N. Jb. Geol. Palãont. Abh . 256/2 , 183- 193Stutlgart, published online March 2010

Article

Morphological variation and distribution of the Tournaisian(Early Mississippian) miospore Waltzispora lanzonii DAEMON 1974

Geoffrey Playford, Brisbane and José Henrique G. Melo, Rio de Janeiro

With 5 figure s

PLAYFORO. G. & MELO, 1. H. G. (20 IO): Morphological variatioo and distribution of the Touroaisiao(Early Mississippian) miospore Waltzispora lanzonii OAEMON 1974. - N. Jb. Geol. Palãont., Abh.,256: 183- 193; Stuttgart .

Ab st ract: Mississippian strata of lhe Amazonas Basin , northem Brazil , conta in abunda m and diverseasse mblages of miospores, many of considerable stratigraphic importance. These inc lude ao acavatetrilete species - Waltzispora lanzonii - which is lhe subject of the present account. Bascd 00nurnerous speci mens from the Toumaisian (Lower Mississippian) Orix iminá Formation, lhe morpho­logy of lhe more typical and also lhe relatively rare, aberrant representatives of W lanzonii isillustrated aod described io detail. Current aod previously published data iodicate that the speciesis restricted palaeageagraph ically to north-northwest Gondwana; i.e., incorporating parts af present­day oorthero Brazil, Peru, aod lhe Sahara regioo of Algeria and Libya. Chronostratigraphically,W lanzcnií is characteristic of and perhaps entirely confined to Tournaisian strata, although thepossibili ty exists that it may extend slightly higher, into the Visean . However, rare acc urrenccs of thespecies in upper Visean strata are considered to be a probable consequence of rewarking.

Key words: Miospores, Amazonas Basin, biostratigraphy, palaeogeography, Mississippian.

1. Introduction

Ouring ongoing investigalions directed at the detailedpalynological zooation of Mississippian (Lower Car­boniferous) strata of northem Brazilian sedimentarybasins, numerous specimens of a highly distinctiveform of acavate trile te miospores have been en­countered. This form - originally described under thebinomen Waltzispora lanzon ii by OAEMON (1974) ­has been recognized as being of part icular strati­graphic significance within the Mississippian succes­sion of the Amazonas Basin (OAEMON 1974; MELO& LOBOZIAK 2003; see Fig. 1 for locat ion). Theavailability of relatively large numbe rs of advan­tageously preserved spores - from the ToumaisianOriximiná Formation of the Amazonas Basin and

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c1early attributable to this species - discloses anapprec iably greater morpholog ical variation thaohas hitherto been reported. Thus, the objective ofthe present account is to describe and iIIustrate suchvariation, thereby enhancing its characterization andbiostratigraphic utilization.

2. Material and methods

The core samples forming the material basis of thisstudy are fraro petroleum exploratory wells drilled byPetróleo Brasileiro S.A. (Petrobras) in the AmazonasBasin; specifically (Fig. 2) in its constituent NorthPlatform (l -CM- \ -PA and adjacent l -CM-2-PA) andCentral Basin ( I-RX-3-AM) . The stratigra phy and

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palynostratigraphic subdivision of the Devonian andCarboniferous succession of the Amazonas Basin, aspenetrated by numerous Petrobras wells, have beendetailed by MELO & LOBOZIAK (2003, and literaturecited therein).

The present samples, consisting largely of car­bonaceous siltstones, are from the sandy/peliticOriximiná Formation and are variously representativeof the three, stratigraphically successive, Tournaisian(Early Mississippian) miospore biozones definedby MELO & LOBOZIAK (2003: 190-192). In ascendingstratigraphic order, these zones are: Radiizonatesarcuatus- Waltzispora lanzonii [AL] Interval Zone;Spelaeotriletes balteatus -Neoraistrickia loganii [BL]Interval Zone; and Spelaeotriletes pretiosus-Colati­sporites decoros [PD] Interval Zone. Sampling detailsare as follows:

• l-CM-I-PA, core 2 (3/5) at 242-243 m, core 3 (3/4) at247-248 m [BL Zone]; core 4 (2/4) at 249.7-254.0 m, core 5(1/4) at 254.1-258.4 m, core 6 (1/4) at 258.4-263.1 m and258.9 m, core 7 (1/3) at 263.1-267.3 m, core 9 (2/3 and 313)at 270.4-275.6 m [AL Zone] . See MELO & LOBOZIAK(2003: 190-191; figo 13B).

