Silurian cryptospores and miospores from the type Llandovery area, south-west Wales

25 Pages·1991·9.9 MB·English

by  N D Burgess

SILURIAN CRYPTOSPORES AND MIOSPORES
FROM THE TYPE LLANDOVERY AREA,
SOUTH-WEST WALES
by N. D. BURGESS
Abstract. The oldest cryptospores and miospores have great significance in studies of the evolution ofland
plants: the former may represent the earliest direct evidence of such organisms and the latter may provide
evidence for rhyniophytoid land plants as they have been recovered from the sporangia ofCooksoniapertoni
Langin thelateSilurian. In thetype Llandoveryarea, twodistinctsporomorph assemblagesaredescribed from
a composite section through uppermost Ordovician, Rhuddanian, Aeronian and basal Telychian strata. The
older(latestOrdovician to lateAeronian)compriseseight generaand fourteen speciesofcryptospores (tetrads,
pseudodyads, true dyads and monads). The younger (late Aeronian to Telychian) is dominated by smooth-
walled trilete miospores of species of the genus Ambitisporites. Both assemblages have strong similarities
to those described from similar horizons around the world. Specimens of Ambitisporites dilutus from the
sedgwickii Graptolite Biozone of the Aeronian/Telychian type boundary section are the oldest known with
unequivocal dating. They are used to define the base of the Ambitisporites dilutus- A. avitus Sporomorph
Zone. Three new cryptospore genera Rimosotetras Segestrespora and Velatitetras) are erected and
( ,
Tetrahedraletes is emended and synonomized with Nodospora. Six new cryptospore species (Velatitetras
laevigata, V. reticulata, V. rugulata, Rimosotetrasproblematical SegestresporalaevigataandS. rugosa),andtwo
new varieties (T. medenensis vars medenensis and parvus) are described, and two combinations are made.
Currently sporomorphs provide the most convincing evidence for the existence of land plants
in the late Ordovician and early Silurian (Gray 1985), the best being those with the greatest
similarity to ones from living plants, or fossils of proven land-plants (Gray et al. 1982; Edwards
et al. 1983; Gray 1985; Richardson 1985; Fanning et al. 1988). Spores in almost all such plants are
produced in tetrads which separate on maturity to yield four trilete miospores. There are few
records of trilete miospores in the Llandovery (Hoflfmeister 1959; Cramer 1968, 1969; Pratt et al.
1978; Aldridge et al. 1979; Hill et al. 1985; Gray 1988; Richardson 1988). However, recent work
has demonstrated the presence of permanently fused tetrads, dyads and monads, all ofwhich can
be enclosed within an envelope, in the late Ordovician and early Llandovery (e.g. Gray and Boucot
1971 Strother and Traverse 1979; Gray et al. 1982; Miller and Eames 1982; Gray 1985; Johnson
;
1985; Vavrdova 1982, 1984, 1988). As these unusually constructed cryptospores sensu Richardson
(
et al. 1984) may represent the oldest evidence of land-plants they have become important in the
debate on the appearance and subsequent evolution of such organisms (Gray and Boucot 1977;
Taylor 1982; Richardson 1985; Gray 1985, 1988).
Despite their importance, few assemblages of Ordovician and Llandovery sporomorphs have
been described from well-dated sections, or ones covering a long stratigraphic interval (Pratt et al.
1978; Strother and Traverse 1979; Miller and Eames 1982; Johnson 1985; Richardson 1988). This
has hindered interpretation of the biostratigraphic and evolutionary significance of the
sporomorphs. This paper presents full descriptions and stratigraphic ranges for those cryptospores
and trilete miospores recovered from a composite section through the uppermost Ordovician and
the majority of the Llandovery Series in the type Llandovery area (Cocks et al. 1984).
IPalaeontology, Vol. 34, Part3, 1991, pp. 575-599, 2 pls.| © The Palaeontological Association
576 PALAEONTOLOGY, VOLUME 34
text-fig. I. Location ofthe type Llandovery area in
southern Wales.
text-fig. 2. Locations ofsamples collected from the
late Ordovician Scrach Formation to the early
Llandovery Bronydd and Crychan formations in the
type Llandovery area: Forestry track F.33, Crychan
Forest near Llandovery (Ordnance survey map refer-
ence SO 84583964 at sample LI = locality 24 of
Cocks et al. 1984).
text-fig. 3. Locations ofsamples collected from the
Rhuddanian/Aeronian type boundary section in the
type Llandovery area: Forestry track F.33, Crychan
Forest, near Llandovery (Ordnance survey map
reference SO 83913960 at sample LI I = locality 38 of
Cocks et al. 1984).
