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Anthetic anthers. Transverse section series from top, downwards. Morphological surfaces drawn with thick continuous lines; pollen sacs shown with thick broken lines; vascular bundles drawn with thin continuous lines and associated resin canals with thin broken lines. A-G, Burseraceae. A, Beiselia mexicana. B, Protium morii. C, Protium obtusifolium. D, Bursera sp. E, Commiphora caudata. F, Canarium caudatum. G, Santiria cf. apiculata. H-P, Anacardiaceae. Spondiadoideae. H, Dracontomelon dao. I, Spondias dulcis. J, Spondias purpurea. K, Pleiogynium solandri. L, Pseudospondias longifolia. M, Tapirira sp. N, Buchanania arborescens. O, Solenocarpus philippinensis. P, Campnosperma squamatum. Q-U, Anacardiaceae, Anacardioideae. Q, Anacardium occidentale. R, Mangifera indica. S, Blepharocarya involucrigera. T, Schinus molle. U, Semecarpus australiensis. Scale bars, 100 mm.
Source publication
Anacardiaceae and Burseraceae are traditionally distinguished by the number of ovules (1 vs. 2) per locule and the direction of ovule curvature (syntropous vs. antitropous). Recent molecular phylogenetic studies have shown that these families are sister groups in Sapindales after having been separated in different orders for a long time. We present...
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... All stamens have a thick round filament base, which narrows towards the constricted tip, and a sagittate anther (Fig. 43A). The anthers are dorsifixed in their lower half and have a thin and narrow connective and a deep dorsal and a ventral longitudinal furrow. Their dorsal side is broader than the ventral side and the anthers are thus introrse (Fig. 44A). The dehiscence lines extend from the tip of the thecae down to their base and encompass their lower shoulders (Fig. 43A′). In our material, the stamens of morphologically bisexual flowers are sterile and the flowers are thus function- ally female (Fig. 3). A lobed intrastaminal nectary disc is present. In female flowers it surrounds ...
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... antepetalous stamens appear slightly longer and larger than the antesepalous ones. All stamens have a broad flattened filament base, which becomes rounder and narrows towards the constricted tip and a slightly sagittate (P. morii, Fig. 44B) or X-shaped anther (P. obtusifolium, Fig. 43B, 44C). The anthers are basally dorsifixed (P. morii, Fig. 44B) or dorsifixed in the lower half (P. obtusifolium). They have a con- nective of medium (P. morii, Fig. 44B) thickness or a thin connective (P. obtusifolium, Figs 43B, 44C) and a dorsal and a deep ventral longitudinal furrow ...
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... antepetalous stamens appear slightly longer and larger than the antesepalous ones. All stamens have a broad flattened filament base, which becomes rounder and narrows towards the constricted tip and a slightly sagittate (P. morii, Fig. 44B) or X-shaped anther (P. obtusifolium, Fig. 43B, 44C). The anthers are basally dorsifixed (P. morii, Fig. 44B) or dorsifixed in the lower half (P. obtusifolium). They have a con- nective of medium (P. morii, Fig. 44B) thickness or a thin connective (P. obtusifolium, Figs 43B, 44C) and a dorsal and a deep ventral longitudinal furrow (Fig. 44B, C). Their dorsal side is broader than the ventral side and the anthers are slightly introrse (Fig. ...
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... a broad flattened filament base, which becomes rounder and narrows towards the constricted tip and a slightly sagittate (P. morii, Fig. 44B) or X-shaped anther (P. obtusifolium, Fig. 43B, 44C). The anthers are basally dorsifixed (P. morii, Fig. 44B) or dorsifixed in the lower half (P. obtusifolium). They have a con- nective of medium (P. morii, Fig. 44B) thickness or a thin connective (P. obtusifolium, Figs 43B, 44C) and a dorsal and a deep ventral longitudinal furrow (Fig. 44B, C). Their dorsal side is broader than the ventral side and the anthers are slightly introrse (Fig. 44B, C). The dehiscence lines extend from the tip of the thecae down to their base (Fig. 43B′). In our ...
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... morii, Fig. 44B) or X-shaped anther (P. obtusifolium, Fig. 43B, 44C). The anthers are basally dorsifixed (P. morii, Fig. 44B) or dorsifixed in the lower half (P. obtusifolium). They have a con- nective of medium (P. morii, Fig. 44B) thickness or a thin connective (P. obtusifolium, Figs 43B, 44C) and a dorsal and a deep ventral longitudinal furrow (Fig. 44B, C). Their dorsal side is broader than the ventral side and the anthers are slightly introrse (Fig. 44B, C). The dehiscence lines extend from the tip of the thecae down to their base (Fig. 43B′). In our material of P. morii, the anthers are devoid of pollen and the flowers are functionally female and, in that of P. obtusifolium, the ...
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... (P. morii, Fig. 44B) or dorsifixed in the lower half (P. obtusifolium). They have a con- nective of medium (P. morii, Fig. 44B) thickness or a thin connective (P. obtusifolium, Figs 43B, 44C) and a dorsal and a deep ventral longitudinal furrow (Fig. 44B, C). Their dorsal side is broader than the ventral side and the anthers are slightly introrse (Fig. 44B, C). The dehiscence lines extend from the tip of the thecae down to their base (Fig. 43B′). In our material of P. morii, the anthers are devoid of pollen and the flowers are functionally female and, in that of P. obtusifolium, the gynoecium is more or less reduced and the flowers functionally male. A thick lobed intrastaminal nectary disc ...
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... stamens have a broad flattened filament base, which narrows towards the constricted tip, and a sagittate anther (Figs 7G′, H′, 44D). The anthers are basally dorsifixed and the attachment zone is almost hidden in a pseudopit (Fig. 44D). ...
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... stamens have a broad flattened filament base, which narrows towards the constricted tip, and a sagittate anther (Figs 7G′, H′, 44D). The anthers are basally dorsifixed and the attachment zone is almost hidden in a pseudopit (Fig. 44D). They may be ver- satile. They have a thin and narrow connective and a dorsal and a ventral median longitudinal furrow (Fig. 44D). Their dorsal side is broader than the ventral side and the anthers are slightly introrse. The dehiscence lines extend from the tip of the thecae down to their base. A lobed intrast- aminal nectary disc is ...
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... have a broad flattened filament base, which narrows towards the constricted tip, and a sagittate anther (Figs 7G′, H′, 44D). The anthers are basally dorsifixed and the attachment zone is almost hidden in a pseudopit (Fig. 44D). They may be ver- satile. They have a thin and narrow connective and a dorsal and a ventral median longitudinal furrow (Fig. 44D). Their dorsal side is broader than the ventral side and the anthers are slightly introrse. The dehiscence lines extend from the tip of the thecae down to their base. A lobed intrast- aminal nectary disc is present on the floral cup ( Fig. ...
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... in the floral base ( Fig. 7G′-M′). Petals have a median vascular bundle extending along their entire length and asso- ciated at the base with one to two smaller lateral bundles, which extend downwards into the floral cup as two or three traces (Fig. 7G′-M′). Stamens have a single vascular bundle with the resin canal distinctive only in the anther (Fig. 44). In the carpels, a median dorsal vascular bundle is present below the stigmatic region ( Fig. 7C-E). Further down, the dorsal bundle splits into two branches (Fig. 7F). The inner portions of those bundles form synlaterals, whereas the peripheral portions form a dorsal arc of bundles ( Fig. 7G-J). The synlaterals are unusual in that ...
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... which becomes rounder and narrows towards the non-constricted tip, and a sag- ittate anther (Fig. 43C, D). The anthers are dorsifixed in the lower half with a broad and thick connective and only a ventral longitudinal furrow (Figs 43C, D, 44E). The dorsal side of the thecae is longer and broader than the ventral side and the anthers are introrse (Fig. 44E). The dehiscence lines extend from the tips of the thecae almost down to the base (Fig. 43C′, D′). In our material, either male or female organs are sterile and the flowers are thus function- ally unisexual. Within the floral cup and calyx tube, the instrastaminal nectary disc is wider in function- ally male flowers than in female ...
