Taxonomic notes on Caribbean Neosiphonia and Polysiphonia ...

Botanica Marina 54 (2011): 269–292 � 2011 by Walter de Gruyter • Berlin • Boston. DOI 10.1515/BOT.2011.036

2010/101

Article in press - uncorrected proof

Taxonomic notes on Caribbean Neosiphonia and Polysiphonia

(Ceramiales, Florideophyceae): five species from Florida, USA

and Mexico

Nadya R. Mamoozadeh and D. Wilson Freshwater*

Center for Marine Science, UNCW, 5600 Marvin MossLane, Wilmington, NC 28409, USA,e-mail: [email protected]

* Corresponding author

Abstract

Molecular-assisted identification using plastid-encoded rbcLand mitochondrion-encoded COI loci identified five speciesof Polysiphonia sensu lato from 16 Florida and CaribbeanMexico samples. Morphological character states were exam-ined and used to identify these species as Neosiphonia baja-cali comb. nov., N. echinata comb. nov., N. sphaerocarpa,N. tepida, and Polysiphonia anomala. Descriptions are pro-vided and the phylogenetic relationships of the five specieswere determined through maximum likelihood analyses ofrbcL and nuclear-encoded SSU sequence data. Neosiphoniabajacali and N. echinata had a combination of characterstates described for Neosiphonia: rhizoids cut-off from peri-central cells, lateral branch or trichoblast initials on everysegment in a spiral pattern, tetrasporangia in spiral series,and spermatangial stichidia developing as bifurcations oftrichoblasts, and these new combinations are proposed.Examination of N. echinata, P. fracta and North Carolinaspecimens identified as P. breviarticulata revealed no signif-icant morphological differences. Polysiphonia fracta is pro-posed as a synonym of N. echinata and the presence of P.breviarticulata within the western Atlantic is questioned.This is the first report of N. bajacali from the Caribbean andthe first report of N. echinata from Caribbean Mexico.

Keywords: Caribbean Mexico; Florida; molecular-assistedidentification; Neosiphonia; Polysiphonia.

Introduction

Polysiphonia Greville (Rhodomelaceae) is a large (ca. 200species) red algal genus whose species have a nearly globaldistribution. Species of Polysiphonia sensu lato (s.l.) have adelicate habit composed of filamentous main axes andbranches that are segmented and polysiphonous. A widerange of morphological variability occurs among the manyspecies of Polysiphonia s.l. This variation has led to muchdebate on the classification of species within the genus. Torevise the classification of Polysiphonia s.l., Kim et al.(2000) suggested the use of ‘‘independent comparative

evidence’’ to determine which morphological characters arerelevant to species identification. The recent coupling ofmorphological characters with DNA sequence data is helpingestablish an accurate classification of Polysiphonia based onnatural relationships.

Molecular-assisted identification (MAI) through use of theplastid-encoded ribulose-1,5-bisphosphate carboxylase/oxy-genase large subunit gene (rbcL) has proven useful for dis-criminating species of Polysiphonia s.l. Previous studies ofrbcL sequence data in Polysiphonia s.l. detected compara-tively low intraspecific and high interspecific sequencedivergences, with values ranging from 0 to 1.3% (with aninstance of 2.13%) and 2.6–14.12%, respectively (McIvor etal. 2001, Kim et al. 2004, Kim and Yang 2005). A relativelydistinct break between intra- and interspecific sequence var-iation makes rbcL a useful tool for aiding species identi-fications.

The 59 end of the mitochondrion-encoded cytochrome coxidase subunit I gene (COI) is another region of DNA thatis employed for MAI. Previous studies of red algae haveshown COI to be successful in distinguishing closely relatedand cryptic species as well as geographic groups within aspecies (Saunders 2005, Robba et al. 2006, Yang et al. 2008,Clarkston and Saunders 2010, Le Gall and Saunders 2010).The utility of COI barcoding within Polysiphonia s.l. has yetto be established.

Phylogenetic relationships among Polysiphonia s.l. speciescan also be determined using molecular data. The nuclear-encoded 18S rDNA (SSU) gene has limited intraspecificsequence variation and provides better resolution of taxo-nomic relationships above the species level. Analyses of SSUsequence data showed that Polysiphonia s.l. is polyphyleticwith respect to other genera (Choi et al. 2001). The rbcLlocus has a faster rate of evolution than SSU allowing greaterresolution of species-level relationships. The use of rbcL todetermine species relationships has been common in recentstudies of Polysiphonia s.l. (McIvor et al. 2001, Kim et al.2004, 2005, Stuercke and Freshwater 2008, 2010). Phylo-genetic relationships within Polysiphonia s.l. can be thor-oughly evaluated when analyses of SSU and rbcL arecompared with one another (Stuercke 2006).

Stuercke and Freshwater (2008) examined many of themorphological characters used to distinguish Polysiphonias.l. species in an integrated molecular-morphological study.Little to no intraspecific variation was found in the characterstates for pericentral cell number, rhizoid-pericentral cellconnection, lateral branch-trichoblast relationship, sperma-tangial axis development, tetrasporangial arrangement, tri-choblasts/scar cell occurrence and pattern, holdfast type, and

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270 N.R. Mamoozadeh and D.W. Freshwater: Five Neosiphonia/Polysiphonia species from the Caribbean


cicatrigenous branch formation. The genus NeosiphoniaM.-S. Kim et I.K. Lee was segregated from Polysiphoniabased on a combination of character states including the fol-lowing from those listed above: rhizoids that are cut-off frompericentral cells, spirally arranged trichoblasts or scar cellson every segment, three-celled carpogonial branches, sper-matangial axes developing as a bifurcation of trichoblasts,and spirally arranged tetrasporangia (Kim and Lee 1999).Polysiphonia s.l. now includes species that are predomi-nantly placed in Neosiphonia or Polysiphonia, although lessspeciose genera are also included.

Study of the Caribbean marine algal flora has resulted inreports of at least 28 distinct species of Polysiphonia s.l.(e.g., Wynne 1998). Early reports of Polysiphonia s.l. byHarvey (1853) and Børgesen (1918), described several spe-cies from Florida, USA and the West Indies, respectively.More recent reports are the result of floristic assessmentsmade throughout the Caribbean (e.g., Taylor 1929, 1941,1942, 1945, 1960, 1969, Almodovar and Ballantine 1983,Ballantine and Aponte 1997, Littler and Littler 2000, Mateo-Cid et al. 2006). Caribbean studies specific to Polysiphonias.l. include reports of species from Belize (Kapraun and Nor-ris 1982), Colombia and Venezuela (Kapraun et al. 1983).

The purpose of this study was to complete integratedmolecular and morphological analyses of Polysiphonia s.l.samples collected from Caribbean Mexico and Florida.Examination of morphology and MAI using rbcL and COIsequence data were used to identify four Neosiphonia andone Polysiphonia species within samples while phylogeneticanalyses of rbcL and SSU sequence data were used to deter-mine the relationships of these species.

