Charles A. Dana Professor of Biology - Trinity College, Hartford, CT

History of Seaweed Work in Bermuda

Reports of benthic (attached) marine red, green, and brown algae from Bermuda began to appear in the literature in the last half of the 19th century based upon collections made on just a few voyages, including the “Challenger Expedition,” to the islands (Kemp 1857; Rein 1873; Dickie 1874; Hemsley 1884; Murray 1888, 1889). Some early Bermuda specimens were distributed as part of the Algae exsiccatae Americae Borealis (A.A.B., Farlow et al. 1877–1889).


Frank Shipley Collins (1848-1920)
Frank Shipley Collins (1848–1920)
Rev. Alpheus Baker Hervey (1839–1931)

In 1917, Frank Shipley Collins and the Rev. Alpheus Baker Hervey produced the first comprehensive marine flora of the islands and distributed the bulk of their specimens in five fascicles entitled ‘The Algae of Bermuda’ as part of Phycotheca Boreali-Americana (P.B.-A., Collins et al. 1912–1917). Collins was an accountant for the Boston Rubber Shoe Company, and in his spare time, an ardent amateur phycologist. Although his work with marine and freshwater algae was accurate and scholarly, Collins frequently conferred and corresponded with Wm. Gilson Farlow (1844–1919), the prominent cryptogamic botanist at Harvard University who had earlier headed the effort to distribute the A.A.B. (Taylor 1945). Like Collins, Hervey was an amateur phycologist, variously having positions as rector of a Universalist Church and President of St. Lawrence University.


William Randolph Taylor (1895-1990)
William Randolph Taylor (1895–1990)
Marshall Avery Howe (1867-1936)
Marshall Avery Howe  (1867–1936)

A year after Collins & Hervey’s (1917) Algae of Bermuda appeared, Marshall Avery Howe (1918) contributed the section, ‘Algae’, in Nathaniel Lord Britton’s Flora of Bermuda, who only included “the more common and more conspicuous algae occurring in the islands.” For many years, this was the last floristic report of Bermuda marine algae. In 1949, Wm. Randolph Taylor of the University of Michigan first visited Bermuda with his student, Albert J. Bernatowicz (Taylor 1952), and he included their data along with previous collectors in his comprehensive Marine Algae of the Eastern Tropical and Subtropical Coasts of the Americas (1960). Later, Taylor & Bernatowicz (1969) produced an annotated list of the most common shallow water macroscopic seaweeds of the Bermudas. Since then and prior to our work, additions to the Bermuda marine flora were published only sporadically.


Richard B. Searles
Richard B. Searles

Richard B. Searles, now Emeritus Professor of Botany at Duke University, was a student of George F. Papenfuss at UC Berkeley receiving his Ph.D. in 1965. I initiated my graduate studies with Rick at Duke in 1970, finishing my Ph.D. in 1975, then taking up my present post at in the Biology Department at Trinity College, Hartford. After we completed our studies on the seaweeds of the southeastern United States concentrated in the Carolinas, we had two deep-water diving expeditions in Bermuda. Using Scuba and Surface-Supplied Air (SSA), we collected seaweeds from 12-50 m depths around the islands on the NOAA funded R/V Seahawk cruises of 1983 and 1985 (Searles & Schneider 1987). These offshore reefs had been neglected phycologically for the most part prior to that time. W.R. Taylor (1952) found the bordering reefs to be “seldom accessible because of the long boat trips involved and the rarity of calm days safe for the arduous work involved.” He further stated that “dredging from deep water contributed little to [his Bermuda] study, a regrettable lack from the floristic standpoint.” In the only deep water study of note, John Joseph Frederick’s dissertation (under Taylor, University of Michigan, 1963) compiled a list of the seaweeds found on the Challenger and Argus Banks offshore of the Bermuda platform, an area unlike most others in its lack of coral pinnacles. Frederick’s specimens at the University of Michigan were collected in the early 1960s by lockout Navy divers working on the seamount platforms. My work with Rick in Bermuda started the “Notes on the marine algae of the Bermudas series” (Schneider & Searles 1997a, 1997b, 1998a, 1998b).

