Influence of stomatal traits on the ecological classification of selected plants in the lateritic belt

 Corresponding author: Sudipa Nag Copyright © 2025 Author(s) retain the copyright of this article. This article is published under the terms of the Creative Commons Attribution Liscense 4.0. Influence of stomatal traits on the ecological classification of selected plants in the lateritic belt Nisha Das 1 and Sudipa Nag 2, * 1 B.Sc. Botany Student, Rampurhat College, Birbhum, West Bengal, India 2 Associate Professor, Botany Department, Rampurhat College, Birbhum, West Bengal, India World Journal of Advanced Research and Reviews, 2025, 26(02), 3995–4012 Publication history: Received on 19 April 2025; revised on 25 May 2025; accepted on 27 May 2025 Article DOI: https://doi.org/10.30574/wjarr.2025.26.2.2055 Abstract Stomatal morphology and distribution exhibit remarkable diversity across monocot and dicot species, reflecting their evolutionary adaptations to varied ecological niches. This study investigates the stomatal diversity in selected taxa, highlighting its taxonomic significance and functional implications. In monocots, the presence of brachyparacytic stomata in Acorus calamus represents a primitive trait, while the tetracytic condition in Kaempferia galanga and Tradescantia spathacea suggests advanced gas exchange mechanisms. Hypostomatic species like Costus speciosus and Tradescantia spathacea indicate adaptations for water conservation, whereas amphistomatic taxa such as Acorus calamus, Caladium bicolour, Dracaena marginata, and Kaempferia galanga thrive in humid environments. Dracaena marginata and Polianthes tuberosa exhibit anomocytic stomata, linking them to basal monocot lineages. The unique dumbbell-shaped guard cells of Cynodon dactylon exemplify Poaceae’s optimization of transpiration control. Dicot species exhibit a spectrum of stomatal adaptations aligned with their ecological conditions. Bryophyllum calycinum demonstrates anisocytic stomata along with CAM metabolism, enhancing water-use efficiency. Calotropis procera displays diverse stomatal types, suggesting specialized xeric adaptations. Anomocytic stomata in Euphorbia neriifolia, Rauwolfia serpentina, and Tabernaemontana divaricata indicate a basal evolutionary trait balancing transpiration. Hypostomatic conditions in Ficus microcarpa and Rauwolfia serpentina promote water conservation, while sunken stomata in Nerium oleander and Ficus microcarpa reduce transpiration. Additionally, stomatal multiplicity in both dicots and monocots were observed. Novel types, including laterocytic, stephanocytic, diacytic, cyclocytic, amphipseudoholoparacytic, stomata were also reported. Scanning electron microscopy revealed sunken stomata and non-contiguous stomatal clusters. These findings underscore the interplay of genetic and environmental factors in stomatal development. Further molecular and physiological studies will help us to understand how stomatal diversity evolved and adapted. This will improve our knowledge of plant responses to climate change and water-use efficiency. Keywords: Stomatal Diversity; Evolutionary Adaptations; Taxonomic Significance; Environmental Factors; Ecological Condition 1. Introduction Stomata, the microscopic pores on plant leaf surfaces, play a pivotal role in regulating gas exchange between plants and the atmosphere [1]. This exchange is essential for photosynthesis, where carbon dioxide (CO2) is taken up, and in transpiration, where water is released [2]. The efficiency of these processes is significantly influenced by stomatal World Journal of Advanced Research and Reviews, 2025, 26(02), 3995–4012 3996 density (SD), stomatal index (SI), and stomatal morphology as key determinants of plant adaptation to diversified environmental condition [3]. Stomata first emerged between the late Silurian and early Devonian periods [4], marking a crucial milestone in plant evolution. According to few scientists [4], the fundamental structure of stomata has remained remarkably consistent over approximately 400 million years, making it one of the most evolutionarily stable features of land plants. Among modern plants, the earliest stomata are observed in Bryophyta, though they appear only in the sporophytic stage [5]. Vascular plants universally possess abundant stomata [6], with grasses exhibiting the most advanced and efficient form—characterized by their distinctive dumbbell shape. The relationship between stomatal traits and environmental factors is crucial for ecological grouping, which involves classification of