophycidae. Taxon 12: 108–110.

    Article  Google Scholar 

  • Doweld, A. (2001) Prosyllabus Tracheophytorum. GEOS, Moscow, USSR.

    Google Scholar 

  • Drew, K.M. (1949) Conchocelis-phase in the life-history of Porphyra umbilicalis (L.) Kütz. Nature 164: 748–749.

    Article  Google Scholar 

  • Fan, K.C. (1960) On pit-connections in Bangiophycidae. Nova Hedwigia 1: 305–307.

    Google Scholar 

  • Freshwater, D.W., Fredericq, S., Butler, B.S., Hommersand, M.H. and Chase, M.W. (1994) A gene phylogeny of the red algae (Rhodophyta) based on plastid rbcL. Proc. Natl. Acad. Sci. USA 91: 7281–7285.

    Article  PubMed  CAS  Google Scholar 

  • Fritsch, F.E. (1945) The Structure and Reproduction of the Algae, Vol. 2. Cambridge University Press, Cambridge, UK.

    Google Scholar 

  • Gabrielson, P.W., Garbary, D.J. and Scagel, R.F. (1985) The nature of the ancestral red alga: inferences from a cladistic analysis. BioSystems 18: 335–346.

    Article  PubMed  CAS  Google Scholar 

  • Gabrielson, P.W., Garbary, D.J., Sommerfeld, M.R., Townsend, R.A. and Tyler, P.L. (1990) Phylum Rhodophyta, In: L. Margulis, J.O. Corliss, and M. Melkonian (eds.) Handbook of Protoctista. Jones & Bartlett, Boston, MA, pp. 102–118.

    Google Scholar 

  • Gantt, E., Scott, J. and Lipschultz, C. (1986) Phycobiliprotein composition and chloroplast structure in the freshwater red alga Compsopogon coeruleus (Rhodophyta). J. Phycol. 22: 280–484.

    Article  Google Scholar 

  • Garbary, D.J. and Gabrielson, P.W. (1990) Taxonomy and evolution, In: K.M. Cole and R.G. Sheath (eds.) Biology of the Red Algae. Cambridge University Press, Cambridge, UK, pp. 477–498.

    Google Scholar 

  • Garbary, D.J., Hansen, G.I. and Scagel, R.F. (1980) A revised classification of the Bangiophyceae (Rhodophyta). Nova Hedwigia 33: 145–166

    Google Scholar 

  • Geesink, R. (1973) Experimental investigations on marine and freshwater Bangia (Rhodophyta) from the Netherlands. J. Exp. Mar. Biol. 11: 239–247.

    Article  Google Scholar 

  • Glockner, G., Rosenthal, A. and Valentin, K. (2000) The structure and gene repertoire of an ancient red algal plastid genome. J. Mol. Evol. 51: 382–390.

    PubMed  CAS  Google Scholar 

  • Grossman, A.R. (2005) Paths towards algal genomics. Plant Physiol. 137: 410–427.

    Article  CAS  Google Scholar 

  • Hawkes, M.W. (1988) Evidence of sexual reproduction in Smithora naiadum (Erythropeltidales, Rhodophyta) and its evolutionary significance. Br. Phycol. J. 23: 327–336.

    Article  Google Scholar 

  • Karsten, U., West, J.A., Zuccarello, G.C., Engbrodt, R., Yokoyama, A., Hara, Y. and Brodie, J. (2003) Low molecular weight carbohydrates of the Bangiophycidae (Rhodophyta). J. Phycol. 39: 584–589.

    Article  CAS  Google Scholar 

  • Lapidot, M., Raveh, D., Sivan, A., Arad, S. and Shapira, M. (2002) Stable chloroplast transformation of the unicellular red alga Porphyridium species. Plant Physiol. 129: 7–12.

