Phycologia (2006) Volume 45 (1), 1-9
075
Published 3 January 2006
,
I
\I!
Creation of the subgenus Testeria Faust subgen. nov. Protoperidinium Bergh from the SW Atlantic Ocean: Protoperidinium novella sp. nov. and
Protoperidinium concinna sp. nov. Dinophyceae
MARIA A. FAUST'"
Department of Botany, US National Herbarium. Smithsonian Institution, 4210 Silver Hill Road, Suitland, Maryland 20746, USA
M.A. F'\\·ST. 2006. Creation of the subgenus Tcsteri« Faust subgcn. nov. Protoperidinium Bergh from the SW Atlantic Ocean: Protoperulinitnn novella sp. nov. and Protope ridinium concinna sp, nov. Dinophyceae. Ph ycologia45: 1-9. DOl:
10.2216/04-62.1
Two new heterotrophic Protopcridinium species arc described from oceanic waters from The Gulf Stream, S\V Atlantic Ocean and Belizean Atlantic Barrier Reef Ecosystems, Caribbean Sea. Protoperidiuium novella Faust sp. nov. and P.
concinna FaustSf'. nov. show a plate formula of 4', La, 7", 4C(3+t), 6S, 5"', and 2"", which is atypical for the genus. It is charucrerizcd by a miniscule flat closure on the extreme anterior of the pointed apical horn. Apical pore complex is absent.
Apical ventral plate I' disconnects from the tip of the apical horn and connects directly 10 the anterior sulcal plate (Sa) along a straight line of the longitudinal axis of the cell's sagittal plane. The shape and position of the intercalary plate (1a) is another distinctive feature fur these species, the presence of only one Ja plate. The thecal plate features ofP. novellaand P. concinnajustifies the establishment of the new subgenus, Testeria Faustsubgen. nov. The relationship with otherCOIl- generic species and theposition within the genus Protopcridiniumare discussed.
Kr.v\VORIJS: Belize. coral reef-mangrove pond, dinofiagellutes. Dinophyceae , ecology. Subgenus Teste ria subgen nov., Protoperidinium novellasp.nov ..Protoperidinium concinna
sr.
nov .. scanning electron microscopy,taxonomyDedicated/0 Dr PatriciaA. Testerill recognition of her significant contributions to The dynamic's ofKarcnina brevis (Davis) G. Hansen
& 0.Moestrup algal blooms.
INTRODUCTION
Protopcridiuium Bergh is a large genus of armoured dinofla- gellatcs usually without chloroplasts. Bergh (1881) realized that thecal plates always exhibit a certain basic pattern com- mon within a genus. Schutt (1895) noted the importance of morphology identifying dinoflagellates. JOrgensen (1912) em- phasized the importance of thecal plate pattern for species identification in the genus and presented new groups: e.g. Per- idinium, whose divisions were based on the shape of the first apical I' plate, and the position and shape of the dorsal epi- thecal plates, particularly the second intercalary 2a plate and the presence of three intercalary plates. Jorgensen (J912) es- tablished a second genus Archaeoperidiniuni with only two anterior intercalary plates. Balech (1974) transferred 231 spe- cies of marinePeridiniumto the genusProtoperidinium which included species with ortho, meta and para apical plate 1', intercalary plate 2a with hexa, penta and quadra-rype, four cingular plates, and six sulcal plates, and left Peridinium to include only those with ortho-type of I' plate, five or six cin- gular plates and five or six sulcal plates. Balech (1980) rec- ommended that the morphology of cingular and sulcal plates, the shape of apical plate 1', and the intercalary plates be used in species identification.
~Corresponding author ([email protected]).
The use of a scanning electron microscope provided addi- tional new information on the (inc morphology of thecal plates (Gocht& Netzel 1974; Baleen 1975, 1999; Taylor 1976; An- dreis et al. 1982; Dodge 1983, 1985; Netzel & Dilrr 1984;
Lewis & Dodge 1990; Delgado & Fortuno 1991; Hansen &
Larsen 19<)2: Tori u mi & Dodge 19(3). Toriurni & Dodge (I <)93) found that the apical pore structures of Protoperidi-
lIiUII/ species were species specific, enclosed by up to six col- lar plate sections and the narrow canal plate X, and varied in morphology. The genus Protoperidinium identified by the typ- ical thecal plate formula: Po, X, 4', 2 or 3a, 7", 4C(3+t), 6S, 5"', and 2NN (e.g. Steidinger& Tangen J(96). Classification of Protoperidinium today are based on prominent morphological features including cell size and shape, horns or spines, apical plate J', intercalary plates, the apical pore complex (APC), ornamentation of thecal plates, displacement of cingulum and six sulcal plates (e.g. Toriumi & Dodge (1993)).
The growing recognition that armoured dinoflagellates are important members of marine ecosystems has accentuated the need for detailed taxonomic studies (Yamaguchi & Horiguchi 2005). Information is Iirnited on the biodiversity ofProtoper- idinium Bergh (I881) dinoflagellates from the SW Atlantic Ocean and the Belizean Atlantic barrier coral reef ecosystems.
This paper illustrates the thecal plate morphology of two new Protoperidinium in recent marine collections. Both species are heterotrophic, exhibit similar cell shape, and a slender apical horn characteristic for the sectionOceanica Jorgensen (1 <) I 2), the genus Protoperidinium. Protoperidinium novella sp. nov.
and P.concinna sp. nov., however, differ in thecal plate mor- phology by the absence of an APC, the apical ventral plate l ' that disconnects from the tip of the apical horn, and the pres-
2 Phy cologiu , Vol. 45 (I), 2006
Table 1. Comparison of mor phological features of Pro toperidinium novella sp,nov. andP. concinn u sp. nov. and selected Prot op cridinium species.'