plants based on their functional adaptations and ecological roles [7]. Plants adjust stomatal size and density to adapt to different moisture, temperature and light regimes [3]. By analyzing these adjustments, it can be achieved the procedure of obtaining plant’s water use efficiency (WUE) and carbon recycling in response to environmental pressures [2]. Stomatal density is the number of stomata per unit area, and it is a critical parameter affecting the rate of gas exchange. Higher stomatal density can potentially increase CO2 uptake for photosynthesis, but it also raises the risk of water loss through transpiration [8]. Therefore, plants in water-stress condition often exhibit lower stomatal densities as an adaptation to conserve water. Stomatal index (SI) represents the proportion of stomata relative to epidermal cells and provides a normalized measure of stomatal abundance. This index helps to account for variations in epidermal cell size, offering a more robust comparison of stomatal frequency across different species or environmental conditions [9]. Stomatal morphology, including pore size and shape, also influences gas exchange efficiency [1]. Smaller, more numerous stomata can respond more quickly to environmental changes, permeating for finer control over transpiration and CO2 uptake. On the other hand, larger stomata may facilitate higher gas exchange rates under favourable conditions [8]. Light intensity and quality are significant factors influencing stomatal development and function. Plants grown under high light intensity often exhibit higher stomatal densities to support increased photosynthetic rates [2]. The spectral composition of light also plays a role, with blue light, for instance, affecting stomatal formation and nutrient absorption in plants [10]. Water availability is a primary driver of stomatal adaptation. In drought-prone environments, plants tend to develop lower stomatal densities and smaller stomatal sizes to minimize water loss. The ability to close stomata rapidly in response to water stress is also a crucial adaptation for survival in arid conditions [11]. Carminati et al.,2020 [12] showed that soil water availability is a key factor in controlling stomatal response to drought. Atmospheric CO2 concentration also influences stomatal development [13]. Elevated CO2 levels can lead to decreased stomatal density, as plants require fewer stomata to capture sufficient CO2 for photosynthesis. This response can have implications for plant water use efficiency and overall ecosystem productivity. Ecological grouping based on stomatal traits involves classifying plants into functional groups that share similar adaptations to specific environmental conditions [4]. This approach helps to understand the ecological roles of different plant species and predict their responses to environmental changes [14]. Plants can be broadly classified into hydrophytes (aquatic plants), mesophytes (plants in moderately moist environments), and xerophytes (plants in arid environments). Hydrophytes often have fewer stomata, primarily on the upper leaf surface, while xerophytes exhibit adaptations such as sunken stomata and thick cuticles to reduce water loss. Mesophytes typically have a moderate stomatal density and distribution, reflecting their adaptation to balanced water availability. Understanding the extent to which plants can modify their stomatal traits in response to short-term environmental changes is essential for predicting their resilience to climate change [10]. Studies on stomatal plasticity and acclimation can reveal the limits of adaptation and identify species that are more vulnerable to environmental stress. Developing mechanistic models that integrate stomatal physiology, morphology, and environmental factors is crucial for predicting plant responses at larger scales [14]. These models can help to assess the impact of climate change on plant productivity, water use efficiency, and ecosystem dynamics. Considering the above views, the present work has been done with the following Objectives: World Journal of Advanced Research and Reviews, 2025, 26(02), 3995–4012 3997 • To Analyze Stomatal Adaptations: Explore how stomatal density, index, and morphology contribute to the adaptation of plants in lateritic soil conditions. • To Understand Ecological Grouping: Examine the role of stomatal traits in classifying plants into specific ecological groups within the lateritic belt. • To Investigate Environmental Responses: Study how stomatal characteristics influence plant responses to abiotic stresses typical of lateritic regions, such as low water retention and nutrient deficiency. • To Support Conservation Efforts: Provide insights into the ecological roles of plant species in lateritic ecosystems to aid in biodiversity conservation. 