Palynological analysis of representatives of Hippeastrum Herb. (Amaryllidaceae: Amaryllidoideae), with implications for systematics

Palynological analysis of representatives of Hippeastrum Herb. (Amaryllidaceae: Amaryllidoideae), with implications for systematics | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Palynological analysis of representatives of Hippeastrum Herb. (Amaryllidaceae: Amaryllidoideae), with implications for systematics Renata Suzano Cândido, Rosana Conrado Lopes, Cláudia Barbieri Ferreira Mendonça, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6785943/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 5 You are reading this latest preprint version Abstract Native to the American continent, the genus Hippeastrum Herb. is distributed from Mexico to Argentina. In Brazil, it is represented by approximately 30 species. This study examined the pollen grains of 20 taxa within the genus to determine whether differences in pollen morphology could support a clearer delimitation and characterization of subgenera and subordinate species. Pollen grains were treated with 40% lactic acid, measured, described, and photomicrographed. Non-acetolyzed pollen grains were examined using scanning electron microscopy. The analyzed taxa have large to very large pollen grains, shed as monads, monosulcate, with an elliptical shape in polar view and a reticulate or retipilate sexine. Species with a retipilate sexine could be distinguished based on the characteristics and organization of pila, whereas those with a reticulate sexine could be differentiated by murus, lumen, and columella characters. Thus, it was possible to develop a palynological key. Qualitative and quantitative analyses revealed differences in pollen morphology among species. However, pollen characters were generally uniform among subgenera. Overall, pollen characters proved informative for delimiting and describing species within the genus Hippeastrum . Asparagales Morphology Pollen Taxonomy Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 1. Introduction Asparagales is a monophyletic order comprising 14 families, 1122 genera, and approximately 36,265 species. One of the families belonging to Asparagales is Amaryllidaceae, which includes about 73 genera and 1605 species distributed in three subfamilies (Agapanthoideae, Allioideae, and Amaryllidoideae). The grouping of these subfamilies is supported by a shared morphological character: the presence of a scapose umbellate inflorescence surrounded by a pair of bracts (Stevens 2001, APG IV 2016). Amaryllidoideae comprises 15 tribes, 6 of which are native to the American continent, including the widely distributed tribe Hippeastreae. Hippeastrum Herb. is one of the few genera within the tribe that has been found to be monophyletic. It was initially described as comprising 15 species, which, according to Herbert (1821), exhibited considerable morphological variation. In Brazil, Hippeastrum is represented by approximately 39 species, 31 of which are endemic. These species occur in all vegetation types, from forests to grasslands, and on a variety of substrates (Dutilh et al. 2025). A recent investigation analyzed relationships among Brazilian species to assess the monophyly of Hippeastrum and its subgenera based on DNA sequence data. The study found no correlation between the traditionally proposed subgenera, concluding that they are polyphyletic (Oliveira 2012). Few palynological studies have been conducted on species of Hippeastrum . Alves-Araújo and Santos (2007) conducted a palynological analysis of two species of Hippeastrum occurring in northeastern Brazil. The authors stated that the studied taxa exhibited valuable characters that can aid in their taxonomic delimitation. In another study, five species of Hippeastrum found in the restingas of Rio de Janeiro State were successfully distinguished based on pollen morphology, particularly exine ornamentation (Candido et al. 2013). Although palynological studies on Hippeastrum are scarce, existing research consistently emphasizes the importance of pollen characters in the taxonomy of the genus. Therefore, this study aimed to evaluate whether species of Hippeastrum exhibit differences in pollen morphology that can contribute to a more precise delimitation and characterization of the taxa under investigation. 2. Material and methods 2.1. Pollen material This study examined pollen grains of 20 taxa within the genus Hippeastrum (Appendix 1). The pollen material was obtained from fertile anthers of flowers at anthesis and/or buds at pre-anthesis, collected from exsiccatae deposited in the following herbaria: HB, HUEFS, IAC, ICN, PACA, R, RB, RFA, UB, and UEC (acronyms according to Thiers, continuously updated). 2.2. Light microscopy For analysis by light microscopy, the material was processed according to the acetolysis method of Raynal and Raynal (1971), which uses 60% lactic acid. This method was chosen because pollen grains exhibited poor resistance to traditional acetolysis. Acetolyzed pollen grains were photomicrographed using a Canon Power Shot G6 digital camera coupled to a Zeiss Axiostar Plus binocular microscope equipped with a 100× objective lens. Microscope slides were deposited in the pollen collection of the Álvaro Xavier Moreira Laboratory of Palynology, Department of Botany, Nacional Museum, Federal University of Rio de Janeiro, Brazil. 2.3. Measurements and statistical analysis For quantitative analysis, measurements were taken from polar and equatorial views. Pollen grains were randomly selected from a minimum of three slides to ensure sample homogenization (Salgado-Labouriau 1973). A total of 25 measurements were performed of the largest and smallest diameters in polar view. Additionally, 10 measurements were taken, when possible, of the polar and equatorial diameters in equatorial view, sexine and nexine thickness, aperture length and width, and reticulum lumen diameter (ornamentation). Acetolyzed pollen grains were measured within three days of preparation to prevent changes in pollen size (Melhem et al. 2003). Variables with 25 measurements were statistically treated to determine the arithmetic mean ( x ), standard deviation of the mean ( s x ), 95% confidence interval (95% CI), and range of variation. The other variables are expressed as arithmetic means. 2.4. Scanning electron microscopy (SEM) Anthers were ground to release pollen grains, which were then dusted in their non-acetolyzed form onto aluminum stubs coated with double-sided carbon tape (Melhem et al. 2003). Samples were transferred to a vacuum pump and metalized with a thin layer (200 Å) of gold-palladium. Subsequently, samples were analyzed using a JEOL JSM-6390LV system at the Invertebrate Electron Microscopy Laboratory, National Museum, Federal University of Rio de Janeiro, and a Zeiss DSM 960 system at the Hertha Meyer Cellular Ultrastructure Laboratory, Institute of Biophysics, Federal University of Rio de Janeiro, Brazil. 2.5. Terminology The terminologies adopted for the description of pollen size, shape, and sexine ornamentation patterns were those of Erdtman (1952) and Punt et al. (2007). 2.6. Multivariate analysis PC-ORD software version 5.31 (McCune & Mefford 2011) was used for exploratory analysis. The pollen characters (8 variables) of each species were organized into a matrix, resulting in better organization of the data and facilitating the generation of graphs. Species names were abbreviated to the first three letters of specific epithets, and pollen characters were abbreviated to their initial letters. The variables included in multivariate analyses were largest diameter (LD), smallest diameter (SD), polar diameter (PD), equatorial diameter (ED), sulcus width (SW), sulcus length (SL), lumen diameter (LD), and muri width (MW). The matrix used for principal component analysis (PCA) and hierarchal cluster analysis was transformed by the square root of x + 0.5 to standardize measurable data. PCA was performed to assess whether the analyzed species could be grouped based on pollen characters. The variance–covariance matrix was generated using the mean values of morphometric data and coordinates on a biplot graph based on Euclidean distances. The results are shown on a two-dimensional plot representing the first and second principal components. Vector loadings on each axis and the total cumulative variance are presented in tables. Hierarchical cluster analysis (HCA) was performed to group species based on similarities in pollen morphology. Two criteria were adopted when analyzing the data: the percentage of information (variables) required to form groups and the final number of groups obtained. A dendrogram was constructed using Euclidean distances (Caccavari et al. 2008) and Ward's linkage method. 3. Results A total of 20 species of the genus Hippeastrum were examined. The palynological description is organized according to the following characters: dispersion unit, polarity, shape, size, aperture type and number, and sexine ornamentation pattern. 3.1. Dispersion unit, polarity, shape, and size All analyzed species of Hippeastrum have pollen grains shed as monads, heteropolar, and elliptical in polar view. In equatorial view, the proximal face is flat (Figs. 1 h, 2 e, 3 g, and 4 i) or convex (Fig. 1 a, h), whereas the distal face is always convex (Figs. 1 g, 1 h, and 3 g). Most of the analyzed species have large pollen grains, with very large grains observed only in H. aulicum , whose largest diameter in polar view was 104.7 µm. In large pollen grains, the lowest value of the largest diameter in polar view was recorded in H. santacatarina (55.9 µm) and the highest in H. calyptratum (91.5 µm). In equatorial view, the largest polar diameter was found in H. aulicum (ca. 54.7 µm) and the smallest in H. angustifolium (ca. 34.8 µm). The largest equatorial diameter was found in H. aulicum (ca. 106.7 µm) and the smallest in H. santacatarina (ca. 52.9 µm) (Tables 1 and 2). 3.2. Aperture All species have monosulcate grains, with long sulci having sharp or rounded ends. The greatest sulcus length was recorded in H. aulicum (ca. 101.2 µm) and the shortest in H. reticulatum (ca. 41.0 µm). The apertural membrane was psilate in H. blossfeldiae (Fig. 1 i), H. brasilianum , H. cipoanum , H. stylosum , and H. vittatum (Table 3) and difficult to distinguish in the other species. 3.3. Sexine stratification and ornamentation patterns The sexine was as thick as the nexine in most species but thicker than the nexine in H. puniceum , H. reticulatum , and H. striatum (Table 3). The exine ornamentation pattern ranged from intectate reticulate to semitectate reticulate. A retipilate sexine was observed in H. elegans (Fig. 2 l), H. puniceum (Fig. 3 i), H. reginae (Fig. 3 l), and H. stapfianum (Fig. 4 e). Pila have a smooth surface and gradually decrease in diameter, with the lumina becoming more densely grouped toward the ends and near the aperture (Figs. 2 k, 3 k), except in H. elegans (Fig. 3 g, h). Pila are connected by the upper part (head), sparsely grouped, forming small strands that create closed meshes resembling a reticulum in some regions (Figs. 2 j, 3 j, 4 k). In H. puniceum , pila are rounded, vary in diameter (Fig. 3 h, i), and are arranged in double rows, with sparse granules in lumina. In H. reginae , sexine bridges join pila at the base (Fig. 3 l), forming strands organized into reticula with well-defined lumina. A reticulate heterobrochate sexine was observed in most species. The reticulate surface with large luminal diameters is restricted to a well-demarcated median band in the pollen grain. The ends exhibit microreticulate ornamentation in H. aulicum (Fig. 1 e, f), H. calyptratum (Fig. 2 d, e), H. papilio (Fig. 3 e), and H. stylosum (Fig. 4 h, i). Muri are narrow (ca. 0.6–1.0 µm) in all taxa, straight in most species and sinuous in H. blossfeldiae (Fig. 1 j), with sparse perforations. Columellae are not apparent or visible in H. aulicum (Fig. 2 g), H. calyptratum (Fig. 2 f), H. glaucescens (Fig. 3 b), H. morelianum (Fig. 3 d), H. papilio (Fig. 3 f), H. striatum (Fig. 4 g), H. stylosum (Fig. 4 j), or H. vittatum (Fig. 4 l). Smaller lumina surrounding larger ones were recorded in H. angustifolium (Fig. 1 c), H. aulicum (Fig. 1 g), H. blossfeldiae (Fig. 1 j), H. brasilianum (Fig. 1 l), H. breviflorum (Fig. 2 c), H. calyptratum (Fig. 2 f), H. morelianum (Fig. 3 d), H. papilio (Fig. 3 f), H. reticulatum (Fig. 4 b), H. santacatarina (Fig. 4 c), H. striatum (Fig. 4 g), and H. stylosum (Fig. 4 j). Lumina may or may not have ornamentation (Figs. 1 j, l, 2 g, i, l, 3 f, 4 b, j). When present, the ornamentation consists of inconspicuous and sparse granules in most species or densely organized and conspicuous granules in H. angustifolium (Fig. 1 c), H. glaucescens (Fig. 3 b), and H. morelianum (Fig. 3 d). 