Craig W. Schneider (L) & Christopher E. Lane (R)
Craig W. Schneider (L) & Christopher E. Lane (R)

My former undergraduate student, Chris Lane (B.S., Trinity College ‘99), has been working with me in Bermuda since 1999 and during that time frame earned his Ph.D. on macroalgal molecular phylogenetics at the University of New Brunswick in Gary Saunders’ seaweed lab. Chris is currently professor of molecular genetics at the University of Rhode Island. Another former student of mine, Dan McDevit (B.S., Trinity College ‘01; Ph.D. ‘10 also from Gary Saunders’ UNB lab), joined us on two collecting trips helping us and then URI grad student, Thea Popolizio (Ph.D. 2015, now an assistant professor at Salem State University), with barcoding all of the marine macrophytes of Bermuda and comparable material from Key West and St. Croix. Dan is currently an Asst. Professor of Biology at the University of Delaware. Due to mutual interests in regional floristics and phylogenetics, I began working with Gary Saunders in 2006 and we have jointly published a swatch of seaweed papers both on Bermuda samples as well as those from many regions including New England, Canada and Australasia. With all of the collaborators mentioned, as well as many of my Trinity undergraduate thesis and research students, we have collectively published papers that include well over 100 new records, new genera and species from offshore, as well as inshore, Bermuda habitats (Dixon et al. 2015; Popolizio et al. 2013, 2015; Richards et al. 2019; Saunders et al. 2006; Schneider 2000, 2004; Schneider & Lane 2005, 2007, 2008; Schneider et al. 2006, 2010, 2011a, 2011b, 2012, 2014a, 2014b, 2015, 2016a, 2016b, 2017, 2018a, 2018b, 2018c, 2019a, 2019b, 2019c, 2020; Schneider & Wynne 2009; Schultz et al. 2015; Wynne & Schneider 1996). Some of these taxa, newly reported for Bermuda, are widespread at least in the warm portions of the Atlantic Ocean, especially the Caribbean Sea. In these papers, based upon island collections, new genera include:

  • Archestenogramma
  • Crassitegula
  • Frikkiella
  • Howella

and new species:  

  • Antithamnionella bermudica
  • Archestenogramma profundum
  • Asteromenia bermudensis
  • Botryocladia bermudana
  • B. exquisita
  • B. flookii
  • Centroceras arcii
  • C. illaqueans
  • Chondracanthus saundersii
  • Crassitegula laciniata
  • C. walsinghamii
  • Champia hasselbringii
  • C. insularis
  • Crouania elisiae
  • Cryptonemia abyssalis
  • C. antricola
  • C. atrocostalis
  • C. lacunicola
  • C. perparva
  • Dasya cryptica
  • Dichotomaria huismanii
  • Digenea arenahauriens
  • Ethelia umbricola
  • Frikkiella searlesii
  • Galene leptoclados
  • Griffithsia aestivana
  • Halopeltis pellucida
  • Helminthocladia kempii
  • Howella gorgoniarum
  • Liagora nesophila
  • Lobophora declerckii
  • L. payriae
  • Meredithia crenata
  • Meristotheca odontoloma
  • Nothocladia erosa
  • Polysiphonia plectocarpa
  • Psaromenia septentrionalis
  • Sporolithon mesophoticum
  • Tepoztequiella muriamans
  • Woelkerlingia sterreri
  • Yamadaella grassyi

All new records collected prior to 2003 were compiled with all historical reports into a checklist of the Bermuda red, green and brown seaweeds (Schneider 2003). Much of this work has been accomplished by molecular-assisted alpha taxonomy (MAAT), and a summary of those practices is outlined by Cianciola et al. (2010).