Materials and methods

Collections and vouchers

Samples of Neosiphonia and Polysiphonia were collectedfrom intertidal and subtidal substrata in Caribbean Mexicoand Florida by snorkeling (Appendix 1). Samples were driedin silica gel desiccant (Chase and Hills 1991) and depositedin the silica collection at the Center for Marine Science, Uni-versity of North Carolina-Wilmington. Examinations of mor-phological characters and taxonomic guides were used toidentify species (Børgesen 1918, Setchell and Gardner 1930,Hollenberg 1942, 1958, 1961, 1968a,b, Taylor 1945, 1960,Dawson 1964, Abbott and Hollenberg 1976, Hollenberg andNorris 1977, Kapraun 1977, 1980, Womersley 1979, 2003,Kapraun and Norris 1982, Kapraun et al. 1983, Adams 1991,Schneider and Searles 1991, Abbott 1999, Littler and Littler2000, Dawes and Mathieson 2008, D. Kapraun unpublishedmanuscript). Specimens were stained with aniline blue solu-tion (Millar and Wynne 1992) and permanent slide vouchersmade following Tsuda and Abbott (1985) for deposit in theUniversity of North Carolina Wilmington (WNC) herbarium.Additional specimens from the United States National Her-barium (US) were also studied. All herbarium abbreviations

follow the Index Herbariorum (http://sciweb.nybg.org/sci-ence2/IndexHerbariorum.asp).

Morphological data

Specimens and slides were observed using an Olympus SZHdissecting microscope (Olympus America Inc., CenterValley, PA, USA) and a Nikon Labophot-2 compound micro-scope (Nikon Inc., Melville, NY, USA). Images were cap-tured using a Zeiss Axio Imager.Z1 compound microscopefitted with an AxioCam MRc 5 camera system (Carl ZeissMicroimaging Inc., Thornwood, NY, USA) or an OlympusBX41 compound microscope fitted with a Roper ScientificPhotometrics� CoolSnapTM camera (Photometrics, Tucson,AZ, USA). Species descriptions were written based on obser-vations of specimens collected in this study and at the US,and with information from the literature included for char-acter states that were not directly observed.

DNA extraction and sequencing

DNA was extracted from specimens according to Hughey etal. (2001) with an additional cleaning step using the One-Step� PCR Inhibitor Removal Kit (Zymo Research, Orange,CA, USA). SSU, rbcL, and COI were amplified followingthe basic PCR recipe outlined in Freshwater et al. (2005) butusing GOTaq DNA polymerase and buffer (Promega, Mad-ison, WI, USA). The thermocycling protocol followed Fresh-water et al. (2000) but with 35 cycles of denaturing,annealing at 40, 45, or 508C, and an elongation period of90 s. Amplification products were cleaned with a StratageneStrataPrep� PCR Purification Kit (Stratagene, La Jolla, CA,USA) and used as templates in BigDye� Terminator v3.1sequencing reactions (Applied Biosystems, Foster City, CA,USA). Sequences were edited and assembled using Sequen-cher� (version 4.9, GeneCodes Corporation, Ann Arbor, MI,USA). Primers utilized in amplification and sequencing reac-tions are listed in Table 1.

rbcL and COI sequences were generated for as many sam-ples as possible. These loci are appropriate for the exami-nation of inter- and intraspecific relationships and were usedas barcodes to objectively assign samples to species (e.g.,Millar and Freshwater 2005, Clarkston and Saunders 2010).An SSU sequence was generated for one sample per species.Both rbcL and SSU sequence data were used to identifygroupings of species (clades) within Polysiphonia s.l.

Molecular data analyses

Sequences of Neosiphonia and Polysiphonia generated inthis study were combined with rbcL and SSU sequencesavailable from GenBank and with some from unpublishedstudies (J. Kelly and D.W. Freshwater unpublished) and ini-tially aligned using MacClade (v.4, Maddison and Maddison2000). SSU sequence data were also aligned using theClustalW multiple sequence alignment feature of the pro-gram Molecular Evolutionary Genetics Analysis (MEGA)(www.megasoftware.net; Tamura et al. 2007, Kumar et al.



N.R. Mamoozadeh and D.W. Freshwater: Five Neosiphonia/Polysiphonia species from the Caribbean 271


Table 1 Primer sequences utilized in amplification and sequencing reactions of rbcL, SSU and COI.

Locus/primer name Primer sequence References

rbcLFrbcLstart 59-ATGTCTAACTCTGTAGAAG-39 Freshwater and Rueness (1994)F87 59-GGATCCTAAYTACGTAAYTAAAG-39 This studyF577 59-GTATATGAAGGTCTAAAAGG-39 Freshwater and Rueness (1994)F753 59-GGAAGATATGTATGAAAGAGC-39 Freshwater and Rueness (1994)R893 59-GAATAAGTTGARTTWCCIGCAC-39 Stuercke and Freshwater (2008)R1415 59-CTACRAAGTCAGCTGTATCTG-39 This studyRrbcSstart 59-GTTCCTTGTGTTAATCTCAC-39 Freshwater and Rueness (1994)

SSUA.2 59-AGACTAAGCCATGCAAGTGC-39 D.W. Freshwater (unpublished)A.3 59-ACA(AT)CGAAACTGCGAATGG-39 D.W. Freshwater (unpublished)CerD 59-GCAAGTCTGGTGCCAGCAG-39 Hommersand et al. (2005)CerE 59-CTATTATTCCATGCTAATGTATTC-39 Hommersand et al. (2005)CerG 59-AGCCTGCGGCTTAATTTGAC-39 Hommersand et al. (2005)CerH 59-TAACCAGACAGATCACTCCAC-39 Hommersand et al. (2005)CerJ 59-TCTCCTTCCTCTAAGTGATAA-39 Hommersand et al. (2005)GO1 59-CACCTGGTTGATCCTGCCAG-39 Saunders and Kraft (1994)GO4 59-CAGAGGTGAAATTCTTGGAT-39 Saunders and Kraft (1994)GO7 59-ATCCTTCTGCAGGTTCACCTAC-39 Saunders and Kraft (1994)GO8 59-GAACGGCCATGCACCACCACC Saunders and Kraft (1994)

COICerR1 59-CCAAAAAATCAAAATARRTG-39 Saunders (unpublished)GazF1 59-TCAACAAATCATAAAGATATTGG-39 Saunders (2005)GHalF 59-TCAACAAATCATAAAGATATYGG-39 Saunders (2008)Mam1R 59-CCICCWCCIGCWGGATCAA-39 This studyMam2R 59-GTATTAAAATTWCKATCWGTTA-39 This study

2008) followed by manual adjustment. Characteristics of theDNA sequence data sets were determined using MacCladeand PAUP (v.4, Swofford 2002). MEGA was used to performUnweighted Pair Group Method with Arithmetic mean(UPGMA) and Neighbor Joining (NJ) cluster analyses onCOI and rbcL sequence data for MAI. Simple mean dis-tances (p-distances) were used in all MAI analyses. ML anal-yses were performed on reduced (identical sequencesremoved) rbcL and SSU sequence data separately using theprogram Genetic Algorithm for Rapid Likelihood Inference(GARLI) (www.bio.utexas.edu/faculty/antisense/garli/Garli.html; Zwickl 2006). The ML analyses included 10 separatesearches from random starting trees that used default par-ameters, including 10,000 generations without improvingtopology and allowing model estimation during the run(models available from the second author). GARLI was alsoused to perform a total of 1003 ML bootstrap (BS) replicateson the rbcL dataset and 1000 ML BS replicates on the SSUdataset.

