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LM micrographs of all Sclerosperma pollen types. A-E. Sclerosperma mannii (from Cameroon, coll. van der Burgt, 1958 [K]), same grain in polar and equatorial view. F-J. Sclerosperma profizianum Type A (from DR Congo, coll. Gillet, 279a [WAG]), same grain in polar and equatorial view. K-O. Sclerosperma profizianum, Type B (from Angola, coll. Grobbelaar, s.n. [K]), same grain in polar and equatorial view. P-T. Sclerosperma profizianum, Type C (from Ghana, coll. Hall & Enti, GC 36150 [K]), same grain in polar and equatorial view. U-Y. Sclerosperma profizianum, Type C (from R Congo, coll. Profizi, 841 [K]), same grain in polar and equatorial view. Z-D′. Sclerosperma walkeri (from DR Congo, coll. Leonard, 1614 [BR]), same grain in polar and equatorial view. A, C, F, H, K, M, P, R, U, W, Z, B′. Polar view, high focus. B, D, G, I, L, N, Q, S, V, X, A′, C′. Polar view, optical cross section. E, J, O, T, Y, D′. Equatorial view, upper in high focus and lower in optical cross-section. Scale bars-10 µm.

LM micrographs of all Sclerosperma pollen types. A-E. Sclerosperma mannii (from Cameroon, coll. van der Burgt, 1958 [K]), same grain in polar and equatorial view. F-J. Sclerosperma profizianum Type A (from DR Congo, coll. Gillet, 279a [WAG]), same grain in polar and equatorial view. K-O. Sclerosperma profizianum, Type B (from Angola, coll. Grobbelaar, s.n. [K]), same grain in polar and equatorial view. P-T. Sclerosperma profizianum, Type C (from Ghana, coll. Hall & Enti, GC 36150 [K]), same grain in polar and equatorial view. U-Y. Sclerosperma profizianum, Type C (from R Congo, coll. Profizi, 841 [K]), same grain in polar and equatorial view. Z-D′. Sclerosperma walkeri (from DR Congo, coll. Leonard, 1614 [BR]), same grain in polar and equatorial view. A, C, F, H, K, M, P, R, U, W, Z, B′. Polar view, high focus. B, D, G, I, L, N, Q, S, V, X, A′, C′. Polar view, optical cross section. E, J, O, T, Y, D′. Equatorial view, upper in high focus and lower in optical cross-section. Scale bars-10 µm.

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Three currently accepted Sclerosperma species appear to produce four different pollen morphologies. Sclerosperma mannii and S. walkeri pollen share the same distinct reticulate sculpture, but S. profizianum produces three different pollen types (microreticulate, fossulate, and perforate). The pollen morphology suggests that S. mannii and S. walkeri...

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... first LM and SEM micrographs showing pollen of this taxon are by Harley and Hall (1991, plate 4, figures 32 [SEM] and 33 [LM]). The same SEM micrograph is shown in Harley (1999, plate 1, figure 12), and again along with two addi- tional LM micrographs and two attached grains under SEM in Harley (1996, plate 16, figures C and F [SEM], and G and H [LM]). These are all repeated in Harley and Baker (2001, figures 77 and 82 [SEM], 80 and 81 [LM]). ...
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... are all repeated in Harley and Baker (2001, figures 77 and 82 [SEM], 80 and 81 [LM]). A new SEM detail is provided in Harley and Dransfield (2003, figure 11). In total, five or six grains were illustrated using either LM or SEM micrographs, but no TEM micrograph has been presented thus far. ...