Nomad and Nemo at 200 m
Submersibles Nomad and Nemo at 200 m

In August of 2016, Thea Popolizio and I were scientists aboard the R/V Baseline Explorer, where specimens were gathered in the mesophotic zone of Bermuda from 50-178 m using submersibles and technical (rebreather) divers. Our specimens were dried on silica gel for MAAT studies, and we prepared herbarium vouchers for future work and distribution. Some of these specimens have already been reported on after molecular analysis and morphological study in my lab at Trinity College (Richards et al. 2019; Schneider et al. 2018b, 2019a, 2019b, 2020; Stefanoudis et al. 2019). Many of the 2016 samples are the same as those reported by J.J. Frederick from the offshore seamounts in 1963, and molecular analyses allowed for their correct placement in new genera and species. An online field guide to many of the common species of plants and animals we collected on the cruise is available as Stefanoudis et al. (2018). This deep-water expedition was supported by the Nekton Foundation, Oxford, England.


Bermuda Seaweeds as Art

I had the opportunity to highlight several Bermuda seaweeds in an art show of greatly enlarged digital prints entitled “Forms Most Beautiful and Most Wonderful” at a Trinity gallery in September, 2018. See the link below.

Forms Most Beautiful


Literature Cited

  • Cianciola, E.N., T. Popolizio, C.W. Schneider & C.E. Lane (2010). Using molecular-assisted alpha taxonomy to better understand red algal biodiversity in Bermuda. Diversity 2010: 946–958. 
  • Collins, F.S. & A.B. Hervey (1917). The algae of Bermuda. Proceedings of the American Academy of Arts & Sciences 53: 1–195, pls 1-6.
  • Collins, F.S., I. Holden & W.A. Setchell (1912–1917). Phycotheca Boreali-Americana (Exsiccata), Algae of North America – Algae of Bermuda. Malden, Mass.
  • Dickie, G. (1874). I. On the marine algae of St. Thomas and the Bermudas, and on Halophila baillonis, Asch., pp. 311–317. In, Hooker, J.D. Contributions to the botany of the expedition of H.M.S.‘Challenger.’ Journal of the Linnaean Society, Botany 14: 311–390 (1874–1875).
  • Dixon, K.R., G.W. Saunders, C.W. Schneider & C.E. Lane (2015). Etheliaceae fam. nov. (Gigartinales, Rhodophyta), with a clarification of the generitype of Ethelia and the addition of six novel species from warm waters. Journal of Phycology 51: 1158–1171.
  • Farlow, W.G., C.L. Anderson & D.C. Eaton (1877–1889). Algae exsiccatae Americae Borealis. Fascicles I–V. Boston, Mass.
  • Frederick, J.J. (1963). The Marine Algae of the Bermuda Platform. Ph.D. dissertation, Univ. Michigan. v + 89 pp.
  • Hemsley, W.B. (1884). Algae, i, ii, pp.104–128. In, Report of the botany of the Bermudas and various other islands of the Atlantic and southern Oceans. Report of the Scientific Research Exploring Voyage of H.M.S. “Challenger,” 1873–1876. Botany 1 (2): 1–135, 13 pls.
  • Howe, M.A. (1918). Algae, pp. 489–540. In Britton, N.L., Flora of Bermuda. Charles Scribner’s Sons, New York. ix + 585 pp.
  • Kemp, A.F. (1857). Notes on the Bermudas and their natural history, with special reference to their marine algae. Canadian Naturalist and Geologist 2: 145–156.
  • Murray, G. (1888). Catalogue of the marine algae of the West Indian region. I.-Florideae, II.-Phaeophyceae. Journal of Botany, British and Foreign 26: 193–196; 237–243, 303–307; 331–338; 358–363, pl. 284.
