First Reports of Padina antillarum and P. glabra ...faculty.valenciacollege.edu/tklenk/marinecd...First Reports of Padina antillarum and P. glabra (Phaeophyta-Dictyotaceae) from Florida,

First Reports of Padina antillarum and P. glabra(Phaeophyta-Dictyotaceae) from Florida, with a Key to the

Western Atlantic Species of the Genus

MICHAEL J. WYNNE1 AND OLIVIER DE CLERCK2

1Department of Biology, University of Michigan, Ann Arbor, Michigan [email protected]

2Laboratory of Botany, University of Ghent, 9000 Ghent, [email protected]

ABSTRACT.—Padina antillarum (Kutzing) Piccone and P. glabra Gaillard are newly reported for Florida.The first species, which has been widely reported from the Indo-Pacific under the name P. tetrastromaticaHauck, has seldom been reported from the Western Atlantic. The type locality of P. antillarum is Trinidadin the Lesser Antilles, and there are reports of its occurrence from Venezuela and Colombia. Observations areprovided on tetrasporangiate and oogonial plants collected from Brevard County, Florida. Padina glabra, firstdescribed from Senegal, northwest Africa, has been collected on three occasions from Anastasia Island, SaintAugustine, St. Johns County, Florida. The Florida material agrees in most respects with the original descrip-tion of P. glabra except for the larger size of the Florida thalli and the fact the thalli bear plantules in additionto sporangia. A key to the Western Atlantic species of Padina is provided.

INTRODUCTION

Wynne (1998b) presented evidence thatPadina tetrastromatica Hauck, a frequentlyreported species in the Indo-Pacific andtropical West Africa, and P. antillarum (Kut-zing) Piccone, a poorly known taxon, areconspecific and that the latter name has pri-ority. Collections of P. antillarum from P.Patrick Air Force beach, Brevard County,Florida, are herein reported and observa-tions made on the vegetative and reproduc-tive characteristics of this species of brownalgae.

Up to now there have been reports ofPadina glabra Gaillard only from the typelocality in West Africa (Gaillard, 1966) andfrom southern India (Rengasamy & Anand,1986). Thus, the identification of materialfrom the northeast coast of Florida repre-sents the first report of this species from thewestern Atlantic. A comparison with otherspecies of the genus occurring in the west-ern Atlantic with which P. antillarum and P.glabra might be confused is presented.

MATERIALS AND METHODS

The collection of Padina antillarum wasmade by Ted Klenk at P. Patrick Air ForceBase beach (28° 13.98 N, 80° 368 W), 8 miles

south of Cocoa Beach, Brevard County,Florida, on 15.vii.1998. The specimens werefound floating in the drift. The collection ofseveral plants has been deposited in theHerbarium of the University of Michigan(MICH).

Collections made on three occasionsfrom the same locality in Florida are hereinidentified as Padina glabra Gaillard.

Anastasia Island, St. Augustine, St. JohnsCounty, Florida:1) legit Mrs. G. A. Hall, 20.vii.1896, Phyco-

theca Boreali-Americana 580b (two setsin MICH).

2) legit Mary S. Snyder, ca. 1895-6 (MICH).3) legit M. A. Howe no. 1189, 5.x.1902 (in

NY); “on rocks of jetties at l. t. m. [= lowtide mark] and below.”

Specimens that had been identified by M.A. Howe as Padina antillarum (or as thatspecies with a query) were borrowed fromNY and US for re-examination.

RESULTS

Padina antillarum (Kutzing) Piccone(Figs. 1-10)

The Florida collection of P. antillarumcontained tetrasporangial and oogonial

Caribbean Journal of Science, Vol. 35, No. 3-4, 286-295, 1999Copyright 1999 College of Arts and SciencesUniversity of Puerto Rico, Mayaguez

286

thalli. The largest thallus (Fig. 1) measured10.5 cm in height and an overall breadth of11.5 cm, although the thallus was deeplysplit into segments 0.5-2.0 cm in width. The

blades showed no calcification. Cross- andsagittal sections of the blades, both in mid-region and in more basal portions, showeda 4-layered organization (Figs. 4, 5). In tet-

FIGS. 1-3. Padina antillarum. 1. Habit. 2. Surface view of inferior surface of tetrasporangial blade showing soralarrangement. 3. Surface view of superior surface of tetrasporangial blade showing scattered sporangia in theregion between hair bands.