6
BURGESS: LLANDOVERY SPOROMORPHS 577
text-fig. 4. Locations ofsamples collected from the
Aeronian/Teychian type boundarysection in thetype
Llandoveryarea: roadside nearCefnCerigfarm, near
Llandovery (Ordnance survey map reference SO
77543271 at sample LI7 = locality 154 ofCocksetal.
1984).
GEOLOGY, SAMPLING AND TECHNIQUE
The location ofthe type Llandovery area is presented in Text-figure 1. A single sample was collected from the
uppermost Ordovician and twenty-two from the Llandovery Series. Sequences collected spanned the
Ashgill/Rhuddanian boundaryexposedalongForestryTrack F. 33 next totheAfonCrychan riverinCrychan
Forest, c. 7 km north-east of Llandovery (Text-fig. 2), the Rhuddanian/Aeronian type boundary section
(Text-fig. 3) and the Aeronian/Telychian type boundary section (Text-fig. 4) (Cocks et al. 1984). The top
c. 100 m of the sequence in the type area were not sampled: this interval covers the crispus to crenulatus
Graptolite Biozones. The stratigraphical framework of the samples is given in Text-figure 5.
The Llandovery sediments ofthe type area were deposited during a transgression which affected the whole
of the Anglo-Welsh Basin (Cocks et al. 1984; Ziegler et al. 1968). As a consequence the samples collected
duringthisstudyarefromincreasinglydistalmarinefaciesfurtherupthesection(Table 1). However, theremay
be slight shallowing event in the Wormwood Formation.
table 1. Distribution of samples by geological formation with the ages and probable depositional
environment.
Probable depositional environment
Samples Formation Age (Cocks et al. 1984)
L23 Cerig early Telychian Open marine shelf
L19-L22 Wormwood late Aeronian Open marine shelf
L17-L18 Rhydings late Aeronian Open marine shelf
LI —L Trefawr early Aeronian Open marine shelf
1 1
L7-L10 Crychan late Rhuddanian Open marine prograding delta lobe
L2-L6 Bronydd early Rhuddanian Mud-dominated marine shelfwith storms
LI Scrach latest Ashgill Shallow sub-tidal or even intertidal
.
PALAEONTOLOGY, VOLUME
578 34
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FORMATIONS
acrach Bronydd Crychan Trefawr
j~** Rhydings Wormwood Cerig
| | [
text-fig. 5. Biostratigraphic and lithostratigraphic position of samples from the type Llandovery area, in
relation to localities ofCocks et al. (1984).
BURGESS: LLANDOVERY SPOROMORPHS 579
Samples varied from dark mudstones in the Scrach and Bronydd formations through to muddy siltstones
and silty mudstones for the remainder of the sequence. Palynomorphs were extracted using standard
palynological methods (HC1 then HF acids followed by sieving with 10//m filter mesh and separation ofthe
organic fraction usingzinc bromide solution (S.G. 20)). Residueswere oxidized in cold Schultze’s solution for
c. 48 hours. The sporomorphs remain dark, but poor preservation did not allow any further extension ofthe
oxidation time. When this was attempted all the sporomorphs disintegrated. After oxidation, the acidic
residues were neutralized with distilled water.
For light microscope observation, measured volumes ofmaterial were strewed on to glass coverslips, dried,
and the coverslips attached to glass slides with ‘Elvacite’ plastic mounting medium.
Photomicrographs were taken on FP4 film using a Zeiss photomicroscope III (no. 2562) housed in the
Palynology Laboratory ofthe British Museum (Natural History) (BM(NH)), and using Nomarski differential
interference contrast with an orange filter to reduce contrast between the palynomorphs and the background.
SYSTEMATIC PALAEONTOLOGY
Preamble. Sporomorphs are described, where possible, using the standard terminology of Grebe (1971).
Specimen dimensions are presented as the minimum and maximum ofthe range, with the mean in brackets.
Stratigraphicrange referstorangewithinthetype Llandoveryareaonly. Thestratigraphical occurrenceoftaxa
is shown in Text-figure 7 and the stratigraphical distribution and correlation ofthe samples in Text-figure 5.
Specimens are located by means ofstandard England Finder and microscope stage co-ordmates taken from
the Zeiss photomicroscope III (no. 2562) housed in the Palynology Laboratory at the BM(NH). All figured
specimens arealso ringed with a red indeliblepen. Slidesand stubscontainingfigured specimens(prefixed EM)
are stored in the Palynology collection at the BM(NH).