Results and discussion

Molecular assisted identification

The rbcL MAI alignment consisted of 113 taxa and included1081 sites in the analysis, of which 402 (37.19%) were var-iable. Five species from 16 Florida and Caribbean Mexicosamples were resolved in UPGMA and NJ cluster analyses

of rbcL sequence data (Figure 1, only UPGMA cluster dia-gram shown). Species distinctions were based on values ofinter- and intraspecific rbcL sequence divergence from pre-vious studies of Polysiphonia s.l. (McIvor et al. 2001, Kimet al. 2004, Kim and Yang 2005). The COI MAI alignmentconsisted of 74 taxa and included 605 sites in the analysis,of which 248 (40.99%) were variable. Four species from 10Florida and Caribbean Mexico samples were resolved inUPGMA and NJ cluster analyses of COI sequence data; thislocus did not amplify in some samples (Figure 2, onlyUPGMA cluster diagram shown). Species distinctions werebased on the 4.80% COI sequence divergence between sam-ples identified as Polysiphonia subtilissima 1 (NC-24) andP. subtilissima 2 (PHYKOS-3271). These samples differedby 2.22% in the rbcL data, which slightly exceeds the rangeof intraspecific rbcL sequence divergence observed in pre-vious studies of Polysiphonia s.l. (McIvor et al. 2001). Asequence divergence of 2.22% in the P. subtilissima rbcLdata corresponds to 4.80% sequence divergence in the COIdata; the latter value was therefore used as the maximumintraspecific barcoding gap value (Meier et al. 2008) in theCOI data. Values of intraspecific sequence divergence arediscussed in the remarks on each species.

Molecular systematics

Phylogenetic relationships of Florida and Caribbean Mexicospecies within Polysiphonia s.l. were determined throughML analyses of SSU and reduced rbcL sequence data





Figure 1 Unweighted pair group method with arithmetic mean rbcL cluster analysis for 113 Polysiphonia s.l. samples.The 2.13% and 1.3% sequence divergence levels are indicated by vertical dashed lines. Florida and Caribbean Mexico species are shownin boldface type. P., Polysiphonia; N., Neosiphonia.

(Figures 3 and 4). The specific relationships among thesespecies are discussed later in the remarks on each species.The reduced rbcL alignment consisted of 52 taxa and includ-ed 1334 sites in the analysis, 528 (39.58%) of which werevariable. The rbcL topology contains several strongly sup-ported clades as well as groups of clades and species withunresolved relationships. Near the base of the topology aretwo well-supported clades (rbcL bootstrap values wrBxs100)

that contain species typically regarded as Polysiphonia sensustricto (s.s.). These species all have four pericentral cells,rhizoids in open connection with the pericentral cells, tetra-sporangia in straight series, and spermatangial stichidia thatdevelopmentally replace trichoblasts. Species in these cladesinclude the generitype, Polysiphonia stricta (Dillwyn) Gre-ville. Another well-supported clade (rB100) contains a groupof species referred to as the ‘‘multipericentral cell group’’





Figure 2 Unweighted pair group method with arithmetic mean COI cluster analysis for 74 Polysiphonia s.l. samples.The 4.80% sequence divergence level is indicated by the vertical dashed line. Florida and Caribbean Mexico species are shown in boldfacetype. P., Polysiphonia; N., Neosiphonia.

by Choi et al. (2001). These species all have 7q pericentralcells and rhizoids cut-off from pericentral cells. The majorityof species in the rbcL ML topology are placed within a largeunsupported (rB-50) group of species and variously sup-ported clades that contain both Neosiphonia and Polysipho-nia species. Although species of Neosiphonia are scatteredthroughout the topology, a strongly supported clade (rB100)of predominantly Neosiphonia species is apparent within thislarge grouping.

The SSU alignment consisted of 44 taxa and included1602 sites in the analysis, 158 (9.86%) of which were vari-able. The major clades and species resolved in the rbcLtopology were also resolved in the SSU topology. Two cladescontaining Polysiphonia s.s. species are strongly supported(SSU bootstrap values wsBxs99 and 100) and received mod-erate support (sB78) for resolution as a monophyletic group.Another well-supported (sB91) clade contains ‘‘multiperi-central cell’’ species of multiple genera including Polysipho-





Figure 3 Maximum likelihood rbcL tree (lnLs–14603.23387) for 50 Polysiphonia s.l. and two outgroup species. Bootstrap proportionvalues for branches are shown for each node when )50. Florida and Caribbean Mexico species are shown in boldface type.Vertical bars show: A) Polysiphonia sensu stricto clades; B) the ‘‘multipericentral cell’’ clade; C) predominately Neosiphonia clade. Car.,Caribbean; Pac., Pacific; NC, North Carolina; FL, Florida; OR, Oregon; P., Polysiphonia; N., Neosiphonia.

nia, Boergeseniella Kylin, Enelittosiphonia Segi, andVertebrata S.F. Gray. The majority of species are containedwithin a large, strongly supported (sB91) grouping of speciesand clades that include both Neosiphonia and Polysiphoniaspecies. A clade of predominantly Neosiphonia species iscontained within this large grouping and received moderatesupport (sB87).

Taxonomic observations and remarks

Genus Neosiphonia

Neosiphonia bajacali (Hollenberg) N.R. Mamoozadeh

et D.W. Freshwater, comb. nov.

(Figures 5–8)

Basionym Polysiphonia bajacali Hollenberg (1961, p. 347).





Figure 4 Maximum likelihood SSU tree (lnLs–4146.45008) for 43 Polysiphonia s.l. and one outgroup species.Bootstrap proportion values for branches are shown for each node when )50. Florida and Caribbean Mexico species are shown in boldfacetype. Vertical bars show: A) Polysiphonia sensu stricto clades; B) the ‘‘multipericentral cell’’ clade; C) predominately Neosiphonia clade.Car., Caribbean; Pac., Pacific; CA, California; NC, North Carolina; FL, Florida; OR, Oregon; RI, Rhode Island; P., Polysiphonia; N.,Neosiphonia.

Description Plants to 3 (–6) cm tall, chiefly erect fromlimited basal prostrate axes attached to the substratum byrhizoids that are cut-off from pericentral cells (Figure 5);highly branched; branching subdichotomous to alternate(Figure 6); erect axes (150–) 200–300 mm in diameter, pros-trate axes (175–) 225–325 mm in diameter; branchlets bas-ally attenuated; mid axis segments of erect axes mostly 1=

as long as wide; light cortication present in some matureprostrate axes; main axes with four pericentral cells (Figure7); branches replacing trichoblasts in development; tricho-blasts with several dichotomies, to 750 mm in length; scarcells one per segment in 1/4 spiral series; adventitious lateralsrare; tetrasporangia greatly distending segments, developingin short spiral series (Figure 8); spermatangial stichidia





Figures 5–8 Neosiphonia bajacali comb. nov.(5) Prostrate axis with rhizoids that are cut-off from the pericentral cells, MEX04-09, scales0.10 mm, WNC2010-s061. (6) Habit of erectaxes, MEX04-09, scales0.20 mm, WNC2010-s061. (7) Portion of erect axis flattened to show four pericentral cells per segment, centralaxial cell (arrowhead), and scar cells (arrows), MEX04-09, scales0.10 mm, WNC2010-s061. (8) Reproductive branch displaying shortspiral series of tetrasporangia (arrows), MEX04-11A, scales0.10 mm, WNC2010-s056.

developing as a furcation of trichoblasts, without sterile tipcells; cystocarps urceolate to globose.