    Article  PubMed  CAS  Google Scholar 

  • Lynch, M.D.J., Sheath, R.G. and Müller, K.M. (2008) Phylogenetic position and ISSR-estimated intraspecific genetic variation of Bangia maxima (Bangiales, Rhodophyta). Phycologia 47: 599–613.

    Article  CAS  Google Scholar 

  • Magne, F. (1960) Le Rhodochaete parvula Thuret (Bangioidée) et la reproduction sexuée. Cah. Biol. Mar. 5: 407–420.

    Google Scholar 

  • Matsuzaki, M., Misumi, O., Shin-I, T., Maruyama, S., Takahara, M., Miyagishima, S.Y., Mori, T., Nishida, K., Yagisawa, F., Nishida, K., Yoshida, Y., Nishimura, Y., Nakao, S., Kobayashi, T., Momoyama, Y., Higashiyama, T., Minoda, A., Sano, M., Nomoto, H., Oishi, K., Hayashi, H., Ohta, F., Nishizaka, S., Haga, S., Miura, S., Morishita, T., Kabeya, Y., Terasawa, K., Suzuki, Y., Ishii, Y., Asakawa, S., Takano, H., Ohta, N., Kuroiwa, H., Tanaka, K., Shimizu, N., Sugano, S., Sato, N., Nozaki, H., Ogasawara, N., Kohara, Y. and Kuroiwa, T. (2004) Genome sequence of the ultrasmall unicellular red alga Cyanidioschyzon merolae 10D. Nature 428: 653–657.

    Article  PubMed  CAS  Google Scholar 

  • Moreira, D., Guyader, H.L. and Philippe, H. (2000) The origin of red algae and the evolution of chloroplasts. Nature 405: 69–72.

    Article  PubMed  CAS  Google Scholar 

  • Müller, K.M., Cannone, J.J., Gutell, R.R. and Sheath, R.G. (2001a) A structural and phylogenetic analysis of the group IC1 introns in the order Bangiales (Rhodophyta). Mol. Biol. Evol. 18: 1654–1667.

    Article  PubMed  Google Scholar 

  • Müller, K.M., Cannone, J.J. and Sheath, R.G. (2005) A molecular phylogenetic analysis of the Bangiales (Rhodophyta) and description of a new genus and species, Pseudobangia kaycoleia. Phycologia 44: 146–155

    Article  Google Scholar 

  • Müller, K.M., Cole, K.M. and Sheath, R.G. (2003) Systematics of Bangia (Bangiales, Rhodophyta) in North America. II. Biogeographical trends in karyology: chromosome numbers and linkage with gene sequence phylogenetic trees. Phycologia 42: 209–219.

    Article  Google Scholar 

  • Müller, K.M., Oliveira, M.C., Sheath, R.G. and Bhattacharya, D. (2001b) Ribosomal DNA phylogeny of the Bangiophycidae (Rhodophyta) and the origin of secondary plastids. Am. J. Bot. 88: 1390–1400.

    Article  PubMed  Google Scholar 

  • Müller, K.M., Sheath, R.G., Vis, M.L., Crease, T.J. and Cole, K.M. (1998) Biogeography and systematics of Bangia (Bangiales, Rhodophyta) based on the Rubisco spacer, rbcL gene and 18S rRNA gene sequences and morphometric analyses. 1. North America. Phycologia 37: 195–207.

    Article  Google Scholar 

  • Nelson, W.A., Broom, J.E. and Farr, T.J. (2003) Pyrophyllon and Chlidophyllon (Erythropeltidales, Rhodophyta): two new genera for obligate epiphytic species previously placed in Porphyra, and a discussion of the orders Erythropeltidales and Bangiales. Phycologia 42: 308–315.

    Article  Google Scholar 

  • Nelson, W.A., Farr, T.J. and Broom, J.E.S. (2005) Dione and Minerva, two new genera from New Zealand circumscribed for basal taxa in the Bangiales (Rhodophyta). Phycologia 44: 139–145.