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... smaller lateral bundle ( Fig. 8B-D). By repetition of this process, a petal can have up to 11 bundles. Stamens have a single vascular bundle with a resin canal only distinctive in filaments of function- ally female flowers and less so in functionally male flowers. In the distal part of the anthers, the vascular bundle splits into two branches (Fig. 44E). In the carpels, a median vascular bundle and two slightly shorter lateral bundles are present in the stigmatic region (Fig. 9C, D). Toward the base of the style, the number of bundles increases rapidly (Fig. 9E, ...
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... stamens have a broad flattened filament base, which narrows slightly toward the non-constricted tip, and an apiculate and slightly sagittate anther (Fig. 44F). The anthers are basally dorsifixed and have a broad and thick connective (Fig. 44F). The dorsal side of the thecae is much larger and slightly longer than the ventral side and the anthers are introrse (Fig. 44F). The dehiscence lines extend from below the tip of the thecae down to their base (Fig. 44F). In our material, the anthetic ...
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... stamens have a broad flattened filament base, which narrows slightly toward the non-constricted tip, and an apiculate and slightly sagittate anther (Fig. 44F). The anthers are basally dorsifixed and have a broad and thick connective (Fig. 44F). The dorsal side of the thecae is much larger and slightly longer than the ventral side and the anthers are introrse (Fig. 44F). The dehiscence lines extend from below the tip of the thecae down to their base (Fig. 44F). In our material, the anthetic anthers are devoid of pollen and the flowers are thus functionally female. A distinct ...
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... a broad flattened filament base, which narrows slightly toward the non-constricted tip, and an apiculate and slightly sagittate anther (Fig. 44F). The anthers are basally dorsifixed and have a broad and thick connective (Fig. 44F). The dorsal side of the thecae is much larger and slightly longer than the ventral side and the anthers are introrse (Fig. 44F). The dehiscence lines extend from below the tip of the thecae down to their base (Fig. 44F). In our material, the anthetic anthers are devoid of pollen and the flowers are thus functionally female. A distinct intrastaminal nectary disc is lacking but the lobes of the stamen ring are nectarif- erous and alternate with the filaments. ...
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... an apiculate and slightly sagittate anther (Fig. 44F). The anthers are basally dorsifixed and have a broad and thick connective (Fig. 44F). The dorsal side of the thecae is much larger and slightly longer than the ventral side and the anthers are introrse (Fig. 44F). The dehiscence lines extend from below the tip of the thecae down to their base (Fig. 44F). In our material, the anthetic anthers are devoid of pollen and the flowers are thus functionally female. A distinct intrastaminal nectary disc is lacking but the lobes of the stamen ring are nectarif- erous and alternate with the filaments. The lobes are asymmetrical and more developed adjacent to the antesepalous ...
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... and a sepal can have up to nine vascular bundles (Fig. 10B, C). The median vascular trace extends downwards in the floral cup while the lateral ones tend to merge into synlaterals ( Fig. 10D, E). Each petal has a col- laterally double median vascular trace in the floral cup, which merges lower down with the synlateral traces of the sepals ( (Fig. 44F). In the carpels, only a median vascular bundle is present in the stigmas ( Fig. 11C-E). Down- wards, the number of vascular bundles of various size and state of differentiation increases and they form a dense network around the locules ( Fig. 11G- N). A regular pattern is difficult to recognize. Syn- lateral bundles are regularly ...
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... (Fig. 45M) and merging lower down with the petal bases and the floral cup. The antepetalous stamens are slightly shorter and smaller than the antesepalous ones at all stages of development (Figs 12C, D, 45M). All stamens have a broad flattened filament base, which narrows slightly towards the non-constricted tip, and a slightly sagittate anther (Fig. 44G). The anthers are basally dorsifixed and have a broad and thick con- nective and a ventral median longitudinal furrow (Fig. 44G). The dorsal side of the thecae is longer and larger than the ventral one and the anthers are introrse (Fig. 44G). The dehiscence lines extend from the tip of the thecae almost down to their base. In our ...
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... than the antesepalous ones at all stages of development (Figs 12C, D, 45M). All stamens have a broad flattened filament base, which narrows slightly towards the non-constricted tip, and a slightly sagittate anther (Fig. 44G). The anthers are basally dorsifixed and have a broad and thick con- nective and a ventral median longitudinal furrow (Fig. 44G). The dorsal side of the thecae is longer and larger than the ventral one and the anthers are introrse (Fig. 44G). The dehiscence lines extend from the tip of the thecae almost down to their base. In our material, the anthers are devoid of pollen and the flowers are thus functionally female. A nectary covers the inner surface of the ...
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... filament base, which narrows slightly towards the non-constricted tip, and a slightly sagittate anther (Fig. 44G). The anthers are basally dorsifixed and have a broad and thick con- nective and a ventral median longitudinal furrow (Fig. 44G). The dorsal side of the thecae is longer and larger than the ventral one and the anthers are introrse (Fig. 44G). The dehiscence lines extend from the tip of the thecae almost down to their base. In our material, the anthers are devoid of pollen and the flowers are thus functionally female. A nectary covers the inner surface of the floral cup and base of stami- nal ring but does not form a conspicuous ...
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... Vascular bundles have a more or less devel- oped resin canal on the dorsal side of the xylem, except for the ovule bundle ( Figs 12, 13). In the syn- sepalous region, sepals have a median vascular bundle and two smaller lateral ones ( (Fig. 12B-D) and, thus, each petal can have up to nine vascular bundles. Stamens have a single vascular bundle (Fig. 44G). In the carpels, there is no median vascular bundle in the stigmatic region. However, two lateral vascular bundles dif- ferentiate below the stigmatic head ( Fig. 13D-F), whereas a median dorsal vascular bundle only appears toward the base of the style (Fig. 13G). Lower down, the peripheral portions of the lateral bundles FLORAL ...
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... narrows slightly towards the constricted tip, and a slightly X-shaped anther ( Figs 43E, 44H). The anthers are dorsifixed below mid-length and have a thick and broad connec- tive and a shallow ventral median furrow (Figs 43E, 44H). Their dorsal side is slightly larger than the ventral side and the anthers are slightly introrse, almost latrorse (Fig. 44H). The dehiscence lines extend from the tip of the thecae down to their base (Figs 43E′). In our material, the stamens are devoid of pollen in some buds and these flowers are thus func- tionally female. A flat intrastaminal nectary disc is present and has erect lobes, which extend between the filaments (Fig. ...
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... side of the median vascular bundle with the main laterals, which do not form synlaterals in the floral base (Fig. 14E). Petals have a single median vascular bundle, which extends along their entire length, and four large additional lateral bundles, plus one or two smaller bundles toward the margins (Fig. 14). Stamens have a single vascular bundle (Fig. 44H). In the carpels, small lateral bundles differentiate on each side of the ventral slit in the stigmatic region. Downwards they are quickly fol- lowed by a median dorsal bundle, which becomes the largest in the styles (Fig. 15C-E). Towards the ovary, a small median ventral vascular bundle is present (Fig. 15F, G). In the synascidiate ...
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... tip, and an X-shaped anther (Figs 43F, 44I, J). The anthers are dorsifixed in the lower half (S. dulcis, (Fig. 43F) or at the base FLORAL STRUCTURE IN ANACARDIACEAE AND BURSERACEAE 529 (S. purpurea) and they have a relatively broad and thick connective and a shallow dorsal and a ventral (only ventral in S. purpurea) longitudinal furrow (Fig. 44I, J). Their dorsal side is broader (and longer in S. purpurea) than the ventral side and they are slightly introrse (Fig. 44I, J). The dehiscence lines extend from the tip of the thecae almost down to their base (Fig. 43F′). In our material the anthers are devoid of pollen in S. dulcis and S. purpurea and the flowers are thus functionally ...