Cell size(p.m)
Taxa APC Plate
"
a* Length Width References
P. novella no onho la 98- 116 62-7:; Presentstudy
P. concinna no onho 1a 103-137 52-67 Present study
P.steiding e ra e no ortho 3a 117-/1<2 64-1<1< Balcch Il188,p. 188.fig.1<5:1-7
P.oblon gum yes ortho 3<J 130 -/47 70-HO Hansen& Larsen IlIn ,p. 124.figs4.72<J-e
P.ocean icum yes ortho 3a 155-/ 65 71;- 80 Abe 1981.p.324. lig.300-2.
P. cla ud icans yes ortho 3a 105-50 76--48 Dodge IlI1<2.p. 182.fig.200 .
P. ca rlls yes ol1ho 3a 85-93 45-48 Abc 1%1. p.271. tigs 337-4 1
P. compla nu tum yes ortho 3a UO 110 Abe 1981,p.276.figs 249 - 54
P.depre ssum yes ortho 3a 116-200 1/6-144 Dodge 1982.p. 177.lig. 20A
P. va lg us yes ort ho 2a 105-11 3 68-72 Abe 1981,p.318. tig.43:2H2- 84
P.con simili" yes or tho 2a 53-58 45-50 Abe 1981,p.316, tig.42:276- 81
'1' ,apicalplate;a ",referstothe numbers of intercalary plates; urn, micrometre.
ence or only one intercala ry plate 1a. 1 report here, unusual thecal plate paucrnon P.no vella sp. IIOV. and P.concinnasp.
nov. The se Protoperidiniutn species exis t in oce a nic neriti c wate rs in The Gulf Stream off shore Point Lookout, No rth Carolina, andoffsho re Fort Pierce Inlet, Florida,SW Atlantic Ocean, and the Atlantic Barri er Coral Reef Ecosyst ems, Be- lize, Central America.
MATERIAL AND METHODS Sample collections
Dinorlagell ates were co llec ted in warm tropical ocean waters at three distant location s : (I) in The Gulf Stream off sho re Point Lo okout,North Caro lina(34C23' N, 79C56 'W) (2002and 2003) ; (2) off shore Fort Pierce Inlet, Flo rida (2r32' N, 79C54 ' W) (200 1 to 2003 ); and (3) outs id e Douglas Cay (16°4 3'N , 88°13 ' W) and Manatee Cay (16c39'N,
ss-n
'W)(1994 to 1996) . Belize . Cent ra l Ame rica . Douglas Cay and Manatee Cay are oceanic mangroves. biologicall y diverse mangrove ponds within the PelicanCays Archipelago, Belize (Macintyre & Ruetzlcr 2000). The cays are considered Ho - locene lagoon reefs(Purdy 1994)colonized byred mangr ov es , RhizophoramangleLinna eu s.The biolog y andec o logyof this ecosystem is complex ; it sup po rts an uncomm onl y rich faun a and flor a ofmarine, benthic and planktoni c org a nis ms (D.S.
Littler& M.M.Littler 1997; Fau st 2000;Ruetzleretal.2000). Two sampling methods wer e used: (I) ahori zontal tow bel o w the water sur fa ce using 20 III pore size plankton net towed by a boat at low speed for 5 minand (2) a vert ica l tow from lO
to 100 J1l depth betwe en su r face usin g the same 20 m pore size net with an attached we ig ht. Water temperature ranged from 28.5cC to 33.9°C. Salin ity le vel s ranged from 32.3 to 36.7 psu.
Sample preparation
Forscan n ing electron microscopy (S E M) preparation . plank- ton sa m p le s we re conc e ntra ted to lao 1Il1 vo lume and fixed with I% glutaraldeh yde finul concentration(Faust 1990).The fixed dinoflagellate s were isolated using a capillary pipette under a Zeiss stereo microscope.Cell swere concentratedonto a polyc arbonate filter at room temperature. rinsed six to eight times with deionized water, dehydrat ed in a graded series or ethanol conc e ntra tio ns, and critical-point dried. The preparu- tion was coated with ca rbo n foll owed by a layer of gold- pall ad ium (F aus t 1990). Ce ll size est imated Irorn SEM pho- togr aph s (at least 10cells); range s of cellsize sho wn in Table 1. Cell length estim at ed from anterio r to posterio r in gird le view , and cell width est imated from trunx-diamctcr in lateral vie w. Kofoidian nomencl ature was used for identificat ion of species (Kofoid 1909). Dinoflagellute specimens used in this investi gation arc deposited in the US Dinoflagellate Type Col- lection, the US Nation al Herbarium, Department of Botany.
NMNH , Smithsonian Institution , Washington. DC 20013- 7012 , USA.
RESULTS
Phylum: Pyrrophyi a Pash er, 1914 Class;Dinophy cea e Fritsch , 19 29
Figs 1-7.Scanning electron microg raphsofProtope rid inium novellasp. nov.Faust.
Fig. 1. Ven tral view. Cell hasanapical horn and two antapical horns: ventrally depressed. Apicalplate I'ortho disconnected fromthe apical horn(arrows).
Fig. 2.Apicalcellviewillustrates thetip of apical hornas aminiscule flatclosure(arrowhead),and Larectangularinrcrcalaryplate,and I' 2'.& 4' ridged apicalplates.
Fig. 3. Apicalpore complex is absent(arrow) andsurfacesculptured.Anteriorintercalar y plate1aislarge and rectangular.
Fig. 4. Dorsal view.Cell is slender,two antapical hornspointed andequalin lengthand shape. Intercalar y plate Jais large.Postcingular plate 3'" ispenta gonal andvery large.
Fig. 5. Cingulum is equatorial. descend ing 1 X its width. Cingular list is smooth. Sulcus is deep and narrow. bordered by sulcal lists (arrow heads), Sulcalplates are:Sa,Sd,Ss, Sp. Apicalplate I' touchesthesulcal Saplate.
Fig. 6. Apical 4' plateislongand narrow situated between apicnl plate I' andintercalary plate 1a.
Fig. 7. Thecal plate surface rugose,some plateswith a smallround pore situated in the centre(arrowheads) and poresin line on thecingulum.