  • Murray, G. (1889). Catalogue of the marine algae of the West Indian region. III.-Chlorophyceae, IV.-Protophyceae, Geographical distribution. Journal of Botany, British and Foreign 27: 237–242, 257–262, 298–305.
  • Popolizio, T.R., C.W. Schneider, T. Chengsupanimit, G.W. Saunders & C.E. Lane (2013). Notes on the marine algae of the Bermudas. 13. Helminthocladia kempii sp. nov. (Nemaliales, Liagoraceae) based upon H. calvadosii sensu auct. from the western Atlantic. Cryptogamie, Algologie 34: 229–244.
  • Popolizio, T.R., C.W. Schneider & C.E. Lane (2015). A molecular evaluation of the Liagoraceae sensu lato (Nemaliales, Rhodophyta) in Bermuda including Liagora nesophila sp. nov. and Yamadaella grassyi sp. nov. Journal of Phycology 51: 637–658.
  • Rein, J. J. (1873). Über die Vegetations- Verhältnisse der Bermudas-Inseln. Bericht uber die Senckenbergische Naturforschende Gesellschaft in Frankfurt am Main  1872–1873: 131–153.
  • Richards, J.L., P.W. Gabrielson & C.W. Schneider (2018). Sporolithon mesophoticum sp. nov. (Sporolithales, Rhodophyta) from Plantagenet Bank off Bermuda at a depth of 178 m. Phytotaxa 385: 67–76
  • Saunders, G.W., C.E. Lane, C.W. Schneider & G.T. Kraft (2006). Unraveling the Asteromenia peltata species complex with clarification of the genera Halichrysis and Drouetia (Rhodymeniaceae, Rhodophyta). Canadian Journal of Botany 84: 1581–1607.
  • Schneider, C.W. (2000). Notes on the marine algae of the Bermudas. 5. Some Delesseriaceae (Ceramiales, Rhodophyta), including the first record of Hypoglossum barbatum Okamura from the Atlantic Ocean. Botanica Marina 43: 455–466.
  • Schneider, C.W. (2003). An annotated checklist and bibliography of the marine macroalgae of the Bermuda islands. Nova Hedwigia 37: 275–361.
  • Schneider, C.W. (2004). Notes on the marine algae of the Bermudas. 6. Some rare or newly reported Ceramiales (Rhodophyta), including Crouania elisiae sp. nov. Phycologia 43: 563–578.
  • Schneider, C.W., E.N. Cianciola, T.R. Popolizio, D.S. Spagnuolo & C.E. Lane (2015). A molecular-assisted alpha taxonomic study of the genus Centroceras (Ceramiaceae, Rhodophyta) in Bermuda reveals two novel species. Algae 30: 15–33.
  • Schneider, C.W., T. Chengsupanimit & G.W. Saunders (2011). A new genus and species from the North Atlantic, Archestenogramma profundum (Phyllophoraceae, Rhodophyta), with taxonomic resolution of the orphaned Leptofauchea brasiliensis. European Journal of Phycology 46: 442–452.
  • Schneider, C.W., C.T. Flook & P.M. Rouja (2016b). An abundant sand flat meadow of Halimeda lacrimosa M.Howe off Bermuda, a species previously unknown in the island flora. Notulae algarum 17: 1–4
  • Schneider, C.W., D.W. Freshwater & G.W. Saunders (2012). First report of Halopeltis (Rhodophyta, Rhodymeniaceae) from the non-tropical Northern Hemisphere: H. adnata (Okamura) comb. nov. from Korea, and H. pellucida sp. nov. and H. willisii sp. nov. from the North Atlantic. Algae 27: 95–108.
  • Schneider, C.W., M.K. Griffith, C.E. Lane & G.W. Saunders (2018c). Notes on the marine algae of the Bermudas. 16. Two new epiphytic species of Champia (Champiaceae, Rhodymeniales), C. hasselbringii and C. insularis. Cryptogamie, Algologie 39: 431–447.