Anteturma cryptosporites Richardson et a/., 1984
1. Cryptospore tetrads
Thisgroupcomprises tetradsand dyads which are not found separated into individual components sporesand
which arebelieved tobe‘permanently’fused. Theymayalsobeenclosedwithinasculpturedornon-sculptured
envelope.
Genus tetrahedraletes Strother and Traverse, 1979 emend.
Type species. Tetrahedraletes medinensis Strother and Traverse, 1979 from the Tuscarora Formation,
Pennsylvania, USA.
Emended diagnosis. Permanent tetrahedral cryptospore tetrads, permanently fused. Tetrads sub-
circular to circular in outline and composed of four laevigate, crassitate, sub-triangular ‘spores’.
Crassitudes of individual ‘spores' +equatorial, fused or discrete. Distal ‘spore’ walls tend to
invaginate, but can remain inflated.
Discussion. Tetrahedraletes Strotherand Traversewaserected toencompass ‘tetradsofinaperturate,
sub-triangular spores or spore-like palynomorphs arranged in tightly adhering tetrahedron
configuration, with the spore walls collapsed towards the centre’ (Strother and Traverse 1979, p. 8).
Nodospora, a similar genus, was defined for ‘tetrads of inaperturate spores or spore-like
palynomorphs, spherical to sub-spherical in outline, which are arranged in a cross configuration'
Strother and Traverse (1979, p. 10). Data collected here and in other recent publications (e.g. Gray
et cd. 1985; 1986; Gray 1985; 1988) indicate that the type specimens of Tetrahedraletes (T.
N
medinensis) and Nodospora burnhamensis) should be placed in synonomy, the two taxa
( .
representing different compressional morphologies of otherwise identical tetrads. Hence, in this
publication all such tetrads are placed in Tetrahedraletes and the diagnosis of this genus emended
accordingly.
a
PALAEONTOLOGY, VOLUME
580 34
Tetrahedraletes medinensis Strother and Traverse, 1979 emend.
This species has been subdivided into two varieties based on size because the original specimens (Strother and
Traverse 1979) aremuch larger than those recorded from the late Ordovician and early Llandovery (e.g. Gray
1988).
Tetrahedraletes medinensis Strother and Traverse, 1979 var. medinensis var. nov.
Holotype andtype locality. Strother and Traverse, 1979, pi. 1, fig. 5: Harvard Paleobotanical Collections no.
60289, slide no. 75-4/A3, location on slide 34-8 mmx 107-9 mm, reference point 13-3 mm x 109-2 mm. Samples
75-4, Tuscarora Formation; Section - Mann Narrows along route 322, north-west of Burnham, Mifflin
County, Pennsylvania, USA.
Description. As for Strother and Traverse (1979).
Comparison. Cryptospores of Tetrahedraletes medinensis var. parvus var. nov. are < 35 pm in diameter, but
otherwise identical.
Remarks. This variety of T. medenensis is common in the basal Wenlock (Burgess and Richardson
1991) but is not known from the uppermost Ordovician or early Llandovery ofthe type area where
the following variety is found (also see Gray 1988).
Tetrahedraletes medinensis var. parvus var. nov.
Plate 1, figs 1-4
1985 Tetrahedraletes cf. T. medinensis Gray, fig. 5 F-H.
EXPLANATION OF PLATE
1
All figures x 1000 unless stated otherwise.
FM
Figs 1-4. Tetrahedraletes medinensis Strother and Traverse, 1979 emend, var. parvus var. nov. 1, 187,
hBoilozootnyep,e (xsl2i0de00L.la2,nF8BM/121,881(7s8li1de17L0l;aEn.F8.B/n8o,:0S6417/1128),3s;aEm.pFl.enLo5:,FB5r8o/n2y),ddsaFmoprlmeatLi5o,n,BraocninyadcdesFGorrampattoiloint,e
FM
acinaces Graptolite Biozone. 3, 92 (slide Llan 5/2, 130 960; E.F. no: N26/1), sample L16, Trefawr
Formation, triangulatusGraptolite Biozone. 4, FM 189 (slide Llan 1/1, 112 1204; E.F. no: K50/4), sample
LI Trefawr Formation, cyphus Graptolite Biozone.
Figs 51,and 6. Velatitetras laevigatagen. et sp. nov. 5, FM 190 (slide Llan 8B/11, 179 1294; E.F. no: S59/S60),
sample L5, Bronydd Formation, acinaces Graptolite Biozone. 6, FM 191, holotype (slide Llan 2/1, 050
1215; E.F. no: E51/4, sample LI2, Trefawr Formation, cyphus Graptolite Biozone.
