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Falcatifolium taxoides. Fig. 46, Inner surface, region of the stomatal apparatus showing two subsidiary cells; 1,400. Fig. 47, Inner surface, stomatal rows; x 150. Fig. 48, Inner view, cuticle on guard and subsidiary cell surfaces; x 3,000.
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Cuticle micromorphology of leaves from all five species of the Southern Hemisphere conifer genus Falcatifolium de Laubenfels (Podocarpaceae) was studied with scanning electron microscopy, Both herbarium and preserved specimens were examined and showed no differences in micromorphology. External and internal features of abaxial and adaxial cuticles...
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... inner sunken ringlike zone corresponds to the position of the Florin ring in other Falcati- folium species and the deep crease in subsidiary cell cuticle internally. The outer ring corresponds to the outer edge of the subsidiary cell wall cuticle internally (e.g., fig. 46). The leaves observed in this study show deep creases in the subsidiary cell wall cuticle and, therefore, what might be inter- preted as a partially sunken Florin ring externally ( fig. ...
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... In other words, Florin rings are "conspicuous thickened rings of cutin overlying the accessory cells and surrounding the stomata" [43]. They are associated with the subsidiary cells observed in both living and fossil gymnosperms, such as P. strobus or Papuacedrus [44,45], and were also described in Falcatifolium (Podocarpaceae) [46]. ...
Using a scanning electron microscope, the micromorphologies of needle primordia and the young needles of seven pine species (Pinus cembra, P. mugo, P. nigra, P. rigida, P. sylvestris, P. strobus, and P. uncinata) were analyzed at phenological stages B2 and B3 (according to Debazac). In B2, needle tips were rounded or pointed, depending on the species. In P. cembra and P. strobus, teeth were noted on the tips. Teeth were also visible on the margins in P. mugo, P. cembra, and P. strobus. Stomata became visible in the late B2 phase (P. sylvestris, P. mugo, and P. nigra) near the needle tips and were arranged in rows. In the B3 phase, needle tips were pointed. Only in P. strobus was the needle tip slightly rounded. The teeth on the margin in all the species were pointed. In P. strobus, their size and density along the margin decreased basipetally. In B3 for all the species, numerous stomata were visible. In P. sylvestris, P. cembra, and P. strobus, Florin rings were also observed. These observations could be useful in pine systematics but also in palaeobotanical or physiological studies. To the best of our knowledge, this is the first study on the micromorphology of very young needles in representatives of the genus Pinus.
... In both species examined here, stomata are surrounded by well-developed, high Florin rings that originate from the subsidiary cells (Fig. 7). Florin rings are common in xeromorphic conifer leaves and may reduce stomatal transpiration (Buchholz & Gray, 1948;Florin, 1951;Oladele, 1983;Yoshie & Sakai, 1985;Hill & Pole, 1992;Stockey, Ko & Woltz, 1992;Kunzmann et al., 2006;Kunzmann, 2007;Kim et al., 2011). Florin rings have also been recorded in extremely wet environments, where they assist in diverting fungal hyphae away from the stomatal pore, a feature that may be important in retaining the photosynthetic function of the long-lived phylloclades (see Mohammadian, Hill & Watling, 2009 for a similar argument in the conifer genus Agathis Salisb.). ...
The structure of phylloclades and true leaves in Phyllocladus was investigated with emphasis on function, ecology and evolution. Only in the earliest ontogenetic stages are true, needle leaves developed. The earliest phylloclades comprise a fusion product of a shoot axis of the first order and its inserted leaves. Later, this is augmented by the expansion of axillary short shoots from the base of some of the fused leaves. Phylloclades of mature individuals are entire short shoot systems, including leaves and shoots, becoming fused with photosynthetic tissue. These phylloclades represent one of the adaptations for increasing functional photosynthetic area that are present in Podocarpaceae. Such adaptations were probably needed because Podocarpaceae are usually medium-sized trees faced with competition from the ever-expanding angiosperm forests at mid to high latitudes in the Southern Hemisphere from the Late Cretaceous, but especially during the Palaeocene–Eocene, when dense and diverse angiosperm-dominated rainforests were common. The data obtained from living species were compared to those from fossil material of Phyllocladus. Phyllocladus with fully developed phylloclades have been in place since at least the early Cenozoic.