  • Schneider, C.W., B.F. Hamzeh, C.E. Lane & G.W. Saunders (2018a). A new species of Digenea (Rhodomelaceae, Ceramiales) based upon a molecular assessment and morphological observations of plants historically known as D. simplex in Bermuda. Phytotaxa 338 (1): 90–98.
  • Schneider, C.W., D.W. Lam & H. Verbruggen (2019c) [‘2020’]. DNA sequencing and anatomy demonstrate that Pacific Codium simulans is a genetically variable species found in the floras of Bermuda and Florida. Phycological Research 68: 98­–102.
  • Schneider, C.W. & C.E. Lane (2005). Notes on the marine algae of the Bermudas. 7. Additions to the flora, including Chondracanthus saundersii sp. nov. (Rhodophyta, Gigartinaceae) based on rbcL sequence analysis. Phycologia 44: 72–83.
  • Schneider, C.W. & C.E. Lane (2007). Notes on the marine algae of the Bermudas. 8. Further additions to the flora including Griffithsia aestivana sp. nov. (Ceramiaceae, Rhodophyta) and an update on the alien Cystoseira compressa (Sargassaceae, Heterokontophyta). Botanica Marina 50: 128–140.
  • Schneider, C.W. & C.E. Lane (2008). Notes on the marine algae of the Bermudas. 9. The genus Botryocladia (Rhodophyta, Rhodymeniaceae), including B. bermudana, B. exquisita and B. flookii spp. nov. Phycologia 47: 614–629.
  • Schneider, C.W., C.E. Lane & G.W. Saunders (2006). Crassitegula walsinghamii (Sebdeniaceae, Halymeniales), a new red algal genus and species from Bermuda based upon morphology and SSU rDNA sequence analyses. European Journal of Phycology 41: 115–124.
  • Schneider, C.W., C.E. Lane & G.W. Saunders (2010). Notes on the marine algae of the Bermudas. 11. More additions to the benthic flora and a phylogenetic assessment of Halymenia pseudofloresii (Halymeniales, Rhodophyta) from its type locality. Phycologia 46: 154–168.
  • Schneider, C.W., C.E. Lane & G.W. Saunders (2018b). A revision of the genus Cryptonemia (Halymeniales, Rhodophyta) in Bermuda, western Atlantic Ocean, including five new species and C. bermudensis (Collins et M. Howe) comb. nov. European Journal of Phycology 53: 350–368.
  • Schneider, C.W., D.C. McDevit, G.W. Saunders & C.E. Lane (2011). Notes on the marine algae of the Bermudas. 12. A phylogenetic assessment of Nemastoma gelatinosum M. Howe (Rhodophyta, Nemastomatales) from its type locality. Cryptogamie, Algologie 32: 313–325.
  • Schneider, C.W., E.S. Peterson & G.W. Saunders (2020). Two new species of Solieriaceae (Rhodophyta, Gigartinales) from the euphotic and mesophotic zones off Bermuda, Meristotheca odontoloma and Tepoztequiella muriamans. Phycologia 59: 177–185.
  • Schneider, C.W., T.R. Popolizio, L.G.K. Kraft & G.W. Saunders (2019b). New species of  Galene and Howella gen. nov. (Halymeniaceae, Rhodophyta) from the mesophotic zone off Bermuda. Phycologia 59: 690–697.
  • Schneider, C.W., T.R. Popolizio & C.E. Lane (2014a). Notes on the marine algae of the Bermudas. 14. Five additions to the benthic flora, including a distinctive second new species of Crassitegula (Rhodophyta, Sebdeniales) from the western Atlantic Ocean. Phycologia 53: 117–126.
  • Schneider, C.W., T.R. Popolizio & G.W. Saunders (2019a). Collections from the mesophotic zone off Bermuda reveal three species of Kallymeniaceae (Gigartinales, Rhodophyta) in genera with transoceanic distributions. Journal of Phycology 55: 415–424.
  • Schneider, C.W., T.R. Popolizio, D.S. Spagnuolo & C.E. Lane (2016a). Notes on the marine algae of the Bermudas. 15. Dichotomaria huismanii (Galaxauraceae, Rhodophyta), a new species in the D. marginata complex from the western Atlantic. Botanica Marina 59: 13–29.
