Botanica Marina Vol. 46, 2003, pp. 357–365 © 2003 by Walter de Gruyter · Berlin · New York

Introduction

The benthic marine algal flora of the Sultanate ofOman has proved to be rich in undescribed taxa, withseveral new species and some new genera recently re-ported (Nizamuddin and Campbell 1995,Wynne 1998,1999a,b, 2001, 2002a,b, 2003a,b, Wynne and de Jong2002, Schils and Coppejans 2002, Schils et al. 2003). Apicture is emerging that shows the benthic marine algalflora to be a heterogeneous mix of tropical,subtropicaland warm temperate components. There are floristicaffinities with such diverse regions as Japan, SouthAfrica, Brazil, and Atlantic Europe as well as with thetropical Indo-Pacific (Wynne 2000,2003c).

The summer monsoon is a period of 4–5 months ofconstant upwelling, associated with relatively cold(16–19 °C), nutrient-rich waters along with pro-nounced wave action. On rocky shores of Dhofar, thesouthern region of Oman, there is a tremendous pro-liferation of marine algae (Salm 1993), some formsgrowing 1.5–2 m in length in this period. This season-al upwelling is regarded as a dominant factor in con-tributing to the ecological richness of this habitat.Benthic algal biomass peaks during the monsoonseason (June–August) and is significantly reducedduring the non-monsoon season (Barratt et al. 1984).This paper reports a new species of red algal genusLeveillea (tribe Polyzonieae of the Rhodomelaceae),which appears to be endemic to the monsoon-im-pacted rocky shoreline of the Sultanate of Oman.

Members of the tribe Polyzonieae have a dorsiven-tral organization. Although earlier workers (Falken-berg 1901, Scagel 1953) placed this tribe in the sameevolutionary lineage that gave rise to the ‘Herposi-

phonia series’, that is, in subfamily Polysipho-nioideae, Hommersand (1963) assigned this tribe tothe subfamily Rhodomeloideae. His arguments werethe following: the production of spermatangia andprocarps is not associated with trichoblasts, and theproduction of tetrasporangia typically occurs in re-duced, stichidia-like indeterminate branches, thesetetrasporangia being positioned in a longitudinaldorsal row in these modified branches. According toHommersand (1963) these features are in accordwith members of the Rhodomeloideae rather thanthose of the Polysiphonioideae. Five genera are cur-rently assigned to the Polyzonieae: Cliftonaea Harv.,Dasyclonium J. Agardh, Echinosporangium Kylin,Leveillea Dec., and Polyzonia Suhr (Scagel 1962a,b,Hommersand 1963, Adams 1994). Members of thePolyzonieae are prostrate, at least in part, and attach-ment is by means of multicellular holdfasts. Determi-nate short shoots (laterals) are of variable form,laterally or dorsally placed, and are formed exoge-nously, whereas the indeterminate long shoots areformed endogenously (Kylin 1956). The genus Lev-eillea is distinguished from the other genera in thistribe by its main (long) shoots being terete or slightlyangular, uncorticated, having 7 pericentral cells persegment, and having bladelike determinate shootswith entire margins. The new species is in conformitywith these criteria except for the number of pericen-tral cells being higher.

Material and Methods

Most of the collections reported in this paper weremade in September 2000 and September 2001 inDhofar, Oman, as part of the Algal Biodiversity Pro-

Leveillea major sp. nov. (Rhodomelaceae, Rhodophyta) from theSultanate of Oman1

Michael J. Wynne

Department of Ecology and Evolutionary Biology and Herbarium, University of Michigan, Ann Arbor, MI 48109, USA,e-mail: mwynne@umich.edu

A new species of the red algal genus Leveillea, L. major M.J. Wynne, is described from the Sultanate of Oman,the northern Arabian Sea. This species is distinguished from L. jungermannioides (Hering et Martens) Har-vey, a species widely reported from the Indian and tropical western Pacific Oceans, by its larger stature, itserect habit, its bladelets, which are larger, more attenuate, and strongly overlapping, and the absence of tendrilformation by the distal portions of axes. Tetrasporangiate thalli are described in the new species, which isknown only from the Sultanate of Oman. Leveillea jungermannioides also occurs in Oman, and the twospecies co-occur in the Al Sharqia region.

Keywords: Leveillea; L. major sp. nov.; northern Arabian Sea; Rhodophyta; Sultanate of Oman.