2. Materials and Methods 2.1. Plant material Leaves from 7 dicotyledons and 8 monocotyledons species (Table 1) were collected randomly during the present investigation from plants grown in Lateritic belt of Birbhum District, West Bengal, India. Table 1 Plant Species Selected for the study Monocotyledons Dicotyledons Species Family Species Family Acorus calamus L. Araceae Bryophyllum calycinum Salisb. Crassulaceae Caladium bicolor (Aiton) Vent Araceae Calotropis procera (Aiton) W.T.Aiton Asclepiadaceae Costus speciosus (Koen. ex. Retz.) Sm. Costaceae Euphorbia neriifolia L. Euphorbiaceae Dracaena marginata V and. ex L. Asparagaceae Ficus microcarpa L. Moraceae Kaempferia galanga L. Zingiberaceae Nerium oleander L. Apocynaceae Polianthes tuberosa L. Amaryllidaceae Rauwolfia serpentina (L.) Benth. ex Kurz · Apocynaceae Tradescantia spathacea Sw. Commelinaceae Tabernaemontana divaricata (L.) R. Br. ex Roem. & Schult. Apocynaceae Cynodon dactylon (L.) Pers. Poaceae 2.2. light microscopy Leaves are labelled in different microscope slides. Young and mature leaves were fixed in FAA (Formalin-AceticAlcohol) and stored in 70% ethanol. Epidermal peels were taken from the central leaf portion. Dry specimens were boiled in 5% KOH for 15 minutes before peeling. Peels were stained with 1% safranin in 50% ethanol and mounted in 50% glycerol. Data was based on 3 observations; stomatal size was measured from 4 observations on 4 samples per taxon. Stomata area was calculated as πr2 (π = 3.14, r = mean radius of the field). Observed under LABOMED VISION 2000 L.E.D compound light microscope at 100x or 400x magnification. For leaves with thick waxy deposits, the waxy layer was removed by treating the leaves with acetone or chloroform before applying the replica method. The methodology involves calibrating the ocular micrometer and then using it to measure Stomatal density, stomatal index and length & breadth of stomatal aperture etc. 2.3. Calibration of Ocular Micrometer The calibration process is essential to ensure accurate measurements. The following steps are involved: Stage Micrometer Placement: A stage micrometer (SM), which has a calibrated scale, is placed on the microscope stage. A typical stage micrometer has 100 divisions within 1 mm, so each division equals 0.01 mm (10 µm). World Journal of Advanced Research and Reviews, 2025, 26(02), 3995–4012 3998 The stage micrometer is focused under a specific magnification (e.g., 40x). An ocular micrometer (OM) is placed in one of the microscope's eyepieces. While observing through the eyepieces, the scale of the ocular micrometer is aligned with the scale of the stage micrometer. The number of divisions on the ocular micrometer that coincide with a known number of divisions on the stage micrometer is counted. We found 7.0 divisions of the ocular micrometer align with 10 divisions of the stage micrometer, then 1.0 OD divisions = 10/7X0.01mm =0.0143mm. 2.4. Stomatal Density The number of stomata within a known area (defined by the ocular micrometer) is counted. Multiple fields of view were counted, and the average stomatal density was calculated as stomata per mm². 2.5. Stomatal Index The number of stomata and the total number of epidermal cells (including stomata) within a known area were counted. The stomatal index is then calculated using the formula: Stomatal Index = Stomata / (Epidermal cells + Stomata)X100 2.6. Stomatal and Cell Dimensions The length and width of stomatal apertures, guard cells were measured using the calibrated ocular micrometer. Multiple measurements were taken for each structure, and the average dimensions were calculated. 2.7. Stomatal and epicuticular wax morphology visualized by scanning electron microscopy (SEM) Field Emission Scanning Electron Microscopy (FESEM) was used to study the topography of the objects, and it worked with electrons rather than light sources (Fig.9). The dehydrated isolated epidermal peels from different plant species collected from field survey were observed under a FESEM (Zes Gemini 2, Germany) after a gold coating using an ion sputter (Quorum Sputter Coaters and SEM and TEM Carbon Coaters, UK). The field emission source generates electrons, which were accelerated toward a high electrical field gradient. 