Type locality Isla Guadalupe, Baja California.

Other sources Abbott and Hollenberg 1976, Hollenberg1961.

Specimens studied Mexico: US-42350, Isotype,extreme south tip of Isla Guadalupe, 18 Dec 1949, E.Y. Daw-son; WNC2010-s055 to s57 (MEX04-11A), Blue Bay Marina,North of Cancun, Yucatan, D.W. Freshwater, 29 Feb 2004;WNC2009-s58 to s061 (MEX04-09), Blue Bay Marina, Northof Cancun, Yucatan, D.W. Freshwater, 29 Feb 2004.

Molecular vouchers GenBank accession numbersHM573572 (rbcL); HM560659 (SSU); HM573526 (COI).

Remarks Hollenberg (1961) originally described Neosi-phonia bajacali (as Polysiphonia bajacali) as ecorticate butin a later description indicated that light basal cortication

may be present (Abbott and Hollenberg 1976). Samples inthis study infrequently displayed cortication in limited por-tions of some mature prostrate axes. The original and sub-sequent descriptions of N. bajacali do not indicate whetherrhizoids are in open connection or are cut-off from pericen-tral cells (Hollenberg 1961, Abbott and Hollenberg 1976,Stewart 1991). Samples in this study had rhizoids cut-offfrom pericentral cells (Figure 5).

Neosiphonia bajacali displays a combination of characterstates unique to Neosiphonia. These include rhizoids cut-offfrom pericentral cells, lateral branch or trichoblast initialsone per segment in a spiral pattern, tetrasporangia developingin spiral series, and spermatangial stichidia developing asfurcations of trichoblasts.

Two samples of Neosiphonia bajacali were collected fromCancun, Mexico (Figures 1, 2). Neosiphonia bajacali isresolved within a strongly supported (rB100; sB87) clade ofpredominantly Neosiphonia species in ML analyses of rbcLand SSU sequence data (Figures 3, 4). Neosiphonia bajacaliis most closely related to N. tongatensis in the SSU phylog-eny, but this relationship received only moderate support





Figures 9–14 Neosiphonia echinata comb. nov.(9) Portion of erect axis flattened to show four pericentral cells per segment, FL09-40, scales100 mm, WNC2009-s112. (10) Apical partof main axis showing a lateral branch developing in the axil of a trichoblast (arrow), FL05-07, scales50 mm, WNC2010-s085. (11) Habitshowing adventitious laterals (arrows) arising every segment to every few segments, FL09-42, scales0.20 mm, WNC2009-s119. (12)Reproductive branch displaying long spiral series of tetrasporangia (arrows), FL09-42, scales100 mm, WNC2009-s118. (13) Branch apexwith spermatangial stichidium developing as a furcation of the trichoblast (arrow), MEX04-8, scales50 mm, WNC2010-s054. (14) Mainaxis bearing a short-stalked cystocarp, MEX04-10, scales50 mm, WNC2010-s062.

(sB78) and is not present in the rbcL phylogeny. Neosiphoniabajacali and N. tongatensis share many character states butcan be distinguished by a smaller diameter of erect and pros-trate axes (to 150 mm and 250 mm, respectively), no corti-cation, and segments 1–2= as long as wide in N. tongatensis.

Neosiphonia echinata (Harvey) N. Mamoozadeh et

D.W. Freshwater, comb. nov.

(Figures 9–14)

Basionym Polysiphonia echinata Harvey (1853, p. 38).

Synonym Polysiphonia fracta Harvey (1853, p. 38).

Misapplied name Polysiphonia breviarticulata sensuStuercke and Freshwater.

Description Plants to 3 cm tall, entirely erect from singlebasal holdfast of rhizoids cut-off from pericentral cells; main

axes sparsely branched in a subdichotomous pattern; erectmain axes (275–) 375–500 mm in diameter, basal axes(375–) 500–625 (–750) mm in diameter; mid axis segmentsof erect axes mostly 0.5–1= as long as wide; light to mod-erate cortication present in some mature prostrate axes andat the bases of older adventitious laterals; main axes withfour pericentral cells (Figure 9); branches forming in theaxils of trichoblasts (Figure 10); trichoblasts with severaldichotomies, to 500–800 mm in length; scar cells one persegment in 1/4 spiral series; adventitious laterals abundant,every segment to every few segments on main axes givingplants a coarse appearance (Figure 11), linear in shape, var-iable in length from 0.5 to 1.5 mm long; tetrasporangia littledistending segments, developing in short spiral series (Figure12); spermatangial stichidia developing as furcations of tri-choblasts, without sterile tip cells (Figure 13); cystocarpsglobose to subglobose, on short stalk (Figure 14).

Type locality Key West, Monroe County, Florida, USA.





Other sources Harvey 1853, Taylor 1960, Dawes andMathieson 2008.

Specimens studied Neosiphonia echinata (Harvey) N.Mamoozadeh et D.W. Freshwater, Florida: US-66789 (slidenumber US-3323, 3324), Isotype, Key West, Monroe County,W.H. Harvey, Feb 1850; US-22409, West Summerland Key,Monroe County, J. Brunson, 21 Mar 1976; US-22404, Alli-gator Point, Franklin County, H.J.H., 16 Apr 1950; US-22403, Alligator Point, Franklin County, H.J.H., 21 Mar1950; WNC2009-s112 to s115 (FL09-40), Lake Surprise,Key Largo, Monroe County, N. Mamoozadeh, 09 Mar 2009;WNC2009-s117 to s119 (FL09-42), Lake Surprise, Key Lar-go, Monroe County, N. Mamoozadeh, 09 Mar 2009;WNC2009-s123 to s127 (FL09-44), Lake Surprise, Key Lar-go, Monroe County, N. Mamoozadeh, 09 Mar 2009;WNC2009-s137 to 140 (FL09-75), KML Mangrove, LongKey, Monroe County, N. Mamoozadeh, 10 Mar 2009;WNC2009-s141 to s144 (FL09-76), KML Mangrove, LongKey, Monroe County, N. Mamoozadeh, 10 Mar 2009;WNC2010-s085 (FL05-7), West Summerland Key, MonroeCounty, B. Stuercke, 28 Feb 2005; Mexico: WNC2010-s051to s054 (MEX04-8), Blue Bay Marina, Cancun, D.W. Fresh-water, 29 Feb 2004; WNC2010-s062 to s064 (MEX04-10),Blue Bay Marina, Cancun, D.W. Freshwater, 29 Feb 2004;North Carolina: WNC2005-s039 (NC-1), Snead’s Ferry, NewRiver Inlet, Onslow County, D.W. Freshwater and F. Mont-gomery, 11 Jul 2005; WNC2005-s053, s054 (NC-3), CORMPSite OB-1, Onslow Bay, D.W. Freshwater and K. Johns, 19Jul 2004; WNC2005-s001, s059 (NC-5), CORMP Site OB-27, Onslow Bay, J. Souza and J. Dorton, 02 Jan 2005;WNC2005-s067 (NC-7), CORMP Site OB-27, Onslow Bay,J. Souza and J. Dorton, 12 Jan 2005; WNC2005-s026, s104(NC-14), Bank’s Channel (Site NH-M), New Hanover Coun-ty, D.W. Freshwater, 11 May 2005; WNC2005-s031 (NC-18),Bogue Sound, Corkey’s house, Carteret County, D.W. Fresh-water, 26 Mar 2005; WNC2005-s008 (NC-20), Bogue Sound,Corkey’s House, Carteret County, D.W. Freshwater, 26 Dec2003; WNC2005-s124, s127 (NC-25), CORMP Site OB-27,J. Souza, 12 May 2005; Polysiphonia fracta Harvey, Florida:US-66791 (slides US-3320 to 3322), Isotype, Key West,Monroe County, W.H. Harvey, Feb 1850; Texas: US-2329,near Aransas Pass, Corpus Christi, E.Y. Dawson, 12 Dec1957; Bahamas: US-14293, Eleuthra, M.E. Hay, 10 May1981; US-14294, Eleuthra, M.E. Hay, 08 May 1981; Poly-siphonia hapalacantha Harvey, Florida: US-66793 (slidesUS-3325, 3326), Isotype, Key West, Monroe County, W.H.Harvey, Feb 1850; Puerto Rico: US-202615, Boqueron, CaboRojo, M. Dıaz-Piferrer, 26 Feb 1964; US-40250, Boqueron,Cabo Rojo, M. Dıaz-Piferrer, 26 Feb 1964; US-78248,Boqueron, Cabo Rojo, M. Dıaz-Piferrer, 26 Feb 1964.