1 Submitted 6 January 2003; accepted 10 April 2003.

ject of Oman (1999–2002) funded by the BritishGovernment’s Darwin Initiative grant for the Sur-vival of Species. A few other collections were madeby individuals and are cited as such. One collectionwas made by unnamed personnel of the Tropical Ma-rine Research Unit of the University of York, UnitedKingdom, and is cited ‘TMRU’. Specimens weregathered both in the attached state by SCUBA andalso from the drift. Material was usually processed asherbarium mounts soon after collecting. Some speci-mens were preserved in 5% Formalin/sea-water.Small portions of wet-preserved material and rehy-drated portions of pressed specimens were mountedon glass slides for observation with a Zeiss researchmicroscope. Portions of main axes were hand-sec-tioned using a single-edged razor blade. Line-draw-ings were made with a camera lucida. Images werecaptured with a Nikon D1 digital camera mountedon a photo-stand. The digital images were then as-sembled into plates using Adobe Photoshop™ ver-sion 7.0.

For purposes of comparison, camera-lucida draw-ings and measurements were made of the followingcollections of Leveillea jungermannioides in MICH:Seychelles. Anse aux Pins, Mahe Island: 26.x.1964,leg. M.Wynne 3S-10, epiphytic on Dictyota. – Sri Lan-ka. Galbolka Point, Colombo: 19.xi.1964, leg. M.Wynne 5A-4A, epiphytic. Also, an Omani collectionof L. jungermannioides [cited in the Results and Ob-servations] was used for Figure 4. In addition, typematerial of two other species, Leveillea comosa andL. pectinata, were received on loan from PC.

Co-ordinates were obtained in the field by usingseveral GPS devices, primarily a model made byGarmin Etrex Summit™. Herbarium abbreviationsare according to Holmgren et al. (1990).

Leveillea major M.J. Wynne sp. nov.(Figs 1–3, 5–8, 14–17)

Diagnosis: Aliis speciebus in genere multum robus-tiore stature thalli (axes indeterminati principalesusque ad 11 cm alt. et rami laterales indeterminatiusque ad 7 cm alt.) distincta; forma elongatiore et am-plitudine major foliolis determinatibus distincta, quaesunt 3.0–4.0 mm in long. et usque ad 2.0 mm lat.,atque apicibus acutis eorum potius quam foliolis ro-tundatis ovatis in L. jungermannioides; foliola alterna-tim et distichiter multum imbricata producta; axeshaptera multicellulares non regulariter producti; sic,habitus est plerumque erectus potius quam prostratus;novem cellulae pericentrales per segmentam axis inde-terminati typice productae.

Distinguished from other species in the genus bythe much more robust stature of the thalli (main axesto 11 cm in height and lateral indeterminate branchesto 7 cm in length); also distinguished by the moreelongate shape and the larger size of the determinatebladelets, which are 3.0–4.0 mm long and to 2.0 mm

358 M.J. Wynne

wide, and by their pointed tips contrasting with therounded oval-shaped bladelets of L. jungerman-nioides; the alternately and distichously producedbladelets greatly overlapping one another; axes notregularly producing multicellular holdfasts; thus, thehabit is generally erect rather than prostrate; usually9 pericentral cells produced per segment of an inde-terminate axis.

Holotype: Sultanate of Oman. First cove east of Sadh(17.05755 ° N, 55.08544 ° E), Dhofar: 19.ix.2000, leg.T.Collins 19092000-13-23, attached and in drift; speci-men deposited in MICH (Fig. 1).

Isotypes: Deposited in BM and ON.

Etymology: The specific epithet, L. major, greater, al-ludes to the greater stature of this species.

Additional collections (all from Oman)

1) Hatom Cove (16.96091 ° N, 54.82795 ° E), east ofMirbat, Dhofar: 16.ix.2000, leg. G. Richards and G.Minton 16092000-09-12 (BM, MICH, ON); 17.ix.2000, leg. G. Richards and G. Minton 17092000-10-27 (BM, MICH, ON); 25.ix.2001, leg. G. Richardsand H. Ford 25092001-19-22 (MICH).

2) cove adjacent to Abalone Research Laboratory(16.96888 ° N, 54.70151 ° E), Mirbat, Dhofar: 22.ix.2000, leg. G. Richards and G. Minton 22092000-15-30 (MICH).

3) Kah’f al Marnaif, Al Mughsayl (16.88333 ° N,53.78333 ° E), Dhofar: 19.ix.1983, leg. TMRU(MICH).

4) Raysut (16.93417 ° N, 53.92250 ° E), Dhofar: 20.ix.1983, leg. L. Barratt (MICH).