2.8. Study Area The study was conducted in the Rampurhat sub division, located in the lateritic belt of Birbhum District, West Bengal (Fig.1). The area is characterized by seasonal climatic variations and diverse vegetation, including dicot and monocot plants. 2.9. Statistical analyses All data were analyzed statistically and standard error (SE) were measured as per Panse and Sukhatme (1967) [15] Figure 1 Rampurhat Sub division of Birbhum, West Bengal 2.10. Identification of stomatal complexes We categorized the types of stomata, their distribution patterns, and their orientation based on the available literature [1,7,23,] World Journal of Advanced Research and Reviews, 2025, 26(02), 3995–4012 3999 3. Results and discussion 3.1. Monocots plant material The monocot taxa studied (Fig. 2) along with their stomatal features were summarized in Table 2. The arrangement of the families and the taxa under each family is described alphabetically. The diversity of Monocot taxa studied was displayed in fig. 4 and the stomatal density and stomatal index was graphically represented in fig. 3. Figure 2 Leaf shapes in some monocots with their family [(A) Acorus calamus (Araceae) , (B) Caladium bicolor (Araceae) , (C) Costus speciosus (Costaceae) (D) Dracaena marginata (Asparagaceae) , (E) Kaempferia galanga (Zingiberaceae) , (F) Polianthes tuberosa L. (Amaryllidaceae) , (G) Tradescantia spathacea (Commelinaceae), (H) Cynodon dactylon (L.) Pers.( Poaceae)] World Journal of Advanced Research and Reviews, 2025, 26(02), 3995–4012 4000 Table 2 Monocot taxa studied and their stomatal features No. Species Family Stomata Type Density (stomata / mm²) in lower surface Index (%) in lower surface Adaptation Group 1 Acorus calamus Araceae Brachyparacytic Amphistomatic 384.8±1.2 28.59±1.4 Aromatic adaptations favour moderate water regulation Semiaquatic 2 Caladium bicolor Araceae Paracytic Amphistomatic 249.01±1.9 18.5±2.5 Advantageous in shaded, humid environments Acaulescent herb, 3 Costus speciosus Costaceae Paracytic, Hypostomatic 273.2±.25 20.3±2.2 The condition in Costus speciosus is indicative of advanced specialization Tropical herb 4 Dracaena marginata Asparagaceae Anomocytic Amphistomatic 363.4±1.33 27.3±1.3 Xerophytic Succulent shrubs 5 Kaempferia galanga Zingiberaceae Tetracytic Amphistomatic 134.2±1.4 10.1±.55 The tetracytic configuration facilitates water-use efficiency and carbon assimilation Tropical aromatic herb 6 Polianthes tuberosa Amaryllidaceae Anomocytic Amphistomatic 255±1.77 19.9*±1.33 Wide adaptability Tropical and temperate herb 7 Tradescantia spathacea Commelinaceae Tetracytic Hypostomatic 215.09±1.4 16.43±2.6 advanced among stomatal types Succulent herb 8 Cynodon dactylon Poaceae Paracytic Amphistomatic 297.87±1.3 22.13±1.6 Adapted in open areas where there are frequent disturbances such as grazing, flooding, and fire. Warm temperate to tropical herb World Journal of Advanced Research and Reviews, 2025, 26(02), 3995–4012 4001 3.2. Monocot taxa studied and their stomatal features The diversity of monocot taxa studied was displayed in Fig. 4 and their diversity and index was graphically represented in Fig 3. 3.2.1. Family: Acoraceae (Acorus calamus): It had been found that the plant was semi-aquatic herb, amphistomatic, brachyparacytic with one pair of subsidiary cells. The guard cells were oriented parallel to the longitudinal axis of the leaf. Epidermal cells were nearly isodiametric in outline. The Brachyparacytic stomata were considered evolutionarily primitive within monocots. The amphistomatic condition was advantageous in shaded, humid environments where light penetration and water availability influenced stomatal functionality and intermediate evolutionary state between terrestrial and aquatic adaptations. 3.2.2. Family: Araceae (Caladium bicolour) The result showed that the plant is a acaulescent herb and stomata were paracytic and amphistomatic. Guard cells were elongated and kidney-shaped with thick inner and outer ledge. Subsidiary cells were almost rectangular with thin walls. Epidermal cells were pentagonal & hexagonal with thick walls. Guard cell contents were prominent and granular materials. The amphistomatic condition was advantageous in shaded, humid environments where light penetration and water availability influence stomatal functionality. 3.2.3. Family: Costaceae (Costus speciosus) The study revealed that the adaxial epidermis of the plant composed of single-layered polygonal cells with paracytic stomata, Abaxial epidermis consised of bluntly angled epidermal cells. The hypostomatic condition in Costus speciosus was indicative of advanced specialization. The stomatal type in Costus speciosus underscored evolutionary advancements toward optimizing physiological processes in tropical ecosystems. 