• l-CM-2-PA, core 12 (1/5) at 264.95 m, and (2/5) at266.10 m, core 13 (7/7) at 274.95 m [PD Zone]; core 14(2/6) at 275.80 m [BL Zone]. See MELO & LOBOZIAK(2003: 191; figs. 8, 13B).

• l-RX-3-AM, core 26 (1/2) at 2212.70 m, core 27 (1/2) at2219.50 m and 2220.80 m, core 28 (212) at 2223.42 m, and(1/2) at 2228.80 m [alI BL Zone]; present study, cf. MELO& LOBOZIAK (2003: 191; figo 13B).

Conventional physico-chemical procedures were utilized inthe Petrobras/Cenpes laboratories for the extraction andconcentration of palynomorphs from the subject samples;

The Tournaisian (Early Mississippian) miospore Waltzispora lanzonii DAEMON 1974 185

Fig. 2. Location map, Amazonas Basin, northem Brazil (adapted from MELO & LOBOZIAK 2003: figo I), showing positionsof Petrobras wells that yielded the miospores described herein. Geographic coordinates are as follows: I-CM-I-PA,Lat. 1° 19' 1O.037"S, Long. 55°55' 14.863"W; l-CM-2-PA, Lat. 1° 19' 15.036"S, Long . 55°56 '7.865"W; and l-RX-3-AM,Lat. 4°22'45.825"S, Long. 58°15'29.527"W.

see, e.g ., PLAYFORD & DINO (2000: 10, 12). Palynomorphyields were generally high, with quality of preservationvarying from good to relatively poor. Strew slides wereexamined under transmitted light by means of a ZeissAxioplan binocular microscope equipped with Zeiss Axio­Cam MRc/Axiovision 4.6.3 for digital photomicrographyat Petrobras/Cenpes; and, at The University of Queensland,School ofEarth Sciences, using an OIympus BH2 binocularmicroscope with attached Leica DFC320 digital cameraand complementary image-acquisition computer software(Leica IM50) .

3. Systematic palaeontology

All figured specimens (Figs. 3-4) are housed at theBiostratigraphy and Paleoecology Management ofPetrobras Research Center (Petrobras/Cenpes/Pdexp/Bpa), Ilha do Fundão, Rio de Janeiro, RJ, Brazil.Curatorial details are provided in the figure explana­tions.

Anteturma Proximegerminantes R. POTONIÉ 1970Turma Triletes REINSCH 1881 emendo DETIMANN

1963Suprasubturma Acavatitriletes DETIMANN 1963

Subturma Azonotriletes LUBER 1935 emendoDETIMANN 1963

Infraturma Apiculati BENNIE & KIOSTON 1886emendo R. POTONIÉ 1956

Subinfraturma Granulati DYBOVÁ & JACHOWICZ1957

Genus Waltzispora STAPLIN 1960

Type species : Waltzispora lobophora (WALTZ in LUBER& WALTZ 1938) STAPLIN 1960; by original designation.

Discussion : The genus is accepted as originally defined(STAPLIN 1960: 18) to embrace radiosymmetric, acavate ,trilete spores featuring a subtriangular, more-or-Iess tri­lobate amb modified at each apical extremity to producea variably developed, "reflexed or expanded, mushroom,saddle, or T-shaped outline" . STAPLIN (1960) characterizedthe exine as being either finely sculptured or laevigate.Because the type species Waltzispora lobophora was de­scribed by WALTZ (in LUBER & WALTZ 1938: 12; pI. 1, figo5; pI. A, fig o8) as having a "fine grained" exine (see alsoPLAYFORD 1962: 581-582; pI. 79, fig o7), the genus is appro­priately assigned, as above, to the subinfraturma Granulati(cf. PLAYFORD 1962; SULLIVAN 1964: 361). Most describ edspecies of Waltzispora are variousl y confined stratigraph­ically within the Touma isian through Serpukhovian or lowerBashkirian intervaI.