3.4. Pollen key to species of the genus Hippeastrum 1. Sexine retipilate 2. Lumina well-defined, pila joined at the base by exine bridges............................. H. reginae 2. Lumina undefined, pila not joined at the base 3. Pila arranged in double rows........................................................................... H. puniceum 3. Pila not arranged in double rows 4. Pila sparsely clustered forming small strands................................................... H. elegans 4. Pila densely clustered without forming small strands................................. H. stapfianum 1. Sexine reticulate heterobrochate 5. Muri with sparse interruptions, lumina with sparse, inconspicuous granules... H. cipoanum 5. Muri without interruptions 7. Reticulum with large lumina restricted to a well-demarcated median band of the pollen and ends with microreticulate ornamentation 8. Lumina > ca. 6.0 µm............................................................................... H. calyptratum 8. Lumina < 5.0 µm 9. Pollen very large.......................................................................................... H. aulicum 9. Pollen large.................................................. H. angustifolium , H. papilio , H. stylosum 7. Reticulum with lumina gradually decreasing in diameter toward the ends 11. Muri sinuous, presence of smaller lumina surrounding larger ones.............................. ........................................................................................... H. blossfeldiae , H. breviflorum 11. Muri straight 13. Columella apparent........ H. glaucescens , H. morelianum , H. striatum , H. vittatum 13. Columella not apparent.............. H. brasilianum , H. canastrense , H. santacatarina 3.5. Multivariate analyses 3.5.1. HCA Analysis of the relationships between species of Hippeastrum yielded a dendrogram with a linkage level of 14.71%. When 50% of the remaining information (variables) was considered, three groups were identified. Group 1 contained the species H. angustifolium , H. elegans , H. reginae , H. stylosum , H. puniceum , H. reticulatum , H. blossfeldiae , H. breviflorum , H. striatum , H. canastrense , and H. santacatarina . Group 2 comprised the species H. aulicum , H. glaucescens , H. brasilianum , H. papilio , H. cipoanum , H. morelianum , H. vittatum ), and H. stapfianum . Finally, H. aulicum and H. calyptratum formed group 3. An increase in the number of variables analyzed (75% of remaining information) resulted in modifications to the clusters. Group 1 was subdivided into three other groups: 1′ ( H. angustifolium , H. elegans , H. reginae , H. blossfeldiae , H. stylosum , H. breviflorum , H. puniceum ), 2′ ( H. canastrense , H. santacatarina and H. reticulatum ), and 3′ ( H. striatum ). Group 2 comprised groups 4′ ( H. glaucescens , H. brasilianum , H. papilio , H. cipoanum , H. morelianum , and H. vittatum ) and 5′ ( H. stapfianum ). Group 6 remains unchanged and is a repeat of group 3 (Fig. 5 ). 3.5.2. PCA PCA resulted in two principal components, which together explained 75.99% of the total variance. The first component accounted for 63.06% of the variance, and the second 12.93%. Sulcus length, largest diameter, and equatorial diameter were the most influential variables for the first principal component. By contrast, lumen diameter, murus width, and polar diameter contributed the most to the second principal component (Table 4). On axis 1, species were broadly distributed across both positive and negative sides. Along axis 2, most species clustered on the positive side, with the exception of H. stapfianum , which was isolated at the extreme negative end due to its low lumen diameter. The dispersion of species on the PCA biplot may facilitate the identification of groups, even though species differed by only a few morphological characters (Fig. 6 ). Initially, species were separated into three groups. The first group, located near the negative pole of axis 1, comprised H. angustifolium , H. canastrense , H. reginae , H. reticulatum , H. santacatarina , and H. striatum . The group was less dispersed and characterized by low values of sulcus length. The second group, positioned to the left of axis 2, was formed by H. elegans , H. blossfeldiae , H. breviflorum , H. morelianum , H. stylosum , and H. vittatum . These species clustered together because of their similar values of equatorial diameter and largest diameter. The third group was plotted to the right of axis 2, composed of H. brasilianum , H. cipoanum , H. glaucescens , H. papilio , and H. puniceum . This group was more internally dispersed and distinct from the others, primarily due to differences in polar diameter. H. aulicum and H. calyptratum were polarized in relation to the others because they exhibited the highest values of largest diameter, smallest diameter, polar diameter, equatorial diameter, and sulcus length among the sample (Fig. 6 ). 4. Discussion The findings allowed developing a pollen key, initially divided into two major groups based on exine ornamentation. Subsequent taxon identification was guided by ornamentation elements and pollen size. Other aspects of pollen morphology, such as aperture number and type (monosulcate), dispersion unit (monads), and polarity (heteropolar) were consistent across the sample and therefore not included in the key. Some species could not be differentiated using the proposed key due to shared pollen characteristics, including H. angustifolium , H. papilio , and H. stylosum ; H. blossfeldiae and H. breviflorum ; H. glaucescens , H. morelianum , H. striatum , and H. vittatum ; and H. brasilianum , H. canastrense , H. reticulatum , and H. santacatarina . Pollen size, however, was useful for separating H. aulicum (very large) from the other species, which exhibited large pollen. These results are partially consistent with those of Alves-Araújo and Santos (2007), who described large to very large pollen grains in H. puniceum and H. stylosum . In the current study, both species were classified as having large pollen. Exine ornamentation was a key diagnostic character, with retipilate sexine observed in H. elegans , H. puniceum , H. reginae , and H. stapfianum and reticulate sexine identified in the other species, as detailed in the pollen key. According to Walker & Doyle (1975), the architecture of the pollen wall is an important source of phylogenetic information. The evolutionary trends suggested by the authors indicate imperforate tectate exine as the least derived, followed by semitectate and intectate as the most derived. Thus, among the studied species, H. elegans , H. puniceum , H. reginae , and H. stapfianum had more derived pollen grains. Species with retipilate sexine were distinguished based on pilum characteristics and organization. Among those with a reticulate exine, H. aulicum , H. calyptratum , H. papilio , and H. stylosum were notable for their large lumina, which were confined to the well-defined median region of the pollen grain, and poles with microreticulate pattern. The remaining taxa with reticulate exine were differentiated by features of their muri, lumina, and columellae. This dimorphic pattern of reticulum organization was first reported by Erdtman (1952) in some genera of the family Amaryllidaceae, such as Hymenocallis Salisb. and Lycoris Herb. Meerow and Dehgan (1985) also observed a similar pattern, characterized by a coarse reticulum in the median region and a finer structure at the extremities (dimorphic reticulum), in Caliphruria Herb. and Eucharis Planch. & Linden. They proposed an evolutionary trend within subgenera of Hymenocallis , suggesting a transition from a dimorphic to a homogeneous reticulum, with the latter representing a derived character. In their investigation of H. stylosum , Alves-Araújo and Santos (2007) found that the heterobrochate reticulate exine formed a psilate-perforate cap in the acute equatorial region, thereby composing a dimorphic reticulum at the extremities of the equatorial region. The present study confirmed the dimorphic pattern in H. stylosum but recorded a different type of ornamentation, namely microreticulate at the extremities (equator) of the pollen. This discrepancy can be attributed to differences in material quality or variations in sample preparation procedures for SEM analysis. Additionally, Alves-Araújo and Santos (2007) reported H. puniceum pollen as having reticulate semitectate exine and transparent tectum; however, the material was not analyzed by SEM. Here, H. puniceum was observed under both light microscopy and SEM. The exine ornamentation was found to be retipilate and intectate, with pila organized in double rows, corroborating the analysis of Candido et al. (2013) for this species. HCA revealed groups composed of species from different subgenera. The scattering of species on the PCA biplot allowed further individualization of taxa. Nevertheless, it was difficult to delimit groups along the ordering axes, given that species shared pollen attributes, even those of different subgenera. These findings raise questions about the current subgeneric classification of Hippeastrum . Most species belonging to Hippeastrum subg. Aschamia (Salisb.) Baker were grouped into the same HCA cluster (Group 1) but were not plotted close to one another on the PCA biplot. The dendrogram revealed that only H. puniceum was separated from the other species of the group. This finding corroborated PCA results, in that H. puniceum was plotted on the positive side of axis 1, far from the other species of the subgenus. This separation was attributed to the high values of largest diameter, equatorial diameter, and sulcus length of H. puniceum compared with other members of the subgenus. Species of the other subgenera of Hippeastrum were dispersed across HCA clusters and the PCA biplot, indicating that Hippeastrum subgenera cannot be delimited by pollen morphology. Species composing Group 2 (HCA) belonged to different subgenera, as follows: Hippeastrum glaucescens and H. morelianum ( H. subg. Caephaleaeon Traub.); H. brasilianum and H. stapfianum ( H. subg. Macropodastrum Baker); H. papilio ( H. subg. Omphalissa (Salisb.) Baker); and H. vittatum ( H. subg. Lais (Salisb.) Baker). The species were quite dispersed on the PCA biplot. Of note, H. stapfianum was plotted far from the other species because of its low lumen diameter. Group 3 (HCA) was formed by H. aulicum and H. calyptratum , both subordinate to the subgenus Omphalissa . They were plotted far from the others on the positive pole of axis 1 (PCA), owing to their high largest diameter, smallest diameter, polar diameter, equatorial diameter, and sulcus length. Although other types of pollen have been described for Amaryllidaceae, such as disulcate aperture ( Amaryllis belladonna L., Crinum L., Nerine Herb, Strumaria Jacq.) with intectate-columellate exine (Erdtman 1952; Meerow and Dehgan 1985; Meerow and Snijman 1998), monosulcate pollen with semitectate exine can be considered a synapomorphy for the family (Dönmez and Isik 2008). The external morphology of H. aulicum and H. morelianum is similar. H. calyptratum and H. aulicum are also similar with regard to leaf morphology and presence of corona distributed in plates (Oliveira 2012). However, the palynological morphology of H. aulicum differed from that of H. morelianum . Pollen of the former species were characterized by very large size, presence of reticulum with large lumina restricted to the median portion of the pollen and ends with microreticulate ornamentation. H. calyptratum was more similar to H. aulicum , differing only in lumen diameter (ca. 6.0 µm), as indicated in the pollen key. Palynological proximity between H. aulicum and H. calyptratum was confirmed by HCA. These species formed group 3 and were located far from the other species on the PCA biplot. Hippeastrum blossfeldiae is known to be morphologically similar to H. striatum (Oliveira 2012). However, pollen analysis revealed some differences between these species. Reticulum muri were sinuous in H. blossfeldiae and straight in H. striatum . Additionally, the species were placed in different groups in AHC, namely H. blossfeldiae in group 1′ and H. striatum isolated in group 3′. On the PCA biplot, the species were plotted far from one another, because H. blossfeldiae exhibited higher values of largest diameter and polar diameter. According to Oliveira (2012), H. brasilianum is very similar to H. elegans . In living specimens, the species can be distinguished by flower color; in herbarium material, by the shape of the floral tube, which is more open in H. brasilianum . H. elegans also exhibits color variation and morphological overlap with H. stapfianum and H. vittatum . In the current study, H. elegans and H. stapfianum differed from H. brasilianum and H. vittatum , as the first two had a retipilate exine. By contrast, H. brasilianum and H. vittatum exhibited a reticulate exine, differing with regard to muri. H. elegans and H. stapfianum differed in pilum organization, as described in the pollen key. PCA showed that the species were widely dispersed along the positive and negative sides of axis 2, influenced by lumen diameter. Hippeastrum puniceum exhibited intermediate forms of external morphology, being sometimes indistinguishable from H. reginae . It usually has a proportionally longer hypanthial tube and distinct color patterns (Oliveira 2012). Both species exhibited retipilate exine, indicative, in theory, of their proximity. However, they were distinguished in the pollen key by differences in ornamentation elements. Such proximity was confirmed by HCA, in which species were placed in the same group (1′), and had similar values of polar diameter and sulcus length. It should be noted that the pollen morphology of the following species was characterized for the first time in the present study: Hippeastrum angustifolium , H. aulicum var. glaucophylum , H. blossfeldiae , H. brasilianum , H. breviflorum , H. calyptratum , H. canastrense , H. cipoanum , H. elegans , H. stapfianum , H. morelianum , H. papilio , H. reginae , H. santacatarina , and H. vittatum . The results of this study hold taxonomic and systematic value, as they allowed distinguishing some taxa by specific pollen characters, including closely related species. However, the findings do not corroborate the delimitation of the subgenera proposed for Hippeastrum. 5. Conclusion In conclusion, the pollen grains of species of Hippeastrum exhibit notable morphological variation, particularly in exine ornamentation, sculpture elements, and pollen size, which allowed distinguishing some taxa. Two main groups were identified regarding exine ornamentation: retipilate and reticulate. Therefore, pollen morphology provides valuable information for improving species circumscription and characterization. Palynological data do not support the current delimitation of Hippeastrum into subgenera. Declarations Conflict of interest No potential conflict of interest was reported by the authors. Author contributions All authors contributed to the study conception and design. Material preparation, data collection, and analysis were performed by VGE, CBFM, and RCL. All authors commented on previous versions and read and approved the final manuscript. Acknowledgments VGE and CBFM are grateful to the Brazilian National Council for Scientific and Technological Development (CNPq) for the research grants (grants Nos. 307276/2023-6 and 311618/2021-9). All authors thank the Rio de Janeiro State Research Foundation (FAPERJ) (grant Nos. E-26/210674/2023 and 260003/015254/2021). We also thank the herbarium curators and staff for providing access to their collections. References Alves-Araújo A and Santos FAR (2007) Caracterização palinológica das espécies de Amaryllidaceae sensu stricto ocorrentes no nordeste brasileiro. Acta Bot Bras 21:967-976 APG IV (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants. Bot J Linn Soc 181: 1-20 Caccavari MA, Naab OA, Tamame MA (2008) Palynology And Physicochemical Characteristics Of Three Unifloral Honey Types From Central Argentina. Span J Agric Res 6: 566-576 Candido RS, Fourny ACS, Gonçalves-Esteves V, Lopes RC (2013) Hippeastrum species in áreas of restinga in the state of the Rio de Janeiro, Brazil: pollen characters. Acta Bot Bras 27(4):661-668 Dönmez EO and Isik S (2008) Pollen morphology of Turkish Amaryllidaceae, Ixioliriaceae and Iridaceae. Grana 47(1):15-38 Dutilh JHA, Campos-Rocha A, Oliveira RS, Garcia N, Streher NS, Giussani LM, Semir J (in memoriam), Meerow AW, Sassone AB. Hippeastrum in Flora e Funga do Brasil. Jardim Botânico do Rio de Janeiro. Available at:. consulta.publica.uc.citacao.acesso.em06 mai. 2025 Erdtman G (1952) Pollen morphology and plant taxonomy - Angiosperms. Almqvist & Wiksell, Stockholm Herbert WH (1821) An Appendix. Bot. Mag. James and Ridway Sons. London. McCune B, Mefford MJ (2011) PC-ORD. Multivariate Analysis of Ecological Data. Version 6. MjM Software, Gleneden Beach, Oregon, U.S.A. Meerow AW and Dehgan B (1985) The auriculate pollen grain of Hymenocallis quitoensis Herb. (Amaryllidaceae) and its systematic implications. American Journal of Botany 72(4):540-547 Meerow AW and Snijman DA (1998) Amaryllidaceae. In K. Kubitzki (ed.). The families and genera of vascular plants. Monocotyledons – Lilianae (except Orchidaceae). Hamburg, Germany. 83-110p Melhem TS, Cruz-Barros MAV, Corrêa AMS, Makino-Watanabe H, Silvestre-Capelato MSF, Gonçalves-Esteves V (2003) Morfologia polínica em plantas de Campos do Jordão (São Paulo, Brasil). Bol. Inst. Bot. 16:1-104. Oliveira RS (2012) O gênero Hippeastrum Herb. (Amaryllidaceae) no Brasil: evidência de evolução reticulada e análise de caracteres florais. Campinas, São Paulo, Universidade Estadual de Campinas, Tese Punt W, Blackmore S, Nilsson S, Le Thomas A (2007) Glossary of pollen and spore terminology. Rev. Paleobot. Palynol. 143: 1-81 Raynal A and Raynal J (1971) Une technique de preparation des grains de pollen fragilis. Adansônia 11(1): 77-79 Salgado-Labouriau ML (1973) Contribuição à palinologia dos cerrados. Academia Brasileira de Ciências. Rio de Janeiro Stevens P F (2001). Angiosperm Phylogeny Website. Version 14, July 2017 [and more or less continuously updated since]." Acessed 20 july 2024.http://www.mobot.org/MOBOT/research/APweb/.Traub, H. P. 1963. The genera of Amaryllidaceae. 8pp. Ed. 1. The American Plant. LifeSociety. Jolla, California. Thiers B Index herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden's Virtual Herbarium. http://sweetgum.nybg.org/ih/.2023. Walker JW and Doyle JA (1975) The bases of angiosperm phylogeny: Palynology. Ann. Missouri Bot. Gard. 62: 664-723. Tables Table 1 Measurements (µm) of pollen grains in equatorial view of Hippeastrum species. Abbreviation Largest Diameter Smallest Diameter Species Range x ± sx CI 95% Range x ± sx CI 95% H. angustifolium ang 52.5-60.0 56.2±0.4 55.3-57.1 32.5-42.5 38.4±0.5 37.4-39.5 H. aulicum aul 100.0-110.0 104.7±0.6 103.5-105.9 50.0-60.0 53.0±0.6 51.8-54.2 H. blossfeldiae blo 65.0-68.7 66.7±0.3 66.1-67.4 37.5-43.7 41.5±0.3 40.8-42.2 H. brasilianum bra 70.0-75.0 71.3±0.2 70.7-71.8 46.3-56.3 50.2±0.5 49.2-51.2 H. breviflorum bre 67.5-72.5 70.3±0.3 69.5-71.0 32.5-37.5 34.7±0.3 34.1-35.3 H. calyptratum cal 87.5-93.7 91.5±0.3 90.8-92.1 55.0-61.3 57.4±0.4 56.6-58.2 H. canastrense can 50.0-60.0 56.5±0.4 55.5-57.4 42.5-52.5 47.6±0.4 46.7-48.6 H. cipoanum cip 65.0-71.2 68.5±0.3 67.9-69.2 43.7-50.0 47.5±0.4 46.7-48.3 H. elegans ele 62.5-67.5 64.2±0.3 63.6-64.8 37.5-41.2 39.0±0.2 38.5-39.5 H. glauscesens gla 72.5-77.5 74.3±0.4 73.5-75.1 45.0-50.0 47.4±0.4 46.6-48.2 H. morelianum mor 60.0-70.0 65.0±0.5 64.0-66.0 45.0-52.5 48.4±0.4 47.6-49.3 H. papilio pap 78.7-82.5 81.0±0.2 80.4-81.5 52.5-47.5 49.6±0.3 49.0-50.3 H. puniceum pun 75.0-82.5 79.3±0.5 78.3-80.3 32.5-42.5 36.9±0.6 35.7-38.1 H. reginae reg 55.0-60.0 58.1±0.3 57.5-58.7 37.5-43.7 40.5±0.3 39.7-41.2 H. reticulatum ret 62.5-70.0 65.5±0.4 64.7-66.3 40.0-46.2 42.2±0.4 41.4-43.0 H. santacatarina san 50.0-57.5 55.9±0.4 55.1-56.7 45.0-50.0 46.5±0.3 47.2-45.9 H. stapfianum sta 75.0-82.5 77.3±0.4 77.1-79.0 45.0-55.0 48.3±0.6 46.9-49.6 H. striatum str 57.5-62.5 60.6±0.3 60.0-61.2 40.0-43.7 41.8±0.3 41.2-42.4 H. stylosum sty 62.5-68.8 64.9±0.4 64.1-65.8 37.5-45.0 40.7±0.4 39.8-41.5 H. vittatum vit 62.5-70.0 67.2±0.4 66.3-68.0 42.5-50.0 46.3±0.5 45.3-47.3 x- arithmetic mean; sx – standard deviation of the mean; IC- confidence interval. Table 2 Measurements (µm) ( n = 10) of pollen grains in polar view of Hippeastrum species. Specie Polar Diameter Equatorial Diameter Range x Range x H. angustifolium 30.0-37.5 34.8 60.0-62.5 60.8 H. aulicum 50.0-57.5 54.7 105.0-110.0 106.7 H. blossfeldiae 37.5-43.7 41.5 65.0-70.0 67.3 H. brasilianum 42.5-47.5 45.7 67.5-72.5 70.9 H. breviflorum 33.8-37.5 35.9 67.5-72.5 70.6 H. calyptratum 47.5-55.0 52.2 87.5-95.0 92.2 H. canastrense 42.5-46.2 43.9 55.0-58.7 56.5 H. cipoanum 45.0-47.5 46.2 66.2-70.0 68.1 H. elegans 37.5-40.0 39.4 65.0-70.0 67.0 H. glauscesens 42.5-47.5 45.7 70.0-75.0 72.7 H. morelianum 40.0-50.0 44.2 65.0-70.0 67.2 H. papilio 45.0-55.0 48.6 77.5-80.0 79.2 H. puniceum 32.5-37.5 35.7 70.0-80.0 73.6 H. reginae 37.5-42.5 39.9 55.0-58.8 56.3 H. reticulatum 37.5-42.5 40.0 65.0-70.0 67.4 H. santacatarina 45.0-47.5 46.0 50.0-55.0 52.9 H. stapfianum 33.8-40.0 36.1 72.5-82.5 80.0 H. striatum 32.5-40.0 37.5 60.0-65.0 62.0 H. stylosum 35.0-40.0 38.3 62.5-70.0 66.7 H. vittatum 45.0-50.0 47.1 65.0-70.0 67.8 Table 3 Measurements (µm) (n = 10) of apertures and exine layers of pollen grains of Hippeastrum species. Species Sulcus Exine thickness Lumen Muri Length Width Exine Sexine Nexine Diameter Width H. angustifolium 56.0 9.6 2.0 1.0 1.0 4.7 0.7 H. aulicum 101.2 8.2 2.0 1.0 1.0 3.5 0.6 H. blossfeldiae 59.5 9.5 2.0 1.0 1.0 3.2 1.0 H. brasilianum 66.4 7.4 2.0 1.0 1.0 2.8 1.0 H. breviflorum 64.5 6.7 2.0 1.0 1.0 3.7 0.9 H. calyptratum 85.5 7.5 2.0 1.0 1.0 6.0 0.9 H. canastrense 48.5 8.4 2.0 1.0 1.0 1.2 1.0 H. cipoanum 64.5 9.0 2.0 1.0 1.0 3.5 0.7 H. elegans 57.6 10.1 2.0 1.0 1.0 3.4 ____ H. glauscesens 70.0 7.5 2.0 1.0 1.0 4.6 0.6 H. morelianum 61.6 11.1 2.0 1.0 1.0 4.1 1.0 H. papilio 75.0 9.2 2.0 1.0 1.0 4.3 0.8 H. puniceum 70.0 10.0 2.4 1.4 1.0 5.9 ___ H. reginae 52.7 