Figs 7-9. Velatitetras reticulata gen. et sp. nov. 7, FM 91 (slide Llan 1/2, 232 1110: E.F. no: Y41/2), sample
FM
Lll, Trefawr Formation, cyphus Graptolite Biozone. 8, 197 holotype (slide Llan 8B/9, 128 1253: E.F.
no: N55/2), sample L5, Bronydd Formation, acinaces Graptolite Biozone. 9, FM 192 (slide Llan 8B/8, 106
1304; E.F. no: K60/61/L60/61 sample L5, Bronydd Formation, acinaces Graptolite Biozone.
), FM
Fig. 10. Velatitetras rugulata gen. et sp. nov. 194, holotype (slide Llan 8B/12, 128 1220; E.F. no:
M52/3/M51/4), sample L5, Bronydd Formation, acinaces Graptolite Biozone.
Fig. 11. Velatitetras sp. A. FM 195 (slide Llan 8B/15, 118 1277; E.F. no: M58/1), sample L5, Bronydd
Formation, acinaces Graptolite Biozone.
FM
Figs 12, 14, 15. Rimosotetrasproblematic gen. et sp. nov. 12, 196 (slide Llan 8B/12, 240 1080; E.F. no:
Y38/3), sample L5, Bronydd Formation, acinaces Graptolite Biozone. 14, FM 198, holotype (slide Llan
FM
8B/6, 138 1228; E.F. no: 053/1), sample L5, Bronydd Formation, acinacesGraptolite Biozone. 15, 199
(slide Llan 8B/8, 147 1089; E.F. no: P38/2), sample L5, Bronydd Formation, acinaces Graptolite Biozone.
FM
Figs 13, 16, 17. Pseudodyadospora cf. laevigata Johnson, 1985. 13, 197 (slide Llan 8B/26, 035 1250; E.F.
no: C55/1/2), sample L5, Bronydd Formation, acinaces Graptolite Biozone. 16, FM 200 (slide Llan 8B/22,
226 1112; E.F. no: X41/I), sample L5, Bronydd Formation, acinacesGraptolite Biozone. 17, FM 201 (slide
Llan 8B/12, 240 1095; E.F. no: Y39/4), sample L5, Bronydd Formation, acinaces Graptolite Biozone.
PLATE
1
fee '1
aW
BURGESS, Llandovery sporomorphs
582 PALAEONTOLOGY, VOLUME 34
1985 Tetrahedraletes cf. T. medinensis Gray et a/., p. 524, fig. 5 F-H.
1986 Tetrahedraletes cf. T. medinensis Gray et al., p. 451, fig. 7, items 1-7.
1988 ‘Smooth walled tetrads’. Gray, p. 355, pi. 1, figs 1 and 5.
1988 Tetrahedraletes sp., Richardson, pi. 19, fig. 1.
FM
Holotype and type locality. 187, PI. 1, fig. I, slide Llan 8B/12, 178 1170, E.F. no: S47/1, sample L5
(Text-fig. 2), Forestry Track F.33, Bronydd Formation, acinaces Graptolite Biozone, Rhuddanian.
Paratypes. FM 188, PI. 1, fig. 2, slide Llan 8B/8, 061 1283, samples L5; Forestry Track F.33, Bronydd
Formation, acinaces Graptolite Biozone; FM92, PI. I, fig. 3, slide Llan 5/2, 130 0960, sample L20,
Aeronian/Telychian boundary section. Wormwood Formation, sedgwickiiGraptolite Biozone; FM 189, PI. 1,
fig. 4, slide Llan 1/1, 020 1204, sample L1 1, Rhuddanian/Aeronian boundary section, Trefawr Formation,
cvphus Graptolite Biozone.
Derivation ofname. Latinparvus small.
,
Diagnosis. A variety of Tetrahedraletes medinensis < 35 pm in diameter with low, narrow, rounded
and unfused equatorial crassitudes.
Description. Laevigate obligate tetrads, sub-circular to circular in outline and preserved in a variety of
compressional morphologies depending on the degree ofrotation ofthe tetrad from an apical view prior to
compression. Individual ‘spores’ laevigate, amb sub-circular to sub-triangular, joined by unfused equatorial
crassitudes 1-5 pm wide. Proximal surface not observed. Distal exine 1-2pm thick and invaginated in most
specimens.