Waltzispora lanzonii DAEMON 1974Figs. 3A-T, 4A-K

1969 Waltzispora sp. 2890. - LANZONI & MAGLOIRE, pI.4, figo 13. [No description]

186 G. Playford and J . H. G. Melo

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Fig.3 (Legend see p. 187)

The Tournaisian (Early Mississippian) miospore Waltzispora lanzonii DAEMON 1974 187

1969 Waltzispora sp. 2890 forme "b", - LANZONI &MAGLOlRE, pI. 4, figo 14. [No description]

1974 Waltzispora lanzonii DAEMON, p. 561; pI. 7, figs.2-3.

Original diagnosis: "Trilete spores with triangularamb; sides of triangle are straight to slightly concave, andapices are mushroom-shaped; edges of mushroom structuremay be rounded (fig. 2) or extended as appendages (fig. 3);ornament on distal surface consisting of small spinae orsticks [bacula] that are sparsely distributed, mainly on theapices. Trilete mark 1/3 to 1/2 of spore radius.

"Average dimensions: Total diameter ca. 55 to 65 um;exine [thickness] ca. 1 um or less; ornament [height ofsculptural elements] 1 um or less; total width of apicalregion ca. 29 um; length of apical appendage (fig. 3) ca.II um," [Translated from DAEMON (1974: 561), includingreference to pI. 7, figs. 2-3].

Amplification of diagnosis : Spores radial, trilete.Amb typically subtriangular with straight to slightly con­cave sides; each amb apex variably expanded tangentiallyand, to a lesser extent, radially to form a strongly to slightlydeveloped, mushroom- or shoulder-like structure havingrounded or acutely to obtusely pointed lateral margins; inlatter case, each edge commonly extending as spine-like orconical projection, up to ca. 8 um long (usually less), havingpointed or blunted apex. Laesurae distinct to perceptible,simple, straight, length usually ca. one-quarter to one-halfof spore radius. Exine 1-2 um thick, proximally laevigate;distai surface commonly bearing scattered, discrete, minutebacula and conilspinae especially on and about apicalregions.

Dimensions (75 specimens): Equatorial diameter 35 (53)75 um, Width (tangential) of apical structure 18-33 um,length (radial) 11-15 um,

Holotype (DAEMON 1974: pI. 7, figo 2): Slide 6780A/DESUL, No. 5229-1M.N., England Finder coordinates R42;

deposited in Departamento de Geologia e Paleontologiado Museu Nacional, in Rio de Janeiro, RJ, BraziI. Our re­examination of the holotype (in October 2007) indicatedits deterioration through fungai invasion and hence un­suitability for re-photography. However, DAEMON'S originalphotomicrograph, cited above, is of good quality and iscertainly well-representative of the species, as is his pI. 7,figo 3. Several of the specimens figured herein (e.g., Fig.3Q, 3H) conform closely to the two figured by DAEMON(1974: pI. 7, figs. 2-3, respectively).

Type locality and stratum : Amazonas Basin, NorthPlatform, well 2-NA-I -PA, core 21, 1285.1-1286.8 m;Oriximiná Formation (BL Interval Zone), Lower Missis­sippian (Tournaisian): DAEMON (1974: 561, 565), withadditions and revisions from MELO & LOBOZIAK (2003:159,191 , figs. 1, 13B). Note that DAEMON (1974: 565) sta­ted in his PI. 7, figo 2 explanation that the holotype is fromwell "a-NA-I-PA" (an evident misprint for well 2-NA-l­PA); anel, moreover, that its host lithostratigraphic unit wasthe Faro Formation. However, according to MELO & LOBO­ZIAK (2003: 159, figo 13), core 21 ofwe112-NA-I-PA is nowconsidered representative of the Oriximiná Formation(specifical1y its lower upper part).