7.2 2.0 1.0 1.0 3.7 ____ H. reticulatum 41.0 9.7 2.6 1.6 1.0 2.4 0.8 H. santacatarina 44.4 9.4 1.0 1.0 1.0 2.0 0.6 H. stapfianum 74.5 14.5 2.0 1.0 1.0 ____ ____ H. striatum 51.3 20.5 2.2 1.2 1.0 4.9 0.7 H. stylosum 61.2 7.4 2.0 1.0 1.0 2.7 0.7 H. vittatum 62.9 13.5 2.0 1.0 1.0 4.6 0.7 Table 4 Cumulative variance and vector values of principal component analysis (PCA). Principal Axis 1 2 VARIANCE (%) 63.06 12.93 Characters Axis Largest diameter 0.5160 -0.0884 Smallest diameter 0.1900 0.1281 Polar diameter 0.1934 0.3489 Equatorial diameter 0.5114 -0.1433 Sulcus length 0.6225 -0.0875 Sulcus width 0.0746 -0.3179 Muri width 0.0183 0.5369 Lumen diameter 0.0727 0.6607 Supplementary Files Appendix.doc Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Minor revisions 04 Jul, 2025 Reviewers agreed at journal 09 Jun, 2025 Reviewers invited by journal 09 Jun, 2025 Editor assigned by journal 05 Jun, 2025 First submitted to journal 31 May, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6785943","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":468777006,"identity":"af8839aa-93f8-4f27-aec9-c73c7a3698be","order_by":0,"name":"Renata Suzano Cândido","email":"","orcid":"","institution":"Universidade Federal do Rio de Janeiro","correspondingAuthor":false,"prefix":"","firstName":"Renata","middleName":"Suzano","lastName":"Cândido","suffix":""},{"id":468777007,"identity":"bce2a413-aba9-43c5-8ea6-ac09a0adb3f1","order_by":1,"name":"Rosana Conrado Lopes","email":"","orcid":"","institution":"Universidade Federal do Rio de Janeiro Instituto de Biologia","correspondingAuthor":false,"prefix":"","firstName":"Rosana","middleName":"Conrado","lastName":"Lopes","suffix":""},{"id":468777008,"identity":"bd047fbc-bed9-4912-adde-7d27bb8f6552","order_by":2,"name":"Cláudia Barbieri Ferreira Mendonça","email":"","orcid":"","institution":"Universidade Federal do Rio de Janeiro Museu Nacional","correspondingAuthor":false,"prefix":"","firstName":"Cláudia","middleName":"Barbieri Ferreira","lastName":"Mendonça","suffix":""},{"id":468777009,"identity":"2f7dd7b4-89bb-4f42-bed3-4b1fe93ea160","order_by":3,"name":"Vânia Gonçalves-Esteves","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA30lEQVRIiWNgGAWjYHAD5gNAQkKGGJUwBlsCSAsPKVp4DMAkQQ38/OcPPuapqGMwuH3m86sbNRY8DOyHj27Ap0VyRjKzMc+ZwwwG53K3WeccAzqMJy3tBj4tBjeY2SRnth1gkOzh3WacwwbUIsFjhleL/fnD7D9n/qsDauF5ZpzzjwgtBgzJbAwfG5gZ+Hl4mB/nthGhReJGsrHEh2OHefh52MyYc/skeNgI+YW//+DDDwk1dXJsQEs+53yrk+NnP3wMrxYYAEUHmwSIxUaMchhg/kCK6lEwCkbBKBg5AAAB3T3x0XOdgQAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0002-2803-6027","institution":"Universidade Federal do Rio de Janeiro Museu Nacional","correspondingAuthor":true,"prefix":"","firstName":"Vânia","middleName":"","lastName":"Gonçalves-Esteves","suffix":""}],"badges":[],"createdAt":"2025-05-30 15:15:08","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6785943/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6785943/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":84389447,"identity":"8d0a9934-a04a-49c5-b563-0b3ebefa0f35","added_by":"auto","created_at":"2025-06-11 11:03:21","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1332543,"visible":true,"origin":"","legend":"\u003cp\u003ePhotomicrographs and scanning electron micrographs (SEM) of selected species of \u003cem\u003eHippeastrum\u003c/em\u003e. \u003cem\u003eH. angustifolium\u003c/em\u003e \u003cstrong\u003ea\u003c/strong\u003e polar view, \u003cstrong\u003eb\u003c/strong\u003e distal face in polar view: aperture (SEM), and \u003cstrong\u003ec\u003c/strong\u003e surface detail (SEM). \u003cem\u003eH. aulicum\u003c/em\u003e \u003cstrong\u003ed \u003c/strong\u003edistal face in polar view (SEM), \u003cstrong\u003ee\u003c/strong\u003e equatorial view (SEM), \u003cstrong\u003ef\u003c/strong\u003eproximal face in polar view (SEM), and \u003cstrong\u003eg\u003c/strong\u003esurface detail (SEM). \u003cem\u003eH. blossfeldiae\u003c/em\u003e\u003cstrong\u003e h\u003c/strong\u003e equatorial view,\u003cstrong\u003e i\u003c/strong\u003e distal face in polar view: aperture (SEM) and \u003cstrong\u003ej\u003c/strong\u003e surface detail. \u003cem\u003eH. brasilianum\u003c/em\u003e\u003cem\u003e\u003cstrong\u003e k\u003c/strong\u003e\u003c/em\u003e distal face in polar view: aperture (SEM) and \u003cstrong\u003el\u003c/strong\u003esurface detail (SEM). Scale bar: 10 mm (a, b, d-f, h, k); 2 mm (c, g, i, j, l).\u003c/p\u003e","description":"","filename":"Fig.1.tif.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6785943/v1/e908e9d23ec404dc86f80fef.jpg"},{"id":84389448,"identity":"040f53b7-119d-400a-a7bf-55ec1fce002b","added_by":"auto","created_at":"2025-06-11 11:03:21","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1466566,"visible":true,"origin":"","legend":"\u003cp\u003ePhotomicrographs and scanning electron micrographs (SEM) of selected species of \u003cem\u003eHippeastrum\u003c/em\u003e. \u003cem\u003eH. breviflorum \u003c/em\u003e\u003cstrong\u003ea\u003c/strong\u003e distal face in polar view: aperture (SEM), \u003cstrong\u003eb\u003c/strong\u003e equatorial view (SEM), and \u003cstrong\u003ec\u003c/strong\u003e surface detail (SEM). \u003cem\u003eH. calyptratum\u003c/em\u003e \u003cstrong\u003ed\u003c/strong\u003e equatorial view, \u003cstrong\u003ee\u003c/strong\u003eproximal face in polar view (SEM), and \u003cstrong\u003ef \u003c/strong\u003esurface detail (SEM). \u003cem\u003eH. canastrense\u003c/em\u003e \u003cstrong\u003eg\u003c/strong\u003e surface detail (SEM). \u003cem\u003eH. cipoanum\u003c/em\u003e\u003cstrong\u003e h \u003c/strong\u003eequatorial view (SEM), \u003cstrong\u003ei\u003c/strong\u003e surface detail (SEM). \u003cem\u003eH. elegans\u003c/em\u003e\u003cstrong\u003e j\u003c/strong\u003e proximal face in polar view (SEM) and\u003cstrong\u003e k\u003c/strong\u003e equatorial view: aperture (SEM) and \u003cstrong\u003el\u003c/strong\u003e surface detail (SEM). Scale bar: 10 mm (a, b, d, e, g, h, j, k); 2 mm (c, f, i, l).\u003c/p\u003e","description":"","filename":"Fig.2.tif.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6785943/v1/3b43ae7f3f2462fe0b453434.jpg"},{"id":84389449,"identity":"45200392-6167-49e6-a8a1-6351819a4f77","added_by":"auto","created_at":"2025-06-11 11:03:21","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1609086,"visible":true,"origin":"","legend":"\u003cp\u003ePhotomicrographs and scanning electron micrographs (SEM) of selected species of \u003cem\u003eHippeastrum\u003c/em\u003e. \u003cem\u003eH. glaucescens\u003c/em\u003e \u003cstrong\u003ea\u003c/strong\u003eequatorial view: aperture (SEM) and \u003cstrong\u003eb\u003c/strong\u003esurface detail (SEM). \u003cem\u003eH. morelianum\u003c/em\u003e \u003cstrong\u003ec \u003c/strong\u003eequatorial view (SEM) and \u003cstrong\u003ed\u003c/strong\u003e surface detail (SEM).\u003cem\u003e H. papilio\u003c/em\u003e \u003cstrong\u003ee\u003c/strong\u003e distal face in polar view: aperture (SEM) and\u003cstrong\u003e f\u003c/strong\u003e surface detail (SEM). \u003cem\u003eH. puniceum\u003c/em\u003e \u003cstrong\u003eg \u003c/strong\u003eequatorial view, \u003cstrong\u003eh\u003c/strong\u003e equatorial view (SEM), and\u003cstrong\u003e i\u003c/strong\u003e surface detail (SEM). \u003cem\u003eH. reginae\u003c/em\u003e \u003cstrong\u003ej\u003c/strong\u003e distal face in polar view: aperture (SEM), \u003cstrong\u003ek\u003c/strong\u003e proximal face in polar view (SEM), and \u003cstrong\u003el \u003c/strong\u003esurface detail (SEM). Scale bar: 10 mm (a, c, e, g, h, j, k); 2 mm (b, d, f, i, l).\u003c/p\u003e","description":"","filename":"Fig.3.tif.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6785943/v1/1db2cb04b92cd09765724ea9.jpg"},{"id":84389450,"identity":"ecd8b81a-e7c7-4b08-8d19-24424797a0d7","added_by":"auto","created_at":"2025-06-11 11:03:21","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":1442910,"visible":true,"origin":"","legend":"\u003cp\u003ePhotomicrographs and scanning electron micrographs (SEM) of selected species of \u003cem\u003eHippeastrum\u003c/em\u003e. \u003cem\u003eH. reticulatum\u003c/em\u003e \u003cstrong\u003ea\u003c/strong\u003e distal face in polar view: aperture (SEM) and \u003cstrong\u003eb\u003c/strong\u003esurface detail (SEM). \u003cem\u003eH. santacatarina\u003c/em\u003e \u003cstrong\u003ec \u003c/strong\u003esurface detail (SEM). \u003cem\u003eH. stapfianum\u003c/em\u003e \u003cstrong\u003ed\u003c/strong\u003e distal face in polar view: aperture (SEM) and \u003cstrong\u003ee\u003c/strong\u003esurface detail (SEM). \u003cem\u003eH. striatum\u003c/em\u003e \u003cstrong\u003ef\u003c/strong\u003e distal face in polar view: aperture (SEM) and \u003cstrong\u003eg\u003c/strong\u003e surface detail (SEM). \u003cem\u003eH. stylosum\u003c/em\u003e \u003cstrong\u003eh\u003c/strong\u003e distal face in polar view: aperture (SEM) and \u003cstrong\u003ei \u003c/strong\u003esurface detail (SEM). \u003cem\u003eH. vittatum\u003c/em\u003e \u003cstrong\u003ej\u003c/strong\u003e distal face in polar view (SEM),\u003cstrong\u003e k\u003c/strong\u003e equatorial view (SEM), and \u003cstrong\u003el\u003c/strong\u003esurface detail (SEM). Scale bar: 10 mm (a, d, f, h, i, k); 2 mm (b, c, e, g, j, l).\u003c/p\u003e","description":"","filename":"Fig.4.tif.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6785943/v1/68fe4a26a9401a64f7bc8d85.jpg"},{"id":84389483,"identity":"3df2f733-8cfd-40ba-a650-904d7defb86e","added_by":"auto","created_at":"2025-06-11 11:03:24","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":445978,"visible":true,"origin":"","legend":"\u003cp\u003eDendrogram generated by agglomerative hierarchical clustering of quantitative characters of species of \u003cem\u003eHippeastrum\u003c/em\u003e. Taxa are indicated by the first three letters of the specific epithet.\u003c/p\u003e","description":"","filename":"Fig.5.tif.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6785943/v1/da16886c00d2347b2750142a.jpg"},{"id":84390360,"identity":"685e3f72-472f-4b17-b67d-e554a9e611d4","added_by":"auto","created_at":"2025-06-11 11:19:21","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":1502643,"visible":true,"origin":"","legend":"\u003cp\u003ePrincipal component analysis biplot representing species of the genus \u003cem\u003eHippeastrum\u003c/em\u003e\u003c/p\u003e","description":"","filename":"Fig.6.tif.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6785943/v1/32af9237c286a7c63b3fddd4.jpg"},{"id":84390940,"identity":"c871e4ef-daa0-41ea-a8be-5ea2a7c08e98","added_by":"auto","created_at":"2025-06-11 11:27:24","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":8921098,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6785943/v1/25a22dd3-c924-49ab-b071-ea08e3d0bcd6.pdf"},{"id":84389461,"identity":"2ac5da3c-39ff-4f49-bc22-8ad97ba80e08","added_by":"auto","created_at":"2025-06-11 11:03:22","extension":"doc","order_by":15,"title":"","display":"","copyAsset":false,"role":"supplement","size":31232,"visible":true,"origin":"","legend":"","description":"","filename":"Appendix.doc","url":"https://assets-eu.researchsquare.com/files/rs-6785943/v1/9ab26a831c2f1c82e9319fb8.doc"}],"financialInterests":"","formattedTitle":"Palynological analysis of representatives of Hippeastrum Herb. (Amaryllidaceae: Amaryllidoideae), with implications for systematics","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eAsparagales is a monophyletic order comprising 14 families, 1122 genera, and approximately 36,265 species. One of the families belonging to Asparagales is Amaryllidaceae, which includes about 73 genera and 1605 species distributed in three subfamilies (Agapanthoideae, Allioideae, and Amaryllidoideae). The grouping of these subfamilies is supported by a shared morphological character: the presence of a scapose umbellate inflorescence surrounded by a pair of bracts (Stevens 2001, APG IV 2016).