Faust: Protop eridiniunino vella sp. nov. Faust and Protoperidinium conc inna sp.nov. Faust 3
4 Phycol ogi a, Vol. 45 (I), 2006
Protoperidinium novella Faust,sp. nov.
Subgenus: Testeria Faust
Cellulae arrna tue,98-116IJ-m lon guecr62-75IJ-m larue,py riformes. Dispositi o disc orum:4', La,7",4C (3+t). 65.5'". et 2"".Valva ap- ical is J'ortho.Valvaapiea lis(I')orthoetva lvaintrc ca laris Lahexa. APC abs q ue . Cingulum equa to ria lc cxca vutum dcrnoturn IX ejux luritudinis. Sulcu s profundus et marginutus.Hyporhecu ventraliter dcprcs sa, Spinae duae in valiv ibus antapica les I"" er2 "": Valvae rugosae. Chrornato phorcsabsq ue,Flagellae duac.
Cell s armou red. 98-116 urn long (L), and 62- 75 u.m wide (W) sha pe pyriform.Theplate tab u latio n: 4' . la, 7".4C(3+t),6S,5"',
2>NO. Sha pe ofapica lplate I' isort ho. Intercal ar yplate is Ia,and
sha pe of plate Iaquud r a .APC is abse nt.Cing u lu mequa to rial with cing u lar lists, excavated,disp laced 1X its width. Sulc us is deep . borderedby sulcallist.Hypotheea ve ntra lly slightl y.depressed . Two hornsare present onanrapica l plat es 1""an d2 "":Thec al sur face is rugose.Chro matophoresareabse nt. Fla gella arc two.
TYPE:Fig. I,collec te dby Faus t.Sam p le no.2.W~-200I.
ISOT YPES: Figs 2-6,sample no.2488-2002: Fig .7,samp le no.2367- 20 02.
Figs 1-12
11
C{ : . .. ~~ . _ . . _ ._~~ .:~._ . .. .. ~:;""
: 4' '.
:. 6" 2' 2" :
... / r ....
•• 7" 1" ...
10 - .
TYPESPEC IES: Protop eridinium no vellasp . no v.Fa ust.
Subgenus: Testeria Faust, subgen.nov.
ETYMOLOUY: Named afte r Dr Patr icia A.Testerin rec og niti on of her significa ntcontr ib utio nsto understandingthe dynamicsofKarenina brevis(Da v is) G.Hansen& 0. Me strupharmful alga l blooms.
Dinoflagell atac arma tae. Epithe c a unicorni s er hypoth ec a bic orni s aeq ua lis. Valva apica lis (I')symmetricalis "ortho ' et cornu apic a li separate.APC abs q ue. Disc i inte rca lis (Ia) quadra. Dispo sitodis- corum:4' ,3a.7",4C(3+t), 65,5'". et 2"",Pagin arugosa thecae.
ET YMO LOGY: No vella (Lat in) mean sunfamili ar structure.
DISTRJ B1JTJON: Know n fromwa te rsTheGu lfStrea mocea nic curre nt offshore fromPoint Lookout.NorthCarolina.andoutsi de Ma nate e Cay (J6°3 9 'N. 8~olJ'W). and Dou glas Cay (16°43' N. l)~oI3' W), Belize. and in The Gulf Stream off shore Point Looko ut. North Caroli na (34'13 'N.79°56'W). USA .
TYPE l.OCALlTY: Gulf Stream off sho re ron Pierce Inlet, Florida (27°32'N. 7yo54'Wl,Southwes t Atlant ic Oce an, USA. Samplepre- served in gilitma idehy de, curr e nt ly held in the US Dinotiag ellate Type Coll ec tio n. US Nati on al Herba rium, Department of Bot an y , NMN H , Sm ith s oni an Instituti on , Washing to n DC, 200 13-70 12.
USA .
Cells of P. no vell a sp. no v.armo ured. pyrifo rm insha pe with a po int ed apica l hornandtwo ant up ica lhorns (Fig s 1,4, X-12).Ep ithec u and hypotheca are eq ua l in size. Cells are 98-1J6 IJ.I1J lon g (L) (X
104.2 ::: 8.3;n = 9)and62-75u.mwide (W) (X70.3:t 5.6 :n = 9),
LfW = 1,48 (Table 2). The shapeofapica l pla te I' is ort ho ,andiris disconnecte d by 25 rn from the point ed and slender apic a l horn (Fig . I), but ap ica l I' plate in direct co ntact withanterior sulcal (Sa) plate (Figs I,2, 5. 8). APC is abse nt; is cha rac terized by a miniscul e tJat closure on the extreme anter io r ofthe pointed apica l horn (Figs 1-4 , 6).The anterior inte rcalary plate la is quau ra situated do rsallyadja- cent apical 2' to 4' plates (Figs 2-4,9, JO).Surfac e ofP.no vell ais rugose , covered by polygonal intricate ridged surfac e pattern and small uneven lydistributed round por es (Figs1-7).Cellsventra lly de- pre ssed (Figs I, 5, 10 , 11). Ch lorop lasts are absent. Cel l conte nt is colo urlessorpale pink.
The cing u lum isdeep,equat orial ,descending I X itxwid th (Fig s I, 5). It bears promin ent smooth cing ular lists (Figs I,2,4, 5, 7).
Four cingu lar plat es are pre sen t4C (3+t): cingu lar plat e s ICplat e is cons ide redatran s ition al pla te(t),C2andC4 platesarc rathe r narrow, whereas.C3 plate very broad and consti tute a major part of the cin- gulum(Figs8,9). Row of sma ll round poreslinethecingu lu m surface (Fig.7) .
no v. Faus t.
Armoured dinoflagellate. Epitheca one-horned are equal in size to the two-ho rned hypotheca.Ap ica l plate (1') issy mmetrical,ort ho, and discon nec ted from the apic al ho rn .APC is absent.With inter- calary plat es Ia quadra.Arrangemen tofplates :4', la,7",4C( 3 +t), 6S, 5"', and2"". Thecal surface isrugo se.