Molecular vouchers GenBank accession numbersHM573561, HM573559, HM573558, HM573560,HM573557 (rbcL); HM560658 (SSU); HM573503,HM573506, HM573504, HM573505 (COI).

Remarks Harvey (1853) originally described Neosipho-nia echinata (as Polysiphonia echinata), P. fracta Harvey,

and P. hapalacantha Harvey from Key West, Florida, USA.A report and description of P. breviarticulata (C. Agardh)Zanardini were also included among these original speciesdescriptions (Harvey 1853). These species all have erect thal-li with four pericentral cells, segment length-width ratiosgenerally F1, and numerous adventitious laterals that oftengive the plants a coarse appearance. The character state com-bination of rhizoids cut-off from pericentral cells, sperma-tangia developing as a furcation of trichoblasts, tetra-sporangia in spiral series, and trichoblasts or scar cells in aspiral pattern on every segment observed in the specimensof N. echinata studied is characteristic of the genus Neosi-phonia. This same combination of character states may bepresent in P. breviarticulata and P. hapalacantha, but thesecharacters were not all observed in the specimens availablefor study and recent descriptions do not include this infor-mation (Taylor 1960, Athanasiadis 1987, Dawes and Mathie-son 2008), making it impossible to determine whether newcombinations are warranted for these species as well.

Harvey’s original description of Polysiphonia hapalacanthaincluded no remarks on its similarity to Neosiphonia echinata,P. fracta, or P. breviarticulata. P. hapalacantha has traditionallybeen distinguished from these species by its adventitious lat-erals to 2–4 mm in length. The type material of P. hapalacan-tha at US is in poor condition, and an assessment of the speciesfrom this material was not possible. Material at US identifiedas P. hapalacantha from Puerto Rico displayed a habit consis-tent with the original species description and seemed to exem-plify true P. hapalacantha. These specimens showed longeradventitious laterals that gave the specimens a more lax andless coarse habit than N. echinata and P. fracta, allowing thesespecies to be distinguished.

Harvey (1853) indicated that, while Neosiphonia echinataclosely resembled Polysiphonia fracta, a more robust habitand adventitious laterals that were shorter, more abundant,and ‘‘more equally inserted on all sides of the branches’’could distinguish N. echinata. Examination of type materialfor N. echinata and P. fracta suggests that these two namesrepresent one morphological species as no satisfactory dif-ference was observed. It seems possible that Harvey’s mate-rial of P. fracta is simply a diminutive form of N. echinata,and this possibility was also suggested by D. Kapraun (unpub-lished manuscript). Although both names were introduced inthe same publication, N. echinata appears to be more widelyutilized in species reports and is therefore conserved.

Harvey’s reports of Polysiphonia breviarticulata in KeyWest, Florida, USA and Vera Cruz, Mexico seem to be thefirst reports of this species in the western Atlantic (Harvey1853). Agardh (1824) originally described P. breviarticulatafrom the Adriatic Sea as Hutchinsia breviarticulata C.Agardh. Prior to Harvey (1853), P. breviarticulata wasknown only from the Adriatic and the Mediterranean. In hisdescription of P. breviarticulata, Harvey noted that the Flor-ida material appeared more robust and with more lateralbranches than Mediterranean material of P. breviarticulata(Harvey 1853). Kutzing (1863) described and illustrated aspecimen identified as P. breviarticulata, from Vera Cruz,Mexico that may be the same Liebman specimen that Harvey





Figures 15–20 Neosiphonia sphaerocarpa.(15) Prostrate axis with rhizoids (arrows) that are cut-off from the pericentral cells, FL05-06, scales0.10 mm, WNC2010-s083. (16) Habitof erect axes, FL05-5B, scales0.20 mm, WNC2010-s073. (17) Portion of erect axis flattened to show four pericentral cells per segment,central axial cells (arrowheads), and scar cells in spiral pattern (arrows), FL05-5B, scales0.10 mm, WNC2010-s071. (18) Apical portionof erect axis bearing many trichoblasts (arrows), FL05-5B, scales50 mm, WNC2010-s071. (19) Reproductive branch displaying long spiralseries of tetrasporangia (arrows), FL05-5B, scales0.10 mm, WNC2010-s073. (20) Main axis bearing cystocarp (flattened during preparation)on short stalk, FL05-5B, scales0.10 mm, WNC2010-s072.

studied. It differs in branching pattern and segment size fromthe Adriatic P. breviarticulata that Kutzing also describedand illustrated (Kutzing 1863).

The next report of Polysiphonia breviarticulata in thewestern Atlantic (Kapraun and Searles 1990) identified thisas the species in intense macro-algal bloom along the coastof North Carolina. Kapraun and Searles’ (1990) descriptionof North Carolina P. breviarticulata appears identical todescriptions and material of Mexico and Florida Neosiphoniaechinata, except for the presence of basal attenuation in thelateral branches of P. breviarticulata specimens. Examinationof North Carolina specimens identified as P. breviarticulataby Stuercke and Freshwater (2008) revealed basal attenua-tion to be a variable character state that is present in somebranchlets, particularly ones that are tetrasporangial, but notothers. The basal attenuation observed in tetrasporangialbranchlets could be the result of mid axis segments that aredistended due to the presence of tetrasporangia and in turngive basal segments a slightly constricted appearance. Noother morphological difference was observed between NorthCarolina material of P. breviarticulata examined by Stuercke

and Freshwater (2008) and Florida and Mexico material ofN. echinata. North Carolina material in WNC labeled as‘‘P. echinata, 01 Dec 1982, Masonboro Bay’’ most likelyrepresents specimens published under the name P. breviar-ticulata in Kapraun and Searles (1990), who indicated thattheir initial material of P. breviarticulata was collected ‘‘inthe sound behind Masonboro Island in December 1982’’.This material, apart from basal attenuation in some bran-chlets (particularly ones that are tetrasporangial) also appearsindistinguishable from Florida and Mexico specimens. Basedon type localities and morphology, it seems likely that NorthCarolina material previously identified as P. breviarticulataby Stuercke and Freshwater (2008) and perhaps Kapraun andSearles (1990) actually represents N. echinata.