\u003c/p\u003e \u003cp\u003eAmaryllidoideae comprises 15 tribes, 6 of which are native to the American continent, including the widely distributed tribe Hippeastreae. \u003cem\u003eHippeastrum\u003c/em\u003e Herb. is one of the few genera within the tribe that has been found to be monophyletic. It was initially described as comprising 15 species, which, according to Herbert (1821), exhibited considerable morphological variation. In Brazil, \u003cem\u003eHippeastrum\u003c/em\u003e is represented by approximately 39 species, 31 of which are endemic. These species occur in all vegetation types, from forests to grasslands, and on a variety of substrates (Dutilh et al. 2025). A recent investigation analyzed relationships among Brazilian species to assess the monophyly of \u003cem\u003eHippeastrum\u003c/em\u003e and its subgenera based on DNA sequence data. The study found no correlation between the traditionally proposed subgenera, concluding that they are polyphyletic (Oliveira 2012).\u003c/p\u003e \u003cp\u003eFew palynological studies have been conducted on species of \u003cem\u003eHippeastrum\u003c/em\u003e. Alves-Ara\u0026uacute;jo and Santos (2007) conducted a palynological analysis of two species of \u003cem\u003eHippeastrum\u003c/em\u003e occurring in northeastern Brazil. The authors stated that the studied taxa exhibited valuable characters that can aid in their taxonomic delimitation. In another study, five species of \u003cem\u003eHippeastrum\u003c/em\u003e found in the restingas of Rio de Janeiro State were successfully distinguished based on pollen morphology, particularly exine ornamentation (Candido et al. 2013). Although palynological studies on \u003cem\u003eHippeastrum\u003c/em\u003e are scarce, existing research consistently emphasizes the importance of pollen characters in the taxonomy of the genus. Therefore, this study aimed to evaluate whether species of \u003cem\u003eHippeastrum\u003c/em\u003e exhibit differences in pollen morphology that can contribute to a more precise delimitation and characterization of the taxa under investigation.\u003c/p\u003e"},{"header":"2. Material and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1. Pollen material\u003c/h2\u003e \u003cp\u003eThis study examined pollen grains of 20 taxa within the genus \u003cem\u003eHippeastrum\u003c/em\u003e (Appendix 1). The pollen material was obtained from fertile anthers of flowers at anthesis and/or buds at pre-anthesis, collected from exsiccatae deposited in the following herbaria: HB, HUEFS, IAC, ICN, PACA, R, RB, RFA, UB, and UEC (acronyms according to Thiers, continuously updated).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2. Light microscopy\u003c/h2\u003e \u003cp\u003eFor analysis by light microscopy, the material was processed according to the acetolysis method of Raynal and Raynal (1971), which uses 60% lactic acid. This method was chosen because pollen grains exhibited poor resistance to traditional acetolysis. Acetolyzed pollen grains were photomicrographed using a Canon Power Shot G6 digital camera coupled to a Zeiss Axiostar Plus binocular microscope equipped with a 100\u0026times; objective lens. Microscope slides were deposited in the pollen collection of the \u0026Aacute;lvaro Xavier Moreira Laboratory of Palynology, Department of Botany, Nacional Museum, Federal University of Rio de Janeiro, Brazil.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3. Measurements and statistical analysis\u003c/h2\u003e \u003cp\u003eFor quantitative analysis, measurements were taken from polar and equatorial views. Pollen grains were randomly selected from a minimum of three slides to ensure sample homogenization (Salgado-Labouriau 1973). A total of 25 measurements were performed of the largest and smallest diameters in polar view. Additionally, 10 measurements were taken, when possible, of the polar and equatorial diameters in equatorial view, sexine and nexine thickness, aperture length and width, and reticulum lumen diameter (ornamentation). Acetolyzed pollen grains were measured within three days of preparation to prevent changes in pollen size (Melhem et al. 2003).\u003c/p\u003e \u003cp\u003eVariables with 25 measurements were statistically treated to determine the arithmetic mean (\u003cem\u003ex\u003c/em\u003e), standard deviation of the mean (\u003cem\u003es\u003c/em\u003e\u003csub\u003e\u003cem\u003ex\u003c/em\u003e\u003c/sub\u003e), 95% confidence interval (95% CI), and range of variation. The other variables are expressed as arithmetic means.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.4. Scanning electron microscopy (SEM)\u003c/h2\u003e \u003cp\u003eAnthers were ground to release pollen grains, which were then dusted in their non-acetolyzed form onto aluminum stubs coated with double-sided carbon tape (Melhem et al. 2003). Samples were transferred to a vacuum pump and metalized with a thin layer (200 \u0026Aring;) of gold-palladium. Subsequently, samples were analyzed using a JEOL JSM-6390LV system at the Invertebrate Electron Microscopy Laboratory, National Museum, Federal University of Rio de Janeiro, and a Zeiss DSM 960 system at the Hertha Meyer Cellular Ultrastructure Laboratory, Institute of Biophysics, Federal University of Rio de Janeiro, Brazil.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003e2.5. Terminology\u003c/h2\u003e \u003cp\u003eThe terminologies adopted for the description of pollen size, shape, and sexine ornamentation patterns were those of Erdtman (1952) and Punt et al. (2007).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e2.6. Multivariate analysis\u003c/h2\u003e \u003cp\u003ePC-ORD software version 5.31 (McCune \u0026amp; Mefford 2011) was used for exploratory analysis. The pollen characters (8 variables) of each species were organized into a matrix, resulting in better organization of the data and facilitating the generation of graphs. Species names were abbreviated to the first three letters of specific epithets, and pollen characters were abbreviated to their initial letters. The variables included in multivariate analyses were largest diameter (LD), smallest diameter (SD), polar diameter (PD), equatorial diameter (ED), sulcus width (SW), sulcus length (SL), lumen diameter (LD), and muri width (MW).\u003c/p\u003e \u003cp\u003eThe matrix used for principal component analysis (PCA) and hierarchal cluster analysis was transformed by the square root of \u003cem\u003ex\u003c/em\u003e\u0026thinsp;+\u0026thinsp;0.5 to standardize measurable data. PCA was performed to assess whether the analyzed species could be grouped based on pollen characters. The variance\u0026ndash;covariance matrix was generated using the mean values of morphometric data and coordinates on a biplot graph based on Euclidean distances. The results are shown on a two-dimensional plot representing the first and second principal components. Vector loadings on each axis and the total cumulative variance are presented in tables.\u003c/p\u003e \u003cp\u003eHierarchical cluster analysis (HCA) was performed to group species based on similarities in pollen morphology. Two criteria were adopted when analyzing the data: the percentage of information (variables) required to form groups and the final number of groups obtained. A dendrogram was constructed using Euclidean distances (Caccavari et al. 2008) and Ward's linkage method.\u003c/p\u003e \u003c/div\u003e"},{"header":"3. Results","content":"\u003cp\u003eA total of 20 species of the genus \u003cem\u003eHippeastrum\u003c/em\u003e were examined. The palynological description is organized according to the following characters: dispersion unit, polarity, shape, size, aperture type and number, and sexine ornamentation pattern.\u003c/p\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e3.1. Dispersion unit, polarity, shape, and size\u003c/h2\u003e \u003cp\u003eAll analyzed species of \u003cem\u003eHippeastrum\u003c/em\u003e have pollen grains shed as monads, heteropolar, and elliptical in polar view. In equatorial view, the proximal face is flat (Figs.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eh, \u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ee, \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eg, and \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ei) or convex (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea, h), whereas the distal face is always convex (Figs.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eg, \u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eh, and \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eg). Most of the analyzed species have large pollen grains, with very large grains observed only in \u003cem\u003eH. aulicum\u003c/em\u003e, whose largest diameter in polar view was 104.7 \u0026micro;m. In large pollen grains, the lowest value of the largest diameter in polar view was recorded in \u003cem\u003eH. santacatarina\u003c/em\u003e (55.9 \u0026micro;m) and the highest in \u003cem\u003eH. calyptratum\u003c/em\u003e (91.5 \u0026micro;m). In equatorial view, the largest polar diameter was found in \u003cem\u003eH. aulicum\u003c/em\u003e (ca. 54.7 \u0026micro;m) and the smallest in \u003cem\u003eH. angustifolium\u003c/em\u003e (ca. 34.8 \u0026micro;m). The largest equatorial diameter was found in \u003cem\u003eH. aulicum\u003c/em\u003e (ca. 106.7 \u0026micro;m) and the smallest in \u003cem\u003eH. santacatarina\u003c/em\u003e (ca. 52.9 \u0026micro;m) (Tables\u0026nbsp;1 and 2).\u003c/p\u003e\u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003e3.2. Aperture\u003c/h2\u003e \u003cp\u003eAll species have monosulcate grains, with long sulci having sharp or rounded ends. The greatest sulcus length was recorded in \u003cem\u003eH. aulicum\u003c/em\u003e (ca. 101.2 \u0026micro;m) and the shortest in \u003cem\u003eH. reticulatum\u003c/em\u003e (ca. 41.0 \u0026micro;m). The apertural membrane was psilate in \u003cem\u003eH. blossfeldiae\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ei), \u003cem\u003eH. brasilianum\u003c/em\u003e, \u003cem\u003eH. cipoanum\u003c/em\u003e, \u003cem\u003eH. stylosum\u003c/em\u003e, and \u003cem\u003eH. vittatum\u003c/em\u003e (Table\u0026nbsp;3) and difficult to distinguish in the other species.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003e3.3. Sexine stratification and ornamentation patterns\u003c/h2\u003e \u003cp\u003eThe sexine was as thick as the nexine in most species but thicker than the nexine in \u003cem\u003eH. puniceum\u003c/em\u003e, \u003cem\u003eH. reticulatum\u003c/em\u003e, and \u003cem\u003eH. striatum\u003c/em\u003e (Table\u0026nbsp;3). The exine ornamentation pattern ranged from intectate reticulate to semitectate reticulate.\u003c/p\u003e \u003cp\u003eA retipilate sexine was observed in \u003cem\u003eH. elegans\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003el), \u003cem\u003eH. puniceum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ei), \u003cem\u003eH. reginae\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003el), and \u003cem\u003eH. stapfianum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ee). Pila have a smooth surface and gradually decrease in diameter, with the lumina becoming more densely grouped toward the ends and near the aperture (Figs.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ek, \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ek), except in \u003cem\u003eH. elegans\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eg, h). Pila are connected by the upper part (head), sparsely grouped, forming small strands that create closed meshes resembling a reticulum in some regions (Figs.