Order :Peridiniales Haeckel. 1894 Family:PeridiniaceueEhren be rg , 1832
Genu s :Prot opcridinium Bergh ,emend Balech, 1974 Subgenus : Tcsteri a subge n. no v. Faust , 2005
Fi~s 8-12.Line drawing of Protop eridinium no vella sp.
Morphologyofthe calplat e s illu st rat edin var ious views.
Fig.8. Ventral.
Fig.9. Dor sal.
Fig. 10. Apical.
Fig. II. Anrap ical, Fig. 12.Sulca l.
Table 2. Comparison ofthe mor pholog ica l fea tures of Prot operidiniumnov el/a andP. concinna.'
IL,length;W, width ;L'W, ratioof length/Width; la, intercalar yplat e Ia.
L W
Cellsize (p.m)
6.5 :::03 7.3:!: 0.3 la Plate
L W
2l.9 1.1 26. 5 :t 2.0 ::t1.0
:t 0.8 w 15.8 J3.4 Antapical horn L
28.9 ::t 1.2 30.5 ::t1.9 9.3 :t0.8
7.4 :t 0.7
L W
Apica lhorn
19.5 :::: 2.2 24.9 :!: 2.2 1.4 8
2. 13 Rati o L/W :!:5.6
5.0 70.3 58.2 104.2 :!: 8.3
124.0 :!:11.4 P.no vella
P.concinna Ta xa
Faust: Protoperidiniurn novella sp. nov. Faust and Protoperidinium concinna sp. nov. Faust 5
The hypothcca is oblong and ventrally depressed (Figs I, 5. 10.
1J). Postcingular plates arc short, but 3'" plate is the largest (Figs 4.
9). The postcingular listcontinuously follows along the deep and nar- row sulcus (Figs I. 5. X). Sulcus bordered by widened postcingular sulcal list (Figs I. 5. 8). It is bearing six sulcal plates. where Sa plate extends anteriorly into the epithecu (Fig. 5). Two dimorphic flagella present, but not shown. Two antapical horns arc 40 ml.eac h with tins associated with aniapical plates Iun and 2 "" (Figs I. 4. 5. 8.9. II).
Plate fonnula: 4'. la,7".4C(3+t),6S, 5'''. and 2"".
HABITAT: Protoperidinium novclla sp. nov .•is a neritic. heterotro- phic dinoflagellate species. identified from The Gulf Stream oceanic current oil shore from Point Lookout. and off shore Fort Pierce Inlet.
Florida (27C 3 2 ' N ,
79°54'W). and SW Atlantic Ocean, USA; and the Belizean coral reef-mangrove ecosystems outside Manatee Cay and Douglas Cay. Caribbean Sea.
ASSOCI/\TEIlALC;/\I': In the collections P. novella sp. nov. were as- sociated with coxmopoliran and world wide distributed oceanic spe- cies. The biodiversity ofdinotlngcllatc species varied in each collec- tion. Photosynthetic species were:ProrocentrumCO/ll.preSSlII11(Bailey) Abe ex Dodge, P. gracile Schutt, P. lim« (Ehrenberg) Dodge & P.
micans (Ehrenberg), Pvropluuus steinii (Schiller) Wall & Dale and Pyrodinium bahamense Plate var, bahamense. Heterotrophic species were: Phalachrorna 1'<11'(1Jorgensen and P. rotundaturn (Claparade &
Lachmann) Kofoid & Mitchener. Dinophvsis caudutu Saville-Kent.
Ornithocercus thumii (Schmidt) Kofoid & Skogsberg and O. splcn- didus Schurr. Crrarocorvs horrida Stein. and Palcophalachroma rrn- icinctuni Schiller.
Subgenus: Testeria Faust
Protoperidinium concinna Faust,SjJ. nov.
Figs 13-25
Cellulae armatac. 103-137 fun longae ct 52-67 fun latac,pyrifor- mcs. Dispositio discorum: 4'. Ia. 7". 4C(3+I) .6S. 5''', et 2 "", Valva apicalis I' ortho. Valva apicalis (I') ortho et valva intrccalaris ta quadra. APC absque. Cingulum equatoriale excavatum demotum IX ejus latitudinis. Sulcus profundus et marginatus. Hypotncca ven- traliter deprcssa. Spinae duae in valivibus antnpicales I "" et 2 "".
Valvae rugosae. Chrornatophores absque. Flagellac duac.
TVpJ·:: Fig. 13. collected by Faust. Sample no. 2:198-200l.
ISOTYPES: Figs 14-25.sample no. 2488-2002; Figs 7. 8, sample no.
2367-2002.
TYPE LOCALITY: The Gulf Stream off shore Point Lookout, North Carolina (34°23'N, 79"56'W). SW Atlantic Ocean. USA. Sample preserved in glutaraldehyde. currently held in the National Dinofla- gellate Type Collection. US National Herbarium, Department of Botany, Smithsonian Institution, Wasbington DC. 200 13-70 12.
USA.
ETYMOLOC;Y: Concinnu (Latin) means harmony and proportion.
DISTHIlll iTION: Known from waters outside Manatee Caymangrove pond (16°39'N, 88°II 'W ). and Douglas Cay (l6°43'N. 88°J 3'W).
Belize, and Gulf Stream off shore Fort Pierce Inlet. Florida (2r32'N, 79°54'W). SW Atlantic Ocean, USA.