... The term 'Florin ring' was first used Buchholz and Gray (1948) for circular thickenings above the guard cells in younger conifers. It occurs also in Podocarpaceae (Stockey et al., 1992) and Araucariaceae (Stockey and Atkinson, 1993 Derivatio nominis: the epitheton carpaticus was chosen because this species was firstly described by Florjan and Żołdani (1999) from the Carpathian region. ...
The adpression fossil record from the Westphalian of the Intrasudetic Basin (Czech Republic) contains diversified medullosan prepollen organs. Langsettian bell-shaped prepollen organs were found in the Lampertice Member in the Žacléř area: Whittleseya elegans Newberry, W. delphinae Stockmans et Willière, W. silesiaca Pacyna et Zdebska and Givesia media (Stockmans) Stockmans et Willière. These fructifications may have been produced by plants with fronds of the fossil-genera Neuralethopteris Cremer and Alethopteris Sternberg. Parent plants of small Langsettian Aulacotheca sp. are uncertain. Duckmantian medullosan male fructifications from the Malé Svatoňovice area are elongated capsules described as Aulacotheca idae Šimůnek sp. nov. and were produced by plants with Alethopteris idae Šimůnek fronds. The prepollen grains and cuticles were isolated from Whittleseya elegans, W. delphinae, Givesia media and Aulacotheca idae. All prepollen grains are oval with two longitudinal grooves and a monolete suture, and are classified as Monoletes ovatus Schopf. There is great overlap in the dimensions of the prepollen of Whittleseya elegans, W. delphinae and Givesia media – 155–230 μm long and 100–180 μm wide. Prepollen of Aulacotheca idae are much bigger – 330–375 μm long and 190–245 μm wide. The inner cuticular layer is gently cutinised with polygonal cells. Cuticles from the prepollen organ surface has elongated cells in Whittleseya elegans (with stomata) and Givesia media (with trichome bases), and has polygonal cells in Whittleseya delphinae (with periclinal cell wall thickenings) and Aulacotheca idae (with hollow papillae on periclinal cell walls). The question, as to whether Whittleseya elegans and W. delphinae are conspecific is discussed.
... Florin rings are common features in leaves of extant gymnosperms (Florin 1931(Florin , 1951Buchholz & Gray 1948;Oladele 1983;Yoshie & Sakai 1985;Stockey et al. 1992;Kim et al. 2011) and are also well preserved in the fossil record (Hill & Pole 1992;Kunzmann et al. 2006;Kunzmann 2007). They are raised ring-like structures or cuticular wall protrusions, which originate from the subsidiary cells of the stomata. ...
The foliar shift from juvenile needle leaves to the mature scale leaves was investigated in five Callitris species. Their habitats range from rainforest to the semi‐arid. In C. macleayana from the most mesic environment the change from needle to scale leaves is variable and juvenile leave are even retained on mature trees. In the other four species there is a rapid change in size and structure to scale leaves. Anatomically the photosynthetic tissue is condensed and stomatal area is limited compared to species with conventional leaves. A hypodermis is most prominent on the abaxial side of the leaves and may play a role in protection from high solar radiation as most species grow in high light areas. While stomata in the four taxa from wet or moderate rainfall areas are freely exposed, those of C. gracilis from semi‐arid habitats are well protected in longitudinal furrows between the decurrent leaves. The reduced leaf area of all species allows close association between the water‐conducting xylem and the stomata that will facilitate rapid leaf conductance. This links with the anisohydric physiology of the genus and the shallow rooting that can take advantage of brief rainfall events for species in arid climates.
... The random stomatal orientation and distribution observed for H. laubenfelsii are compatible with most of the genera within the scale-leaved podocarp clade (sensu Andruchow-Colombo et al. 2019; Fig. 8), wherein all its members, except Phyllocladus and Lepidothamnus, have randomly oriented and distributed stomata (Florin 1931;Hill 1989;Wells and Hill 1989;Stockey et al. 1995;Andruchow-Colombo et al. 2019; Table S1). Neither stomatal orientation nor distribution was recovered as synapomorphic for the scale-leaved clade by Andruchow-Colombo et al. (2019), but they were suggested as putatively synapomorphic (Lepidothamnus and Phyllocladus were interpreted as having changed independently). ...