5) Sadh (17.04366 ° N, 55.08050 ° E), Dhofar: 20.ix.1994, leg. J. Stirn, 1 m. depth, tetrasporangiate(MICH).

6) Al Ashkarah (21.83333 ° N, 59.57500 ° E), south ofSuwayh, Al Sharqia: 6.x.1992, leg. B. Jupp (MICH).

Results and Observations

Vegetative organization

Thalli are dark red in color, with erect main axes upto 11 cm in height and indeterminate branches up to7 cm in length (Fig. 1). Even though the organizationis dorsiventral, the habit is predominantly erect. In-determinate axes bear distichously arranged deter-minate bladelets that arise alternately and disti-chously, strongly overlapping one another (Figs 2–3,5–6). Cross-sections reveal the main axes to be tereteto somewhat angular in profile, the angularity beingthe result of the frequent emergence of the laterals.These indeterminate axes are 380–440 µm in diame-ter and are uncorticated. The pericentral cells arenon-uniform in size and appear to number 9 per seg-ment, although counts of 7 to 10 have been made.

Leveillea major sp. nov. from the Sultanate of Oman 359

Figs 1–4. Leveillea.Figs 1–3. L. major. Fig. 1. Holotype (in MICH). Fig. 2. Erect axis with closely overlapping bladelets. Fig. 3. Several erectaxes. Fig. 4. L. jungermannioides. Creeping axis with rounded bladelets. [Figs 2 and 3: leg. Stirn at Sadh; Fig. 4: leg. Wynne10897-A.].

The determinate bladelets (Figs 14–16) arise exoge-nously at the apex and become 3.0–4.0 mm long, to2.0 mm wide. They are more elongate and pointedthan those of L. jungermannioides, which are round-

360 M.J. Wynne

ed to oval (Figs 4, 9–13). Bladelets arise singly fromevery other segment of an indeterminate axis, alter-nating to the right and then to the left. The bladeletsin section are 80–95 (–126) µm in thickness. Tri-

Figs 5–13. Leveillea.Figs. 5–8. L. major. Figs 5 and 6. Portions of erect axes. Figs 7 and 8. Tetrasporangial stichidia. Figs 9–13. L. jungerman-nioides. Creeping axes. [Figs 5 and 6: leg. Richards -2000-10-27; Figs 7 and 8: leg. Stirn at Sadh; Figs 9 and 10: leg. Wynne 3S-10; Figs 11–13: leg. Wynne 5A-4A.].

Leveillea major sp. nov. from the Sultanate of Oman 361

Figs 14–17. Leveillea major.Fig.14.Distal region of an erect axis.Figs 15 and 16.Vegetative erect axes.Fig.17.Erect axis bearing tetrasporangial stichidia.[Fig.14: leg.Jupp at Al Ashkarah;Fig.15: leg.Richards –2000-15-30;Fig.16: leg.Collins -2000-13-23;Fig.17: leg.Stirn at Sadh.].

choblasts, which may be produced at the tips of thebladelets in L. jungermannioides (Scagel 1953, Huis-man 2000), were not observed in the Omani speci-mens. Thalli are attached by short lengths (less than1 cm) of axes bearing much reduced determinatebladelets and frequent multicellular holdfasts com-prised of consolidated masses of rhizoids. Erect inde-terminate axes do not produce multicellular hold-fasts.

Reproductive structures

The only reproductive stage that was observed in thenew species was tetrasporangiate. In fertile thalli,tetrasporangia are produced in short, modified inde-terminate branches, or stichidia, that arise endoge-nously in the anterior axil of every consecutive deter-minate bladelet (Fig. 17). The stichidia are stronglyarching outward and downward, resulting in a round-ed aspect (Figs 7 and 8). The most mature sporangiain a single stichidium appear most distal because ofthe overarching aspect of the fertile branch. Usuallyonly a single stichidium arises in an axil, but occa-sionally the fertile axis is branched, with two or threestichidia arising in a tight cluster, giving a bi-lobed ortri-lobed aspect. The stichidia are somewhat protect-ed by adjacent bladelets. Each stichidium consists ofabout 11–13 segments. A single tetrasporangium isproduced by a pericentral cell from each tier of thestichidium, always on the dorsal side, forming a longi-tudinal series. Typically, about 4–6 tetrasporangia invarious stages of acropetal maturation can be seenper stichidium, but closer scrutiny reveals additionalvery small sporangial primordia. So, each stichidiumappears to have the potential to produce up to 8 spo-rangia. Mature tetrasporangia are tetrahedrally di-vided and at maturity measure 184 to 235 µm in di-ameter. Two cover cells are initially produced, butthese undergo several divisions to produce smallercover cell derivatives. The original large cover cellsand the smaller cover cell derivatives all lie in thesame plane arching over the expanded mature spo-rangium.