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... or at the base FLORAL STRUCTURE IN ANACARDIACEAE AND BURSERACEAE 529 (S. purpurea) and they have a relatively broad and thick connective and a shallow dorsal and a ventral (only ventral in S. purpurea) longitudinal furrow (Fig. 44I, J). Their dorsal side is broader (and longer in S. purpurea) than the ventral side and they are slightly introrse (Fig. 44I, J). The dehiscence lines extend from the tip of the thecae almost down to their base (Fig. 43F′). In our material the anthers are devoid of pollen in S. dulcis and S. purpurea and the flowers are thus functionally female. A thick lobed intrastaminal nectary disc is present and surrounds the lower half of the ovary (Fig. 41D, ...
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... vascular bundle, which further up gives off two smaller lateral bundles (Figs 16C, D, 18C, D). Additional smaller vascular bundles connect with the median and lateral bundles and a petal can have up to seven vascular bundles in S. dulcis (Fig. 16A, B) and only up to four to five in S. purpurea (Fig. 18A, B). Stamens have a single vascular bundle (Fig. 44I, J). In the carpels, a dorsal vascular bundle differentiates below the stigmas and extends down- wards into the floral base (Figs 17D-L, 19B-L). It is followed in the style by two lateral bundles (Figs 17D-H, 19C-F). Branches of the lateral bundles form synlaterals in the upper ovary and converge towards its centre, whereas the peripheral ...
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... antepetalous stamens are shorter than the antesepalous ones at all stages of development and their anthers are apiculate vs. non-apiculate ( Figs 41I, J, 43G, H). All stamens have a broad flat- tened filament base, which further up becomes rounder and narrows towards the constricted tip, and a sagittate anther (Fig. 44K). The anthers are dorsi- fixed in the lower half (Fig. 43G, H). The connective is intermediate in thickness and the antepetalous anthers have a longitudinal furrow on the ventral side (Fig. 44K). In antesepalous stamens the dorsal parts of the thecae curve backwards and form a pseudopit around the filament tip (Fig. 44K). The dorsal ...
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... H). All stamens have a broad flat- tened filament base, which further up becomes rounder and narrows towards the constricted tip, and a sagittate anther (Fig. 44K). The anthers are dorsi- fixed in the lower half (Fig. 43G, H). The connective is intermediate in thickness and the antepetalous anthers have a longitudinal furrow on the ventral side (Fig. 44K). In antesepalous stamens the dorsal parts of the thecae curve backwards and form a pseudopit around the filament tip (Fig. 44K). The dorsal side of the anthers is broader and longer than the ventral side and they are thus introrse (Fig. 44K). The dehiscence lines extend from the tip of the thecae almost down to their base (Fig. 43G′, ...
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... and a sagittate anther (Fig. 44K). The anthers are dorsi- fixed in the lower half (Fig. 43G, H). The connective is intermediate in thickness and the antepetalous anthers have a longitudinal furrow on the ventral side (Fig. 44K). In antesepalous stamens the dorsal parts of the thecae curve backwards and form a pseudopit around the filament tip (Fig. 44K). The dorsal side of the anthers is broader and longer than the ventral side and they are thus introrse (Fig. 44K). The dehiscence lines extend from the tip of the thecae almost down to their base (Fig. 43G′, H′). A thick lobed intrastaminal nectary disc is present and sur- rounds the base of the superior ...
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... is intermediate in thickness and the antepetalous anthers have a longitudinal furrow on the ventral side (Fig. 44K). In antesepalous stamens the dorsal parts of the thecae curve backwards and form a pseudopit around the filament tip (Fig. 44K). The dorsal side of the anthers is broader and longer than the ventral side and they are thus introrse (Fig. 44K). The dehiscence lines extend from the tip of the thecae almost down to their base (Fig. 43G′, H′). A thick lobed intrastaminal nectary disc is present and sur- rounds the base of the superior ...
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... entire length and they merge in the floral base (Fig. 20B, C). Further up, each lateral bundle branches, forming an additional smaller lateral bundle toward the petal margin and, by repetition of this process, the petals can have up to nine vascular bundles. Stamens have a single vascular bundle, which in the anther comprises a large resin canal (Fig. 44K). In the carpels, the dorsal vasculature is poorly differentiated in the ovary wall (Fig. 21G-J), whereas, below the stigma, a median dorsal vascular bundle is present and branches, forming a lateral smaller bundle on each side of the ventral slit towards the base of the short style (Fig. 21B-D). In the upper end of the synascidiate ...
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... becomes rounder and narrows toward the constricted tip, and a slightly X-shaped anther (Figs 43I, 44L). The anthers are dorsifixed at mid-length of the thecae and have a thin and narrow connective and a dorsal and a deep ventral longitu- dinal furrow (Figs 43I, 44L). Their dorsal side is larger than the ventral side and they are slightly introrse (Fig. 44L). The dehiscence lines extend from the tip of the thecae down to their base (Fig. 43I′). A thick lobed intrastaminal nectary disc is present and surrounds the base of the superior ...
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... stamens have a broad and thick filament base, which becomes rounder and narrows toward the con- stricted tip, and a sagittate anther with a short con- nective tip (Figs 43J, 44M). The anthers are basally dorsifixed and have a thin and narrow connective and a dorsal and a deep ventral longitudinal furrow (Fig. 44M). They are slightly introrse and the dehis- cence lines extend from the tip of the thecae down to their base (Fig. 43J′). A thick lobed intrastaminal nectary disc is present and surrounds the base of the superior ovary (Fig. ...
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... have a flattened filament, which narrows toward the constricted tip, and a sagittate anther (Figs 24D, 41Q). The anthers are basifixed and may be versatile (Fig. 41S) and the attachment zone is almost hidden in a pseudopit (Figs 41Q, 44N). The connective is intermediate in thickness and there is a deep dorsal and ventral longitudinal furrow (Fig. 44N). The pollen sacs do not extend into the free basal parts of the thecae (Fig. 44N). The anthers are almost latrorse and the dehiscence lines extend from the tip of the thecae down to where the pollen sacs end (Fig. 44N). In our material, most anthers are devoid of pollen and the flowers are functionally female. A thick lobed ...
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... anther (Figs 24D, 41Q). The anthers are basifixed and may be versatile (Fig. 41S) and the attachment zone is almost hidden in a pseudopit (Figs 41Q, 44N). The connective is intermediate in thickness and there is a deep dorsal and ventral longitudinal furrow (Fig. 44N). The pollen sacs do not extend into the free basal parts of the thecae (Fig. 44N). The anthers are almost latrorse and the dehiscence lines extend from the tip of the thecae down to where the pollen sacs end (Fig. 44N). In our material, most anthers are devoid of pollen and the flowers are functionally female. A thick lobed intrastaminal nectary disc is present and is much thicker toward the centre than at the ...
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... pseudopit (Figs 41Q, 44N). The connective is intermediate in thickness and there is a deep dorsal and ventral longitudinal furrow (Fig. 44N). The pollen sacs do not extend into the free basal parts of the thecae (Fig. 44N). The anthers are almost latrorse and the dehiscence lines extend from the tip of the thecae down to where the pollen sacs end (Fig. 44N). In our material, most anthers are devoid of pollen and the flowers are functionally female. A thick lobed intrastaminal nectary disc is present and is much thicker toward the centre than at the periphery so that only the connivent styles (more or less reflexed distally) and stigmas are visible of the gynoecium (Fig. ...
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... rounder further up and narrows towards the constricted attachment zone, and a sagittate anther. The anthers are dorsifixed in the lower half of the thecae and may be versatile and caducous. The connective is extremely broad and thick and the dorsal side of the thecae curves back- ward and almost forms a pseudopit around the attachment zone (Fig. 44O). In our material, in late anthetic flowers all anthers were fallen and in the single bud we could study they were latrorse (slightly introrse), with longitudinal dehiscence lines extending from the tip of the thecae down to their base (Fig. 26A, B). A thick lobed intrastaminal nectary disc surrounds the base of the superior ovary (Fig. ...