PADINA ANTILLARUM AND P. GLABRA FROM FLORIDA 287

rasporangial and oogonial plants, repro-ductive organs occurred as pairs of concen-tric bands, each pair closely abutting acommon hair-line (Fig. 2). These soralbands were well developed on the bladesurface away from the circinnately inrolledapex of the thallus, as is characteristic of thegenus. This surface was referred to as thelower side by Trono (1969) and Womersley(1987) and the inferior surface by Gaillard(1967). Gaillard’s terminology is followedin this paper. In sporangial plants, sporan-

gia were also sometimes present on the su-perior side, but they were loosely scatteredin the mid-region between the soral bandson the opposite (inferior) side (Fig. 3). Thesporangia were not associated with hairbands, contrary to Gaillard’s (1967) obser-vation that they may be associated withscattered hairs on the superior surface. Fig-ure 10 represents a schematic view of a sec-tioned sporangial blade. In female thalli, anindusium covering the oogonia (Figs. 6, 7)is present, as depicted for this species by

FIGS. 4-10. Padina antillarum. 4. Sagittal-section in mid-region of blade. 5. Cross-section in basal region ofblade. 6-7. Sagittal-sections of female blade with oogonial sorus. 8-9. Sagittal-section of blade bearing sporangia.10. Schematic representation of soral pattern in relation to hair-lines.

M. J. WYNNE AND O. DE CLERCK288

Gaillard (1967). An indusium was not ob-served in tetrasporangial thalli (Figs. 8, 9).

Specimens in NY and US that had beenassigned by M. A. Howe to Padina antilla-rum (sometimes with a “?”) were re-examined. None was found to be P. antilla-rum, but they were re-determined as P.boergesenii, P. haitiensis, P. sanctae-crucis,and P. boryana.

Padina glabra Gaillard (Figs. 11-21)

In working over collections of Padina inNY, the first author found a collection ofhairless blades from Florida. This collectionconsisted of numerous entangled speci-mens mounted on a single herbarium sheet.Individual thalli measure 10-12 cm inheight. Most thalli are deeply divided, in-dividual segments being 2-3 cm wide (Fig.11). There is no evidence of calcification. Insectional view blades consist of 3 to 4 celllayers (Figs. 17, 21). Cells on the superiorand inferior thallus face are 48-62 mm long,24-30 mm broad, and 25-27 mm high (Fig.19). The central row(s) of cells possess thesame width and length as the other cells,but their height varies. If 3 cell layers arepresent, the middle layer is approximately50 mm high; if 4 cell layers are present, themiddle cell seems to be divided into 2 cells,each 25 mm high. Sporangia are bornemainly on the superior face of the blade;less frequently sporangia are borne on theinferior face, but they do not occur in denseconcentric sori. The sori lack indusia, andthe absence of hairs is noteworthy (Figs. 16,18). The uppermost sori contain sporangia(Fig. 20), but in a proximal direction thesori contain plantules apparently by a di-rect germination of the tetrasporangialmother cells (Figs. 12-15). These plantulesare sometimes arranged as a pair of closelyplaced concentric rows (Fig. 12). Towardthe base of the blade there is a gradual in-crease in the size of the plantules.

Our attention was drawn to Dangeard(1952) [see below] and the existence of ahairless Padina distributed as PhycothecaBoreali-Americana no. 580, specifically580b, (“Padina durvillaei Bory”). In the twosets of PB-A 580b in MICH both specimensare hairless. These specimens (Fig. 11) bear

sporangia in the sori located in the upper-most portion of the blade, and there is agradual transition to the production ofplantules in the older sori, as observed incollections made some years later by Howefrom the same locality. These sporangiaand resultant plantules often occur in a pairof closely associated concentric rows. Thelack of calcification, the deeply dividedblades, and the blade thickness of 3-4 celllayers point to the identity of PB-A 580b asP. glabra.

DISCUSSION

Padina antillarum and P. tetrastromaticawere included in a recent checklist of ben-thic marine algae of the tropical and sub-tropical Western Atlantic (Wynne, 1998a).Gaillard (1967) published a detailed ac-count of vegetative and reproductive obser-vations on P. tetrastromatica, based both onthree specimens in the type collection inLeiden and on recent specimens fromDakar, Senegal, West Africa. Gaillard (1967,fig. 1) designated one of the three speci-mens as the lectotype of P. tetrastromatica.The first account of P. antillarum was itsdescription as Zonaria antillarum by Kut-zing (1859). Piccone (1886) transferred thespecies to Padina. The fact that Kutzing didnot provide information on its provenancemay explain why Taylor (1960) did not in-clude it in his marine benthic algal flora ofthe tropical western Atlantic. Wynne(1998b) examined the type specimen inMEL (National Herbarium of Victoria, Mel-bourne, Australia) and observed that thetype locality was Trinidad in the Lesser An-tilles, as the epithet “antillarum” suggests.Wynne also presented evidence that P. an-tillarum is taxonomically identical to P. tet-rastromatica Hauck (1887), with a type local-ity of Meith, Somalia.