Dimensions. Tetrads 19 (26) 30//m in diameter (150 specimens measured, see Text-figure 6 for range of
measurements).
text-fig. 6. Size frequency distribution ofa hundred
Tetrahedraletes medinensis var. parvus tetrads from
sample L5, ForestryTrackF.33, BronyddFormation,
acinaces Graptolite Biozone, Rhuddanian.
MICRONS
Biostratigraphical range. Latest Ordovician (persculptus Graptolite Biozone), to early Telychian (turriculatus
Graptolite Biozone; Text-fig. 7).
Comparisons. Tetrahedraletes medinensis var. medinensis is larger (> 35pm). Specimens described as
Tetrahedraletes cf. T. medinensis (Gray 1988) from various sections spanning the Ordovician/Silurian
boundary around the world have essentially identical measurements to those described above.
Remarks. Gray (1988) has presented data showing that tetrads gradually increase in size over the
Ordovician/Silurian boundary and states that this increase continues through the Llandovery. In
the Anglo-Welsh Basin, T. medinensis var. parvus var. nov. is the dominant variety in the latest
Ordovician and early Llandovery, whereas T. medinensis var. medinensis Strother and Traverse,
1979 dominates in the basal Wenlock (Burgess and Richardson 1991).
BURGESS: LLANDOVERY SPOROMORPHS 583
Genus velatitetras gen. nov.
Type species. Velatitetras laevigata sp. nov.
Thisgenusiserected to accommodate tightlyadherent cryptospore tetradsenclosed within anenvelope. Some
specimens of Nodospora Strother and Traverse are believed to possess an envelope (e.g. N. retimembrana).
However, because the type species of Nodospora (N. burnhamensis) lacks an envelope and is considered
synonomous with the type species of Tetralredraletes (T. medinensis) the genus Nodospora cannot be used for
enveloped forms. Stegambiquadrella Johnson was erected for palynomorphs with four loosely-attached
inaperturate vesiclesenclosed within anenvelope. Thisgenuscannot be used toaccommodate tightly-adherent
tetrahedral tetrads enclosed within an envelope.
Derivation ofname. Latin velatus. covered; tetras tetrad.
,
Diagnosis. Obligate cryptospore tetrads composed of tightly-adherant laevigate sub-triangular to
sub-circular ‘spores’ with low and rounded +fused equatorial crassitudes. Tetrads enclosed within
a closely adherent to completely separated, +ornamented envelope; when envelope tightly-
adpressed any ornamentation passes over ‘spore’ contacts uninterrupted.
Velatitetras laevigata sp. nov.
Plate 1, figs 5 and 6
1985 ‘Obligate tetrad with smooth perispore’, Gray, p. 177, pi. 1, figs 2 and 3.
1988 ‘Spore tetrad with smooth or possibly degraded reticulate envelope’. Gray, p. 355, pi. 1, fig. 3.
1988 Nodospora sp. B, Richardson, p. 94.
Holotvpe andtype locality. FM 191, PI. I, fig. 6, slide Llan 2/1, 050 1215, E.F. no: E51.4, sample L12 (Text-
fig. 3), Rhuddanian/Aeronian type boundary section, Trefawr Formation, cyphus Graptolite Biozone, late
Rhuddanian.
Paratype. FM 190, PI. 1, fig. 5, slide Llan 8B/11, 179 1294, sample L5, Forestry Track F.33, Bronydd
Formation, acinaces Graptolite Biozone.
Derivation ofname. Latin laevigatus smooth.
,
Diagnosis. A Velatitetras with cryptospore tetrad enclosed within a thin, laevigate and often folded
envelope.
Description. Obligate tetrads, sub-circular to circular in outline, totally enclosed within an envelope. Tetrads
composed oflaevigate, sub-triangular ‘spores’ with unfused, low and rounded equatorial crassitudes 1-3 pm
wide; distal polar exine c. 1 pm thick. Envelope laevigate, diaphanous, folded and < 1 //m thick; varies from
completely separated, to closely adpressed to tetrad.
Dimensions. Enclosed tetrads 21 (27) 34/im in diameter (22 specimens measured).
Biostratigraphic range. Latest Ordovician (persculptus Graptolite Biozone), to late Rhuddanian (cyphus
Graptolite Biozone; Text-fig. 7).
Comparisons. Gray (1985, pi. 1, figs 2 and 3; 1988, pi. 1, fig. 3), illustrated small (c. 25 pm) and
essentially identical tetradsenclosed within smooth envelopes from the Late Ordovician ofAmerica.
Biostratigraphic
Note
and
that
the
lithostratigraphic
majority
of
the
ranges
Telychianof
was
sporomorphs
not
sampled.from
the
type
Llandovery
area. SAMPLES