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... have a median and two lateral main vascular traces, which extend along their entire length, and one or two smaller lateral bundles intercalated between the three main ones, which merge basally with the median vascular bundle (Fig. 26A-D). Stamens have a single vascular bundle which splits into two smaller bundles in the upper half of the anther (Fig. 44O). In the carpel, a dorsal vascular bundle with a conspicu- ous resin canal extends from the stigma down to the ovary base (Fig. 27C-R). Smaller lateral vascular bundles are also present below the stigma and they merge to form a single lateral bundle on each side of the ventral slit ( Fig. 27C-I). At the transition from the style to the ...
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... more arrowhead-shaped anthers (Fig. 42B). All stamens have a large and broad filament base, which narrows slightly toward the attachment zone, but are not constricted, and have a slightly X-shaped anther (Figs 42B, 43K, 44P). The anthers are basally dorsi- fixed with a connective of intermediate thickness and a deep ventral longitudinal furrow (Fig. 44P). Their dorsal side is larger than the ventral side and they are introrse (Fig. 44P). The dehiscence lines extend from the tip of the thecae almost down to their base (Fig. 43K′). Anthers of functionally female flowers are devoid of pollen. A thick and lobed intrastaminal nectary disc is present and surrounds the base of the superior ...
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... base, which narrows slightly toward the attachment zone, but are not constricted, and have a slightly X-shaped anther (Figs 42B, 43K, 44P). The anthers are basally dorsi- fixed with a connective of intermediate thickness and a deep ventral longitudinal furrow (Fig. 44P). Their dorsal side is larger than the ventral side and they are introrse (Fig. 44P). The dehiscence lines extend from the tip of the thecae almost down to their base (Fig. 43K′). Anthers of functionally female flowers are devoid of pollen. A thick and lobed intrastaminal nectary disc is present and surrounds the base of the superior ovary (Fig. ...
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... (Fig. 30B), whereas later this difference is no longer present (Fig. 47A). The filaments are round and often have a distal annular thickening just below the sagittate and slightly apiculate anthers (Figs 43L, 44Q). The basal extensions of the thecae are sterile. The anthers are basally dorsifixed (Fig. 43L) and have a broad and thick connective (Fig. 44Q). Their dorsal side is slightly broader than the ventral side and curved backwards around the constricted attachment, which is almost hidden in a pseudopit (Fig. 44Q). The anthers are slightly introrse, almost latrorse and the dehiscence lines extend from the tip of the thecae almost down to where the pollen sac ends (Figs 43L′, 44Q). ...
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... and slightly apiculate anthers (Figs 43L, 44Q). The basal extensions of the thecae are sterile. The anthers are basally dorsifixed (Fig. 43L) and have a broad and thick connective (Fig. 44Q). Their dorsal side is slightly broader than the ventral side and curved backwards around the constricted attachment, which is almost hidden in a pseudopit (Fig. 44Q). The anthers are slightly introrse, almost latrorse and the dehiscence lines extend from the tip of the thecae almost down to where the pollen sac ends (Figs 43L′, 44Q). When the thecae dehisce, the anther tilts and then seems to sit transversally on the filament (Fig. 43L′). Later, it abscises above the annular thick- ening of the ...
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... synlaterals (Fig. 30D). Petals have a median and two main lateral vascular bundles, which extend almost along their entire length (Fig. 30A-C). They fuse at the base of the petal (Fig. 30D). Additional smaller lateral bundles toward the petal margins are also present. A petal can have up to seven bundles. Stamens have a single vascular bundle (Fig. 44Q). In the carpel, a large dorsal and two smaller main lateral vascular bundles are present below the stigma (Fig. 31C). Additional lateral bundles appear lower down (Fig. 31D-F) and, in the upper end of the ovary, the dorsal and lateral bundles form an arc of vasculature on the dorsal side of the carpel (Fig. 31G). In the ovary wall, the ...
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... the constricted tip (Fig. 43M). The anther is slightly X-shaped. It is dorsifixed at mid-length and FLORAL STRUCTURE IN ANACARDIACEAE AND BURSERACEAE 537 the attachment zone is hidden in a pseudopit formed by the dorsal side of the thecae (Figs 43M, 44R). It has a broad and thick connective and a deep ventral and a dorsal longitudinal furrow (Fig. 44R). The dorsal side is larger than the ventral side and the anther is introrse (Fig. 44R). The dehiscence lines extend from the tip of the thecae down to the base furrow (Fig. 43M′). An intrastaminal nectary disc is ...
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... at mid-length and FLORAL STRUCTURE IN ANACARDIACEAE AND BURSERACEAE 537 the attachment zone is hidden in a pseudopit formed by the dorsal side of the thecae (Figs 43M, 44R). It has a broad and thick connective and a deep ventral and a dorsal longitudinal furrow (Fig. 44R). The dorsal side is larger than the ventral side and the anther is introrse (Fig. 44R). The dehiscence lines extend from the tip of the thecae down to the base furrow (Fig. 43M′). An intrastaminal nectary disc is ...
Context 46
... which extend almost along their entire length (Fig. 32C). Additional smaller lateral bundles are present and petals may have up to eight bundles in the distal part (Fig. 32B). The main lateral bundles merge with the median bundle only in the floral base, where there is a single vascular trace (Fig. 33D, E). Stamens have a single vascular bundle (Fig. 44R). In the carpel, a massive lateral vascular bundle is present below the stigma on each side of the ventral slit ( Fig. 33D-I). The dorsal bundle is weak and the lateral bundles extend into the ventral side of the ovary wall. The weak dorsal vascular bundle extends downwards into the base of the style (Fig. 33E-I); in the ovary it is ...
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... stamens are shorter than the antesepalous ones at all stages of development. All stamens have a broad and thick filament base, which narrows slightly toward the constricted tip, and an X-shaped anther (Fig. 43N). The anthers are basally dorsifixed and have a thin and narrow connective and a deep ventral and a dorsal longitudinal furrow (Fig. 44S). Their dorsal side is larger than the ventral side and the anthers are slightly introrse (Fig. 44S). The dehiscence lines extend from the tip of the thecae almost down to their bases (Fig. 43N′). In female flowers, remnants of highly reduced stamens are sometimes present on the petal radii. A thick lobed intrastaminal nectary disc is ...
Context 48
... have a broad and thick filament base, which narrows slightly toward the constricted tip, and an X-shaped anther (Fig. 43N). The anthers are basally dorsifixed and have a thin and narrow connective and a deep ventral and a dorsal longitudinal furrow (Fig. 44S). Their dorsal side is larger than the ventral side and the anthers are slightly introrse (Fig. 44S). The dehiscence lines extend from the tip of the thecae almost down to their bases (Fig. 43N′). In female flowers, remnants of highly reduced stamens are sometimes present on the petal radii. A thick lobed intrastaminal nectary disc is present in bisexual flowers, whereas in female flowers it is smaller and is located between the petal ...
Context 49
... and, depending of their position, a shorter lateral bundle, which merges with the median one at the base (Fig. 34). Petals have a median and two main lateral vascular bundles, which extend almost along their entire length, and one trace. Stamens have a single vascular bundle, which may split into two branches in the distal part of the anther (Fig. 44S). In the carpel, a dorsal vascular bundle extends from the stigma to the ovary (Fig. 35B-K). In the lower part of the style, two shorter lateral bundles differentiate on each side of the ventral slit (Fig. 35E). In the upper end of the ovary, the laterals extend toward the ventral side of the carpel (Fig. 35F). Below the placenta they ...
Context 50
... stamens throughout development. All stamens have a broad flattened filament base, which becomes rounder and narrower towards the con- stricted tip, and a slightly X-shaped anther (Figs 43O, 44T). The anthers are basally dorsifixed and may be versatile. They have a thin and narrow connective and a deep ventral and a deep dorsal longitudinal furrow (Fig. 44T). Their dorsal side is not much larger than the ventral side and the anthers are almost latrorse. Distally they are slightly introrse (Fig. 44T). The dehiscence lines extend from the tip of the thecae down to their bases and encompass the lower shoulders (Fig. 43O′). In our material, either male or female organs are sterile and the ...