Protoperidinium concinna sp. nov. a heterotrophic marine species, slender in cell shape, dorso-ventrally compressed with characteristic long apical and antapical horns (Figs 13-15, 22, 23). Cell size ranges between 103-137 u.m (L) (X 124 :+: 11.4;n
=
9) and 52-67 urn (W) (X58.2 :+: 5.0; n = 9; Table 2). Epitheca and hypothec a equal in length. Thecal surface polygonally ornamented with intricate shallow ridged polygonal surface pattern and small unevenly distributed round pores (Fig. 21), and recognized as faint striations under the light mi- croscope. Cells lack the APC, characterized by a miniscule flat closure on the extreme anterior of the pointed apical horn (Figs I3-l7, 22, 23). Apical plate I' ortho (Figs 13, 16. 18, 22, 24), disconnected from pointed and slender apical horn but in direct contact with the anterior sulcal (Sa) plate (Fig. 20). Anterior intercalary plate Ia is quadra (Figs 14, \6, 23. 24) situated dorsally between apical plates 2' to 4' (Figs 16,23,24).Cingulum is deep, equatorial and displaced about I X its width (Figs 13- J5, 18, 19, 22),Itbears prominent smooth cingular lists (Figs 13-16, 18-20).Four cingular 4C(3+t) plates are present: IC is a small transitional plate (t) (Fig. 20), 2C and 4C plates are rather narrow, whereas.C3 plate is very broad and constitutes a major part of the cingulum (Figs22. 23).
The hypotheca is oblong and ventrally depressed (Figs 1.1,IS, 18, I'J). Poxtcingularplates arc short and 3'" plate is the largest(Figs 14, 23). The postcingular list follows along the deep and narrow sulcus (Figs 13. 18, I'J. 22). Two dimorphic Ilagella present. but not shown.
Aruapical hornseach with tins, associated with ant apical plates I ,,,, and 2"" (Figs 13-15. 19,22,23).
Sulcal list smooth (Figs 16. 18-20) bordered by widened postcin- gular sulcal list (Figs 13, 19). Sulcus is deep and narrow composed of six sulcal plates (Fig. 26). The postcingular list continuously fol- lows along thc narrow sulcus. The anterior sulcal plate Sa extends into the epitbcca and touches the apical I' plate at the cingular-sulcai junction (Figs 18-20). Nucleus situated centrally with radiating pro- toplasmic strands. Protoplasm is clear. Thecal plate formula ofP.con- cinna is: 4'. Ja.7",4Cn +t), 65, 5"'. and 2 '",
H,\IlITAT:Populations of P. concinna cells were associated with ne- ririe oceanic species present in coastal marine waters, coral reef-man- grovc habitats in the western Caribbean Sea: pond Manatee Cay and Douglas Cay. Pelican Cays Archipelago, Belize (Faust 2000), The Gulf Stream off shore Point Lookout. North Carolina,and off shore from the Fort Pierce Inlet. Florida. SW Atlantic Ocean (Faust& Tester 20(5). Balcch (I 'J88) recognized a morphologkally similar species, P. stcidingcrac Balech, in samples collected in both warm waters of southern Braz.il (3:1.2-35.4°C) and in cold waters of the Southeastern Atlantic Ocean (24.7-27.YC).
i\SSOCI/\r1 ,rJ ,\I.c;I\I-:: In the collections P. <:onC;1II1<1 from Manatee Cay wereassociatedwith Diplopsalis lenticula Bergh. Diplopsalopsis sp.,Dinophysis rot undataClaparedc& Lachman,Diplopeltasp.,Lin- golodinium polvcdrum (Stein) Dodge, and Prorocentrum elegans Faust. Harmful species included:P. lima (Ehrenberg) Dodge, and P.
hoffmannianumFaus t.
DISCUSSION
According to the subdivision of the genus proposed by Balech (1974, 1975, 1979. 1988. 19(9) and Abe (1936, 1(81), Pro- toperidinium novella and P. concinna can be attributed to sec- tion Oceanica in the genus Protoperidinium. Some unique morphological features, however, characterize P. novella and P. concinna different from species in the section Oceanica.
Morphology of two new species
Comparison of morphological features of P. novella and P.
concinna and selected Protoperidinlum. species in the section Oceanica are listed in Table 1. Protoperidinium novella and P. concinna differ in morphology from other congeneric spe- cies based on three morphological characters: (1) the extreme anterior of the apical horn is pointed characterized by a mi- nuscule flat closure, absence of APC (Figs 1-3, 16, 17; Table 1); (2) the apical ventral plate l ' disconnects from the tip of the apical horn; and (3) presence of two intercalary plate la.
The plate formula of P. novella and P. con cinna is as follows:
4', l a, 7", 4C, 6S, 5"', and 2'm. The unique plate structures on the episome ofP. novella and P. concinna arc supported by SEM pictures (Figs 1-7, 13-21). The morphological features that characterize them are stable characters and not aberrant forms of certain 'normal Protoperidinium':
Comparing the presence or absence of APC from line draw- ings is troublesome. Balech (l979, 1988) and Steidinger &
Tangen (l996) reported no APC on P. steidingcrae. All spe-
6 Phycol ogia. Vol.45 (1).200 6
Figs 13-17.Scann ing electronmic rographs ofProtop e rid ini um concinnasp.nov.,Faust.
Fig. 13. Ventral view, Cell is slende r with a long pointedapic al horn and two antupic al horns.Apica l plate)' ortho disconnect ed from the apica l horn (arrow he ads) ,Cingulum is displac ed widthby I X.APC is absen t. Left su lcalJist is wide .
Fig. 14. Dorsalview.Apical horn pointed.Anterio rin tercalar y (Ia)plate is la rge.Cing ulum is equa to rialwithsmoothcingu la r lists .Postcin- gu larplate (3"') is very lar ge.Antapical horns equal in leng th.broad proximall y,pointeddistall y.
Fig. IS. Epitheca exca vate d ventra lly.Loc a tion ofap icalplat e 4' is long and na rro w, situate dadjacent inte rc ala ry plate luand apicalplate I".
Fig.16. Apicalview of cpithe ca.Ap ical horn pointed, and APC abse nt (arro wh ea d). Int erc alaryplate Jais la rge.Interc ala ryband slightly striate d.
Fig. 17. Tipofapicalho rn is a minisc ule fiat clos u re, void of an apicalpor e (arrowhead). Thecal surface rug ose with scattered po res.
cies in the section Oceanica show typi calthecal mo rp ho logy.