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... C (Hall & Enti, GC36150 [K]; Profizi, 841 [K]) ( Figures 1P-Y, 5; Table III) Description. -Pollen, monad, heteropolar, P/E ratio oblate, outline straight-triangular to slightly convex-triangular in polar view, bean-shaped in equatorial view (convex distal face versus concave proximal face); equatorial diameter 37-42 µm in LM, 31-39 µm in SEM, polar axis 10-16 µm in LM; triporate, pori positioned sub-apically on the distal polar face, pori circular to elliptic, 4.0- 6.5 µm in diameter, pori equipped with opercula; exine 1.7-2.5 µm thick in LM, nexine thinner than sexine; pollen wall tectate; sculpture scabrate in LM, perforate, rugulate/verrucate and fossulate in SEM; distal face perforate, perforations elliptic to slit-like, perforations often aligned in sinuous rows, rows of perforations outlining irregular shaped rugulae/ver- rucae (SEM); proximal face perforate and fossulate, perforations elliptic to slit-like, perforations often aligned in sinuous rows, rows of perforations and fossulae outlining irregular shaped rugulae/verrucae, sculpture becoming microrugulate to nanorugulate/ verrucate and perforate towards apices; opercula with nanoverrucate to granulate sublayer and perfo- rate supra-layer (SEM). ...
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... same two SEM micrograph are repeated in Harley and Baker (2001, figures 78 and 79). A single TEM showing the aperture region and operculum is presented in Harley and Drans- field (2003, figure 18). All previously illustrated pol- len grains belonging to this taxon (pollen Type C) were formerly assigned to Sclerosperma mannii (see Table IV). ...
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... previously illustrated pol- len grains belonging to this taxon (pollen Type C) were formerly assigned to Sclerosperma mannii (see Table IV). Figures 1D', 6; Table III) Description. -Pollen, monad, heteropolar, P/E ratio oblate, outline straight-triangular to slightly concave-triangular in polar view, bean-shaped in equatorial view (convex distal face versus concave proximal face); equatorial diameter 35-40 µm in LM, 30-35 µm in SEM, polar axis 15-19 µm in LM; triporate, pori positioned sub-apically on the distal polar face, pori elliptic, 5.0-8.0 µm in dia- meter, pori equipped with opercula; exine 1.7- 2.5 µm thick in LM, nexine thinner than sexine; pollen wall semitectate; sculpture reticulate in LM, reticulate to perforate in SEM; distal face reticulate with broad muri and elliptic to triangular to polygo- nal lumina, 16-25 lumina per 100 µm 2 at central distal face, 0-6 nanogemmae free-standing columel- lae per lumina (SEM); proximal face reticulate to perforate, lumina/perforations elliptic to triangular to polygonal, 0-6 nanogemmae free-standing colu- mellae per lumina; central polar areas and interaper- tural areas reticulate, sculpture becoming microreticulate to perforate towards apices; opercula with nanoverrucate to granulate sublayer and distinct microreticulate supra-layer (SEM). ...
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... -Two LM micrographs showing pollen of this taxon are provided by Sowunmi (1972, plate 3, figure 8 and plate 4, figure 1), but assigned to S. mannii (Table IV). ...
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... LM only, Sclerosperma pollen can be divided into reticulate (including S. manni, S. profizianum Type A, and S. walkeri) and non-reticulate (includ- ing S. profizianum Type B and C; Table III). The reticulate pollen are further divided into coarsely reticulate (including S. mannii and S. walkeri) versus finely reticulate (S. profizianum Type A; compare Figure 1A and 1Z with Figure 1F). Our measure- ments indicate the coarsely reticulate pollen of S. mannii and S. walkeri can be set apart using the length of their polar axis, which is longer in the pollen of S. walkeri (15-19 µm) than in S. mannii (9-15 µm). ...
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... LM only, Sclerosperma pollen can be divided into reticulate (including S. manni, S. profizianum Type A, and S. walkeri) and non-reticulate (includ- ing S. profizianum Type B and C; Table III). The reticulate pollen are further divided into coarsely reticulate (including S. mannii and S. walkeri) versus finely reticulate (S. profizianum Type A; compare Figure 1A and 1Z with Figure 1F). Our measure- ments indicate the coarsely reticulate pollen of S. mannii and S. walkeri can be set apart using the length of their polar axis, which is longer in the pollen of S. walkeri (15-19 µm) than in S. mannii (9-15 µm). ...
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... first LM and SEM micrographs showing pollen of this taxon are by Harley and Hall (1991, plate 4, figures 32 [SEM] and 33 [LM]). The same SEM micrograph is shown in Harley (1999, plate 1, figure 12), and again along with two addi- tional LM micrographs and two attached grains under SEM in Harley (1996, plate 16, figures C and F [SEM], and G and H [LM]). These are all repeated in Harley and Baker (2001, figures 77 and 82 [SEM], 80 and 81 [LM]). ...
