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Plicate Species of the Diatom Genus Thalassiosira(Bacillariophyta)

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Plicate Species of the Diatom Genus Thalassiosira(Bacillariophyta) FAU Institutional Repository http://purl.fcla.edu/fau/fauir This paper was submitted by the faculty of FAU’s Harbor Branch Oceanographic Institute. Notice: ©2011 Academy of Natural Sciences. This manuscript may be cited as: Prasad, A.K.S.K., Nienow, J. A., & Hargraves, P. (2011). Plicate species of the diatom genus Thalassiosira (Bacillariophyta) from the Atlantic and Gulf coasts of southeastern United States, with the description of T. livingstoniorum sp. nov. Proceedings of the Academy of Natural Sciences of Philadelphia, 161(1), 1-34. doi: 10.1635/053.161.0101 ISSN 0097-3157 PROCEEDINGS OF THE ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA 161: 1-34 OCTOBER 2011 (NOVEMBER 1, 2011) Plicate species of the diatom genus Thalassiosira (Bacillariophyta) from the Atlantic and Gulf coasts of southeastern United States, with the description of T. livingstoniorum sp. nov. AKSHINTHALA K. S. K. PRASAD* Department of Biological Science, Florida State University, Tallahassee FL 32306-4370 USA E-mail: [email protected] JAMES A. NIENOW Department of Biology, Valdosta State University, Valdosta GA 31698-1500, USA PAUL HARGRAVES Graduate School of Oceanography, University of Rhode Island, Narragansett RI 02882-1197; Harbor Branch Oceanographic Institute, Florida Atlantic University, FL, USA; and Smithsonian Marine Station, Ft. Pierce, FL, USA *Corresponding author ABSTRACT.—Long-term observations of a marine planktonic plicate Thalassiosira species, T. cedarkeyensis Prasad, from the Gulf coasts of Florida, Alabama, and Mississippi and the Atlantic coasts of Florida and Georgia demonstrate its wide RFFXUUHQFHLQWKHVRXWKHDVWHUQ8QLWHG6WDWHVDQGLWVDELOLW\WRIRUPH[WHQVLYHEORRPV:HDOVRUHSRUWIRUWKHÀUVWWLPHLWVDELOLW\ to form typical Thalassiosira chains, linking sibling cells by threads of chitin. A closely related and co-occurring diatom, T. livingstoniorum, is described on the basis of investigations conducted during 2000–2011 as a new species from many localities in Apalachee Bay on the Gulf coast and Indian River Lagoon on the Atlantic coast of Florida. It differs from T. cedarkeyensis in areola density, presence of continuous cribra on loculate areolae, arrangement and distribution of valve processes, and the number of satellite pores surrounding the valve-face fultoportulae. We have not yet found any evidence of chain formation in T. livingstoniorum. Thalassiosira cedarkeyensis and T. livingstoniorum can be easily distinguished in Naphrax-mounted preparations in light microscopy (LM), and they represent two different groups (lineages?) of plicate species with reference to internal cribrum structure of the loculate areolae. Their differences may justify placement in two different genera. Although T. cedarkeyensis (which has individual cribra on the proximal siliceous layer like T. hyperborea) is abundant and widespread on the Gulf and Atlantic coasts of the southeastern United States, T. livingstoniorum (which has continuous cribra like T. lacustris) has been found, thus far, only in Florida coastal waters. Comparisons are made between these two species and the other morphologically similar extant and extinct plicate species. Thalassiosira livingstoniorum and T. cedarkeyensis, although widespread and frequently encountered during warmer months, may be easily overlooked sources of primary production in the nutrient-rich northeastern Gulf of Mexico. Key words: Thalassiosira cedarkeyensis, T. livingstoniorum sp. nov., plicate valves, Indian River Lagoon, Gulf coast of Florida, diatom EORRPV)HQKROORZD\HVWXDU\(FRQÀQDHVWXDU\ INTRODUCTION on the valve serve as diagnostic features within the genus and emended its description. At the same time she Cleve (1873) described the centric diatom genus resurrected the family Thalassiosiraceae Lebour 1930 Thalassiosira from a surface bloom of T. nordenskioeldii DQG H[SDQGHG LWV GHÀQLWLRQ WR LQFOXGH DOO JHQHUD ZLWK in the Davis Strait. Although the genus was still considered fultoportulae (strutted processes; Hasle, 1973). Glezer and monotypic in Van Heurck (1896), by 1935, 23 species Makarova (1986) elevated Hasle’s concept of the family were included in the genus (Mills, 1935). By 1971 Thalassiosiraceae to ordinal level and separated out forms (Makarova, 1971a, 1971b) the number of species and with radial rows of areolae separated by hyaline rays into a LQIUDVSHFLÀFWD[DLQFOXGHGLQWKHJHQXVKDGJURZQWR separate family of the order, Stephanodiscaceae Makarova 32 in the seas of the former U.S.S.R. alone. Hasle (1968, LQ *OH]HU DQG 0DNDURYD $GGLWLRQDO UHÀQHPHQWV 1973) suggested that the type and pattern of processes IROORZHG VR WKDW FXUUHQWO\ ÀYH IDPLOLHV DUH UHFRJQL]HG 2 A.K.S.K. PRASAD, J.A. NIENOW & P. HARGRAVES in the order—Thalassiosiraceae, Stephanodiscaceae, subgroup of the genus Thalassiosira by Hasle and Lange Lauderiaceae (Schütt) emend Medlin and Kaczmarska WKH\ GLG QRW KRZHYHU DFFRUG DQ\ WD[RQRPLF (Kaczmarska et al., 2005), Skeletonemataceae Lebour status to the group. A later phenetic analysis by Julius (1930), and Ectodictyonaceae Khursevich and Cherniaeva and Tanimura (2001) of nineteen fossil and living plicate (1989). These are distinguished primarily on the basis of species supported the hypothesis that the plicate species areolar structure and arrangement and process structure form a natural group containing at least two major clades. and arrangement. The Thalassiosiraceae, as currently These groups will be discussed in more detail later. XQGHUVWRRGFRQWDLQVDSSUR[LPDWHO\WZHQW\YDOLGO\QDPHG The nine previously described living plicate species genera (Wolfe & Siver, 2009), including the recently include forms inhabiting freshwater, brackish-water, or established Shionodiscus Alverson, Kang et Theriot marine environments: T. australiensis (Grunow) Hasle 2006, Spicaticribra Johansen, Kociolek et Lowe 2008, et Lange (1989), T. bramaputrae (Ehrenberg) Håkansson Conticribra Stachura-Suchoples et Williams 2009 and et Locker (1981), T. cedarkeyensis Prasad in Prasad, Livingstonia Prasad et Nienow 2011. )U\[HOOHW/LYLQJVWRQ T. gessneri Hustedt (1956), Thalassiosira remains the most diverse and T. hyperborea (Grunow) Hasle in Hasle and Lange H[WHQVLYHO\ VWXGLHG JHQXV ZLWKLQ WKH 7KDODVVLRVLUDFHDH (1989), T. lacustris *UXQRZ +DVOH HW )U\[HOO with a world-wide distribution in fresh, brackish and T. patagonica Maidana (1999), T. plicata Schrader marine waters, and in polar, temperate and tropical oceans. 6FKUDGHU 0DNDURYD 6DQFHWWD DQG The vast majority of the species have at least one ring T. plicatoides (Simonsen) Akiba et Yanagisawa (1986). of marginal fultoportulae, and one or more central (or T. cedarkeyensis, a small marine species with occluded near central) fultoportulae; many also have additional SURFHVVHV ZDV RULJLQDOO\ GHVFULEHG IURP DUWLÀFLDO UHHI fultoportulae arranged variously between the center and the material collected from a single location on the Gulf coast marginal zone. Live cells in most species of Thalassiosira of northern Florida, USA (Prasad et al., 1993). This led to form chains with sibling cells linked by threads of chitin the assumption that T. cedarkeyensis was a benthic form. (McLachlan et al., 1965, Herth and Barthlott, 1979); the Since then T. cedarkeyensis has been observed in plankton rest are found in gelatinous colonies, or more rarely, solitary samples collected from many localities in coastal waters +DVOH6FKUDGHU)U\[HOOHWDO 7KHUH of the southeastern United States. We, therefore, take this is usually one or more rimoportula, typically located near RSSRUWXQLW\WRUHÀQHRXUGHÀQLWLRQRIWKHVSHFLHVEDVHGRQ the valve mantle. Areolation varies from radial to eccentric these new observations. We also describe the structure of RUOLQHDU )U\[HOO EXWLQDOOFDVHVWKHDUHRODHDUH a new plicate species of Thalassiosira from the region and closed internally by some sort of cribrum. Patterns of compare it with T. cedarkeyensis and resident populations fultoportulae, rimoportulae, areolae, and colony formation of the plicate species T. gessneri and T. lacustris. The are widely used in delineation of species (Rivera, 1981; SUHVHQFHRIRIWKHQRZNQRZQH[WDQWVSHFLHVRISOLFDWH 0DNDURYD0DKRRGHWDO3UDVDG Thalassisiosira provides grounds for comparisons to be et al., 1993). Both the Index Nominum Algarum website drawn among these species and other morphologically maintained at the University of California, Berkeley (http:// VLPLODUH[WDQWDQGH[WLQFWSOLFDWHVSHFLHV ucjeps.berkeley.edu/ina) and the Catalogue of Diatom names maintained at the California Academy of Sciences MATERIAL AND METHODS (http://research.calacademy.org/izg/research/diatom) list LQWKHUDQJHRI²WD[DLQFOXGLQJUHFHQWIRVVLODQG Materials used and the coordinates of the locations WUDQVIHUUHGWD[DDQGWKHLUV\QRQ\PV where T. cedarkeyensis and the new species were found are The genus Thalassiosira is well represented in listed in the appropriate section under each species. Most the coastal waters of the southeastern United States, of the phytoplankton samples containing T. cedarkeyensis, including the Atlantic and the Gulf coast states. At least T. lacustris DQG WKH QHZ WD[RQ ZHUH FROOHFWHG IURP twenty species of Thalassiosira have been recorded from locations on the Gulf coast of Florida, particularly the WKH *XOI RI 0H[LFR &RQJHU HW DO /LFHD 'XUDQ (FRQÀQDDQG)HQKROORZD\HVWXDULHVDVSDUWRIPXOWL\HDU 1999; Krayesky et al., 2009; personal observation), with HFRORJLFDO LQYHVWLJDWLRQV H[WHQGLQJ IURP WKURXJK a similar number from the Atlantic coasts of Florida and /LYLQJVWRQ D E %DUU\ 9LWWRU Georgia (Hargraves, 2002; Prasad and Nienow, 2003; 2005, 2006, 2007).
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