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ej, \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ej, \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ek). In \u003cem\u003eH. puniceum\u003c/em\u003e, pila are rounded, vary in diameter (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eh, i), and are arranged in double rows, with sparse granules in lumina. In \u003cem\u003eH. reginae\u003c/em\u003e, sexine bridges join pila at the base (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003el), forming strands organized into reticula with well-defined lumina.\u003c/p\u003e \u003cp\u003eA reticulate heterobrochate sexine was observed in most species. The reticulate surface with large luminal diameters is restricted to a well-demarcated median band in the pollen grain. The ends exhibit microreticulate ornamentation in \u003cem\u003eH. aulicum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ee, f), \u003cem\u003eH. calyptratum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ed, e), \u003cem\u003eH. papilio\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ee), and \u003cem\u003eH. stylosum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eh, i). Muri are narrow (ca. 0.6\u0026ndash;1.0 \u0026micro;m) in all taxa, straight in most species and sinuous in \u003cem\u003eH. blossfeldiae\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ej), with sparse perforations. Columellae are not apparent or visible in \u003cem\u003eH. aulicum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eg), \u003cem\u003eH. calyptratum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ef), \u003cem\u003eH. glaucescens\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb), \u003cem\u003eH. morelianum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ed), \u003cem\u003eH. papilio\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ef), \u003cem\u003eH. striatum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eg), \u003cem\u003eH. stylosum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ej), or \u003cem\u003eH. vittatum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003el). Smaller lumina surrounding larger ones were recorded in \u003cem\u003eH. angustifolium\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ec), \u003cem\u003eH. aulicum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eg), \u003cem\u003eH. blossfeldiae\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ej), \u003cem\u003eH. brasilianum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003el), \u003cem\u003eH. breviflorum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ec), \u003cem\u003eH. calyptratum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ef), \u003cem\u003eH. morelianum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ed), \u003cem\u003eH. papilio\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ef), \u003cem\u003eH. reticulatum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eb), \u003cem\u003eH. santacatarina\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ec), \u003cem\u003eH. striatum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eg), and \u003cem\u003eH. stylosum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ej). Lumina may or may not have ornamentation (Figs.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ej, l, \u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eg, i, l, \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ef, \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eb, j). When present, the ornamentation consists of inconspicuous and sparse granules in most species or densely organized and conspicuous granules in \u003cem\u003eH. angustifolium\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ec), \u003cem\u003eH. glaucescens\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb), and \u003cem\u003eH. morelianum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ed).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e3.4. Pollen key to species of the genus \u003cem\u003eHippeastrum\u003c/em\u003e\u003c/h2\u003e\u003cp\u003e1. Sexine retipilate\u003c/p\u003e\n\u003cp\u003e2. Lumina well-defined, pila joined at the base by exine bridges............................. \u003cem\u003eH. reginae\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e2. Lumina undefined, pila not joined at the base\u003c/p\u003e\n\u003cp\u003e3. Pila arranged in double rows........................................................................... \u003cem\u003eH. puniceum\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e3. Pila not arranged in double rows\u003c/p\u003e\n\u003cp\u003e4. Pila sparsely clustered forming small strands................................................... \u003cem\u003eH. elegans\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e4. Pila densely clustered without forming small strands................................. \u003cem\u003eH. stapfianum\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e1. Sexine reticulate heterobrochate\u003c/p\u003e\n\u003cp\u003e5. Muri with sparse interruptions, lumina with sparse, inconspicuous granules... \u003cem\u003eH. cipoanum\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e5. Muri without interruptions\u003c/p\u003e\n\u003cp\u003e7. Reticulum with large lumina restricted to a well-demarcated median band of the pollen and ends with microreticulate ornamentation\u003c/p\u003e\n\u003cp\u003e8. Lumina \u0026gt; ca. 6.0 \u0026micro;m............................................................................... \u003cem\u003eH. calyptratum\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e8. Lumina \u0026lt; 5.0 \u0026micro;m\u003c/p\u003e\n\u003cp\u003e9. Pollen very large.......................................................................................... \u003cem\u003eH. aulicum\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e9. Pollen large.................................................. \u003cem\u003eH. angustifolium\u003c/em\u003e, \u003cem\u003eH. papilio\u003c/em\u003e,\u003cem\u003e\u0026nbsp;H. stylosum\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e7. Reticulum with lumina gradually decreasing in diameter toward the ends\u003c/p\u003e\n\u003cp\u003e11. Muri sinuous, presence of smaller lumina surrounding larger ones..............................\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e...........................................................................................\u0026nbsp;H. blossfeldiae\u003c/em\u003e, \u003cem\u003eH. breviflorum\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e11. Muri straight\u003c/p\u003e\n\u003cp\u003e13. Columella apparent........ \u003cem\u003eH. glaucescens\u003c/em\u003e,\u003cem\u003e\u0026nbsp;H. morelianum\u003c/em\u003e,\u003cem\u003e\u0026nbsp;H. striatum\u003c/em\u003e,\u003cem\u003e\u0026nbsp;H. vittatum\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e13. Columella not apparent.............. \u003cem\u003eH. brasilianum\u003c/em\u003e,\u003cem\u003e\u0026nbsp;H. canastrense\u003c/em\u003e,\u003cem\u003e\u0026nbsp;H. santacatarina\u003c/em\u003e\u003c/p\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003e3.5. Multivariate analyses\u003c/h2\u003e \u003cdiv id=\"Sec18\" class=\"Section3\"\u003e \u003ch2\u003e3.5.1. HCA\u003c/h2\u003e \u003cp\u003eAnalysis of the relationships between species of \u003cem\u003eHippeastrum\u003c/em\u003e yielded a dendrogram with a linkage level of 14.71%. When 50% of the remaining information (variables) was considered, three groups were identified. Group 1 contained the species \u003cem\u003eH. angustifolium\u003c/em\u003e, \u003cem\u003eH. elegans\u003c/em\u003e, \u003cem\u003eH. reginae\u003c/em\u003e, \u003cem\u003eH. stylosum\u003c/em\u003e, \u003cem\u003eH. puniceum\u003c/em\u003e, \u003cem\u003eH. reticulatum\u003c/em\u003e, \u003cem\u003eH. blossfeldiae\u003c/em\u003e, \u003cem\u003eH. breviflorum\u003c/em\u003e, \u003cem\u003eH. striatum\u003c/em\u003e, \u003cem\u003eH. canastrense\u003c/em\u003e, and \u003cem\u003eH. santacatarina\u003c/em\u003e. Group 2 comprised the species \u003cem\u003eH. aulicum\u003c/em\u003e, \u003cem\u003eH. glaucescens\u003c/em\u003e, \u003cem\u003eH. brasilianum\u003c/em\u003e, \u003cem\u003eH. papilio\u003c/em\u003e, \u003cem\u003eH. cipoanum\u003c/em\u003e, \u003cem\u003eH. morelianum\u003c/em\u003e, \u003cem\u003eH. vittatum\u003c/em\u003e), and \u003cem\u003eH. stapfianum\u003c/em\u003e. Finally, \u003cem\u003eH. aulicum\u003c/em\u003e and \u003cem\u003eH. calyptratum\u003c/em\u003e formed group 3. An increase in the number of variables analyzed (75% of remaining information) resulted in modifications to the clusters. Group 1 was subdivided into three other groups: 1\u0026prime; (\u003cem\u003eH. angustifolium\u003c/em\u003e, \u003cem\u003eH. elegans\u003c/em\u003e, \u003cem\u003eH. reginae\u003c/em\u003e, \u003cem\u003eH. blossfeldiae\u003c/em\u003e, \u003cem\u003eH. stylosum\u003c/em\u003e, \u003cem\u003eH. breviflorum\u003c/em\u003e, \u003cem\u003eH. puniceum\u003c/em\u003e), 2\u0026prime; (\u003cem\u003eH. canastrense\u003c/em\u003e, \u003cem\u003eH. santacatarina\u003c/em\u003e and \u003cem\u003eH. reticulatum\u003c/em\u003e), and 3\u0026prime; (\u003cem\u003eH. striatum\u003c/em\u003e). Group 2 comprised groups 4\u0026prime; (\u003cem\u003eH. glaucescens\u003c/em\u003e, \u003cem\u003eH. brasilianum\u003c/em\u003e, \u003cem\u003eH. papilio\u003c/em\u003e, \u003cem\u003eH. cipoanum\u003c/em\u003e, \u003cem\u003eH. morelianum\u003c/em\u003e, and \u003cem\u003eH. vittatum\u003c/em\u003e) and 5\u0026prime; (\u003cem\u003eH. stapfianum\u003c/em\u003e). Group 6 remains unchanged and is a repeat of group 3 (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec19\" class=\"Section3\"\u003e \u003ch2\u003e3.5.2. PCA\u003c/h2\u003e \u003cp\u003ePCA resulted in two principal components, which together explained 75.99% of the total variance. The first component accounted for 63.06% of the variance, and the second 12.93%. Sulcus length, largest diameter, and equatorial diameter were the most influential variables for the first principal component. By contrast, lumen diameter, murus width, and polar diameter contributed the most to the second principal component (Table\u0026nbsp;4).