But some published figu res are to some exten t uncertai nas to the exact feature of the APC or intercalar y plates in apical views.Illustrati on s of theAPC in SEM pictu resare limited to a few specksma king comparison with othe rspec ies diffic ult:
P. oblon gum (Hansen & Larsen 1992; Tourimi & Dod ge 1993), P. oceanicum (Tori umi & Dodge (99 3). and P. de- press um (Abe 1981;Dod ge 1985;Toriurn i& Dodge 1993).
Dis tinctive recogni zab lefeature ofP. no vella and P.COII -
cinna concerns the architecture of apicalplate I' which dis- connec ts from the tip of the apical horn si tuated directly alo ng a stra ight line of the lon g itu d inal axis at the cell's sag itta lplane. Plate I' mai nt a ins its typ ica lelo nga tedortho-
shape and seems totally disconn ected fro m the apical ho rn on whol e thecae. Such a feature of plate I' has on ly been disco vered on P. steidin gera e (Balech 1979). The pla te I' not only disc o nnects from the po int ed apic a l horn but is in direct contac t with anteri or sulcalpla teSaand avery nar ro w IC plate located in (he cing ulum-su lc usjunc ture;2Cand 4C plates are also rath e r narro w, where as 3C is ver y broad and constitutes the major part ofthe cingu lum. Altho ugh P.stei- ding erae shares a few im portant characte ris tics with these two new species, itcan bedistinguished from myspe cies by possess ing three apical interca la rypla te s .The apica lpla te I.' of thesectio n Oceanica andotherspec ies inPro top eridin ium (Tabl e J), exte nd fro m the tip of the apica l horn and ends at
Fau st: Protoperi d ini um novella sp, nov . Fau st and Protop eri di ni um concin na sp. no v.Fa us t 7
Figs18-21. Scanningelectron micrographs ofProt operidin ium conci nna sp.nov..Faust.
Fig. 18. Ventralapical view ofepltheca. Apicalplate I' isortho anddisconnectedfrom the apicalhOI11.Cellventrallydepressed.
Fig. 19. Cingulumisdescending IX its width. Cingularlistissmooth. Sulcus deep,narrow, and borderedby smooth sulcal lists.
Fig. 20. Apical plate \' touchesthe sulcal Sa platelet extendedintothecpiihccaand touches transition plate (t) at thecingular-sulcal junction.
Fig. 21. Thecalplatesurfacerugose withsmall unevenly distributed roundpores(arrows).
the sulc uson the ventra lsu rface of the episome (Abe 19 8\: To riurni & Dod g e 1( 9 3 ).
Protoperidin ium no vella and P. concinn a possess only one dorsal intercalary plat e (Tables I, 2). This small dorsal 1a plat e is only rec ogn izabl e on tiltedSEMstub (Figs 2, 16;Abe 1981 ).The morpho logyof plates on the episome on thesene w species is distinc t but com parable in line drawings ofspec ies in the sec tio n Oceanica (Ste id inge r & Williams 197 0; Balech 197 9,1988;Abe 1981).SEM pictu res of spec ies in the sectio n Oceanica are few and often very small to recognize thec al det ails (Delgado & Fort une 1(91) and at times inconsistent with so me features of spe c ies. In fact, P. oblo ngum is illus- tra ted with only one la inte rc a lary plate plus an APC (Ha nse n
& Lar sen 1992), in other publicati ons the same species ex-
hibiti ng 3a plat es plus an APC (Do dge 1985; Toriu mi &
Dod ge 1993).
In concl usion,I demon st rat ed unequi voc all ythat P.novella and P. conci nn a lack contact betw e en apical plat e I' and the tip of apical hom, APC absent,and only la intercalary plate present. I co nc lude tha t P. novell a and P. concinna are dif- feren t fro m all othe r Protop eridin ium.
Separation of new sp ecies
Pro top eri din iu ni novella and P. concinna are se para b le from each other based on cellshape and size, len gth of apica l and
untap ic al horns and dimens ion of la inte rca la ry plate (Table 2). Both spec iesexhib it pol ygonalintric ateridged surfacepat - tern (Figs 5. 19 ). The difference s arc (I) cell sha pe of P.
novel/a is broader with a ratio L/W 1.48 [cell length(L) and ce ll width (W)j, incl ud ing a shorterand wider apical and hy- porheca l horns and a sma ller la interc al ar y plat e;(2) shape ofP.concinna issle nde r,cellsize lon ge r witha rat io of L/W of 2.13. includin g nar rowe r apic al and antapica l horns and lager Ia inte rca lary plat e;(3) sulcalSa plat e ofP. concinna exte ndsinto theepith eca and tou ches apical I' pla teand 'tran- sitio na l' t-plate in the cingulum-sulcus junc tu re (Figs 13, 20);
(4) engen ders change in the plate relati onship of the ventral area ca us ing a distin ct depressi on ve n tra lly (Figs 13, 15, 18, 19) ; and (5) these fe atu res are diffe rent from each other in structuralrelations withi n the ventra l are a and also in the re- lati on s between the ve ntra larea and the forwa rd be ndi ng api- cal horn on P. concinna (Fig. 15) whe n com pa red to upward apic a l horn on P. novella (Fig. 6).
Placement of new species in genusProtoperidinium The morph ol og y and plat e tabulati on of P. no vellu and P.
concin na sugges t tha t these spe cies might be closel y rel at ed to members of the section Ocean ica. The situ ati o n is, ho w- ever,proble ma tic forP.no vella andP.concinna bec ausethese two species are charac terize d by threedistinct features , which
8 Phycologia, Vol. 45 (J), 2006
4"'
Figs 22-26. Line drawings of Protoperidinium concinna sp. nov.
Faust. Morphology of plates illustrated in severalviews.
Fig. 22. Ventral.
Fig. 23. Dorsal.
Fig. 24. Apical.
Fig. 25. Hyporhccal.