Status of Leveillea jungermannioides in the Omani flora

Barratt et al. (1984) reported the presence of Leveil-lea jungermannioides from Dhofar, Oman. All ofthose collections that I have been able to examinebelong to L. major. Yet at least two Omani collec-tions (both in MICH) of L. jungermannioides havebeen identified in the present study: Sur (22.5666 ° N,59.52888 ° E), Al Sharqia: 27.v.1994, leg. B. Jupp, epi-phytic on Acanthophora muscoides (Linn.) Bory.Shinzi (20.55916 ° N, 58.93138 ° E), Masirah Island, AlSharqia: 6.ii.1997, leg. M. Wynne 10897-A, epiphyticon Halimeda.

Thus, in respect to the currently known ranges of

362 M.J. Wynne

these two species in Oman, L. major is known fromtwo regions of Dhofar (southern Oman) and AlSharqia (southeastern Oman), whereas L. junger-mannioides is known only from Al Sharqia. Dhofar isa region much more impacted by the seasonal mon-soon than is Al Sharqia.

Status of Leveillea comosa and L. pectinata

Decaisne assigned several other species to his genusLeveillea, namely, L. comosa and L. gracilis (De-caisne 1839) and L. ciliata and L. pectinata (Decaisne1842, 1846). Harvey (1860) stated that he was “in-clined to unite the L. gracilis, L. pectinata, and L. co-mosa of Dcne. to the L. Schimperi, of which they maybe varying forms, or imperfectly developed fronds”.Silva et al. (1996) treated L. gracilis as conspecificwith L. jungermannioides, but they continued to rec-ognize L. comosa and L. pectinata. Authentic speci-mens of L. comosa and L. pectinata were received onloan from the Museum National d’Histoire Na-turelle, Paris. These types are deposited in the “Herb.Général” of the algal holdings of PC. Both types arerepresented by extremely fragmentary samples. Thatof L. comosa is a fragment on a mica mount, with thelabel: “In Sargasso ilicifolio parasitico ad litus occid.,Novae-Hollandiae. Leschenault.” This specimen isclearly much smaller than the Omani material delin-eated here as L. major. The label of the type of L.pectinata is “Scripsit Cl; Decaisne – in Corall. ancipitiad oras occid. Nov. Holl.”. This label also bears theannotations: “Leveillea pectinata non Leveillea,sed Herposiphonia species ! Suhr.” and “TYPE ofLeveillea pectinata Decaisne HBS Womersly6.12.1952”. I conferred with Prof. Womersley aboutthis specimen, and he confirmed that “it is almost cer-tainly a Herposiphonia, though the indeterminatelaterals on every fourth segment are often absent”.He regarded it as “related to H. calothrix but I thinkdistinct”. Thus, it can also be eliminated from furtherdiscussion in regard to the Omani Leveillea.

Discussion

When the genus Leveillea was established by De-caisne (1839), he introduced the name L. schimperi,which was an illegitimate substitution for Amansiajungermannioides Hering et Martens (Martens andHering 1836). Later, Harvey (1855) effected the bi-nomial L. jungermannioides, which is now acceptedas the correct name for the lectotype of the genus.

In addition to the three names discussed above(Leveillea comosa, L. gracilis and L. pectinata),DeToni (1903) listed some other species in the genusas “Species Inquirendae”. Leveillea ciliata Decaisne(1842) was described as growing epiphytically onDesmarestia from southern Chile. This species, how-ever, was not included by Ramírez and Santelices

Leveillea major sp. nov. from the Sultanate of Oman 363

(1991) in their catalogue of algae from the temperatePacific coast of South America, and it would seemunlikely that this tropical-warm temperate genuswould occur in southern Chile. Leveillea bidentatawas described from “Tshifu” in the China Sea byMartens (1868) growing on fronds of Sargassumthunbergii (Roth) Kuntze [= Myagropsis swartzii]. Itdoes not appear to have been re-collected since theoriginal report. Its bidentate bladelets are unlikethose of other species in the genus. The bladelets asdepicted by Martens (1868, pl. VII, fig. 5) appear tobe very regular in their formation and not the resultof grazing by herbivores or erosion. Its prostratehabit also separates it from L. major.