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... are all repeated in Harley and Baker (2001, figures 77 and 82 [SEM], 80 and 81 [LM]). A new SEM detail is provided in Harley and Dransfield (2003, figure 11). In total, five or six grains were illustrated using either LM or SEM micrographs, but no TEM micrograph has been presented thus far. ...
Context 11
... C (Hall & Enti, GC36150 [K]; Profizi, 841 [K]) ( Figures 1P-Y, 5; Table III) Description. -Pollen, monad, heteropolar, P/E ratio oblate, outline straight-triangular to slightly convex-triangular in polar view, bean-shaped in equatorial view (convex distal face versus concave proximal face); equatorial diameter 37-42 µm in LM, 31-39 µm in SEM, polar axis 10-16 µm in LM; triporate, pori positioned sub-apically on the distal polar face, pori circular to elliptic, 4.0- 6.5 µm in diameter, pori equipped with opercula; exine 1.7-2.5 µm thick in LM, nexine thinner than sexine; pollen wall tectate; sculpture scabrate in LM, perforate, rugulate/verrucate and fossulate in SEM; distal face perforate, perforations elliptic to slit-like, perforations often aligned in sinuous rows, rows of perforations outlining irregular shaped rugulae/ver- rucae (SEM); proximal face perforate and fossulate, perforations elliptic to slit-like, perforations often aligned in sinuous rows, rows of perforations and fossulae outlining irregular shaped rugulae/verrucae, sculpture becoming microrugulate to nanorugulate/ verrucate and perforate towards apices; opercula with nanoverrucate to granulate sublayer and perfo- rate supra-layer (SEM). ...
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... same two SEM micrograph are repeated in Harley and Baker (2001, figures 78 and 79). A single TEM showing the aperture region and operculum is presented in Harley and Drans- field (2003, figure 18). All previously illustrated pol- len grains belonging to this taxon (pollen Type C) were formerly assigned to Sclerosperma mannii (see Table IV). ...
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... previously illustrated pol- len grains belonging to this taxon (pollen Type C) were formerly assigned to Sclerosperma mannii (see Table IV). Figures 1D', 6; Table III) Description. -Pollen, monad, heteropolar, P/E ratio oblate, outline straight-triangular to slightly concave-triangular in polar view, bean-shaped in equatorial view (convex distal face versus concave proximal face); equatorial diameter 35-40 µm in LM, 30-35 µm in SEM, polar axis 15-19 µm in LM; triporate, pori positioned sub-apically on the distal polar face, pori elliptic, 5.0-8.0 µm in dia- meter, pori equipped with opercula; exine 1.7- 2.5 µm thick in LM, nexine thinner than sexine; pollen wall semitectate; sculpture reticulate in LM, reticulate to perforate in SEM; distal face reticulate with broad muri and elliptic to triangular to polygo- nal lumina, 16-25 lumina per 100 µm 2 at central distal face, 0-6 nanogemmae free-standing columel- lae per lumina (SEM); proximal face reticulate to perforate, lumina/perforations elliptic to triangular to polygonal, 0-6 nanogemmae free-standing colu- mellae per lumina; central polar areas and interaper- tural areas reticulate, sculpture becoming microreticulate to perforate towards apices; opercula with nanoverrucate to granulate sublayer and distinct microreticulate supra-layer (SEM). ...
Context 14
... -Two LM micrographs showing pollen of this taxon are provided by Sowunmi (1972, plate 3, figure 8 and plate 4, figure 1), but assigned to S. mannii (Table IV). ...
Context 15
... LM only, Sclerosperma pollen can be divided into reticulate (including S. manni, S. profizianum Type A, and S. walkeri) and non-reticulate (includ- ing S. profizianum Type B and C; Table III). The reticulate pollen are further divided into coarsely reticulate (including S. mannii and S. walkeri) versus finely reticulate (S. profizianum Type A; compare Figure 1A and 1Z with Figure 1F). Our measure- ments indicate the coarsely reticulate pollen of S. mannii and S. walkeri can be set apart using the length of their polar axis, which is longer in the pollen of S. walkeri (15-19 µm) than in S. mannii (9-15 µm). ...