Remarks : It is clear that DAEMON (1974), as the originalauthor, enunciated ali salient features of the species. There­fore, the above diagnostic amplification - based on the largemajority of specimens observed in the present study andillustrated in Fig. 3 - is not to be construed as an emend­ation, although it does extend the size range considerably.A minority of the .specimens studied show slight to quiteappreciable morphological divergence from those dia­gnosed above. While still being regarded as attributable toWaltzispora lanzonii DAEMON 1974, these are considered torepresent aberrancies - deviations from the morphologicalnorm - of the one species. Atypical features, as shown byspecimens il1ustrated in Fig. .4, include: irregularity orasymmetry in amb configuration , some specimens evenbeing quadrangular or nearly so (Fig. 4 I, J), others of

Fig. 3. Photomicrographs of Waltzispora lanzonii DAEMON 1974 from Tournaisian strata of the Oriximiná Formation,Amazonas Basin. Scale bars represent 20 um, except Q (= 50 um). A, slide 9305314, EF (England Finder coordinates)E43/l ; wel1 l-CM-I-PA, core 4 (2/4) , 249.7-254.0 m. B, slide 9305306, EF P42/3; wel1 l-CM-I-PA, core 2 (3/5) ,242-243 m. C, slide 9305321, EF W39/2 ; wel1 l-CM-I-PA, core 6 (1/4), 258.4-263.1 m. D, slide 9305349, EF P31/l;well I-CM-I -PA, core 9 (3/3), 270.4-275.6 m. E, slide 9305311, EF H38/3; well l-CM-I-PA, core 3 (3/4) ,247-248 m.F, slide 9305311, EF 038; well l-CM-I-PA, core 3 (3/4),247-248 m. G, slide 9305311, EF M36/2; well l-CM-I-PA,core 3 (3/4),247-248 m. H, slide 13.595-2, EF E56; wel1 l-CM-2-PA, core 14 (2/6), 275.80 m. I, slide 9305311, EF N34/2;wel1 l-CM-I-PA, core 3 (3/4) , 247-248 m. J, slide 96.0569, EF W40/4; well l-RX-3-AM, core 28 (2/2) , 2223.42 m.K, slide 96.0569, EF M34/3; wel1 l-RX-3-AM, core 28 (2/2),2223.42 m. L, slide 96.0569, EF H34/4 ; well l-RX-3-AM,core 28 (2/2) , 2223.42 m. M, slide 9305317, EF Z40/2; welll-CM-I-PA, core 5 (1/4),254.1-258.4 m. N, slide 96.0566A,EF W40/4; well l-RX-3-AM, core 27 (1/2), 2219.50 m. O, slide 9305317, EF U33/3; well l-CM-I-PA, core 5 (1/4),254.1-258.4 m. P, slide 9305340, EF 1<31; well l-CM-I-PA, core 7 (1/3) , 263.1-267.3 m. Q, slide 9305348, EF M31;well l-CM-I-PA, core 9 (2/3) , 270.4-275.6 m. R, slide 9305348, EF G52/4; wel1 l-CM-I-PA, core 9 (2/3), 270.4-275.6 m.S, slide 9305321, EF B59; well l-CM-I-PA, core 6 (1/4), 258.4-263.1 m. T, slide 9305348, EF U48/4; well l-CM-I-PA,core 9 (2/3), 270.4-275.6 m.

188 G. Playford and J. H. G. Melo

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Fig. 4. Photomi crograph s of Waltzispora lanzonii DAEMo 1974 from Tourn aisian strata of the Oriximiná Forma tion,Amazo nas Basin . Sca le bars represent 20 11m. A, slide 9305349, EF (England Finder coordinates) 043/ 1; well I-CM -I -PA,core 9 (3/3) , 270.4 -275.6 m. B, slide 93053 17, EF G33/1; well I-CM-I -PA, core 5 ( 1/4), 254. 1-258.4 m. C, slide93053 11, EF V48; well I-CM-I -PA, core 3 (3/4) , 247-248 m. D, slide 930532 1, EF T42/1; wcl l I-CM -I -PA, co re 6(1/4 ), 258.4-263.1 m. E, siide 9305306, EF 046; well I-CM-I-PA, co re 2 (315), 242-243 m. F, slide 96.0569, EF N52/2;well I-RX-3-AM, core 28 (2/2), 2223 .42 m. G, slide 93053 17, EF F39/4; well I-CM-I -PA, core 5 (1/4 ), 254.1-258.4 m.H, slide 96.0565, EF X36; well I-R X-3-AM, co re 26 ( 1/2), 22 12.70 m. I, slide 9305348, EF V42/1 ; wcl l I-CM-I-PA,core 9 (2/3) ,270.4-275.6 m.: exine pyr ite-p ittcd . J, slide 930532 1, EF K32/3; wcll I-CM - I- PA, co re 6 (1/4),258.4-263. 1 m.K, siide 960568, EF H48; wcl l I-R X-3-AM, core 28 ( 1/2) ,2228.80 m.