Context 51
... and a slightly X-shaped anther (Figs 43O, 44T). The anthers are basally dorsifixed and may be versatile. They have a thin and narrow connective and a deep ventral and a deep dorsal longitudinal furrow (Fig. 44T). Their dorsal side is not much larger than the ventral side and the anthers are almost latrorse. Distally they are slightly introrse (Fig. 44T). The dehiscence lines extend from the tip of the thecae down to their bases and encompass the lower shoulders (Fig. 43O′). In our material, either male or female organs are sterile and the flowers are thus functionally unisexual. A thick lobed intrastaminal nectary disc surrounds the base of the ...
Context 52
... bundles, which form synlaterals in the floral base ( Fig. 36C- E). Petals also have a single median vascular bundle at the base (Fig. 36C). Further up, it gives off smaller lateral bundles, which may also branch and form additional lateral vascular bundles (Fig. 36A, B). A petal can have up to ten bundles. Stamens have a single vascular bundle (Fig. 44T). In the carpels, two lateral vascular bundles are present below the stigma (Fig. 37C) and, only in the fertile carpel, a dorsal bundle differentiates lower down (Fig. 37D). In the upper part of the style, the laterals of each sterile carpel merge over their ventral side and form an arc of vasculature (Fig. 37E). These two arcs extend ...
Context 53
... (Fig. 42K, L). The stamens have a flattened filament base, which becomes rounder and narrows towards the constricted tip, and a pronouncedly sagittate anther (Figs 43P, 44U). The anthers are dorsifixed at mid-length (Fig. 43P). They may be versatile and are caducous. They have a thin and narrow connective and a deep ventral longitudinal furrow (Fig. 44U). Their dorsal side is distally larger than the ventral side and the anthers are introrse (Fig. 44U). The dehiscence lines extend from the tip of the thecae almost down to their bases and do not encompass the lower shoulders (Fig. 43U). In our material, the flowers are function- ally unisexual. An intrastaminal nectary disc is present ...
Context 54
... the constricted tip, and a pronouncedly sagittate anther (Figs 43P, 44U). The anthers are dorsifixed at mid-length (Fig. 43P). They may be versatile and are caducous. They have a thin and narrow connective and a deep ventral longitudinal furrow (Fig. 44U). Their dorsal side is distally larger than the ventral side and the anthers are introrse (Fig. 44U). The dehiscence lines extend from the tip of the thecae almost down to their bases and do not encompass the lower shoulders (Fig. 43U). In our material, the flowers are function- ally unisexual. An intrastaminal nectary disc is present in all flowers and covers the base of the ...
Context 55
... AND BURSERACEAE 541 entire length, and up to 35 lateral vascular bundles (Fig. 38A, B). In the petal base, the lateral vascular bundles merge successively with the median vascular bundle and they form a single trace in the floral base (Fig. 38C, D). Stamens have a single vascular bundle, which bifurcates in the distal region of the anthers (Fig. 44U). In the carpels, there is a dorsal vascular bundle without a resin canal below the stigma and two lateral bundles with a conspicuous resin canal (Fig. 39B). In the short symplicate zone, the lateral bundles of the fertile carpel and the adjacent lateral bundles of the sterile carpels form synlaterals (Fig. 39C, D), whereas no ...
Context 56
... extreme types of stamen shape with a range of intermediate conditions in both families. In type 1, the anther tends to be shorter than the fila- ment, versatile, caducous and (almost) latrorse (or slightly introrse). The filament tip is constricted and the transition zone from filament to anther extends shortly into a narrow and thin connective (Fig. 44C, D, L, M). In type 2, the filament tends to be shorter and stouter than in type 1, the anther tends to be longer than the filament and is markedly introrse (Fig. 44E- G). The attachment is not constricted and the transi- tion zone from filament to anther extends into a thicker and broader connective. Such anthers tend to be apiculate. ...
Context 57
... and (almost) latrorse (or slightly introrse). The filament tip is constricted and the transition zone from filament to anther extends shortly into a narrow and thin connective (Fig. 44C, D, L, M). In type 2, the filament tends to be shorter and stouter than in type 1, the anther tends to be longer than the filament and is markedly introrse (Fig. 44E- G). The attachment is not constricted and the transi- tion zone from filament to anther extends into a thicker and broader connective. Such anthers tend to be apiculate. Type 1 is most frequently observed in Spondiadoideae (and some Burseraceae: Beiselia, Bursera and Protium), but in core Burseraceae, especially in Canarieae, type 2 is ...
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Citations
... Further studies may reveal whether this tubular corolla involves partial congenital union or only postgenital coherence by petal margins, as described in several anthetic flowers of Galipeineae (Rutaceae, El Ottra et al. 2013). Elsewhere in Sapindales, connected petal margins are reported for Anacardiaceae, Burseraceae and Nitrariaceae but only at the bud stage (Bachelier and Endress 2009;Bachelier et al. 2011). In addition, synorganization of stamens evolved in some Simaroubaceae (see Sect. 4.5 Androecium). ...
Floral features contribute with remarkable additions to morphological studies and are widely used to address questions about the evolution and diversification of several groups of plants. Even though Simaroubaceae are a small monophyletic family, the few detailed structural analyses of reproductive organs and the floral diversity and variations already described in their members stimulate novel structural studies. In this study, we investigate the evolution of reproductive features of Simaroubaceae by means of a combination of original data and a review of the literature, aiming to elucidate which floral characters are most informative for a better understanding of the evolutionary history of the group. We analyzed 21 out of the 23 genera of Simaroubaceae, plus six from Rutaceae and seven from Meliaceae as outgroups. We used a Bayesian method and the Parsimony optimality criterion to reconstruct ancestral reproductive character states using a re-analyzed phylogenetic tree of Sapindales. Here, we combined available molecular sequences to have the largest sample of Simaroubaceae genera. We found that the ancestral flowers of Simaroubaceae were probably polygamous or dioecious plants, with free carpels united only distally, with divergent, elongated stigmas, and with drupaceous, laterally flattened to lenticular fruits. The latter feature plus apocarpous carpels are putative synapomorphies of the family retrieved in this study. Imbricate petals and a diplostemonous androecium were recovered as conditions found in the ancestor of Simaroubaceae but also shared with the ancestors of Meliaceae and Rutaceae. Our findings were mostly in accordance with previous evolutionary studies on genera of Simaroubaceae and with other families of Sapindales. © 2021, The Author(s), under exclusive licence to Botanical Society of Sao Paulo.
... This is supported by recent findings of scattered flowers bearing abortive ovules in some species of these two genera (Franceschinelli and Thomas 2000;Alves et al. 2017;Devecchi et al. 2018a, b). Flowers that are morphologically perfect but functionally unisexual are reported also to some extra-American simaroubaceous genera, as well as in many other groups of Sapindales [e.g., Meliaceae (Styles 1972;Franceschinelli et al. 2015), Anacardiaceae and Burseraceae (Bachelier and Endress 2009), Sapindaceae (Avalos et al. 2019) and Rutaceae (Kubitzki et al. 2011)]. Evolutionary paths of sexual structures and systems in Simaroubaceae and related families are discussed in Alves et al. (2021-this issue) and in Gama et al. (2021). ...
Simaroubaceae are among the families whose circumscription radically changed over time, because phylogenetic analyses undertaken since 1995 demonstrated it was a polyphyletic group in its traditional delimitation. Currently, Simaroubaceae sensu stricto are a mostly pantropical, highly supported monophyletic group composed of 22 genera and approximately 120 species. Growing knowledge about members of the family has allowed several advances over the last couple of decades. The primary center of diversity for Simaroubaceae is in tropical America, and new contributions have been recently made regarding members of the family in the region, including descriptions of several new taxa. Hence, we undertook an updated overview of general information available for the group, with focus on American taxa of Simaroubaceae, and highlighting numerous data published after the 2011 monograph. Besides aiming to contribute to a better knowledge of a family with past controversial limits, we emphasize research topics in which the current scarcity of data demands further investigation.