... The presence of only lateral (and no polar) subsidiary cells, as in H. laubenfelsii, has not been observed previously in any member of the scale-leaved clade (Hill 1989;Wells and Hill 1989;Stockey et al. 1995;Andruchow-Colombo et al. 2019 ; Table S1). However, this pattern is known for some extant and extinct species of Prumnopitys (Stockey and Frevel 1997;Greenwood et al. 2013), which is part of the encompassing prumnopityoid clade (sensu Conran et al. 2000;Biffin et al. 2012;Knopf et al. 2012;Leslie et al. 2012Leslie et al. , 2018Andruchow-Colombo et al. 2019;Fig. ...
Huncocladus laubenfelsii gen. et sp. nov. is described from the early Eocene (52 million years old) Laguna del Hunco site in Patagonia, Argentina, on the basis of a compression fossil with cuticle remains. The taxon has several similarities with Phyllocladus, together with characters that are absent in extant Phyllocladus species but are otherwise typical of the enclosing scale-leaved clade. Consequently, Huncocladus is interpreted as a relative of Phyllocladus, possibly belonging to its stem group. This view is supported by a phylogenetic analysis of Podocarpaceae, which recovers Huncocladus as sister to Phyllocladus within the here-termed phyllocladoid clade (Phyllocladus + Huncocladus). Huncocladus laubenfelsii is the first macrofossil record of the phyllocladoid lineage in South America or anywhere in the western hemisphere, vastly extending its historical range and constituting an additional lineage shared between Eocene Patagonia and extant and extinct Australasian and South-east Asian rainforests. The disappearance of phyllocladoids from South America adds to the general extinction pattern described previously for southern hemisphere Podocarpaceae, associated with the family’s low drought tolerance in the face of climate change (i.e. aridification). Huncocladus is the oldest record of the phyllocladoids, and it represents a new reference point for temporal calibration and biogeographic inference for the evolution of conifers and Australasian rainforests.
... Auraucaria Juss. (Stockey and Taylor, 1978a,b) and Podocarpaceae (Stockey and Ko, 1988;Stockey et al., 1992Stockey et al., , 1995Mill and Schilling, 2009). For cycads, Chamberlain (1935) reported that stomata are found inside crypts for most of the 117 cycad species known at that time (Calonje et al., 2017). ...
Anatomical descriptions can be effective for solving systematic issues, but these studies are relatively scarce for cycads. Therefore, we present here a leaflet and cuticle anatomical study on the genus Dioon, to provide a set of epidermal traits that clarify species delimitation and relationships between species and their habitats. We used standard micro-technique for leaflet sectioning, and cuticular peel preparation for light microscopy. Also, we used the chromium trioxide method for scanning electron microscope observations on cuticles. Measurements were taken on 10 randomly chosen replicates of each cell or tissue type, for each of the leaflets sampled per taxon. Micromorphological variation among species was calculated for each trait. Finally, we reconstructed the ancestral states of the observed epidermal fibre-like cell and pore shapes, by tracing the characters on the species phylogenetic tree of Dioon. We were able to describe the leaflet anatomy, cuticles, and epidermal features for 14 Dioon species. The quantitative analysis was useful to reveal five geographically structured species groups. Character tracing on the phylogenetic tree of Dioon has amplified our current understanding on species relationships with respect to habitats. The presence/absence data suggest that the evolutionary acquisition-deletion of structural shapes is phylogenetically independent, thus climate seems to play a very important role in the variation of cuticular and stomatal traits. Many epidermal traits, especially adaxial cuticle thickness and epistomatal pore width and depth, might be adaptations resulting from a long-term influence of climate, since they appear to have correlation with climatic conditions in relation to their biogeography. We conclude that the variation of all traits are mostly sustained and intrinsic to the species, and are of promising taxonomic value. The combination of the epidermal traits with other characters has potential for taxonomic resolution at species level.
... The method of Alvin & Boulter (1974) has become a standard technique for the isolation of the leaf cuticle. The number of scanning electron microscopy (SEM) studies dealing with the micromorphology of internal and external cuticle surfaces has greatly increased since Alvin & Boulter (1974), mainly for extant gymnosperms, but also for angiosperms (Oladele, 1983a(Oladele, , 1983bHoriuchi & Kimura, 1987;Stockey, Ko & Woltz, 1992;Stockey & Atkinson, 1993;Stockey, Frevel & Woltz, 1998;Kim, Whang & Hill, 1999;Leng et al., 2001;Whang et al., 2001;Ma et al., 2009;Mill & Stark Schilling, 2009;Lumaga et al., 2015;Li et al., 2016). ...