"hump-backed" appearance (Fig . 4C, D); almost impor­ceptible shoulder devclopm ent at amb apices (Fig. 4 F, O,K); and variation in such devclopm ent on a given speci men(Fig. 4 B, C, D, H). A uni fying feature of most of thesespeci mens is their finely, albeit sparscly, apiculate distaiexinc, with the sculptural e lements proj ect ing most dist inct­Iy from the equatoria l radia l regio ns. Howcvcr, ill-pr escr vedspecimens with corroded exines tend to be devoid ofscul ptural projec tions.

Comparison : Waltzispora lanzonii is normally a veryeasil y recogni zable specie s, manifes ted by its characteristic

equatorial contour and by the form and distribution of itsdistai sculptura l elements.

Waltzispora lobophora, originally described from theRussian Mississipp ian (WA LTZ in LUIlER & WALTZ 1938,1941 ), and subsequcntly from the Visean of Spitsbe rgen(PLAYFORD 1962: 58 1-582, pl. 79. figo7), is of fairly com­parable size to Iv. lanzonii, but has a difTerent amb con­fig uration, longer lacsurac, and con spicuously and com­prehensivcly gra nulate exine .

STAPLI 's ( 1960: 18; pl. 4, figs . 2-3) species Iv. alberten­sis, from the Upper Mississippian of Alberta, Canada, andelsewhe re in the northe rn hemisphere (c.g., PLAYFORD

The Tournaisian (Early Mississippian) miospore Waltzispora lanzonií DAEMON 1974 189

1962; FILlPIAK 2004), is readily distinguishable from otherrepresentatives of the genus by a combination of its distinc­tive equatorial contour, small size, and laevigate exine.

The Spitsbergen Visean specie s Waltzispora sagittataPLAYFORD 1962 (pp . 582-583; pl. 79, fig o 12; text-fig. 5c)differs from W lanzonii in its mostly smaller equatorial dia­mete r (averaging ca . 29 um), markedl y concave amb inter­radii , longer laesurae, and granulate exine, and in showinga lesser tendency for amb-api cal expansions to developtangential prolongations at the corn ers. Moreover, it featuressomewhat pointed, obtuse radial extremities.

Incontrast to W lanzon ii, SULLlVAN 'S (1964 : 362; pl. 57,figs . 25-30) species W planiangulata, from the upperVisean of England, has laesurae exceeding three-quarters ofthe spore radius and its distai exine is uniformly granulate­conate. RAVN (1991 : 102) regarded W planiangulata as ajunior synonym of W sagittata, a view not endorsed by us,nor by other authors (e.g., KRAJEWSKI & STEMPIEN-SALEK2003 ; TRZEPIERCZYNSKA 2003; OWENS et aI. 2005) . Inactuality, the single specimen figured by RAVN (1991, pl. 2,figo 18), from the Visean of Alaska's North Slope, appearscongruent with W planiangulata.

Waltzispora polita (HOFFMEISTER, STAPLlN & MALLOY1955) SMITH & BUTTERWORTH 1967 (pp. 159-160 ; pl. 6,figo 14) - widely reported from Upper Mississippian-LowerPennsylvanian strata (e.g., PLAYFORD 1991: 93; PLAYFORDet aI. 2008: 97) - is generally smaller than W lanzonii andit differs further in possessing inconspicuous radial ex­pansions , longer laesurae , and laevigate to very weaklygranulate exine .

Co-occurring with Waltzispora lanzonii in samples be­longing to the Tournaisian BL and PD zones of theAmazonas Basin , Neoraistrickia loganii (WINSLOW 1962)COLEMAN & CLAYTON 1987: 83-84; pl. 2, figs . 5-9; text­fig o5) is chiefly distinguishable by its lack of shoulder-likeprojections at the amb apices and its coarser sculpturalelements.