Fig.26. Sulcal.
are quite uncommon in the genus Protope ridinium. Whether or notto place these two species in the genus Protoperidinium is a worthwhile discussion (Figs 1-3, 16, 17). The variations of epithecal plate arrangement, deviation from the typical plate formula, have been well known in the genus Protoper- idinium. These variant forms are classified in the distinct sub- genus such as Archacperidinium based on the position and shape of the first I' apical plate and those species with only two 2a anterior intercalary plates, e.g. P. consimilis and P.
valgus (Table 1; Abe 1981). Sournia (1986) considered Ar- chaeperidinium tobe taxonomic junior synonym of Protoper- idinium. Fensome et al. (1993) prefer to retain Archaeperi- dinium at generic rank to enable practical subdivision of the large gcnus Protoperidinium.
In the freshwater genus Peridinium, the presence or absence of APC has been regarded as difference at subgeneric level rather than the generic separation (Baleeh 1988). Balech
(1995) separated species in the genus Alexandrium into two
subgenera, Alexandrium and Gessnerium by recognizing syn- onymy in APC architecture and ventral plate I' morphology of species. Furthermore, among Protoperidinium. species, ab- sence of an APC is few, and recognized only in other related dinoflagellate genera, e.g. Gotoius mutsuensis (Abe 1981) and G. abei (Matsuoka 1988) in the diplopsalid group. The situ- ation in P. novel/a and P. concinna is comparable to this prac- tice.
In conclusion,
r
believe that P. novella and P. coucinna are distinct enough from all other species of the genus Protoper- idinium and it is appropriate to classify them as a new sub- genus, I, therefore, propose to establish the subgenus Tcstcria Faust, subgen. nov. in the genus Protoperidinium.r
have se- leered P. novella sp. nov. Faust as the type species, and P.concinna sp. nov. Faust a second species.
ACKNOWLEDGEMENTS
I thank Dr Klaus Rutzler for the use of the Smithsonian's field laboratory facilities at Carrie Bow Cay, Belize, and Sabrina Varnam for collecting water samples from The Gulf Stream, National Ocean Service, NOAA, Beaufort, NC. I extend my gratitude to my colleague Dr Dan Nicolson for preparing the Latin Diagnosis, Dr Takeo Horiguchi, and the reviewers for editorial comments. I convey special thanks to Judit Quasney for editing.Mr Michael R. Carpenter for logistic support; Mrs Susann Braden and Mr Scott Whittaker, SEM laboratory and Mr James DiLoreto, Photo Services, NMNH. This investiga- tion was supported by grants from the Smithsonian Institution.
National Museum Natural History, and the Caribbean Coral Reef Ecosystem Program (CCRE), Publication No. 724.
REFERENCES
ABET.H. 1936. Report of the biological survey oj"Mutsu Ray, 30.
Notes on the protozoa fauna of Mutsu Ray. Genus Pcridiniurn1I1.
Subgenus Protoperidinium. Science Report Tohoku Imperial Uni- vcrsitv Series 11: 19-48.
ABE T.H. 1981. Studies on the family Peridinidue. An unfinished monograph of the armored dinoflagcliara. Sew Marine Biological Laboratory. Special Publication Series 6: 1--413.
ANDREIS C; ClAP] M.D. & ROOINDI G. 1982. The theca! surface of someDinophyceae:a comparative SEM approach. Botunica Marina 25: 225-236.
BALECH E. 1974. EI genero "Protopcridinium" Bergh. 1881. ('"Peri- dinium" Ehrenberg, 1831,partirn). Rcvista del Musco Argentino de Ciencia Naturales "Bernardino Rivadavia." Hidrobiologia 4: J- 79.
BALECHE. J975. Estructurasde Protoperidiniumen microscopia elec- tronica de banido. Neotropica 21: 20-25.
BALECl-JE. J 979. Tres dinoflagelados nucvos0 intcrcsantcsde aguas Brasilenas. Biologi lnstituro de Oceanografia, S. Paulo28: 55-64.
BALECH E. 1980.On thecal morphology ofdinoflagellateswith special emphasis on circular and sulcal plates.Annates CentroCiencia del Marina y Limnology7: 57-68.
BALECH E. 1988. Los Dinojlagelados del Atlnntuo Sudoccidcntal.
Publication Especiales No.1, Instituto Espanol de Occanogrufiu, Madrid, Spain. 310 pp.
BALECH E. 1995. The genus Al exandrium Halim (Dinofiagellata}, Sherkin Island Marine Station. Sherkin Island Co, Cork, Ireland.
15t pp.
BALECH E. 1999. Protoperidinium (Dinoflagellata) nuevas a interc- santes de la Bahia de Manila (Filipinas). Revista del Musro A'XCll- tino Ciencia Naurales I: 165-171.
BERGH R.S. 188!. Del' Org anismus del' Ciliofiagcllcnen.Einc phyJo- genetische Studie. Morphologisches Jahrbueh 7(2): 177-288, pIs
12-16.
DELGADO M. & FORTUNO J.-M. 1991. Atlas de fitoplankton del Mar Mediten·ano.Sciencia Marine 5: 1-133,66 pis.
DODGE J.D. 1983. Ornamentation of thecal plates in Protoperidinium (Dinophyceae) as seen by scanning electron microscopy. Journal Plankton Research5: 1\9-127.
Faust: Protoperidinium novella sp. nov. Faust and Protope ridinium concinna sp. nov. Faust 9
D,'()(iL J.D. 1985. ,111as ot' J)inof/agellales: a scanning electron mi- croscope studv. Ferrand Press, London. I 19pp.
h\ISr M.A. 1990.Morphologic details of six benthicspecies of Pro- roccnt nun(Pyrrophyta)from a mangrove island. Twin Cays, Belize, including two new spccicx. Journal oj'Phycology26: 54~-558.
F,\UST~vI.A. 2000. Dinollagellatc associations in a coral reef-man- grove ecosystem: Pelican and associated Cays. In: Natnral histor-;
of th« PelicanCays, Belize (Ed. by I.G. Macintyre& K. Ructzlcr).
