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Review
. 2019 Mar 25:119:67-95.
doi: 10.3897/phytokeys.119.29555. eCollection 2019.

A morphometric analysis of Tobleriabicuspis, a Voltziales seed cone from the early Permian Jambi palaeoflora, Sumatra (Indonesia)

Affiliations
Review

A morphometric analysis of Tobleriabicuspis, a Voltziales seed cone from the early Permian Jambi palaeoflora, Sumatra (Indonesia)

Isabel M Van Waveren. PhytoKeys. .

Abstract

Tobleriabicuspis, a coniferophyte seed cone, is described from the Jambi Palaeoflora, Sumatra of Asselian (early Permian) age. A morphometric analysis based on cones, paired fertile units, and fertile and sterile scales, demonstrates their close relationship. Small paired fertile units occur mainly in cones. Medium-sized paired fertile units occur mainly on scales. And large paired fertile units are mainly dispersed. The cones are considered female and the paired fertile units are considered to represent the seeds. The cones are composed of helicoidal, bilaterally symmetrical and deeply incised scales with paired seeds. A comparison can be made with the Voltziales female taxon Schizolepis from the Triassic and Jurassic. Tobleria is regarded as having a voltzian Voltziales affinity and dates from approximately 16 to 26 million years before any other such cones.

Keywords: Asselian; Cathaysia; early conifer cones; mesic-xeric.

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Figures

Figure 1.
Figure 1.
Map of the Bangko area showing the outcrops of the Mengkarang Formation along the distributaries of the Merangin River where Tobleriabicuspis was found.
Figure 2.
Figure 2.
The three cones from sample 45311: A cone C1 from 45311 Aa, the black frame indicates the area that is detailed in Figure 5BTobleriabicuspis right edge of cone C2 fragment showing bicuspid scale and seeds CTobleriabicuspis cone C3 fragment showing bicuspid scale. Arrows 1–4 in Figure A indicate scales in side view, arrow 5 indicates the cone axis. The arrow in Figure C indicates a bifid scale. Scale bars: 10 mm (A); 2,5 mm (B); 5 mm (C).
Figure 3.
Figure 3.
Associated seeds and scales A Dispersed seeds and scales (sample 45311a) B Scale with hart shaped scar and scale with darker central area (sample 45471) C Scale seen from adaxial side showing striation pattern and scale seen from adaxial side showing seed remains (sample 45313). Scale bars: 5 mm.
Figure 4.
Figure 4.
Diagram showing empty scale, opaque (A1) or lighter (A2), fertile units as dark discs (B1) or dark circular rims (B2), and fertile scale with lighter fertile units on darker scale (C1) or darker fertile units on lighter scale (C2) (see also Figure 6). It is indicated how length and width of scale and fertile unit are measured.
Figure 5.
Figure 5.
Details from Cone C1: A Facial view of the scale in the cone from sample 45311 Aa (see frame in Figure 2A) B Contours of Facial view of scale and seeds indicated by dashed lines.
Figure 6.
Figure 6.
Details of dispersed seeds, scales and fertile scales: A paired seeds with double wall (sample 45311C) B paired seeds with triangular micropylar protrusion (sample 45311 E) C detail of triangular micropylar protrusion from paired seeds (sample 45311 E) D heart shaped paired seeds/ovules (sample 45311 D) E heart shaped paired seeds/ovules (sample 45311 B) F juxtaposed almond shaped seeds/ovules (sample 45311 B) G bicuspid scale (sample 45311 Aa) H bicuspid scale (sample 45311D) I bicuspid scale (sample 45471) J bicuspid scale with contour of two seeds/ovules (sample 45315) K bicuspid scale with darker organic contour of seed (sample 45310) L bicuspid scale with heart shaped contour of seeds/ovules (sample 45471).
Figure 7.
Figure 7.
Normal probability plot for the seed surface (length x width) proxy (n=263) (green represents seeds in cone, red represents seeds on scales, and black represents dispersed paired seeds).
Figure 8.
Figure 8.
Histogram for the seed surface (length x width) proxy distribution compared to the normal distribution curve (n=263).
Figure 9.
Figure 9.
Comparison between the box charts of the proxy for seeds size (length x width) in cones, on scales and free.
Figure 10.
Figure 10.
Proxy for seed surface (length x width) organised in ascending order with the superimposed polynomial function describing it (n=263).
Figure 11.
Figure 11.
Normal probability plot for the scale length (n=158). Green represents scales in cone, red represents dispersed fertile scales, and black represents dispersed empty scales.
Figure 12.
Figure 12.
Normal probability plot for the scale width (n=158) (green represents scales in cone, red represents dispersed fertile scales, and black represents dispersed empty scales).
Figure 13.
Figure 13.
Histogram for the proxy for scale surface (length x width) distribution compared to the normal distribution curve (n=158).
Figure 14.
Figure 14.
Comparison between the box charts of the proxy for seeds surface (length x width) in cones and dispersed, fertile and empty (n=158).
Figure 15.
Figure 15.
Proxy for scale surface (length x width) organised in ascending order (n=158).
Figure 16.
Figure 16.
Absence of fluorescence in a scale from the Cone C1 from 45311 Aa: A scale under normal light showing the scale as being slightly darker than the rock B identical scale with fluorescent light showing the scale as being opaque while the rock around it shows some fluorescence (arrows indicate the scale).
Figure 17.
Figure 17.
Reconstruction of Tobleriabicuspis. Scale bar: 1 cm.

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