4. Stratigraphic and palaeogeographicdistribution

The pioneering investigation by LANZONI & MAG­LOIRE (1969) of the palynostratigraphy of the sub­surface Upper Devonian through Mississippiansuccession of the Grand Erg Occidental (AIgerianSahara: Central Sahara Platforrn) provided the initialrecord of what was later to be established formally asWaltzispora lanzonii (see synonymy list, above). Theycited it as one of seven miospore species restricted to,and hence diagnostic of, the upper part (Sub-zone M3)of their Convolutispora Biozone. LANZONI & MAG­LOIRE'S (1969: 449) dating of M3 and the underlyingM2 Sub-zone as Tournaisian was supported by asso­ciated marine invertebrates (goniatites and articulatebrachiopods). Subsequently, COQUEL & ABDESSELAM­ROUIGHI (2000) and LOBOZIAK et aI. (2000) concur­rently and independently affirmed a mid to early late

Tournaisian dating of Sub-zone M3. Furthermore,COQUEL & ABDESSELAM-RoUIGHI (2000 : 357; figo 2;pl. I, I) recorded Waltzispora lanzonii from the upper(M3-correlative) part of the "Série argileuse" in theBéchar Basin, northwestern Grand Erg Occidental.BELHAJ (2000: 134) cited the species within a com­posite of 15 palynofloral components of the M'rarForrnation. The latter constitutes the basal unit of theCarboniferous succession in the Ghadamis (or Gha­dames) Basin of the Central Sahara Platform inwestern Libya, and reportedly spans the Tournaisianthrough Serpukhovian interval. However, BELHAJ(2000) did not specify the vertical range of W lanzonii(nor of any of the other listed taxa) within the M'rarFormation . Aside from that uncertainty, the specieshas not been recorded from post-Tournaisian strata inthe North African region (e.g ., PLAYFORD et aI. 2008and references cited therein) .

The presence of Waltzispora lanzonii in Missis­sippian strata of northern Brazil was first reported,and the specific epithet formally established, byDAEMON (1974), in his palynostratigraphic analysisof Upper Devonian and Mississippian subsurfacesuccessions of the Amazonas and Parnaíba basins.DAEMON (1974: 561; table 1) cited W lanzonii asoccurring in the Oriximiná Member (since accordedformational rank) of the Amazonas Basin, within hisbiostratigraphic intervals IX(p. 561)-X (table I), whichhe dated as Tournaisian. MELO & LOBOZIAK (2003:190-192; pl. 8, 23) further emphasized the biostrati­graphic importance of the species , detailing its verti­cal range as encompassing their interval zones AL,BL, and PD (early through early late Tournaisian) .These successively span the Mississippian part of theOriximiná Formation, the lower part ofwhich, in someAmazonas Basin wells, can range down into the"Strunian", i.e., the uppermost Devonian. As stated byMELO & LOBOZIAK (2003 : 191, footnote), DAEMON'S(1974: 561) mention of Waltzispora lanzonii occurringin the upper Visean Poti Formation of the ParnaíbaBasin is problematic and could well be explicableas either a consequence of recycling or by the hostsample or samples being representative of the LongáFormation rather than the overlying Poti Formation . Itis noteworthy that the species is unrecorded in MELO& LOBOZIAK'S (2000) palynological inventory of thePoti Formation, based on four Petrobras wells.

In a section of the Ambo Formation of Pongo deMainique , Peru, Azcuv & DI PASQUO (2005) identi­fied Waltzispora lanzonii in a single sample fromclose to the top of that formation . The same specimen

190 G. Playford and J. H. G. Melo

Fig. 5. Occurrences (star symbols) of Waltzispora lanzonii DAEMON 1974 located on Mississippian (Early Carboniferous)palaeogeographic map adapted from TORSVIK & COCKS (2004: figo 8). Gondwana prescribed by bold dot-dashed line.Key to numbered locations (generalized) and respective sources of data: I, Amazonas Basin, northern Brazil (DAEMON1974; MELO & LOBOZIAK 2003; present study); 2, Parnaíba Basin, northern Brazil (DAEMON 1974); 3, Pongo de Mainique,Peru (Azcuv & DI PASQUO 2005 , 2006); 4, Grand Erg Occidental, AIgeria, including Béchar Basin (LANZONI & MAGLOIRE1969; COQUEL& ABDESSELAM-RoUIGHI 2000); 5, Ghadamis Basin, Libya (BELHAJ 2000). See text for discussion ofin-situand probable reworked occurrences.