Aloll Research BulletinNo. 47.>: 135-152.
F.ILST M.A.& Tlsn':R PA 2005. Harmful dinotlugcllutcs in the Gulf stream and Atlantic barrier coral reel'. Belize. In: Harmful Algae ZOO.? Proceedings of the
xu.
lnternutionol Conferenc« of Harmful Algae (Ed.by K.A. Steidinger, .I.H. Landsberg, C.c. Tomas&G.A.Vargo), pp. 326-.i2X. Florida Fish and Wil<..llik Conservation and IOeof UNESCO, S1.Petersburg. Floridu.
FF:\SO\'IE R.A .. TAYLOR F.I.R .. NORRISG .. S,\RJIA:\T W.AS .. WII'\RTOi\
D.l. & WII.L1A\lS G.L. 199'>. "A classification of living and [ossil dinofiagcllatcs. "Sheridan Press. Hanover,PJ-\, USA. 351 pp.
Gocur H. & NUZI'!. H. 1974. Scanning electron microscopy studies on the theca ofPcridinium(Dinotlagellada).ArchieI' Pro/isle Kunde
1[6: 381-410.
HA:\sl-::\G.& L.\RSE:\ J. ILJ92. Diuotiugcllatcr (Dinophyccae)idanskc Iarvunde.In: Plankton ide indrc danskcfarvandc. Nr. (Ed. by H.A.
Thomsen), vol I I. pp. 45-155.Havforskningfra Mih,jstyrelsen, Mil- jominisrcrict, Kobcnhavn, Denmark.
.Il)R(iE,\iSI'SF. ILJ 12. Bcricht ubcrdie von del'schwcdishcuhydmgra- phisch biologischen Kommission in denschwcdishenGewassen in den Jahreu 1909-10 EingesammcltcnPlanktonprobcn. Svcnskahy- drogratisk-bioloeistt«: kornmissioncns Skriftcr4: 1-20.
KOFO!1l C.A. 1909. On Peridiniurn stciniiJorgensen. with a note on the nomenclature or theskeletonthe Pcridinidac.Archicv Protistcn Kuncle 16: 25-47,pl. I.
Llc"'ls J. & DOl)(iL J.D. 1990. The use of SEM in dinoflagellate tax- onomy. In: Scanning electron microscopy in taxonomic andjune- tional morphology. (Ed, hy D. Cluugher). Svstrmatu: Associatio/l Specia!Volume41: 12~-14K.
Lnu.r« D.S. & LrrrLER lvl.l"l. 1997. An illustrated marine flora of the Pelican Cays, Belize.Hullelill or the Biologica] Sociely of Washillg- ton (): 1-149.
MACINTYRI' I.G. & Rlrrzl.U, K. 2000. Natural history"I'thePeliC1I1I
CIIYS, Beli;«, Atol! Neseureh Bulletin Nos. 466--480. Smithsonian Institution Press, Washington, DC. USA.
:rn
pp.l'vLUSI'oKA K. 199~.Cyst-theca relationship in the Diplopsulidgroup (Pcridinialcs. Diuophycoac ).Review ofPuleobotany and Palynology 56: 95-122.
NLTZLI 1-1. & D(:HH G. 19~4. Dinollagcllate cell cortex. In: Dinofia- ge!lu/es. (E<..I. by D.L. Spector). pp. 4'>-105. Academic Press, Or- lando, FL. USA.
PI:IWY E.G. 1994. Knrst-detcrrnincd facies patterns in British Hon- durus: holocene carbonate sedimentation model. AmericanAssoci- ationoj'PetroleumGeologisls5~: ~25-~55.
RUETLLEH K .. DI,\z M.A .. V()0; SOEST R.W.iVI.. SWL1':Z .. SMITHK.P, ALVARJ,Z B. & WULlT J. 2000. Biodiversity of sponge fauna in mangrove ponds, Pelican Cays, Belize. In: Natural history of the Pelican Cav», Bcli;«, (Ed. by I.G. Macintyre& K. Ruetvler), /vtoll Research Bulletin476: 231-250.
SUIITTF I~95. Die Pcridinccndel' Planktun Expedition. I. Teil. Stu- dien ubcrdie zellcn del' Pcridincen. Ergebnisse del' Plankton-Ex- pcdition dcr Humboldt-Stiftung 4: 1-170, 27 pis. A. Kiel, Lipsius and Teicher.
SOI;RNIi\ A. 19~6. "1\11".1' du Phvtoplnkton Morin." Volume I: lntro- duction, Cl'lIl1oph\'e';es, Dictvochophvccr», Dinophvcces et Raphi- do"hve';es." Editions du Centre National de la Recherche Scienti- rique. Paris, France. 219pp.
STF.JI)IN(iU( K.A. & TN,l(iL:\ K. 1996. Dinollagellatcs. In: Idenlif.l·ing murine diatoms and dinoflagellales (Ed. by C.R. Tomas), pp.'>87- 584.Academic Press, San Diego. CA, USA.
STFlDIi\(iER K.A & WILLIAMSJ. jLJ70. Dinoflagellates.Memoirs of the Hourglass Cruises. Marine Research Laboratory Contributions
148: 1-251.
TA YI.OR F.J.R. j976. Dinollagc!lares from the internatiunal Indian ocean expedition.A reporton material collected by theRV, 'Anton Brunn' /96'>-64. Bibliothrca Botunv 1.>2: 1-234,pis 1--46.
TOt<Jl'\11 S. & DOD(iEOJ. 199'>. Thecal apex structure in thePeridi- niaceac (Dinophyccac ).European Journal of Phvcolog»2~:39-45, YMvV\GI'CHI A. & HORIGliCl1iT.2005, Molecular phylogenetic study of the heterotrophic dinoflagellate genus Protoperidinium (Dino- phyccac ) inferred from small subunit rRNA gene sequences. Phy- cologicu! Research53: 30-42.
Received2 August 2004: accepted 2/ JWIe 2005 Communicating editor: T. Horiguchi