This conclusion is supported by cluster analyses of rbcLand COI sequence data. Four samples of Neosiphonia echi-nata were collected from the Florida Keys and two fromCaribbean Mexico (Figures 1, 2). Samples identified as Poly-siphonia breviarticulata by Stuercke and Freshwater (2008)differed from Florida and Mexico samples of N. echinata by0.093–0.19% and 0.50–1.09% in the rbcL and COI sequence





Figures 21–26 Neosiphonia tepida.(21) Prostrate axis with rhizoid cut-off from the pericentral cell, FL05-02, scales20 mm, WNC2010-s077. (22) Apical portion of erect axesbearing trichoblasts (arrows), FL05-02, scales0.10 mm, WNC2010-s077. (23) Habit of erect axes, FL05-02, scales0.20 mm, WNC2010-s077. (24) Cross-sections of young and mature branch axes showing central axial cell and seven pericentral cells, FL05-02, scales20 mm,WNC2010-s079. (25) Cross-section of mature branch axis showing central axial cell and eight pericentral cells, FL05-02, scales20 mm,WNC2010-s079. (26) Apical part of main axis showing a lateral branch developing in the axil of a trichoblast, FL05-02, scales20 mm,WNC2010-s077.

data, respectively. These values are well within the range ofintraspecific sequence divergence observed in previous stud-ies for these two loci (e.g., McIvor et al. 2001, Yang et al.2008). ML analysis of rbcL sequence data resolves N. echi-nata within a large unsupported clade (rB-50) of speciesand variously supported clades of Polysiphonia s.l. (Figure3). This placement is also supported in the SSU ML phylog-eny (Figure 4). The rbcL ML phylogeny places N. echinataas sister to P. havanensis sensu Børgesen and P. binneyi Har-vey, with this relationship receiving weak support (rB72).These species are all similar in having four pericentral cells,but the latter two have rhizoids in open connection withpericentral cells.

Neosiphonia sphaerocarpa (Børgesen) M.S. Kim et

Lee (1999, p. 280) (Figures 15–20)

Basionym Polysiphonia sphaerocarpa Børgesen (1918,p. 321).

Description Plants to 2.5 cm tall, erect from basal rhi-zoids cut-off from pericentral cells (Figure 15), with somebranches becoming decumbent and attached to substratumby rhizoids; moderately to highly branched in a subdicho-tomous pattern (Figure 16); erect axes 75–150 mm in diam-eter, prostrate axes 225–325 mm in diameter; mid axissegments of erect axes mostly (1–) 1.5–2= as long as wide;cortication lacking; main axes with four pericentral cells(Figure 17); branches replacing trichoblasts in development;trichoblasts long, to 875 mm in length, with several dichot-omies, dense at apices (Figure 18); scar cells one per seg-ment in 1/4 spiral series (Figure 17); adventitious lateralsabsent; tetrasporangia greatly distending segments, in longspiral series near branch tips (Figure 19), 70–90 mm in diam-eter; spermatangial stichidia developing as furcations of tri-choblasts, 40–60=150–180 (–290) mm, with or without onesterile tip cell; cystocarps globose, short stalked (Figure 20),250–375 mm in diameter.

Type locality St. Thomas, Virgin Islands.





Other sources Dawes and Mathieson 2008; as Polysi-phonia sphaerocarpa, Abbott 1999, Børgesen 1918, Hollen-berg 1968a, Kapraun 1977, Kapraun and Norris 1982,Kapraun et al. 1983, Schneider and Searles 1991, Taylor1960.

Specimens studied Florida: WNC2010-s71 to s73(FL05-5B), Keys Marine Laboratory, Long Key, MonroeCounty, B. Stuercke, 27 Feb 2005; WNC2009-s83, s84(FL05-6), Keys Marine Laboratory, Long Key, MonroeCounty, B. Stuercke, 27 Feb 2005.

Molecular vouchers GenBank accession numbersHM573569 (rbcL); HM573527 (COI).

Remarks Neosiphonia sphaerocarpa is part of a greatercomplex of Polysiphonia s.l. species that share severalmorphological character states, potentially leading to spe-cies misidentifications. These species include N. tongaten-sis (Harvey in Kutzing) M.S. Kim et I.K. Lee, N. bajacali,N. beaudettei (Hollenberg) M.-S. Kim et I.A. Abbott, P.acuminata N.L. Gardner, P. japonica var. savatieri (Hariot)Yoon, P. masonii Setchell et N.L. Gardner, and P. mollisJ.D. Hooker et Harvey. All are described as having fourpericentral cells, rhizoids cut-off from pericentral cells,scar cells one per segment in 1/4 spiral series, tetrasporan-gia developing in spiral series, and spermatangial stichidiadeveloping as a furcation of trichoblasts (Setchell andGardner 1930, Hollenberg 1961, Abbott and Hollenberg1976, Hollenberg and Norris 1977, Yoon 1986). Thesecharacter states, among others, are used to define membersof the genus Neosiphonia. N. sphaerocarpa is distin-guished from the aforementioned species by a combinationof the following characters: lack of a central percurrentaxis, smaller habit (typically F1.5 cm), smaller dimen-sions for erect and prostrate axes, lack of cortication, ulti-mate and fertile branches that are not basally attenuated,segments mostly longer than wide (but not typically morethan twice as long as wide), and a subdichotomous branch-ing pattern.

Two samples of Neosiphonia sphaerocarpa were collectedfrom Long Key, Monroe County, Florida, USA (Figures 1,2). Only one rbcL sequence was generated from these sam-ples, but the COI sequences for both samples were identical.SSU sequence data could not be generated for this species.Neosiphonia sphaerocarpa is placed within a strongly sup-ported clade (rB100) of predominantly Neosiphonia speciesin the rbcL ML phylogeny (Figure 3). This topology showsN. sphaerocarpa as most closely related to Polysiphonia for-fex Harvey, with this relationship receiving strong support(rB98). These species are similar in having rhizoids cut-off from pericentral cells, scar cells every segment in spiralseries, branches replacing trichoblasts, and tetrasporangiain spiral series, but P. forfex has (5–) 6 (–7) pericentralcells, segments shorter than wide, basal cortication, andspermatangial stichidia that developmentally replace tricho-blasts.

Neosiphonia tepida (Hollenberg) S.M. Guimaraes

et M.T. Fujii in Guimaraes et al. (2004, p. 171)

Figures 21–26

Basionym Polysiphonia tepida Hollenberg (1958, p. 65).