... Among extant gymnosperms, the large group of the conifers comprises six commonly recognized families, Pinaceae, Araucariaceae, Podocarpaceae s.l., Sciadopityaceae, Taxaceae s.l. and Cupressaceae s.l., which form monophyletic groups and have typical morphological characters (Hart, 1987;Cheng et al., 2000;Quinn, Price & Gadek, 2002;Farjon, 2001Farjon, , 2010Eckenwalder, 2009;Christenhusz et al., 2011;Knopf et al., 2012;Leslie et al., 2012;Lu et al., 2014;Schulz et al., 2014). Studies of cuticular micromorphology have been carried out extensively for the Araucariaceae-Podocarpaceae clade (Stockey & Taylor, 1981;Stockey & Ko, 1988;Stockey et al., 1992Stockey et al., , 1998Stockey & Atkinson, 1993;Mill & Stark Schilling, 2009) and Cupressaceae s.l. (Oladele, 1983a(Oladele, , 1983bLeng et al., 2001;Ma, Li & Li, 2007;Ma et al., 2009). ...
... This might be feasible for morphologically similar species, e.g. the Central Asian species (Möller et al., 2007(Möller et al., , 2013. The internal surfaces of gymnosperm leaf cuticles have been previously reported to be a useful character in conifer investigations (Stockey & Taylor, 1981;Oladele, 1983a;Stockey & Ko, 1988;Stockey et al., 1992Stockey et al., , 1998Stockey & Atkinson, 1993;Leng et al., 2001;Whang et al., 2001;Xiang & Farjon, 2003;Ma et al., 2009;Mill & Stark Schilling, 2009); for the first time, we have applied this method extensively to Taxaceae s.l. using SEM. ...
Among living conifers, Taxaceae s.l. are a small family of six genera and c. 36 species. In this study, the external and the isolated internal surfaces of the leaf cuticles were examined, using scanning electron microscopy (SEM), to find characteristics at the generic or species level. Therefore, we investigated 130 specimens, representing 31 species. Character evolution was reconstructed using Mesquite and the results were evaluated under consideration of the fossil record. For the outer cuticle surfaces, epidermal protuberances and the shape of stomatal pores proved to be the best differentiating characters at the generic level. In addition, elongated protuberances were first documented in some Cephalotaxus spp. Based on the cuticle internal surfaces, the six genera could be distinguished by monocyclic or amphicyclic stomata and the composition of protuberances, which was documented for the first time using SEM. We showed that papillae are hollow or solid in Taxus and Torreya and that protuberances in Cephalotaxus accompany a furrowed cuticle. Stomatal rings in Taxus and Torreya are always composed of hollow papillae, whereas stomatal rings in Austrotaxus are formed entirely by the cuticle. We hypothesize that protuberances may be apomorphic in the evolution of this family, having evolved several times independently. The present study provides cuticular characters that are also relevant in palaeontological studies, since they are often better preserved than other features.
... This can be seen when stained with Sudan III and IV ( Figure 1D) and is reminiscent of Florin rings, for want of a better term. Florin rings are associated with the subsidiary cells observed in both living and fossil gymnosperms such as Pinus strobus by Deckert et al. (2001), Papuacedrus (Wilf et al., 2009) and in Falcatifolium (Podocarpaceae) by Stockey et al. (1992). In Dioon these structures can also seen under epifluorescence microscopy of the abaxial cuticular surface using 400-440 nm exciter filters ( Figure 1E) comparable to the Florin rings reported by Wilf et al. (2009) on fossil Papuacedrus and Paull and Hill (2010) on fossil Cupressaceae. ...
... The outermost (distal) encircling cells have a thickened upper periclinal wall as well as a thickened cuticle and can be seen in the transverse section of the Sudan stained cuticle in figure 1D (arrow) also under fluorescence microscopy in figure 1E (arrow) and illustration (Figure 2). Another ANDREW P. VOVIDES AND SONIA GALICIA difference is that stomatal plugs, as reported for Podocarpaceae by Stockey et al. (1992), were rarely seen in Dioon but present occasionally. The strongly overarching papillate subsidiary cells reported for Cycas revoluta by Pant and Mehra (1964) are unlike the structures in Dioon though their ontogeny may be similar; figure 1G in Pant and Mehra (1964) shows an overarching papilla of a subsidiary cell in a young stoma, and another subsidiary cell dividing to give rise to one encircling cell, and their figure 3F an adult stoma overarched by thickened papillae of subsidiary cells. ...