Padina antillarum is widely distributed inthe Indian Ocean (Silva et al., 1996, as P.tetrastromatica as well as P. antillarum) andthe tropical coast of West Africa (Lawsonand John, 1987, as P. tetrastromatica). Afterthe original description of the species, theonly reports of P. antillarum in the westernAtlantic are those by Schnetter (1976) from


Colombia and by Aponte and Ganesancited by Ganesan (1990) from Venezuela(both records as P. tetrastromatica). Accord-ing to Thivy (1945), the report of P. tetra-stromatica by Setchell and Gardner (1930)

and repeated by Gonzalez-Gonzalez et al.(1996) from Pacific Mexico is most likely P.crispata Thivy in Taylor (1945).

Padina glabra was described by Gaillard(1966) from Pointe de Fann, Dakar, Senegal,

FIGS. 11-13. Padina glabra. 11. Habit of specimen, PB-A 580b (MICH). 12-13. Plantules on surface of blade(Howe no. 1189, NY).


northwest Africa. This is the only reportedcollection of this species from the westcoast of Africa (Price et al., 1978). In a papernot mentioned by Gaillard (1966), Dange-

ard (1952) reported on various collectionsof Padina from Dakar, including some fromPointe de Fann (the type locality of P. gla-bra). He called attention to some specimens

FIGS. 14-17. Padina glabra. (These figures are based on Howe no. 1189). 14. Schematic depiction of transitionfrom sporangia-bearing to plantule-bearing soral bands. 15. Individual plantule. 16. Schematic representation ofsporangia in a sorus. 17. Sagittal section of blade.


of Padina that were four cell layers thick,had sporangial sori arranged in concentriccircles generally associated in pairs, andwere hairless. According to Dangeard, J.Feldmann confirmed his observations andsuggested that a new genus could be

erected for these specimens on the basis ofthe absence of hairs. Although Dangeardmentioned that the four-cell-layered con-struction suggested a relationship with P.tetrastromatica [= P. antillarum], he refrainedfrom assigning a specific name. There is

FIGS. 18-21. Padina glabra. (These figures are based on Howe no. 1189). 18. Surface view of blade bearing soribut lacking hair bands. 19. Surface view of blade. 20. Region of sorus bearing sporangia. 21. Sagittal section ofblade.


little doubt that Dangeard was dealing withthe species later described by Gaillard(1966) from the same location as P. glabra.

Dangeard (1952) called attention to thefact that a specimen of Padina that was dis-tributed as Phycotheca Boreali-Americanano. 580, specifically 580b, (“Padina durvillaeiBory”) was hairless. PB-A 580b was a col-lection made by Mrs. G. A. Hall, 20.vii.1896,from Anastasia Island, the identical loca-tion of the M. A. Howe collection. Anothershared feature is that these specimens bearplantules; the printed label states “Both aand b bear tetraspores; in b, a large part ofthem have germinated in situ, on thefrond.” Collins and Hervey (1917) statedthat PB-A 580b “should be referred to P.variegata”, a name which was later replacedby P. vickersiae Howe (in Hoyt, 1920).

The Florida material being assigned to P.glabra agrees with Gaillard’s (1966) accountof this species in many aspects: the absenceof hairs, the lack of calcification, the (3-)4cell-layered organization and the cell di-mensions, and the non-indusiate nature ofthe sporangial sori. The material differsfrom Gaillard’s (1966) account of P. glabrain the greater size of the blades (10-12 cmtall versus 1.0-1.5 cm for the type collec-tion), the divided nature of the blades incontrast to entire blades, and the produc-tion of plantules. There is a report of P.glabra from Mandapam, southern India byRengasamy and Anand (1986). Their obser-vations minimize the apparent differencesbetween Gaillard’s account and our obser-vations on the Florida material in that thoseauthors noted that the blades were 4.0-5.0cm tall, were divided down to the middleportion, and bore plantules. They alsostress that this is the only species of Padinaknown to lack hairs, a feature shared by theDakar, Indian, and Florida collections. An-other point of agreement between theFlorida and the Indian specimens is thepresence of sporangia on the superior sideof the thallus. Dangeard’s (1952, fig. 7) de-picted hairless “Padina sp.” (from the typelocality of P. glabra and most likely belong-ing to this species) as being 9 cm tall. Thus,the small stature reported in Gaillard’s(1966) account of P. glabra would be at thelow end of the possible range. The produc-