Synonyms Polysiphonia taylorii Hollenberg ex Williams(1949, p. 694); Polysiphonia flabellulata sensu Menez (1964,p. 219) wnon P. flabellulata Harvey (1860, p. 330)x.

Description Plants to 1.5 cm tall, erect branches arisingfrom a prostrate branching system attached to substratum byrhizoids cut-off from pericentral cells (Figure 21); highlybranched in an alternate to subdichotomous pattern (Figures22, 23); erect axes 50–75 mm in diameter, prostrate axes125–175 mm in diameter; mid axis segments of erect axesmostly 1.5= as long as wide; cortication lacking; main axeswith 7–8 pericentral cells (Figures 24, 25), occasionally 5–6in immature axes; branches forming in the axils of tricho-blasts (Figure 26); trichoblasts to 620 mm in length, typicallywith 2–3 dichotomies; scar cells obvious, mostly every threeto four segments and in no particular pattern; adventitiouslaterals absent; tetrasporangia scattered or in short to longstraight or slightly spiral series, 50–95 mm in diameter; sper-matangial stichidia developing as a furcation of trichoblasts,60–80=250 mm, without sterile tip cells; cystocarps sub-globose to urceolate, 160 mm in diameter, stalked, with wideostioles.

Type locality Beaufort, Carteret County, North Carolina,USA.

Other sources Dawes and Mathieson 2008; as Polysi-phonia tepida, Abbott 1999, Hollenberg 1958, Hollenberg1968b, Kapraun 1977, Kapraun 1980, Schneider and Searles1991, Taylor 1960; as Polysiphonia flabellulata, Menez 1964.

Specimens studied Florida: WNC2010-s77 to s79(FL05-02), Sebastian Inlet, Indian River County, B. Stuercke,26 Feb 2005.

Molecular voucher GenBank accession numberHM573552 (rbcL).

Remarks Hollenberg (1958) originally described Neosi-phonoia tepida (as Polysiphonia tepida) as having tetraspor-angia in slightly spiral series but in a later publicationdescribed tetrasporangia as developing in straight series(Hollenberg 1968b). This later description agrees withdescriptions of P. tepida provided by Abbott (1999), Menez(1964, as Polysiphonia flabellulata Harvey), and Schneiderand Searles (1991). Tetrasporangia were not observed in thisstudy but are listed above as developing in straight or slightlyspiral series according to Hollenberg’s descriptions.

Only one sample of Neosiphonia tepida was collectedfrom Sebastian Inlet, Indian River County, Florida, USA.COI and SSU sequence data could not be generated for thissample. Cluster analyses of the N. tepida rbcL sequenceshow it to be similar to that for Polysiphonia isogona Harvey





Figures 27–32 Polysiphonia anomala.(27) Prostrate axis with a rhizoid cut-off from the pericentral cell, FL09-41B, scales20 mm, WNC2010-s048. (28) Habit of erect axes,FL09-78, scales0.20 mm, WNC2009-s149. (29) Portion of erect axis flattened to show four pericentral cells per segment, FL09-41B,scales25 mm, WNC2010-s048. (30) Apex of lateral branch bearing trichoblasts (arrows), FL09-41B, scales0.10 mm, WNC2010-s049.(31) Main axis showing development of adventitious laterals, FL09-77, scales0.20 mm, WNC2009-s145. (32) Reproductive branch dis-playing long spiral series of tetrasporangia (arrows), FL09-77, scales50 mm, WNC2009-s145.

(Figure 1), with sequences differing by only 1.85%. This iscomparable to the maximum intraspecific rbcL sequencedivergence of F2.13% (predominantly F1.3%) that hasbeen observed in previous studies of Polysiphonia (McIvoret al. 2001). Neosiphonia tepida is resolved in a stronglysupported clade (rB100) of multipericentral cell species inML analysis of rbcL sequence data (Figure 3). Neosiphoniatepida is sister to P. isogona in this tree, with the relationshipreceiving strong support (rB100).

Neosiphonia tepida and Polysiphonia isogona share char-acter states of rhizoids cut-off from pericentral cells, scarcells that are variable in pattern and frequency, spermatangialstichidia developing as a furcation of trichoblasts, and sub-globose cystocarps. If N. tepida has tetrasporangia in spiralseries, then this character state would also be shared. P. iso-gona can be distinguished from N. tepida by having erectaxes to 250 (–300) mm in diameter, segments 2–4 (–6)= aslong as wide in mid portions of erect axes, spermatangial

branches with 1–3(–5) sterile tip cells, and branches thatdevelop to the side of trichoblasts (Adams 1991, Womersley1979, 2003).

Neosiphonia tepida and Polysiphonia isogona may alsodiffer in pericentral cell number. Hollenberg originallydescribed N. tepida as ‘‘mostly with 7–8 pericentral cells’’(Hollenberg 1958). Whether the term ‘‘mostly’’ indicatesthat fewer or more pericentral cells were observed is uncer-tain. Harvey did not indicate the number of pericentral cellsin his original description of P. isogona (Harvey in Hooker1855), but this species is described as having 9–10 (–12)pericentral cells by Adams (1991) and (8–) 9–10 pericentralcells by Womersley (1979). Womersley also remarked thatthis species may rarely have seven pericentral cells, as sev-eral samples collected from southern Australia appearedidentical to P. isogona except for this difference in pericentralcell number and were therefore considered ‘‘unusual vari-ants’’ of the species (Womersley 1979). He further com-





mented that comparison of P. isogona and N. tepida (as P.tepida) is required. Specimens examined in this study strictlyhad no more than eight pericentral cells. Thorough compar-ison of type material and molecular and morphological anal-yses of topotype material are needed to clarify the status ofboth species.

Genus Polysiphonia

Polysiphonia anomala Hollenberg (1968, p. 59)

(Figures 27–32)

Description Plants small, to 5 mm tall, forming densetangled mats, erect determinate branches arising from anindeterminate prostrate branching system attached to the sub-stratum by rhizoids cut-off from pericentral cells (Figure 27);erect axes simple or with limited subdichotomous to irregularbranching (Figure 28); erect axes 40–50 (–80) mm in diam-eter, prostrate axes 80–100 mm in diameter; mid axis seg-ments of erect axes mostly 2.5–3= as long as wide;cortication lacking; main axes with four pericentral cells(Figure 29); relationship of branches to trichoblasts unknown;trichoblasts with several dichotomies, long and delicate, to900 mm in length (Figure 30); scar cells obvious, variablein pattern and frequency; adventitious laterals frequent tooccasionally present, linear in shape (Figure 31); tetraspor-angia slightly distending segments, in long spiral series (Fig-ure 32), 45 mm in diameter; spermatangial stichidiadevelopment unknown; cystocarps ovoid to slightly urceo-late, short stalked, 120–140 mm in diameter.

Type locality Bikini Atoll, Marshall Islands.

Other sources Abbott 1999, Dawes and Mathieson2008, Hollenberg 1968a.