of W lanzonii was iIlustrated in Azcuv & DI PASQUO(2005: pl, 1, 4) and in the succeeding, essentiallyreiterative paper (Azcuv & DI PASQUO 2006: figo IA).Associated miospore species in the sample, notablyReticulatisporites magnidictyus PLAYFORD & HELBY1968 (pp. 110-111; pl. 10, figs. 7-10), indicated corre­lation with MELO & LOBOZIAK'S (2003) Mag IntervalZone of the Amazonas Basin and hence a late Viseanage. In that respect, the presence of W lanzonii posessomething of a biostratigraphic conundrum, insofar as

the species extends no higher in the Amazonas Basinthan the Tournaisian AL through PD biozonal interval;i.e., it was unrecorded in MELO& LOBOZIAK'S (2003)similarly well-documented Mag Zone. And it is alsopertinent to note that other South American occur­rences ofthe Mag Zone or its equivalents - as reportedfor instance by CÉSAR! & GUTIÉRREZ (2001) andPÉREZ LOINAZE (2007, 2008 a, b) from Argentina andFASOLO et aI. (2006) from Bolivia - are likewise evi­dent1y devoid of the readily identifiable W lanzonii .

lhe Tournaisian(EarlyMississippian) miospore Waltzisporalanzonii OAEMON 1974 191

Whether Azc uv & DI PASQUO (2005, 2006) encoun­tered only one speeimen of W lanzonii in their Amb osample is a moot po int; assu redly, lhe number wouldbe less, perhaps eonsidera bly less, than 17, judgi ngfrom Azcuv & DI PASQUO (200 5: 176). Moreover, itis clear from their data that their Mag-assignedassemblage eontains a not- insignifieanl proportion ofreworked palynomorphs, and hene e that the specimen(or speeimens?) of W lanzonii eould well belong inlhe latter category, Th e generally robusl morpho­logieal eharaeter of the spee ies woul d of eourse tendlo favour its reeycl ing potentiality.

From lhe above , it is evident that in-situ occ ur­renees of Waltzispora lanzonii are largely if not ex­clusive ly eonfined lo slrata of Early Mississippian(Tournaisian) age . Its stratigraphie ineept ion is eer­tainly post-Oevo nian, as exemplified by its FAO (firstappea ranee daturn) ai lhe lower (but nol neee ssarilylowest) Tournaisian base of lhe AL Interval Zone oflhe Amazonas Basin (MELO & LOBOZlAK 2003 : 190 ;figo 14) and supported also by North Afriean data(primarily LANZONI & MAGLOlRE 1969). In the Ama­zonas Basin, W lanzonii ranges thro ughout the enti reTourna isian seetion of lhe Oriximiná Formation, thusspanning the AL , BL, and PO interval zones (althoughits frequeney declines in PO; MELO & LOBOZIAK2003: 192). Whether lhe species's LAO (lasl app ear­anee datum) approximates to or extends sornewhatbeyond the close of the Tournaisian is une lear, at leastin the Amazona s Basin, wher e an appreeiable hiatusseparares the Oriximiná Formation from the overlyingupper Visean Faro Formation (Mag Interval Zone , anddevoid of demonstrably autoc hthonous speei mens ofW lanzonii). Henee, lhe possibi lity that the speeieseo uld extend into lhe lower part of the Visean can­not objeetively be discounled, although LANZONI &MAGLOIRE'S (1969) AIgerian study does not suggestsueh exte nsion . As diseussed previously, lhe sup­posedly in-situ eredentia ls ofan isolaled oee urrenee ofW lanzonii in a Peruvian sam ple dated as late Viseanmusl, ai this stage, be regarded as questionable. Thesame doubt pertains to the alleged presenee of lhespeeies in the similarly dated Poti Formation (Par­naiba Basin, northern Brazil ; OAEMON 1974: 56 1).