... The family Anacardiaceae belongs to the order Sapindales and includes two subfamilies, the basal and polyphyletic Spondioideae and the monophyletic and more diverse Anacardioideae (Pell et al. 2011;Weeks et al. 2014). The members of this family have, in general, inconspicuous, actinomorphic, dichlamydeous flowers with superior ovary, rarely perigynous or epigynous (Wannan and Quinn 1991;Bachelier and Endress 2009;Pell et al. 2011). The occurrence of pseudomonomerous gynoecia with a single fertile ovule is well documented in the subfamily Anacardioideae (Copeland 1961;Wannan and Quinn 1991;Hormaza and Polito 1996;Gallant, Kemp, and Lacroix 1998;Endress 2007, 2009;Gonzalez 2016). ...
... Despite the large number of studies on the structural features on the gynoecia, it is still difficult to determine if the gynoecium is monomerous or pseudomonomerous is some genera of Anacardiaceae (e.g., Anacardium and Mangifera) (Wannan and Quinn 1991;Bachelier and Endress 2009;Gonzales 2016). In this context, despite the criticisms, floral vasculature has been a useful tool in the interpretation of the potential loss of whorls and/or of individual organs during the evolutionary process in many lineages of plants Schmid 1972;Dickison 2000;Sajo et al. 2012;. ...
... In addition, no evidence of additional carpel primordia was observed in our developmental study. Earlier publications that suspected pseudomonomery in Anacardium and Mangifera did not provide a complete overview of the development and vascularization of the gynoecium, suggesting that an extreme reduction of carpels in both genera occurred since the related genera have a pseudomonomerous gynoecium (Copeland 1961;Wannan and Quinn 1991;Bachelier and Endress 2009). According to Sokoloff et al. (2017) the absence of clear morphological series and traces of sterile carpels are enough to support the classification of the gynoecium as monomerous. ...
Species of Cannabaceae, Moraceae, Urticaceae and Ulmaceae form the group of urticalean rosids. This group shares the presence of small, slightly showy, diclinous (unisexual), achlamydeous or monochlamydeous flowers, that have a bicarpellate pseudomonomerous gynoecium with a single functional ovule. The flower is considered to be reduced in relation to the other rosids. This study shows that development can explain such an uncommon floral construction of urticalean rosids. Our data are based on studies of at least 20 species, summarized as follows: (1) The monochlamydeous flower results from the absence of a petal whorl from the inception. The interspecific variation in the sepal number is also due to the absence of sepal primordia in the floral meristem. (2) The dicliny results from abortion of stamens and carpels, resulting in staminate flowers with a pistillode and pistillate flowers with staminodes. (3) The pseudomonomerous gynoecium results by the arising of one primordium at the center of the floral meristem that differentiates into two carpels; only one develops normally and produces an ovule while the other does not initiate any ovule, although part of its tissues composes the ovary and even the style and the stigma. (4) The association of the floral organ arrangement with wind pollination is remarkable in the staminate flower of some species. The pistillode inflates during flower development, propelling the anthers enveloped by the sepals outward from the floral center, releasing explosively agglutinated pollen. In the entomophylous species, the union of several stigmas of different flowers forming platforms (Ficus), or the supply of pollen and varied secretions to the pollinating insects (Artocarpus, Castila, Dorstenia) guarantee the formation of seeds. Concluding, floral development is conserved and supports the monophyly of the group. On the contrary, floral specializations are responses to the selection pressures exerted by the different pollinators in the group.
... The family Anacardiaceae belongs to the order Sapindales and includes two subfamilies, the basal and polyphyletic Spondioideae and the monophyletic and more diverse Anacardioideae (Pell et al. 2011;Weeks et al. 2014). The members of this family have, in general, inconspicuous, actinomorphic, dichlamydeous flowers with superior ovary, rarely perigynous or epigynous (Wannan and Quinn 1991;Bachelier and Endress 2009;Pell et al. 2011). The occurrence of pseudomonomerous gynoecia with a single fertile ovule is well documented in the subfamily Anacardioideae (Copeland 1961;Wannan and Quinn 1991;Hormaza and Polito 1996;Gallant, Kemp, and Lacroix 1998;Endress 2007, 2009;Gonzalez 2016). ...
... Despite the large number of studies on the structural features on the gynoecia, it is still difficult to determine if the gynoecium is monomerous or pseudomonomerous is some genera of Anacardiaceae (e.g., Anacardium and Mangifera) (Wannan and Quinn 1991;Bachelier and Endress 2009;Gonzales 2016). In this context, despite the criticisms, floral vasculature has been a useful tool in the interpretation of the potential loss of whorls and/or of individual organs during the evolutionary process in many lineages of plants (Puri 1951;Schmid 1972;Gifford and Foster 1989;Dickison 2000;Sajo et al. 2012;Hirao, Monteiro, and Demarco 2019). ...
... In addition, no evidence of additional carpel primordia was observed in our developmental study. Earlier publications that suspected pseudomonomery in Anacardium and Mangifera did not provide a complete overview of the development and vascularization of the gynoecium, suggesting that an extreme reduction of carpels in both genera occurred since the related genera have a pseudomonomerous gynoecium (Copeland 1961;Wannan and Quinn 1991;Bachelier and Endress 2009). According to Sokoloff et al. (2017) the absence of clear morphological series and traces of sterile carpels are enough to support the classification of the gynoecium as monomerous. ...
The diversity of gynoecium morphology is an important topic of evolutionary biology since this floral organ plays a central role in the reproduction of angiosperms. Gynoecia are called pseudomonomerous when they present only one fertile carpel and more or less prominent traces of sterile carpels, having evolved independently multiple times in angiosperm history. Here we revisit this particular gynoecium feature in the subfamily Anacardioideae (Anacardiaceae) and use the results as a case study to demonstrate how different patterns of development lead to a pseudomonomerous gynoecium and how a detailed morphological analysis can improve our understanding about the mechanisms behind the great floral diversity of the angiosperms. We describe and compare gynoecium development and vasculature in three species of Anacardioideae using SEM and LM. A database from published reports was compiled in order to explore the development of pseudomonomerous gynoecia across major clades. Anacardioideae species have an asymmetrical development of the gynoecium, leading to pseudomonomery; however, all carpels are connected by a compitum. In species belonging to genera like Anacardium and Mangifera, numerous vascular traces indicate that there was a reduction in the number of carpels in this subfamily, from three to one. Similar developmental patterns are addressed in other lineages of angiosperms. This may represent a key evolutionary novelty in Anacardioideae. We also discuss the role of pseudomonomerous gynoecium in a phylogenetic and evolutionary context in other lineages of angiosperms.
... 14), we recovered that the ancestor of Meliaceae and Simaroubaceae might have had four to five loculi, with the same antepetalous position, but this number is a homoplastic character state and can vary from 1 to 20 in living meliaceous genera (Fig. 8B). The antepetalous position of the carpels is consistent with obdiplostemony, which is a common condition in Sapindales lineages, as defined by Bachelier & Endress (2009). ...
Meliaceae are a mostly pantropical family in the Sapindales, bearing flowers typically provided with a staminal tube, formed by filaments that are fused partially or totally, although several genera of subfamily Cedreloideae have free stamens, which may be adnate to an androgynophore in some taxa. The fact that the family exhibits a wide diversity of floral and fruit features, as well as of sexual systems and pollination syndromes, offers interesting questions on the evolutionary processes that might have taken place during its history. In this study, we analysed the distribution of 20 reproductive morphological traits of Meliaceae, upon an available molecular phylogenetic framework, using 31 terminals from the family's two main clades (Cedreloideae and Melioideae), plus six Simaroubaceae taxa as outgroup. We aimed to identify and/or confirm synapomorphies for clades within the family, and to develop hypotheses on floral evolution and sexual systems in the group. Our reconstruction suggests that the ancestor of Meliaceae possibly was provided with united stamens and unisexual flowers in dioecious individuals, with a subsequent change to free stamens and monoecy in the ancestor of Cedreloideae. Most characters studied show some degree of homoplasy, but some are unique synapomorphies of clades, such as the haplostemonous androecium. An androgynophore defines the Cedrela‐Toona clade. The comparative approach of our study and the evolutionary hypotheses generated herein reveal several aspects demanding further structural investigation, and possible evolutionary pathways of the reproductive structures along the lineages diversification, mostly related to specialization of sexual systems, floral biology and dispersal strategies. This article is protected by copyright. All rights reserved.