    Article  Google Scholar 

  • Nelson, W.A., Farr, T.J. and Broom, J.E.S. (2006) Phylogenetic relationships and generic concepts in the red order Bangiales: challenges ahead. Phycologia 45: 249–259.

    Article  Google Scholar 

  • Nikaido, I., Asamizu, E., Nakajima, M., Nakamura, Y., Saga, N. and Tabata, S. (2000) Generation of 10,154 expressed sequence tags from a leafy gametophyte of a marine red alga, Porphyra yezoensis. DNA Res. 7: 223–227.

    Article  PubMed  Google Scholar 

  • Oliveira, M.C. and Bhattacharya, D. (2000) Phylogeny of the Bangiophycidae (Rhodophyta) and the secondary endosymbiotic origin of algal plastids. Am. J. Bot. 87: 482–492.

    Article  PubMed  CAS  Google Scholar 

  • Ott, F.D. (1987) A brief review of the species of Porphyridium with additional records for the rarely collected alga Porphyridium sordidum Geitler, 1932 (Rhodophycophyta, Porphyridiales). Arch. Protist. 134: 35–41.

    Article  Google Scholar 

  • Patrone, L.M., Broadwater, S.T. and Scott, J.L. (1991) Ultrastructure of vegetative and dividing cells of the unicellular red algae Rhodella violacea and Rhodella maculata. J. Phycol. 27: 742–753.

    Article  Google Scholar 

  • Pekárková, B., Hindák, F. and Šmards, J. (1988) Morphological characteristics and physiological properties of a coccoid Rhodophycean alga Rhodella grisea from thermal springs at Pieštany, Czechoslovakia. Arch. Protistenk. 135: 69–83.

    Article  Google Scholar 

  • Pinto, G., Ciniglia, C., Cascone, C. and Pollio, A. (2007) Species composition of cyanidiales assemblages in Pisciarelli (Campi Flegrei, Italy) and description of Galdieria phlegrea sp. nov., In: J. Seckbach (ed.) Algae and Cyanobacteria in Extreme Environments. Springer, Dordrecht, The Netherlands, pp. 487–502.

    Google Scholar 

  • Pueschel, C.M. (1990) Cell structure, In: K.M. Cole and R.G. Sheath (eds.) Biology of the Red Algae. Cambridge University Press, Cambridge, UK, pp. 7–41.

    Google Scholar 

  • Ragan, M.A., Bird, C.J., Rice, E.L., Gutell, R.R., Murphy, C.A. and Singh, R.K. (1994) A molecular phylogeny of marine red algae (Rhodophyta) based on the nuclear small-subunit rRNA gene. Proc. Natl. Acad. Sci. USA 91: 7276–7280.

    Article  PubMed  CAS  Google Scholar 

  • Rodríguez-Ezpeleta, N., Brinkmann, H., Burey, S.C., Roure, B., Burger, G., Löffelhardt, W., Bohnert, H.J., Philippe, H. and Lang, B.F. (2005) Monophyly of primary photosynthetic eukaryotes: green plants, red algae, and glaucophytes. Curr. Biol. 15: 1325–1330.

    Article  PubMed  Google Scholar 

  • Rintoul, T.L., Sheath, R.G. and Vis, M.L. (1999) Systematics and biogeography of the Compsopogonales (Rhodophyta) with emphasis on the freshwater families in North America. Phycologia 38: 517–527.

    Article  Google Scholar 

  • Saunders, G.W. and Hommersand, M.H. (2004) Assessing red algal supraordinal diversity and taxonomy in the context of contemporary systematic data. Am. J. Bot. 91: 1494–1507.

    Article  PubMed  Google Scholar 

  • Saunders, G.W. and Kraft, G.T. (1997) A molecular perspective on red algal evolution: focus on the Florideophycidae. Plant Syst. Evol. (Suppl.) 11: 115–138.