3.2.4. Family: Asparagaceae (Dracaena marginata) The plant was a succulent shrub, and amphistomatic, anomocytic stomata type i.e. without any subsidiary cells. The anomocytic stomatal type represented a basal state, linking Dracaena marginata to early monocot lineages allowing the species to thrive in challenging xerophytic habitats. 3.2.5. Family: Zingiberaceae (Kaempferia galanga) The epidermal cells were large and hexagonal to round in shape. Lamina was amphistomatic, with straight to curved epidermal cell wall. Stomata were tetracytic (four annexed cells per stomatic apparatus) and stomatal index was the highest in abaxial surface of K. galanga. The tetracytic stomatal arrangement in Kaempferia galanga signified a derived condition compared to simpler stomatal types (e.g., anomocytic or paracytic), showcasing advanced regulatory mechanisms,reflects the adaptation of the species to tropical climates. 3.2.6. Family: Amaryllidaceae (Polianthes tuberosa) It was found that the plant exhibited anomocytic stomata, i.e., the stomata lack distinct subsidiary cells. Scanning electron microscopy (SEM) revealed that stomata were sunken in the epidermis (Fig. 3.7 C). The stomatal type in Polianthes tuberosa highlighted the retention of primitive features, demonstrating how plants with basal traits could thrive and adapt in cultivated and natural settings. 3.2.7. Family:Commelinaceae (Tradescantia spathacea) The stomatas were hypostomatic type, which indicated that they were only located on the abaxial face of the leaves of Tradescantia spathacea. They were in tetracytic condition. The tetracytic stomata arrangement allowed for dynamic stomatal movement, enabling rapid response to environmental fluctuations. 3.2.8. Family:Poaceae (Cynodon dactylon) The study revealed that abaxial intercostals long leaf cells had thin, sinuous walls. The guard cells were dumbbellshaped, while the subsidiary cells ranged from low to high dome-shaped. The number of rows of long cells between two costal zones varied from 6 to 10, whereas the number of stomatal rows between two costal zones ranged from 1 to 3. World Journal of Advanced Research and Reviews, 2025, 26(02), 3995–4012 4002 Figure 3 Monocot taxa studied and their stomatal features in graphical representation 3.3. Monocot taxa stomata types A B C D E F G H Figure 4 Diversity of monocot stomata [(A) Acorus calamus (Acoraceae), Paracytic stoma with a pair of lateral subsidiary cells, with oblique cell walls, In this “paracytic-oblique” type, the lateral subsidiary cells are only slightly smaller than pavement cells and clearly belong to the adjacent cell file. (B) Caladium bicolor (Araceae) , Brachyparacytic stomata with oblique cell walls (C) Costus speciosus (Costaceae) Paracytic stomata with oblique cell walls (D) Dracaena marginata (Asparagaceae) Anomocytic stomata (lacking subsidiary cells) , (E) Kaempferia galanga (Zingiberaceae) , Tetracytic stomata (F) Polianthes tuberosa L. (Amaryllidaceae) , Anomocytic stomata, (G) Tradescantia spathacea (Commelinaceae) Tetracytic stomata, (H) Cynodon dactylon (L.) Pers.( Poaceae) paracytic stomata with dumbbell shaped subsidiary cells.] World Journal of Advanced Research and Reviews, 2025, 26(02), 3995–4012 4003 3.4. Dicotyledons plant material Figure 5 Dorsiventral Leaves in some dicots [A. Bryophyllum calycinum (Crassulaceae) B. Calotropis procera (Asclepiadaceae), C. Euphorbia neriifolia (Euphorbiaceae), D. Ficus microcarpa (Moraceae ), E. Nerium oleander (Apocynaceae), F. Rauwolfia serpentina (Apocynaceae ), G. Tabernaemontana divaricata (Apocynaceae )] Table 3 Dicot taxa studied and their stomatal features No . Species Family Type Density (stomata /mm²) in lower surface Index (%) Adaptation Group 1 Bryophyllum calycinum Crassulaceae Anisocytic, Amphistomati c 50.54 ±1.2 11.11±2. 