... In members of Sapindales, obdiplostemony is commonly mentioned because it is traditional in the literature of this group (Nair & Joseph, 1957;Nair & Joshi, 1958;Narayan & Sayeeduddin, 1958;Eckert, 1966;Gut, 1966;Simpson, 2010). In some studies, however, such as that of the Rutaceae tribe Ruteae (Wei, Wang & Li, 2012), the androecium is still classified as obdiplostemonous based on the antepetalous position of the carpels, following the definition of Bachelier & Endress (2009), despite observation of the initial development of the antesepalous stamens in the outer whorls. Therefore, a more conclusive verification of patterns found in the androecium of Simaba could be gained from future studies of floral development. ...
... In Simaba, we observed the presence of a short connate region at the base of the ovary of all analysed species, as found in other Simaroubaceae (Nair & Joshi, 1958;Ramp, 1988) and Rutaceae (Gut, 1966;Ramp, 1988;Pirani et al., 2010;El Ottra et al., 2013). In addition, the occurrence of post-genitally united carpel tips in Simaba is not only common in Simaroubaceae and Rutaceae, but also reported for Kirkiaceae and some Anacardiaceae, and outside Sapindales in other core eudicots (Endress et al., 1983;Ramp, 1988;Bachelier & Endress, 2008, 2009Matthews, Amaral & Endress, 2012;El Ottra et al., 2013). ...
... It is possible that they also perform a role in fruit development (Endress, 2008). Although documented for some families, including Anacardiaceae (Bachelier & Endress, 2009), Rutaceae (Zanthoxylum L., Beurton, 1994), Hamamelidaceae, Cunoniaceae (Matthews & Endress, 2002), Monimiaceae (Endress, 1980), Araliaceae (Endress, 2006) and Moraceae (Berg, 1990), the density and length of the indument on the gynophore and ovaries found in species of Simaba spp. are high. ...
Simaroubaceae are a small, monophyletic family with a mostly pantropical distribution. Although the
considerable floral variation found in Simaroubaceae has resulted in some detailed structural studies, no such
studies exist for Simaba, the largest genus in the family. We present data on the floral structure of Simaba using
a comparative approach, including studies of vascularization and histology. We analysed species of two of the
three sections of the genus (Simaba sections Floribundae and Grandiflorae) with the goal of providing characters for studies of taxonomy and phylogeny in the genus. We found several floral structures commonly reported for other genera of Simaroubaceae and closely related groups of Sapindales. We also found some novel structural features in Simaba, such as variation in merism among flowers of the same inflorescence and sexual floral variations related to organ sterility, a condition known in other members of Sapindales, but not previously recorded in Simaroubaceae. We also found partial connation and coherence of the androecium, which forms a pseudotube in species of section Grandiflorae. Finally, the nectariferous feature of the gynophore surface of the studied species was described in detail.
... In members of Sapindales, obdiplostemony is commonly mentioned because it is traditional in the literature of this group (Nair & Joseph, 1957;Nair & Joshi, 1958;Narayan & Sayeeduddin, 1958;Eckert, 1966;Gut, 1966;Simpson, 2010). In some studies, however, such as that of the Rutaceae tribe Ruteae (Wei, Wang & Li, 2012), the androecium is still classified as obdiplostemonous based on the antepetalous position of the carpels, following the definition of Bachelier & Endress (2009), despite observation of the initial development of the antesepalous stamens in the outer whorls. Therefore, a more conclusive verification of patterns found in the androecium of Simaba could be gained from future studies of floral development. ...
... In Simaba, we observed the presence of a short connate region at the base of the ovary of all analysed species, as found in other Simaroubaceae (Nair & Joshi, 1958;Ramp, 1988) and Rutaceae (Gut, 1966;Ramp, 1988;Pirani et al., 2010;El Ottra et al., 2013). In addition, the occurrence of post-genitally united carpel tips in Simaba is not only common in Simaroubaceae and Rutaceae, but also reported for Kirkiaceae and some Anacardiaceae, and outside Sapindales in other core eudicots (Endress et al., 1983;Ramp, 1988;Bachelier & Endress, 2008, 2009Matthews, Amaral & Endress, 2012;El Ottra et al., 2013). ...
... It is possible that they also perform a role in fruit development (Endress, 2008). Although documented for some families, including Anacardiaceae (Bachelier & Endress, 2009), Rutaceae (Zanthoxylum L., Beurton, 1994), Hamamelidaceae, Cunoniaceae (Matthews & Endress, 2002), Monimiaceae (Endress, 1980), Araliaceae (Endress, 2006) and Moraceae (Berg, 1990), the density and length of the indument on the gynophore and ovaries found in species of Simaba spp. are high. ...
Simaroubaceae are a small, monophyletic family with a mostly pantropical distribution. Although the considerable floral variation found in Simaroubaceae has resulted in some detailed structural studies, no such studies exist for Simaba, the largest genus in the family. We present data on the floral structure of Simaba using a comparative approach, including studies of vascularization and histology. We analysed species of two of the three sections of the genus (Simaba sections Floribundae and Grandiflorae) with the goal of providing characters for studies of taxonomy and phylogeny in the genus. We found several floral structures commonly reported for other genera of Simaroubaceae and closely related groups of Sapindales. We also found some novel structural features in Simaba, such as variation in merism among flowers of the same inflorescence and sexual floral variations related to organ sterility, a condition known in other members of Sapindales, but not previously recorded in Simaroubaceae. We also found partial connation and coherence of the androecium, which forms a pseudotube in species of section Grandiflorae. Finally, the nectariferous feature of the gynophore surface of the studied species was described in detail.
... Flowers in which the gynoecium is not isomerous with the other fl oral whorls exhibit a kind of inconspicuous (oblique) monosymmetry (such as in Nitrariaceae, except for Tetradiclis ). Isomerous and polysymmetric fl owers are pentamerous in Biebersteiniaceae (Reiche, 1889 ), tetramerous in Kirkiaceae (Stannard, 1981 ;Bachelier and Endress, 2008 ), and can variously have three, four or fi ve organs per whorl in Anacardiaceae, Burseraceae, Meliaceae, Rutaceae or Simaroubaceae (Bachelier and Endress, 2009 ). Such inconspicuously monosymmetric fl owers with an oblique symmetry also occur in Anacardiaceae and Burseraceae (Bachelier and Endress, 2009 ). ...
... Isomerous and polysymmetric fl owers are pentamerous in Biebersteiniaceae (Reiche, 1889 ), tetramerous in Kirkiaceae (Stannard, 1981 ;Bachelier and Endress, 2008 ), and can variously have three, four or fi ve organs per whorl in Anacardiaceae, Burseraceae, Meliaceae, Rutaceae or Simaroubaceae (Bachelier and Endress, 2009 ). Such inconspicuously monosymmetric fl owers with an oblique symmetry also occur in Anacardiaceae and Burseraceae (Bachelier and Endress, 2009 ). However, pronounced monosymmetry with unequal diff erentiation of certain fl oral sectors is less common and not present in Nitrariaceae. ...
... Flowers with a pentamerous calyx, in which the sepals are initiated in a quincuncial spiral sequence with the second sepal in abaxial (posterior) position, such as in Nitraria and Peganum , are common in Sapindales and other rosids (Ronse De Craene, 2010 ). Th at the corolla protects the inner fl oral organs up to anthesis, with a postgenital coherence between the petals (formed by the papillate epidermis or interlocking hairs), also occurs in other Sapindales (Bachelier and Endress, 2009 ;for details, see Ronse De Craene and Haston, 2006 , p. 468), and in other rosids, especially in malvids Matthews and Endress, 2006 ;Endress, 2010 ). ...