tion of plantules appears to be a variablefeature in some species of Padina and inother Dictyotales. Thivy (1945) referred tothese plantules as both brood buds andpropagula, developing in the place of spo-rangia or from sporangia at a very youngstage (“presumably before reduction divi-sions occurs”). She indicated their occur-rence in P. durvillei Bory and P. distromaticaHauck. These plantules have been notedalso in P. pavonica by Bitter (1899) andKuster (1899) and in P. gymnospora (Kutz.)Sonder by Hoyt (1920, as P. vickersiae).Hauck (1887), Gaillard (1967), and Lawsonand John (1977) reported that thalli of P.antillarum (= P. tetrastromatica) sometimesbear plantules arising in the same positionas the tetrasporangial sori. Feldmann (1937)observed that Mediterranean plants of Dic-tyota [Dilophus] fasciola (Roth) Lamourouxmay bear plantules.

There is some similarity of the FloridaPadina glabra to P. profunda, described byEarle (1969) from the Dry Tortugas ofFlorida, Gulf of Mexico, and also reportedfrom offshore North Carolina (Schneiderand Searles, 1973). According to Earle,blades of P. profunda are non-calcified andhave faint piliferous zones with few hairs.Plants ranged in height from 10 to 25 cm.Blade thickness ranged from 2 cell layersnear the margins to 3-4 cell layers in themid-blade portion, and 3-6 cells layers atthe base. Reproductive plants were not ob-served.

Padina gymnospora (Kutzing) Sonder hasbeen the only species of the genus reportedwith confidence from the east coast ofFlorida (Moffler and Van Breedveld, 1979,as P. vickersiae; see Allender and Kraft,1983). Padina antillarum can be distin-guished from P. gymnospora by the latterspecies’ thicker blades (6-8 layers in thebasal region) and soral bands placed proxi-mal to each successive hair zone.

Schneider and Searles (1998) reviewedthe many reports of marine algae sharing adistribution in the eastern and westerntropical or subtropical Atlantic. This firstreport of Padina glabra from the western At-lantic, thus significantly extending itsrange, is not surprising because many otheralgae have a similar distribution. The rarity


of this species is an interesting question.Whether this apparent rarity of P. glabra isreal or merely the result of its being mis-taken for other species remains unan-swered.

Key to the species of Padina known forthe Western Atlantic1

1. Hairs and hair lines absent -- P. glabra18. Hairs and hair lines present (at times

difficult to detect because they mayhave been shed or are poorly devel-oped, but their formation is re-vealed by the presence of concentriclines)---------------------------------- 2

2. Sporangial sori closely abutting thehair lines on both sides (the lowersporangial band often not well de-veloped in P. pavonica) -------------- 3

28. Sporangial sori not closely abuttingthe hairs lines------------------------ 4

3. Blades 3 cell layers (3-4 in basal re-gion); superior surface of bladeheavily calcified, inferior surfacemoderately calcified; sporangialsori with persistent indusium;gametophytes bisexual ----- P. pavonica

38. Blades 4 cell layers (3 in apical re-gion; 6 in basal region); calcificationabsent or negligible; sporangial sorinot indusiate; gametophytes uni-sexual --------------------- P. antillarum

4. Blades 4 cell layers in mid frond; 6-8cell layers near base; sori present be-tween successive hair bands (alter-nating fertile and sterile zones ab-sent) --------------------- P. gymnospora

48. Blades either 2 or 2-3 cell layersthroughout; alternating fertile andsterile zones present----------------- 5

5. Blades 2 cell layers throughout----- 658. Blades a mixture of 2 and 3 cell lay-

ers in same thalli --------- P. boergesenii

6. Indusium not persistent; sporangialsori located in proximal region offertile zone, just distal to hair lines------------------------- P. sanctae-crucis

68. Indusium persistent; sporangial soriin mid-region of hair lines ---------- 7

7. Superior surface of blade heavilycalcified and inferior surface verylightly calcified; surface cells ofblade 25-38 mm wide; sporangia upto 120 mm in diameter --- P. haitiensis2

78. Both blade surfaces moderately cal-cified except for zones of hairbands; surface cells of blade 30-50mm wide; sporangia up to 170 mm indiameter ----------------- P. perindusiata

Acknowledgments.—We thank Mr. TedKlenk of Orlando, Florida, for donating hiscollections to the senior author. We ac-knowledge William R. Buck of NY andRobert H. Sims of US for arranging the loanof specimens. We are grateful to Dr. Brunode Reviers of PC for his assistance in tryingto locate material of P. glabra.

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First Reports of Padina antillarum and P. glabra ...faculty.valenciacollege.edu/tklenk/marinecd...First Reports of Padina antillarum and P. glabra (Phaeophyta-Dictyotaceae) from Florida,