\u003c/p\u003e \u003cp\u003eOn axis 1, species were broadly distributed across both positive and negative sides. Along axis 2, most species clustered on the positive side, with the exception of \u003cem\u003eH. stapfianum\u003c/em\u003e, which was isolated at the extreme negative end due to its low lumen diameter. The dispersion of species on the PCA biplot may facilitate the identification of groups, even though species differed by only a few morphological characters (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eInitially, species were separated into three groups. The first group, located near the negative pole of axis 1, comprised \u003cem\u003eH. angustifolium\u003c/em\u003e, \u003cem\u003eH. canastrense\u003c/em\u003e, \u003cem\u003eH. reginae\u003c/em\u003e, \u003cem\u003eH. reticulatum\u003c/em\u003e, \u003cem\u003eH. santacatarina\u003c/em\u003e, and \u003cem\u003eH. striatum\u003c/em\u003e. The group was less dispersed and characterized by low values of sulcus length. The second group, positioned to the left of axis 2, was formed by \u003cem\u003eH. elegans\u003c/em\u003e, \u003cem\u003eH. blossfeldiae\u003c/em\u003e, \u003cem\u003eH. breviflorum\u003c/em\u003e, \u003cem\u003eH. morelianum\u003c/em\u003e, \u003cem\u003eH. stylosum\u003c/em\u003e, and \u003cem\u003eH. vittatum\u003c/em\u003e. These species clustered together because of their similar values of equatorial diameter and largest diameter. The third group was plotted to the right of axis 2, composed of \u003cem\u003eH. brasilianum\u003c/em\u003e, \u003cem\u003eH. cipoanum\u003c/em\u003e, \u003cem\u003eH. glaucescens\u003c/em\u003e, \u003cem\u003eH. papilio\u003c/em\u003e, and \u003cem\u003eH. puniceum\u003c/em\u003e. This group was more internally dispersed and distinct from the others, primarily due to differences in polar diameter. \u003cem\u003eH. aulicum\u003c/em\u003e and \u003cem\u003eH. calyptratum\u003c/em\u003e were polarized in relation to the others because they exhibited the highest values of largest diameter, smallest diameter, polar diameter, equatorial diameter, and sulcus length among the sample (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e"},{"header":"4. Discussion","content":"\u003cp\u003eThe findings allowed developing a pollen key, initially divided into two major groups based on exine ornamentation. Subsequent taxon identification was guided by ornamentation elements and pollen size. Other aspects of pollen morphology, such as aperture number and type (monosulcate), dispersion unit (monads), and polarity (heteropolar) were consistent across the sample and therefore not included in the key. Some species could not be differentiated using the proposed key due to shared pollen characteristics, including \u003cem\u003eH. angustifolium\u003c/em\u003e, \u003cem\u003eH. papilio\u003c/em\u003e, and \u003cem\u003eH. stylosum\u003c/em\u003e; \u003cem\u003eH. blossfeldiae\u003c/em\u003e and \u003cem\u003eH. breviflorum\u003c/em\u003e; \u003cem\u003eH. glaucescens\u003c/em\u003e, \u003cem\u003eH. morelianum\u003c/em\u003e, \u003cem\u003eH. striatum\u003c/em\u003e, and \u003cem\u003eH. vittatum\u003c/em\u003e; and \u003cem\u003eH. brasilianum\u003c/em\u003e, \u003cem\u003eH. canastrense\u003c/em\u003e, \u003cem\u003eH. reticulatum\u003c/em\u003e, and \u003cem\u003eH. santacatarina\u003c/em\u003e. Pollen size, however, was useful for separating \u003cem\u003eH. aulicum\u003c/em\u003e (very large) from the other species, which exhibited large pollen.\u003c/p\u003e \u003cp\u003eThese results are partially consistent with those of Alves-Ara\u0026uacute;jo and Santos (2007), who described large to very large pollen grains in \u003cem\u003eH. puniceum\u003c/em\u003e and \u003cem\u003eH. stylosum\u003c/em\u003e. In the current study, both species were classified as having large pollen. Exine ornamentation was a key diagnostic character, with retipilate sexine observed in \u003cem\u003eH. elegans\u003c/em\u003e, \u003cem\u003eH. puniceum\u003c/em\u003e, \u003cem\u003eH. reginae\u003c/em\u003e, and \u003cem\u003eH. stapfianum\u003c/em\u003e and reticulate sexine identified in the other species, as detailed in the pollen key.\u003c/p\u003e \u003cp\u003eAccording to Walker \u0026amp; Doyle (1975), the architecture of the pollen wall is an important source of phylogenetic information. The evolutionary trends suggested by the authors indicate imperforate tectate exine as the least derived, followed by semitectate and intectate as the most derived. Thus, among the studied species, \u003cem\u003eH. elegans\u003c/em\u003e, \u003cem\u003eH. puniceum\u003c/em\u003e, \u003cem\u003eH. reginae\u003c/em\u003e, and \u003cem\u003eH. stapfianum\u003c/em\u003e had more derived pollen grains.\u003c/p\u003e \u003cp\u003eSpecies with retipilate sexine were distinguished based on pilum characteristics and organization. Among those with a reticulate exine, \u003cem\u003eH. aulicum\u003c/em\u003e, \u003cem\u003eH. calyptratum\u003c/em\u003e, \u003cem\u003eH. papilio\u003c/em\u003e, and \u003cem\u003eH. stylosum\u003c/em\u003e were notable for their large lumina, which were confined to the well-defined median region of the pollen grain, and poles with microreticulate pattern. The remaining taxa with reticulate exine were differentiated by features of their muri, lumina, and columellae. This dimorphic pattern of reticulum organization was first reported by Erdtman (1952) in some genera of the family Amaryllidaceae, such as \u003cem\u003eHymenocallis\u003c/em\u003e Salisb. and \u003cem\u003eLycoris\u003c/em\u003e Herb. Meerow and Dehgan (1985) also observed a similar pattern, characterized by a coarse reticulum in the median region and a finer structure at the extremities (dimorphic reticulum), in \u003cem\u003eCaliphruria\u003c/em\u003e Herb. and \u003cem\u003eEucharis\u003c/em\u003e Planch. \u0026amp; Linden. They proposed an evolutionary trend within subgenera of \u003cem\u003eHymenocallis\u003c/em\u003e, suggesting a transition from a dimorphic to a homogeneous reticulum, with the latter representing a derived character.\u003c/p\u003e \u003cp\u003eIn their investigation of \u003cem\u003eH. stylosum\u003c/em\u003e, Alves-Ara\u0026uacute;jo and Santos (2007) found that the heterobrochate reticulate exine formed a psilate-perforate cap in the acute equatorial region, thereby composing a dimorphic reticulum at the extremities of the equatorial region. The present study confirmed the dimorphic pattern in \u003cem\u003eH. stylosum\u003c/em\u003e but recorded a different type of ornamentation, namely microreticulate at the extremities (equator) of the pollen. This discrepancy can be attributed to differences in material quality or variations in sample preparation procedures for SEM analysis.\u003c/p\u003e \u003cp\u003eAdditionally, Alves-Ara\u0026uacute;jo and Santos (2007) reported \u003cem\u003eH. puniceum\u003c/em\u003e pollen as having reticulate semitectate exine and transparent tectum; however, the material was not analyzed by SEM. Here, \u003cem\u003eH. puniceum\u003c/em\u003e was observed under both light microscopy and SEM. The exine ornamentation was found to be retipilate and intectate, with pila organized in double rows, corroborating the analysis of Candido et al. (2013) for this species.\u003c/p\u003e \u003cp\u003eHCA revealed groups composed of species from different subgenera. The scattering of species on the PCA biplot allowed further individualization of taxa. Nevertheless, it was difficult to delimit groups along the ordering axes, given that species shared pollen attributes, even those of different subgenera. These findings raise questions about the current subgeneric classification of \u003cem\u003eHippeastrum\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eMost species belonging to \u003cem\u003eHippeastrum\u003c/em\u003e subg. \u003cem\u003eAschamia\u003c/em\u003e (Salisb.) Baker were grouped into the same HCA cluster (Group 1) but were not plotted close to one another on the PCA biplot. The dendrogram revealed that only \u003cem\u003eH. puniceum\u003c/em\u003e was separated from the other species of the group. This finding corroborated PCA results, in that \u003cem\u003eH. puniceum\u003c/em\u003e was plotted on the positive side of axis 1, far from the other species of the subgenus. This separation was attributed to the high values of largest diameter, equatorial diameter, and sulcus length of \u003cem\u003eH. puniceum\u003c/em\u003e compared with other members of the subgenus. Species of the other subgenera of \u003cem\u003eHippeastrum\u003c/em\u003e were dispersed across HCA clusters and the PCA biplot, indicating that \u003cem\u003eHippeastrum\u003c/em\u003e subgenera cannot be delimited by pollen morphology.\u003c/p\u003e \u003cp\u003eSpecies composing Group 2 (HCA) belonged to different subgenera, as follows: \u003cem\u003eHippeastrum glaucescens\u003c/em\u003e and \u003cem\u003eH. morelianum\u003c/em\u003e (\u003cem\u003eH.\u003c/em\u003e subg. \u003cem\u003eCaephaleaeon\u003c/em\u003e Traub.); \u003cem\u003eH. brasilianum\u003c/em\u003e and \u003cem\u003eH. stapfianum\u003c/em\u003e (\u003cem\u003eH.\u003c/em\u003e subg. \u003cem\u003eMacropodastrum\u003c/em\u003e Baker); \u003cem\u003eH. papilio\u003c/em\u003e (\u003cem\u003eH.\u003c/em\u003e subg. \u003cem\u003eOmphalissa\u003c/em\u003e (Salisb.) Baker); and \u003cem\u003eH. vittatum\u003c/em\u003e (\u003cem\u003eH.\u003c/em\u003e subg. \u003cem\u003eLais\u003c/em\u003e (Salisb.) Baker). The species were quite dispersed on the PCA biplot. Of note, \u003cem\u003eH. stapfianum\u003c/em\u003e was plotted far from the other species because of its low lumen diameter. Group 3 (HCA) was formed by \u003cem\u003eH. aulicum\u003c/em\u003e and \u003cem\u003eH. calyptratum\u003c/em\u003e, both subordinate to the subgenus \u003cem\u003eOmphalissa\u003c/em\u003e. They were plotted far from the others on the positive pole of axis 1 (PCA), owing to their high largest diameter, smallest diameter, polar diameter, equatorial diameter, and sulcus length.\u003c/p\u003e \u003cp\u003eAlthough other types of pollen have been described for Amaryllidaceae, such as disulcate aperture (\u003cem\u003eAmaryllis belladonna\u003c/em\u003e L., \u003cem\u003eCrinum\u003c/em\u003e L., \u003cem\u003eNerine\u003c/em\u003e Herb, \u003cem\u003eStrumaria\u003c/em\u003e Jacq.) with intectate-columellate exine (Erdtman 1952; Meerow and Dehgan 1985; Meerow and Snijman 1998), monosulcate pollen with semitectate exine can be considered a synapomorphy for the family (D\u0026ouml;nmez and Isik 2008).\u003c/p\u003e \u003cp\u003eThe external morphology of \u003cem\u003eH. aulicum\u003c/em\u003e and \u003cem\u003eH. morelianum\u003c/em\u003e is similar. \u003cem\u003eH. calyptratum\u003c/em\u003e and \u003cem\u003eH. aulicum\u003c/em\u003e are also similar with regard to leaf morphology and presence of corona distributed in plates (Oliveira 2012). However, the palynological morphology of \u003cem\u003eH. aulicum\u003c/em\u003e differed from that of \u003cem\u003eH. morelianum\u003c/em\u003e. Pollen of the former species were characterized by very large size, presence of reticulum with large lumina restricted to the median portion of the pollen and ends with microreticulate ornamentation. \u003cem\u003eH. calyptratum\u003c/em\u003e was more similar to \u003cem\u003eH. aulicum\u003c/em\u003e, differing only in lumen diameter (ca. 6.0 \u0026micro;m), as indicated in the pollen key. Palynological proximity between \u003cem\u003eH. aulicum\u003c/em\u003e and \u003cem\u003eH. calyptratum\u003c/em\u003e was confirmed by HCA. These species formed group 3 and were located far from the other species on the PCA biplot.\u003c/p\u003e \u003cp\u003e \u003cem\u003eHippeastrum blossfeldiae\u003c/em\u003e is known to be morphologically similar to \u003cem\u003eH. striatum\u003c/em\u003e (Oliveira 2012). However, pollen analysis revealed some differences between these species. Reticulum muri were sinuous in \u003cem\u003eH. blossfeldiae\u003c/em\u003e and straight in \u003cem\u003eH. striatum\u003c/em\u003e. Additionally, the species were placed in different groups in AHC, namely \u003cem\u003eH. blossfeldiae\u003c/em\u003e in group 1\u0026prime; and \u003cem\u003eH. striatum\u003c/em\u003e isolated in group 3\u0026prime;. On the PCA biplot, the species were plotted far from one another, because \u003cem\u003eH. blossfeldiae\u003c/em\u003e exhibited higher values of largest diameter and polar diameter.\u003c/p\u003e \u003cp\u003eAccording to Oliveira (2012), \u003cem\u003eH. brasilianum\u003c/em\u003e is very similar to \u003cem\u003eH. elegans\u003c/em\u003e. In living specimens, the species can be distinguished by flower color; in herbarium material, by the shape of the floral tube, which is more open in \u003cem\u003eH. brasilianum\u003c/em\u003e. \u003cem\u003eH. elegans\u003c/em\u003e also exhibits color variation and morphological overlap with \u003cem\u003eH. stapfianum\u003c/em\u003e and \u003cem\u003eH. vittatum\u003c/em\u003e. In the current study, \u003cem\u003eH. elegans\u003c/em\u003e and \u003cem\u003eH. stapfianum\u003c/em\u003e differed from \u003cem\u003eH. brasilianum\u003c/em\u003e and \u003cem\u003eH. vittatum\u003c/em\u003e, as the first two had a retipilate exine. By contrast, \u003cem\u003eH. brasilianum\u003c/em\u003e and \u003cem\u003eH. vittatum\u003c/em\u003e exhibited a reticulate exine, differing with regard to muri. \u003cem\u003eH. elegans\u003c/em\u003e and \u003cem\u003eH. stapfianum\u003c/em\u003e differed in pilum organization, as described in the pollen key. PCA showed that the species were widely dispersed along the positive and negative sides of axis 2, influenced by lumen diameter.