Cuticle information and epidermal characters have great potential in systematic studies. However, micromorphology and anatomical studies on cycads are relatively scarce in comparison to similar studies on other gymnosperms. In this study leafet anatomy and cuticles in the genus Dioon have been investigated with bright field microscopy, epifluorescence, and scanning electron microscopy. Structures hitherto not completely studied for the genus are reported such as lignified hypodermis, girder sclerenchyma, especially G-fibers, and Florin ring-like structures associated with the stomatal apparatus.
... If fertile material is present, it is frequently either inaccessible or detached from the tree making it difficult to convincingly associate the fertile and sterile portions. Although sterile material of Podocarpaceae can usually be identified to genus using phyllotaxis and leaf form [3,4], accurate species identification often requires careful microscopic examination of internal [5][6][7][8][9][10][11][12][13][14][15][16][17][18][19][20] and external characteristics [21][22][23][24][25][26][27]. Proper use of the existing identification tools requires training in botanical terminology, skill in microtechnique, and familiarity with Podocarpaceae. ...
We have generated matK, rbcL, and nrITS2 DNA barcodes for 320 specimens representing all 18 extant genera of the conifer family Podocarpaceae. The sample includes 145 of the 198 recognized species. Comparative analyses of sequence quality and species discrimination were conducted on the 159 individuals from which all three markers were recovered (representing 15 genera and 97 species). The vast majority of sequences were of high quality (B 30 = 0.596-0.989). Even the lowest quality sequences exceeded the minimum requirements of the BARCODE data standard. In the few instances that low quality sequences were generated, the responsible mechanism could not be discerned. There were no statistically significant differences in the discriminatory power of markers or marker combinations (p = 0.05). The discriminatory power of the barcode markers individually and in combination is low (56.7% of species at maximum). In some instances, species discrimination failed in spite of ostensibly useful variation being present (genotypes were shared among species), but in many cases there was simply an absence of sequence variation. Barcode gaps (maximum intraspecific p-distance > minimum interspecific p-distance) were observed in 50.5% of species when all three markers were considered simultaneously. The presence of a barcode gap was not predictive of discrimination success (p = 0.02) and there was no statistically significant difference in the frequency of barcode gaps among markers (p = 0.05). In addition, there was no correlation between number of individuals sampled per species and the presence of a barcode gap (p = 0.27).
... The term 'Florin ring' was first used Buchholz and Gray (1948) for circular thickenings above the guard cells in younger conifers. It occurs also in Podocarpaceae (Stockey et al., 1992) and Araucariaceae (Stockey and Atkinson, 1993 Derivatio nominis: the epitheton carpaticus was chosen because this species was firstly described by Florjan and Żołdani (1999) from the Carpathian region. ...
Cordaitalean cuticles represent a great deal of the dispersed cuticular assemblages obtained from coal seams of the Upper Silesian Basin. Cordaitalean cuticles are easy to distinguish having stomatal complexes mostly with two polar and two lateral subsidiary cells. The stomata of the abaxial cuticle usually form stomatal rows or stomatiferous bands. The cuticular types obtained from coal of the Upper Silesian Basin are not identical with any “in situ” cordaitalean cuticles so far described. For the purpose of classifying cordaitalean dispersed cuticles and because it was impossible to correlate adaxial and abaxial cuticles within dispersed cuticular spectra, two new fossil genera are erected — Cordaadaxicutis Šimůnek et Florjan, gen. nov. and Cordaabaxicutis Šimůnek et Florjan, gen. nov. Nine species of Cordaadaxicutis and nine species of Cordaabaxicutis are described. They are from the Westphalian Mudstone Series (Załęże and Orzesze Beds) and the Cracow Sandstone Series (Łaziska and Libiąż Beds). Most species were collection from the Duckmantian/Bolsovian Łaziska Beds. The greatest difference between the cordaitalean cuticles from coal and cuticles from mudstones (“in situ” cuticles) is in presence of papillae on cuticles from coal samples. The papillae occur seldom on cuticles isolated from adpressions.