3 Crassulacean Acid Metabolism (CAM), a physiological adaptation Succulent xerophytes 2 Calotropis procera Asclepiadaceae Anomocytic, Amphistomati c 90.8±2.1 20±3.3 Xeric and semi-xeric Nonsucculent xerophytes 3 Euphorbia neriifolia Euphorbiaceae Anomocytic, Hypostomatic 22.7±1.0 5±2.2 During high temperature s or dry conditions, Succulent xerophytes World Journal of Advanced Research and Reviews, 2025, 26(02), 3995–4012 4010 The existence of stomata with large meristemoids in Bryophyllum suggests prolonged meristematic activity before guard cell differentiation. This could indicate a prolonged developmental phase or an anomaly in the regulatory mechanisms which controls meristem size [28]. Larger meristemoids might contribute to increased stomatal production under favourable conditions is the reflection of developmental lag in epidermal cell division. Figure 10 Abnormal stomata and guard cells of leaves of studied species of the monocots and dicots [A.Non-contiguous stomata in Tabernaemontana, B. Paired stomata in Rauwolfia C.Single guardcell in Tabernaemontana, D. Single guardcell in Dracaena, E. Single guardcell in Polyanthes, F. Single guardcell in Rauwolfia, G.Abnormal guardcell in Calotropis, H. Non-contiguous stomata in Rauwolfia, I. Arrested stomata in Calotropis, J. Stomata with large meristemoid in Bryophyllum, K. Paired stomata in Tabernaemontana] 4. Conclusion The diversity of stomatal types across monocot families reflects evolutionary adaptations to varied ecological conditions. The presence of brachyparacytic stomata in Acorus calamus represents an evolutionarily primitive trait, whereas the tetracytic condition in Kaempferia galanga and Tradescantia spathacea suggests advanced regulatory mechanisms for efficient gas exchange. Hypostomatic species like Costus speciosus and Tradescantia spathacea indicate specialization towards water-use efficiency, while amphistomatic taxa (Acorus calamus, Caladium bicolour, Dracaena marginata, Kaempferia galanga) exhibit traits favouring humid environments. The anomocytic stomata in Dracaena marginata and Polianthes tuberosa link these taxa to basal monocot lineages, underscoring their primitive yet functional adaptations. Meanwhile, Cynodon dactylon exemplifies the unique stomatal morphology of Poaceae, with dumbbellshaped guard cells and dome-like subsidiary cells, optimizing transpiration control in grasses. These variations highlight the intricate evolutionary pathways of monocots, demonstrating how stomatal structure has played a crucial role in their success across different habitats. On the other hand, the presence of anisocytic stomata in dicot plants like Bryophyllum calycinum, coupled with CAM metabolism, represents a dual strategy for optimizing water-use efficiency in arid environments. Similarly, Calotropis procera exhibits a remarkable diversity of stomatal types, with irregularly scattered and sunken stomata, suggesting fine-tuned physiological mechanisms for survival in xeric and semi-xeric habitats. Sunken stomata in Nerium oleander and Ficus microcarpa, often accompanied by cuticular thickening and stomatal crypts, further enhance drought tolerance by reducing transpiration. Overall, the diversity in stomatal types, subsidiary cell arrangements, and stomatal distribution patterns among these dicot species highlights significant evolutionary advancements toward ecological specialization. Detailed investigation of both adaxial and abaxial World Journal of Advanced Research and Reviews, 2025, 26(02), 3995–4012 4011 epidermal surfaces under light microscope revealed five major stomatal types and eleven subtypes. Scanning electron microscopy (SEM) revealed that stomata were sunken in the epidermis. The diverse array of stomatal abnormalities observed in the present study underlines the complexity of stomatal development and variations across different species suggest an interplay of genetic and environmental factors influencing epidermal differentiation. Thus, stomatal traits in monocot and dicots influenced a great in adopting different environmental variations with different ecological classification. Compliance with ethical standards Acknowledgments We would like to thank Prantik Gangopadhyay, Assistant DST PURSE, Visva Bharati, Santiniketan, Birbhum, West Bengal, India for helping FESEM analysis of the epidermal micromorphology. Disclosure of conflict of interest There is no conflict of interest. References [1] Metcalfe, C. R., and L. Chalk. 1979. *Anatomy of the Dicotyledons: Systematic Anatomy of the Leaf and Stem. * Vol. 1. 2nd ed. Oxford: Clarendon Press. [2] Stace, C. 1980. *Plant Taxonomy and Biosystematics. * London: Edward Arnold Publisher Ltd. [3] Ticha, I. 1982. "Photosynthetic Characteristics During Ontogenesis of Leaves, Stomata Density and Sizes." *Photosynthetica* 16: 375–471. [4] Edwards, D., E. M. Morel, F. Paredes, D. G. Ganuza, and A. Zúñiga. 2001. "Plant Assemblages from the Silurian of Southern Bolivia and Their Palaeogeographic Significance." *Botanical Journal of the Linnean Society* 135 (3): 229–250. https://doi.org/10.1006/bojl.2000.0418. 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