Context 16
... LM only, Sclerosperma pollen can be divided into reticulate (including S. manni, S. profizianum Type A, and S. walkeri) and non-reticulate (includ- ing S. profizianum Type B and C; Table III). The reticulate pollen are further divided into coarsely reticulate (including S. mannii and S. walkeri) versus finely reticulate (S. profizianum Type A; compare Figure 1A and 1Z with Figure 1F). Our measure- ments indicate the coarsely reticulate pollen of S. mannii and S. walkeri can be set apart using the length of their polar axis, which is longer in the pollen of S. walkeri (15-19 µm) than in S. mannii (9-15 µm). ...

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... The pollen morphology of Sclerosperma is not only unique within the Arecaceae but within all angiosperms, and its diagnostic features have been pinpointed in numerous publications (e.g., Erdtman and Sing, 1957;Harley and Hall, 1991;Harley, 1996Harley, , 1999Harley, , 2004Harley and Baker, 2001;Harley and Dransfield, 2003;Dransfield et al., 2008). The combined morphological features of extant Sclerosperma pollen observed with light microscopy (LM) and scanning electron microscopy (SEM) were conducted by Grímsson et al. (2019c). The authors showed that the three living Sclerosperma species produce four different pollen morphologies, with S. mannii and S. walkeri sharing a coarsely reticulate pollen and S. profizianum producing three pollen types that are either microreticulate or fossulate or perforate (Grímsson et al., 2019c). ...
... The combined morphological features of extant Sclerosperma pollen observed with light microscopy (LM) and scanning electron microscopy (SEM) were conducted by Grímsson et al. (2019c). The authors showed that the three living Sclerosperma species produce four different pollen morphologies, with S. mannii and S. walkeri sharing a coarsely reticulate pollen and S. profizianum producing three pollen types that are either microreticulate or fossulate or perforate (Grímsson et al., 2019c). Sclerosperma fossil records are rare and comprise only a handful of finds, mostly pollen, depicted from Late Oligocene to Miocene sediments of Africa (Grímsson et al., 2019b;Currano et al., 2020;Ulrich and Grímsson, 2020). ...
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... -The outline of this fossil pollen type in both polar and equatorial view, along with its reti- culate sculpture, distally placed apertures and reti- culate opercula, places it firmly within the extant genus Sclerosperma (Table I). Of the four pollen morphologies produced by extant Sclerosperma (see Grímsson et al. 2018), the S. protomannii sp. nov. ...
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Background Plants develop a variety of pollen morphological features during long-term evolution, which are controlled by genes and are not easily affected by the external natural environment. Therefore, pollen morphology has great significance in plant taxonomy, evolution and identification. However, there is no detailed study on the pollen morphology of the Gossypium genus although some cotton species have been investigated in the scattered reports. In this study, the pollen morphology of 23 cotton species was comprehensively examined using scanning electron microscopy to evaluate the pollen diversity of the genus and its taxonomic significance. Results The common characteristics of Gossypium pollen are spherical, radially symmetrical, echinate, panporate and operculate. The pollen diameter ranges from 62.43µm in G. harknessii to 103.41µm in G. barbadense, revealing that there are great variations in pollen size among cotton species. Moreover, the exine sculpture is echinate. The exine echini shape is mostly conical or sharply conical and occasionally rodlike. Echini density is found maximum in G. incanum (173) and minimum in G.gossypioides (54), meanwhile, echini length varies from 3.53 µm in G. herbaceum to 9.47 µm in G. barbadense. In addition, all cotton species are divided into three clusters based on cluster analysis, which provides new understanding of the evolution and phylogeny of the Gossypium genus. Conclusion Although the pollen characteristics alone are insufficient to reconstruct taxonomic and systematic relationships within the genus Gossypium, our findings can enrich our knowledge of sporopollen morphology and fill the phenological gap of these taxa and will contribute to future systematic and phylogenetic studies of the Gossypium genus.