The type locality of Leveillea jungermannioides isTor, Sinai Peninsula, Egypt (Martens and Hering1836, as Amansia jungermannioides). This species hasa very wide distribution throughout the tropicalIndo-Pacific. A lengthy list of records was providedby Scagel (1953). More recent records include thosefrom Vietnam (Pham-Hoàng Hô 1969), China(Zhang and Xia 1979, Tseng 1983), Mozambique(Seagrief 1980), both eastern (Cribb 1983, Price andScott 1992) and western Australia (Huisman andWalker 1990, Huisman 2000), Indonesia (Verheij andPrud’homme van Reine 1993), the Seychelles(Wynne 1995), the Arabian Gulf (De Clerck andCoppejans 1996), the Philippines (Calumpong andMeñez 1996, Kraft et al. 1999), Japan (Yoshida 1998),Iran (Sohrabipour and Rabii 1999), and the Hawai-ian Islands (Abbott 1999).

Several morphological characters can be cited tojustify the delineation of this Omani material as anew species, distinct from L. jungermannioides. Thehabit of Leveillea jungermannioides is mostly pros-trate, epiphytic on other algae by means of multicel-lular holdfasts produced regularly from the lowersurface of every 5th (–6th) segment of indeterminateaxes. Their formation and potential attachment tohost substrata result in a prostrate habit (Falkenberg1901, Børgesen 1937, Scagel 1953, Price and Scott1992). Such a regular formation of holdfasts is absentin L. major. The bladelets of L. jungermannioides aremore rounded (De Clerck and Coppejans 1996,Huisman 2000) than the elongate and pointedbladelets of L. major. The bladelets of L. major re-main strongly overlapping, whereas those of L.jungermannioides, while being imbricate in the grow-ing regions (Price and Scott 1992, Huisman 2000), be-come more spaced in older portions (Kützing 1865,Scagel 1953, Cribb 1983, Teo and Wee 1983, De-sikachary et al. 1998). The distal ends of the branchesin L. jungermannioides often are reflexed or curledback on themselves, frequently forming tendrils

(Okamura 1912). Such tendril-formation does notoccur in L. major. The production of tetrasporangiain short indeterminate branches arising in the axilfrom the anterior edge of successive determinatebladelets is very similar to that which has been de-picted for L. jungermannioides (Decaisne 1841, Oka-mura 1929, Scagel and Chihara 1968). There appearto be usually 9 pericentral cells per segment pro-duced in L. major, which is in contrast to the numberof 7 reported for L. jungermannioides (Falkenberg1901, Scagel 1953, Kylin 1956). Possibly, this increasednumber is correlated with the greater stature and sizeof L. major. In light of the fact that all the other mor-phological criteria (the dorsiventral organziation, theterete non-corticated main axes, and the leaflike lat-eral bladelets with entire margins) conform to Lev-eillea, it is reasonable to conclude that this Omanimaterial is a new species in the genus. Also, collec-tions from southern Oman had been called ‘L.jungermannioides’ up to now.

Acknowledgement

Some of the material used in this study was obtainedas part of the Algal Biodiversity Project of Oman(1999–2002), funded by a British Government’s Dar-win Initiative grant for the Survival of Species. Theproject was managed by HTS Development Ltd.,U.K., working with the Natural History Museum ofMuscat, Oman, and supported by the Natural HistoryMuseum of London and the Herbarium of the Uni-versity of Michigan. The Darwin Initiative is part ofthe British Government’s Department of Environ-ment Transport and the Regions (DETR). I amgrateful to the following individuals for their contri-butions to this research: Dr Lynne Barratt and MrGlenn Richards, both of whom were personnel ofHTS Development Ltd, and to Dr Henry Ford, MrsGianna Minton, and Mr Tim Collins for their partici-pation on the collecting trips. I also thank Dr JozeStirn at the National Institute of Biology, Universityof Ljubljana, Slovenia, and Dr Barry P. Jupp of Mus-cat for sharing their collections. I thank FrançoiseArdré of the Laboratoire de Cryptogamie, Paris, forarranging a loan from PC. I also thank Dr MakotoYoshizaki and Dr Mitsuo Chihara for sharing theirinsights on variation in Japanese L. jungerman-nioides. Prof H.B.S. Womersley shared with me hisobservations on the type of L. pectinata. Finally, I amindebted to Mr David Bay, Dept. of EEB, Universityof Michigan, for his assistance in assembling the art-work.

364 M.J. Wynne

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