    Article  CAS  Google Scholar 

  • Scott, J.L., Broadwater, S.T., Saunders, B.D. and Thomas, J.P. (1992) Ultrastructure of vegetative organization and cell-division in the unicellular red alga Dixoniella grisea gen. nov. (Rhodophyta) and a consideration of the genus Rhodella. J. Phycol. 28: 649–660.

    Article  Google Scholar 

  • Scott, J.L., Yokoyama, A., Billard, C., Fresnel, J., Hara Y., West K.A. and West, J.A. (2008) Neorhodella cyanea, a new genus in the Rhodellophyceae (Rhodophyta). Phycologia 46: 560–572.

    Article  Google Scholar 

  • Scott, J.L., Barca, B., Ott, F.D. and West, J.A. (2006) Light and electron microscopic observations of Erythrobolus coxiae gen. et sp. nov. (Porphyrdiophyceae, Rhodophyta) from Texas, U.S.A. Algae 21: 407–416.

    Article  Google Scholar 

  • Seckbach, J. (1987) Evolution of eukaryotic cells via bridge algae, the cyanidia connection. Ann. N Y Acad. Sci. 503: 424–437.

    Article  Google Scholar 

  • Sheath, R.G. (2003) Red algae, In: J.D. Wehr and R.G. Sheath (eds.) Freshwater Algae of North America Ecology and Classification. Academic, San Diego, CA, pp. 197–224.

    Google Scholar 

  • Sheath, R.G. and Cole, K.M. (1984) Systematics of Bangia (Rhodophyta) in North America. I. Biogeographic trends in morphology. Phycologia 23: 383–396.

    Article  Google Scholar 

  • Sheath, R.G. and Hambrook, J.A. (1990) Freshwater ecology, In: K.M. Cole and R.G. Sheath (eds.) Biology of the Red Algae. Cambridge University Press, Cambridge, UK, pp. 423–453.

    Google Scholar 

  • Sheath, R.G. and Sherwood, A.R. (2002) Phylum Rhodophyta (Red Algae), In: D.M. John, B.A. Whitton and A.J. Brook (eds.) The Freshwater Algal Flora of the British Isles. Cambridge University Press, Cambridge, UK, pp. 123–143.

    Google Scholar 

  • Silva, P.C., Basson, P.W. and Moe, R.L. (1996) Catalogue of the Benthic Marine Algae of the Indian Ocean. University of California Publications in Botany, Berkeley, CA.

    Google Scholar 

  • Simons, J., Schulp, H. and Stegenga, H. (2001) Erythrocladia setifera sp. nov. (Rhodophyta: Erythropeltidales/Compsopgonales), a new species from freshwater lakes in the Netherlands. Phycologia 40: 475–482.

    Article  Google Scholar 

  • Stiller, J.W. and Hall, B.D. (1997) The origin of red algae: implications for plasmid evolution. Proc. Natl. Acad. Sci. USA 94: 4520–4525.

    Article  PubMed  CAS  Google Scholar 

  • Stiller, J.W. and Waaland, J.R. (1993) Molecular analysis reveals cryptic diversity in Porphyra (Rhodophyta). J. Phycol. 29: 506–517.

    Article  CAS  Google Scholar 

  • Toplin, J.A., Norris, T.B., Lehr, C.R., McDermott, T.R. and Castenholz, R.W. (2008) Biogeographic and phylogenetic diversity of thermoacidophilic Cyanidiales in Yellowstone National Park, Japan, and New Zealand. Appl. Environ. Microbiol. 74: 2822–2833.

    Article  PubMed  CAS  Google Scholar 

  • Ueki, C., Nagasato, C., Motomura, T. and Saga, N. (2008) Reeximination of the pit plugs and characteristic membranous structures in Porphyra yezoenis (Bangiales, Rhodophyta). Phycologia 47: 5–11.