For the last 20 years, the development and improvement of molecular methods, based mostly on the comparison of DNA sequences, have been increasingly successful in reconstructing the phylogenetic tree of plants at all hierarchical levels. Consequently, they have contributed greatly to the recent improvement of angiosperm systematics. In addition, they have shown that earlier classifications, based mostly on plant vegetative and reproductive structures, had sometimes been misled by homoplastic characters, and a number of orders and families have had to be newly circumscribed or even newly established (e.g. APG, 1998, 2003, 2009; Stevens, 2001 onwards). These new results provide a novel basis for comparative structural studies to characterize the newly recognized clades and to evaluate clades that have only limited molecular support. However, because such comparative studies are time-consuming and the systematic classification has different hierarchical levels, they can only be done in a stepwise fashion (for eudicots, e.g. Matthews and Endress, 2002, 2004, 2005a, b, 2006, 2008; von Balthazar et al., 2004; Schönenberger and Grenhagen, 2005; Endress and Matthews, 2006; Ronse De Craene and Haston, 2006; von Balthazar et al., 2006; Bachelier and Endress, 2008, 2009; Janka et al., 2008; Schönenberger, 2009; von Balthazar and Schönenberger, 2009; Schönenberger et al., 2010). As part of such a comparative approach, we studied the floral structure of Nitrariaceae, a small family which has been recently reclassified in Sapindales (APG, 2009). Nitrariaceae comprise four genera and around 15 species (Stevens, 2001 onwards; APG, 2009). They are native to arid and semi-arid regions of the Old World and are small to medium-sized shrubs (Nitraria; Engler, 1896a, b; Bobrov, 1965; Noble and Whalley, 1978), perennial herbs (Peganum and Malacocarpus; Engler, 1896a, 1931; El Hadidi, 1975) or small annual herbs of only a few centimetres height (Tetradiclis; Engler, 1896b, 1931; Hamzaoglu et al., 2005). In earlier classifications, the position and the mutual affinities of these genera varied tremendously, depending on the weight an author gave either to their vegetative or their reproductive features (Takhtajan, 1969, 1980, 1983, 2009; El Hadidi, 1975; Dahlgren, 1980; Cronquist, 1981, 1988; see Sheahan and Chase, 1996 for a detailed review of classifications). Because none of the traditional classifications was entirely satisfactory, however, most authors followed Engler’s influential work (1896a, b, 1931) and Nitraria, Tetradiclis and Peganum (including Malacocarpus) remained for a long time in their own subfamilies in Zygophyllaceae (Nitrarioideae, Tetradiclidoideae and Peganoideae; for more details of the history of classification, see Sheahan and Chase, 1996).
... Flowers in which the gynoecium is not isomerous with the other fl oral whorls exhibit a kind of inconspicuous (oblique) monosymmetry (such as in Nitrariaceae, except for Tetradiclis ). Isomerous and polysymmetric fl owers are pentamerous in Biebersteiniaceae (Reiche, 1889 ), tetramerous in Kirkiaceae (Stannard, 1981 ;Bachelier and Endress, 2008 ), and can variously have three, four or fi ve organs per whorl in Anacardiaceae, Burseraceae, Meliaceae, Rutaceae or Simaroubaceae (Bachelier and Endress, 2009 ). Such inconspicuously monosymmetric fl owers with an oblique symmetry also occur in Anacardiaceae and Burseraceae (Bachelier and Endress, 2009 ). ...
... Isomerous and polysymmetric fl owers are pentamerous in Biebersteiniaceae (Reiche, 1889 ), tetramerous in Kirkiaceae (Stannard, 1981 ;Bachelier and Endress, 2008 ), and can variously have three, four or fi ve organs per whorl in Anacardiaceae, Burseraceae, Meliaceae, Rutaceae or Simaroubaceae (Bachelier and Endress, 2009 ). Such inconspicuously monosymmetric fl owers with an oblique symmetry also occur in Anacardiaceae and Burseraceae (Bachelier and Endress, 2009 ). However, pronounced monosymmetry with unequal diff erentiation of certain fl oral sectors is less common and not present in Nitrariaceae. ...
... Flowers with a pentamerous calyx, in which the sepals are initiated in a quincuncial spiral sequence with the second sepal in abaxial (posterior) position, such as in Nitraria and Peganum , are common in Sapindales and other rosids (Ronse De Craene, 2010 ). Th at the corolla protects the inner fl oral organs up to anthesis, with a postgenital coherence between the petals (formed by the papillate epidermis or interlocking hairs), also occurs in other Sapindales (Bachelier and Endress, 2009 ;for details, see Ronse De Craene and Haston, 2006 , p. 468), and in other rosids, especially in malvids Matthews and Endress, 2006 ;Endress, 2010 ). ...
Introduction
For the last 20 years, the development and improvement of molecular methods, based mostly on the comparison of DNA sequences, have been increasingly successful in reconstructing the phylogenetic tree of plants at all hierarchical levels. Consequently, they have contributed greatly to the recent improvement of angiosperm systematics. In addition, they have shown that earlier classifications, based mostly on plant vegetative and reproductive structures, had sometimes been misled by homoplastic characters, and a number of orders and families have had to be newly circumscribed or even newly established (e.g. APG, 1998, 2003, 2009; Stevens, 2001 onwards). These new results provide a novel basis for comparative structural studies to characterize the newly recognized clades and to evaluate clades that have only limited molecular support. However, because such comparative studies are time-consuming and the systematic classification has different hierarchical levels, they can only be done in a stepwise fashion (for eudicots, e.g. Matthews and Endress, 2002, 2004, 2005a, b, 2006, 2008; von Balthazar et al., 2004; Schönenberger and Grenhagen, 2005; Endress and Matthews, 2006; Ronse De Craene and Haston, 2006; von Balthazar et al., 2006; Bachelier and Endress, 2008, 2009; Janka et al., 2008; Schönenberger, 2009; von Balthazar and Schönenberger, 2009; Schönenberger et al., 2010).
As part of such a comparative approach, we studied the floral structure of Nitrariaceae, a small family which has been recently reclassified in Sapindales (APG, 2009). Nitrariaceae comprise four genera and around 15 species (Stevens, 2001 onwards; APG, 2009). They are native to arid and semi-arid regions of the Old World and are small to medium-sized shrubs (Nitraria; Engler, 1896a, b; Bobrov, 1965; Noble and Whalley, 1978), perennial herbs (Peganum and Malacocarpus; Engler, 1896a, 1931; El Hadidi, 1975) or small annual herbs of only a few centimetres height (Tetradiclis; Engler, 1896b, 1931; Hamzaoglu et al., 2005). In earlier classifications, the position and the mutual affinities of these genera varied tremendously, depending on the weight an author gave either to their vegetative or their reproductive features (Takhtajan, 1969, 1980, 1983, 2009; El Hadidi, 1975; Dahlgren, 1980; Cronquist, 1981, 1988; see Sheahan and Chase, 1996 for a detailed review of classifications). Because none of the traditional classifications was entirely satisfactory, however, most authors followed Engler’s influential work (1896a, b, 1931) and Nitraria, Tetradiclis and Peganum (including Malacocarpus) remained for a long time in their own subfamilies in Zygophyllaceae (Nitrarioideae, Tetradiclidoideae and Peganoideae; for more details of the history of classification, see Sheahan and Chase, 1996).
2023 update! Cole TCH, Bachelier JB (2023) ANACARDIACEAE Phylogeny Poster (AnaPP) © Cole, Bachelier 2023 (CC-BY) • hypothetical tree based on molecular phylogenetic data (chiefly following Joyce et al. 2023) • branch lengths deliberate, not expressing actual time scale • species numbers (in gray) are approximate (from POWO, Plants of the World Online) • Reference: Joyce EM et al. (2023) Phylogenomic analyses of Sapindales support new family relationships, rapid Mid-Cretaceous Hothouse diversification, and heterogeneous histories of gene duplication. Front. Plant Sci. 14:1063174. doi: 10.3389/fpls.2023.1063174
https://www.researchgate.net/publication/369052431_Phylogenomic_analyses_of_Sapindales_support_new_family_relationships_rapid_Mid-Cretaceous_Hothouse_diversification_and_heterogeneous_histories_of_gene_duplication