Specimens studied Marshall Islands: US-48521 (slidenumber US-1100), Holotype, Amen Island, Bikini Atoll, G.J.Hollenberg, 07 July 1948; Florida: WNC2009-s145 to s147(FL09-77), mangrove near Keys Marine Laboratory, LongKey, Monroe County, N. Mamoozadeh, 10 March 2009;WNC2009-s148 to s151 (FL09-78), mangrove near KeysMarine Laboratory, Long Key, Monroe County, N. Mamoo-zadeh, 10 March 2009; WNC2010-s048, s-49 (FL09-41B),Lake Surprise, Key Largo, Monroe County, N. Mamoozadeh,09 March 2009; US-66362 (slide numbers US-2352 to 2354),Panama City, Bay County, M.L. Jones, 17 Jan 1958; US-2355 to 2357, Along highway near NW side of lagoon, LakeSurprise, Key Largo, Monroe County, E.Y. Dawson, 28 May1949.

Molecular vouchers GenBank accession numbersHM573549, HM573550 (rbcL); HM560654 (SSU);HM573502 (COI).

Remarks Hollenberg originally described Polysiphoniaanomala with scar cells occurring one per segment in 1/4

spiral series on both prostrate and erect axes (Hollenberg1968a). Hollenberg also noted that a very similar specimencollected by E.Y. Dawson from Lake Surprise, Key Largo,

Florida (US-2355 to 2357) closely resembled P. anomala andwas therefore identified as such by Hollenberg in the originaldescription of the species. Hollenberg observed slight differ-ences between Dawson’s Florida samples and the Pacificholotype of the species, however, including scar cells that donot occur regularly one per segment on prostrate branchesand tetrasporangia that occur in much longer series in theFlorida specimen. This same scar cell pattern was observedin all other Florida samples examined in this study. Tetra-sporangia were also observed occurring in long spiral seriesin WNC2009-s145 to s147. Scar cell pattern alone is not areliable species identifier as this character is often variablewithin species (Stuercke and Freshwater 2008); it is unclearwhether length of tetrasporangial series can be independentlyused to distinguish species.

Three samples of Polysiphonia anomala were collectedfrom Lake Surprise and Long Key, Monroe County, Florida,USA in this study (Figures 1, 2). Two of the three rbcLsequences generated for these samples are identical and thethird differs by only 0.37%, which is well within the rangeof intraspecific rbcL sequence variation observed in previousstudies of Polysiphonia s.l. (e.g., McIvor et al. 2001). Onlyone COI sequence was generated from these specimens. MLanalyses of rbcL and SSU sequence data resolve P. anomalaas an independent lineage with variable levels of support(rB-50 and 56; sB91 and 69) for its topological position(Figures 3, 4).

Conclusion

Four species of Neosiphonia and one species of Polysiphoniawere identified in 16 samples from Florida and CaribbeanMexico. This is the first report of Neosiphonia bajacali inthe Caribbean and of N. echinata from Caribbean Mexico.Samples of the other three species do not represent newreports from their collection sites, but help confirm theirknown distributions. Further study of Caribbean Neosiphoniaand Polysiphonia is necessary to determine the true genericstatus of the species reported for this widespread area. It alsoseems likely that more than 29 of the nearly 200 Neosipho-nia/Polysiphonia species are present in this diverse region.

Acknowledgements

The authors wish to thank J.C. Bailey for reviewing an early versionof the manuscript, two anonymous reviewers and the editor for help-ful comments and edits, those listed in the Appendix who providedsamples, and J. Kelly and B. Stuercke for the use of unpublishedsequence data. We are grateful to D.H. Freshwater for facilitatingMexico collections, J. Norris for assisting with US specimens,and S. Schmitt for fieldwork help. This research was funded byNSF-BS&I grant �0742437 and the CMS DNA Algal Trust.





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Fres

hwat

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mor

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001)

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AF4

2753

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Mar

1997

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(App

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x1

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ple

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tion,

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Oct

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Oct

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Fres

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and

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mer

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nd,

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land

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5735

7928

Oct

2004

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.W.

Fres

hwat

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nia

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land

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and

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001)

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3960

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olys

ipho

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pani

cula

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Kim

etal

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004)

3Se

noO

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nta

Are

nas,

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leA

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al.

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4)4

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Mile

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ch,

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aC

ruz,

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,U

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Y61

7144

Pol

ysip

honi

ape

ntam

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PHY

KO

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IR

esea

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Stat

ion,

Punt

aG

alet

a,C

olon

,Pa

nam

aH

M57

3563

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5606

44H

M57

3510

Hol

lenb

erg

14M

ay20

09,

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Wys

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rgen

tPH

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5606

4315

Jan

2008

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and

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PHY

KO

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dane

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B.

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enPH

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3531

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iaH

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uas,

Pana

ma

HM

5735

6415

Jan

2008

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and

J.A

lden

PHY

KO

S-35

32Is

laA

fuer

a,V

erag

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Pana

ma

HM

5735

6416

Jan

2008

,B

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and

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ms

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5734

9501

Nov

2004

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hwat

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Paua

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5735

7601

Nov

2004

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Fres

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912

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land

HM

5735

76H

M56

0637

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5734

9502

Nov

2004

,D

.W.

Fres

hwat

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Z04

-309

Ulv

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land

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art

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land

HM

5735

76H

M57

3495

03N

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04,

D.W

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ater

and

M.

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mer

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04-5

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HM

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04,

D.W

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ater

and

M.

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mer

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04-5

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nd,

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land

HM

5735

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M57

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17N

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04,

D.W

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and

M.

Hom

mer

sand

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04-5

57M

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bor,

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thIs

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ndH

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3576

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5734

9517

Nov

2004

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.W.

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(App

endi

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Spec

ies

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ple

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lect

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tion,

date

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dco

llect

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Jul

2008

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KO

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Co.

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Jul

2005

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hwat

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Aug

2009

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5735

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May

2009

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4929

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Mar

2005

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ma

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3582

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5735

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M.

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mer

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NZ

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hIs

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ewZ

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ndH

M57

3582

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3026

Oct

2004

,D

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Fres

hwat

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5835

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5735

3429

Oct

2004

,D

.W.

Fres

hwat

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HM

5735

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Nov

2004

,D

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hwat

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Z04

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Bea

ch,

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3858

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3532

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04,

D.W

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M.

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(App

endi

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Spec

ies

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Nov

2004

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PHY

KO

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2008

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2009

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References

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Abbott, I.A. and G.J. Hollenberg. 1976. Marine algae of California.Stanford University Press, Stanford, CA, USA. pp. 844.

Adams, N.M. 1991. The New Zealand species of Polysiphonia Gre-ville (Rhodophyta). New Zeal. J. Bot. 29: 411–427.

Agardh, C.A. 1824. Systema algarum. Literis Berlingianis, Lund.pp. 312.

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Athanasiadis, A. 1987. A Survey of the seaweeds of the Aegean Seawith taxonomic studies on species of the tribe Antithamnieae(Rhodophyta). University of Gothenburg, Goterna, Kungalv,Sweden. pp. 174qxl.

Ballantine, D.L. and N.E. Aponte. 1997. A revised checklist of thebenthic marine algae known to Puerto Rico. Caribb. J. Sci. 33:150–179.

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Received 18 October, 2010; accepted 23 February, 2011; online first1 June, 2011



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