    Article  Google Scholar 

  • van den Hoek, C., Mann, D.G. and Jahns, H.M. (1995) Algae, An Introduction to Phycology. Cambridge University Press, Cambridge, UK.

    Google Scholar 

  • Vis, M.L. and Sheath, R.G. (1993) Distribution and systematics of Chroodactylon and Kyliniella (Porphyridiales, Rhodophyta) from North American streams. Jpn. J. Phycol. 41: 237–241.

    Google Scholar 

  • Weber, A.P.M., Barbier, G.G., Shrestha, R.P., Horst, R.J., Minoda, A. and Oesterhelt, C. (2007) A genomics approach to understanding the biology of thermo-acidophilic red algae, In: J. Seckbach (ed.) Algae and Cyanobacteria in Extreme Environments. Springer, Dordrecht, The Netherlands, pp. 503–518.

    Google Scholar 

  • West, J.A., Zuccarello, G.C., Scott, J.L., West, K.A. and deGoer, S.L. (2007a) Pulvinus veneticus gen. et sp. nov. (Compsopogonales, Rhodophyta) from Vauatu. Phycologia 46: 237–246.

    Article  Google Scholar 

  • West, J.A., Zuccarello, G.C., Scott, J.L., West, K.A. and Karsten, U. (2007b) Rhodaphnes brevistipitata gen. et sp. nov., a new member of the Stylonematophyceae. Phycologia 46: 440–449.

    Article  Google Scholar 

  • Wujek, D.E. and Timpano, P. (1986) Rufusia (Porphyridiales, Phragmonemataceae), a new red alga from sloth hair. Brenesia 25/26: 163–168.

    Google Scholar 

  • Yokoyama, A., Sato, K. and Hara, Y. (2004) The generic delimitation of Rhodella (Porphyridiales, Rhodophyta) with emphasis on ultrastructure and molecular phylogeny. Hydrobiologia 512: 177–183

    Article  CAS  Google Scholar 

  • Yoon, H.S., Ciniglia, C., Wu, M., Comeron, J.M., Pinto, G., Pollio, A. and Bhattacharya, D. (2006a) Establishment of endolithic populations of extremophilic Cyanidiales (Rhodophyta). BMC Evol. Biol. 6: 78.

    Article  PubMed  Google Scholar 

  • Yoon, H.S., Müller, K.M., Sheath, R.G., Ott, F.D. and Bhattacharya, D. (2006b) Defining the major lineages of red algae (Rhodophyta). J. Phycol. 42: 482–492.

    Article  CAS  Google Scholar 

  • Zemke-White, W.L. and Ohno, M. (1999) World seaweed utilisation: an end-of-century summary. J. Appl. Phycol. 11: 369–376.

    Article  Google Scholar 

  • Zuccarello, G., West, J., Bitans, A. and Kraft, G. (2000) Molecular phylogeny of Rhodochaete parvula (Bangiophycidae, Rhodophyta). Phycologia 39: 75–81.

    Article  Google Scholar 

  • Zuccarello, G.C., West, J.A. and Kikuchi, N. (2008) Phylogenetic relationships within the Stylonematales (Stylonematophyceae, Rhodohyta): biogeographic patterns do not apply to Stylonema alsidii. J. Phycol. 44: 384–393.

    Article  CAS  Google Scholar 

Download references

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Kirsten M. Müller .

Editor information

Editors and Affiliations

Rights and permissions

Reprints and permissions

Copyright information

© 2010 Springer Science+Business Media B.V.

About this chapter

Cite this chapter

Müller, K.M., Lynch, M.D.J., Sheath, R.G. (2010). Bangiophytes: From one Class to Six; Where Do We Go from Here?. In: Seckbach, J., Chapman, D. (eds) Red Algae in the Genomic Age. Cellular Origin, Life in Extreme Habitats and Astrobiology, vol 13. Springer, Dordrecht. https://doi.org/10.1007/978-90-481-3795-4_13

Download citation

Publish with us

Policies and ethics