The palaeogeograph ie distribution of Waltzisporalanzonii oee urrenees - both in situ (of Tournaisianage) and those judged to be a eonsequenee of re­eyeli ng (Iate Visean) - indieale s that the speeies is, oneurrent evidenee, restricted lo the weste rn-n orth­westcrn region of Gond wana (Fig. 5); i. e., to a regionthat broadly includes prescnt-day eenlral South

Ameriea plus North Afriea, whieh were of eourse injuxtaposition during lhe Palaeozoie. Although globaleoverage ofMissis sippian palynofloras is by no meanseomprehensive, investigations have been conducted inmany regions of Eastern and Southern Gondwana andofEurameriea (see, e. g., CLAY10N 1996; PLAYFORO &OINO 2005 ; TRAVERSE 2007) and in these W lanzoniiso far remains unre eorded . (A nd il is pertinenl loemp hasize here that lhe morph ologieal distinctivenessof lhe speeies is sueh that , if pre sent , it would hardlygo unnotieed). Thus, it appea rs that the distrib ution ofW lanzo nii eould well be lirnited lo that seetor ofGondwana, depieted in Fig. 5, whieh ineorporates itspresently known oeeurrenees .

In the Gondwanan Tournaisian palynofloras con­taining undoubtedly in-situ Waltzispora lanzonii, noother representatives of lhe genus have so far beenreported. This cont rasts to younger Mississippianpalynofloras of Eurameriea, especially Spitsbergenand other Aretie/Subarclie regions, in whieh Waltzi­spora is more diversely and abundantly represented.

5. Conclusions

I. The substantial majority of miospores attributablelo Waltzispora lanzonii constitute a distinctive andeohe sive morphospeeies distinguished by a eharae ­leristiea lly trilobate equatorial out líne featu ringvariably expanded apieal regions, and by re lativelysparse apieulate (baeulate, eon ale , spin ose) ex ina lseulpture that tends to be more eoneentrated or eon­spic uous on apica l regions of the distal surfaee. Astrict minority of atypi eaI or aberrant speeimensdeviate morphoIogieally from lhe majority, eh iefly inequatorial outline, but clearI y do not warrant taxo­nomic discrimination.

2. Identifieation of this speeies in Mississippian paly­nofloras of northern Brazi l (Amazonas and Parnaibabasins) and Peru , and of intracratonic basi ns of lheCen tral Sahara Platform of North Afriea, signifiesits palaeogeographie restriction to the western tonorthwestern seetor of Gondwa na.

3. In ehronostratig raphie terms, Waltzispora lanzonii- seemingIy the oldest known represe ntative of thegenus - is eertainly post-Devon ian and is eonf inedlo the Mississipp ian and very likely to its lowerstage (Tournaisian) . Th e in-situ authenticity of POSl­Tourn aisian oceurrenees of this species remains anopen question ; however, it seems probable that lhe

192 G. Playford and J. H. G. Melo

very few such records - from South Ameriean upper

Visean strata - are of reeyeled origino

Acknowledgements

Grateful acknowledgement is extended to the following:ADELl NO TEI XEIRA DIAS (of Petrobras/CenpeslPdexp/Bpa)for drafting assistance; and JOSÉ RICARDO MAIZATIO andMÁRCIA EMILlA LONGHIM (both of Petrobras/Cenpes/Pdexp/Bpa) for reviewing and helpfully eommenting on theoriginal manuscript. The management ofPetr61eoBrasileiroS.A. has granted permissionfor publication.

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Manuscript received: February 18th, 2009.Revised version accepted: February 26th, 2009.

Addresses of the authors:

GEOFFREY PLAYFORD, Schoo1 of Earth Sciences, TheUniversity of Queens1and, Brisbane, Queens1and 4072,Austra1ia;e-mail: g.p1ayford@uq.edu.au

Joss HENRIQUE G. DEMELO, Petrobras/Cenpes/Pdexp/Bpa,Avenida Horácio Macedo, 950, Cidade Universitária, Ilhado Fundão, 21941-915 Rio de Janeiro, RJ, Brazi1;e-mail: jhme1o@petrobras.com.br