  • Schneider, C.W., P.K. Quach & C.E. Lane (2017). A case for true morphological crypsis: Pacific Dasya anastomosans and Atlantic D. cryptica sp. nov. (Dasyaceae, Rhodophyta). Phycologia 56: 359–368. 
  • Schneider, C.W., G.W. Saunders & C.E. Lane (2014b). The monospecific genus Meredithia (Kallymeniaceae, Gigartinales) is species rich and geographically widespread with species from temperate Atlantic, Pacific and Indian Oceans. Journal of Phycology 50: 167–186.
  • Schneider, C.W. & R.B. Searles (1997a). Notes on the marine algae of the Bermudas. 1. New records of Antithamnieae and Dohrnielleae (Ceramiaceae, Rhodophyta), including Antithamnionella bermudica sp. nov. Phycologia 36: 12–23.
  • Schneider, C.W. & R.B. Searles (1997b). Notes on the marine algae of the Bermudas. 2. Some Rhodophyta, including Polysiphonia tongatensis and a discussion of the Herposiphonia secunda/tenella complex. Cryptogamie, Algologie 18: 187–210.
  • Schneider, C.W. & R.B. Searles (1998a). Notes on the marine algae of the Bermudas. 3. Avrainvillea sylvearleae, Discosporangium mesarthrocarpum and Peyssonnelia valentinii. Journal of Phycology 34: 180–188.
  • Schneider, C.W. & R.B. Searles (1998b). Notes on the marine algae of the Bermudas. 4. Additions to the flora, including Polysiphonia plectocarpa sp. nov. Phycologia 37: 24–33.
  • Searles, R.B. & Schneider, C.W. (1987). Observations on the deep-water flora of Bermuda. Hydrobiologia 151/152: 261–266.
  • Schneider, C.W. & M.J. Wynne (2009) [‘2008’]. Notes on the marine algae of the Bermudas. 10. Woelkerlingia sterreri sp. nov. (Rhodophyta, Ceramiaceae), a first record of the genus in the western Atlantic. Caribbean Journal of Science 44: 303–310.
  • Schultz, N.E., C.E. Lane, L. Le Gall, D. Gey, A.R. Bigney, B. de Reviers, F. Rousseau & C.W. Schneider (2015). A barcode analysis of the genus Lobophora (Dictyotales, Phaeophyceae) in the western Atlantic Ocean with four novel species and the epitypification of L. variegata (J.V. Lamouroux) E.C. Oliveira. European Journal of Phycology 50: 481–500.
  • Stefanoudis, P.V., S.R. Smith, C. Schneider, D. Wagner, M. Rivers, G. Goodbody-Gringley, J. Xavier, L.C. Woodall & A.D. Rogers (2018). Deep Reef Benthos of Bermuda: Field Identification Guide. Nekton: Oxford, England. 168 pp., available for download at Figshare:
  • Stefanoudis, P.V., M. Rivers, S.R. Smith, C.W. Schneider, D. Wagner, H. Ford, A.D. Rogers& L.C. Woodall (2019). Low connectivity between shallow, mesophotic and rariphotic reef communities. Royal Society Open Science 6: 190958
  • Taylor, W.R. (1945). William Gilson Farlow. Promoter of phycological research in America. Farlowia 2: 53–70.
  • Taylor, W.R. (1952). Survey of the marine algae of Bermuda. Yearbook of the American Philosophical Society 1951: 167–171.
  • Taylor, W.R. (1960). Marine Algae of the Eastern Tropical and Subtropical Coasts of the Americas. University of Michigan Press, Ann Arbor. xi + 879 pp.
  • Taylor, W.R. & A.J. Bernatowicz (1969). Distribution of marine algae about Bermuda. Bermuda Biological Station, Special Publications 1: 1–42.
  • Wynne, M.J. & C.W. Schneider (1996). Frikkiella gen. nov. (Delesseriaceae, Rhodophyta) from Bermuda and the Caribbean Sea. Systematic Botany 21: 77-84.