\u003c/p\u003e \u003cp\u003e \u003cem\u003eHippeastrum puniceum\u003c/em\u003e exhibited intermediate forms of external morphology, being sometimes indistinguishable from \u003cem\u003eH. reginae\u003c/em\u003e. It usually has a proportionally longer hypanthial tube and distinct color patterns (Oliveira 2012). Both species exhibited retipilate exine, indicative, in theory, of their proximity. However, they were distinguished in the pollen key by differences in ornamentation elements. Such proximity was confirmed by HCA, in which species were placed in the same group (1\u0026prime;), and had similar values of polar diameter and sulcus length.\u003c/p\u003e \u003cp\u003eIt should be noted that the pollen morphology of the following species was characterized for the first time in the present study: \u003cem\u003eHippeastrum angustifolium\u003c/em\u003e, \u003cem\u003eH. aulicum\u003c/em\u003e var. \u003cem\u003eglaucophylum\u003c/em\u003e, \u003cem\u003eH. blossfeldiae\u003c/em\u003e, \u003cem\u003eH. brasilianum\u003c/em\u003e, \u003cem\u003eH. breviflorum\u003c/em\u003e, \u003cem\u003eH. calyptratum\u003c/em\u003e, \u003cem\u003eH. canastrense\u003c/em\u003e, \u003cem\u003eH. cipoanum\u003c/em\u003e, \u003cem\u003eH. elegans\u003c/em\u003e, \u003cem\u003eH. stapfianum\u003c/em\u003e, \u003cem\u003eH. morelianum\u003c/em\u003e, \u003cem\u003eH. papilio\u003c/em\u003e, \u003cem\u003eH. reginae\u003c/em\u003e, \u003cem\u003eH. santacatarina\u003c/em\u003e, and \u003cem\u003eH. vittatum\u003c/em\u003e. The results of this study hold taxonomic and systematic value, as they allowed distinguishing some taxa by specific pollen characters, including closely related species. However, the findings do not corroborate the delimitation of the subgenera proposed for \u003cem\u003eHippeastrum.\u003c/em\u003e\u003c/p\u003e"},{"header":"5. Conclusion","content":"\u003cp\u003eIn conclusion, the pollen grains of species of \u003cem\u003eHippeastrum\u003c/em\u003e exhibit notable morphological variation, particularly in exine ornamentation, sculpture elements, and pollen size, which allowed distinguishing some taxa. Two main groups were identified regarding exine ornamentation: retipilate and reticulate. Therefore, pollen morphology provides valuable information for improving species circumscription and characterization. Palynological data do not support the current delimitation of \u003cem\u003eHippeastrum\u003c/em\u003e into subgenera.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003cstrong\u003eConflict of interest\u003c/strong\u003e \u003cp\u003eNo potential conflict of interest was reported by the authors.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eAuthor contributions\u003c/h2\u003e \u003cp\u003eAll authors contributed to the study conception and design. Material preparation, data collection, and analysis were performed by VGE, CBFM, and RCL. All authors commented on previous versions and read and approved the final manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgments\u003c/h2\u003e \u003cp\u003eVGE and CBFM are grateful to the Brazilian National Council for Scientific and Technological Development (CNPq) for the research grants (grants Nos. 307276/2023-6 and 311618/2021-9). All authors thank the Rio de Janeiro State Research Foundation (FAPERJ) (grant Nos. E-26/210674/2023 and 260003/015254/2021). We also thank the herbarium curators and staff for providing access to their collections.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAlves-Ara\u0026uacute;jo A and Santos FAR (2007) Caracteriza\u0026ccedil;\u0026atilde;o palinol\u0026oacute;gica das esp\u0026eacute;cies de Amaryllidaceae sensu stricto ocorrentes no nordeste brasileiro. Acta Bot Bras 21:967-976\u003c/li\u003e\n\u003cli\u003eAPG IV (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants. \u003cem\u003eBot J Linn Soc\u003c/em\u003e 181: 1-20\u003c/li\u003e\n\u003cli\u003eCaccavari MA, Naab OA, Tamame MA (2008) Palynology And Physicochemical Characteristics Of Three Unifloral Honey Types From Central Argentina. Span J Agric Res 6: 566-576\u003c/li\u003e\n\u003cli\u003eCandido RS, Fourny ACS, Gon\u0026ccedil;alves-Esteves V, Lopes RC (2013) \u003cem\u003eHippeastrum\u003c/em\u003e species in \u0026aacute;reas of restinga in the state of the Rio de Janeiro, Brazil: pollen characters. Acta Bot Bras 27(4):661-668\u003c/li\u003e\n\u003cli\u003eD\u0026ouml;nmez EO and Isik S (2008) Pollen morphology of Turkish Amaryllidaceae, Ixioliriaceae and Iridaceae. Grana 47(1):15-38\u003c/li\u003e\n\u003cli\u003eDutilh JHA, Campos-Rocha A, Oliveira RS, Garcia N, Streher NS, Giussani LM, Semir J (in memoriam), Meerow AW, Sassone AB. \u003cem\u003eHippeastrum\u003c/em\u003e in Flora e Funga do Brasil. Jardim Bot\u0026acirc;nico do Rio de Janeiro. Available at:\u0026lt;https://floradobrasil.jbrj.gov.br/FB4354\u0026gt;. consulta.publica.uc.citacao.acesso.em06 mai. 2025\u003c/li\u003e\n\u003cli\u003eErdtman G (1952) Pollen morphology and plant taxonomy - Angiosperms. Almqvist \u0026amp; Wiksell, Stockholm\u003c/li\u003e\n\u003cli\u003eHerbert WH (1821) An Appendix. Bot. Mag. James and Ridway Sons. London.\u003c/li\u003e\n\u003cli\u003eMcCune B, Mefford MJ (2011) PC-ORD. Multivariate Analysis of Ecological Data. Version 6. MjM Software, Gleneden Beach, Oregon, U.S.A.\u003c/li\u003e\n\u003cli\u003eMeerow AW and Dehgan B (1985) The auriculate pollen grain of Hymenocallis quitoensis Herb. (Amaryllidaceae) and its systematic implications. American Journal of Botany 72(4):540-547\u003c/li\u003e\n\u003cli\u003eMeerow AW and Snijman DA (1998) Amaryllidaceae. In K. Kubitzki (ed.). The families and genera of vascular plants. Monocotyledons \u0026ndash; Lilianae (except Orchidaceae). Hamburg, Germany. 83-110p\u003c/li\u003e\n\u003cli\u003eMelhem TS, Cruz-Barros MAV, Corr\u0026ecirc;a AMS, Makino-Watanabe H, Silvestre-Capelato MSF, Gon\u0026ccedil;alves-Esteves V (2003) Morfologia pol\u0026iacute;nica em plantas de Campos do Jord\u0026atilde;o (S\u0026atilde;o Paulo, Brasil). Bol. Inst. Bot. 16:1-104.\u003c/li\u003e\n\u003cli\u003eOliveira RS (2012) O g\u0026ecirc;nero \u003cem\u003eHippeastrum\u003c/em\u003e Herb. (Amaryllidaceae) no Brasil: evid\u0026ecirc;ncia de evolu\u0026ccedil;\u0026atilde;o reticulada e an\u0026aacute;lise de caracteres florais. Campinas, S\u0026atilde;o Paulo, Universidade Estadual de Campinas, Tese\u003c/li\u003e\n\u003cli\u003ePunt W, Blackmore S, Nilsson S, Le Thomas A (2007) Glossary of pollen and spore terminology. Rev. Paleobot. Palynol. 143: 1-81\u003c/li\u003e\n\u003cli\u003eRaynal A and Raynal J (1971) Une technique de preparation des grains de pollen fragilis. Adans\u0026ocirc;nia 11(1): 77-79\u003c/li\u003e\n\u003cli\u003eSalgado-Labouriau ML (1973) Contribui\u0026ccedil;\u0026atilde;o \u0026agrave; palinologia dos cerrados. Academia Brasileira de Ci\u0026ecirc;ncias. Rio de Janeiro\u003c/li\u003e\n\u003cli\u003eStevens P F (2001). Angiosperm Phylogeny Website. Version 14, July 2017 [and more or less continuously updated since].\u0026quot; Acessed 20 july 2024.http://www.mobot.org/MOBOT/research/APweb/.Traub, H. P. 1963. The genera of Amaryllidaceae. 8pp. Ed. 1. The American Plant. LifeSociety. Jolla, California.\u003c/li\u003e\n\u003cli\u003eThiers B Index herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden\u0026apos;s Virtual Herbarium. http://sweetgum.nybg.org/ih/.2023.\u003c/li\u003e\n\u003cli\u003eWalker JW and Doyle JA (1975) The bases of angiosperm phylogeny: Palynology. Ann. Missouri Bot. Gard. 62: 664-723.\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003e\u003cstrong\u003eTable 1\u003c/strong\u003e Measurements (\u0026micro;m) of pollen grains in equatorial view of \u003cem\u003eHippeastrum\u0026nbsp;\u003c/em\u003especies.\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"84%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003eAbbreviation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"3\" valign=\"top\" style=\"width: 37px;\"\u003e\n \u003cp\u003eLargest Diameter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"3\" valign=\"top\" style=\"width: 32px;\"\u003e\n \u003cp\u003eSmallest Diameter\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003eSpecies \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003eRange\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003ex \u0026plusmn; sx\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003eCI 95%\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003eRange\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003ex \u0026plusmn; sx\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003eCI 95%\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. angustifolium\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003eang\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e52.5-60.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e56.2\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e55.3-57.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e32.5-42.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e38.4\u0026plusmn;0.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e37.4-39.5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. aulicum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003eaul\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e100.0-110.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e104.7\u0026plusmn;0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e103.5-105.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e50.0-60.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e53.0\u0026plusmn;0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e51.8-54.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. blossfeldiae\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003eblo\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e65.0-68.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e66.7\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e66.1-67.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e37.5-43.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e41.5\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e40.8-42.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. brasilianum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003ebra\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e70.0-75.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e71.3\u0026plusmn;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e70.7-71.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e46.3-56.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e50.2\u0026plusmn;0.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e49.2-51.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. breviflorum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003ebre\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e67.5-72.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e70.3\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e69.5-71.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e32.5-37.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e34.7\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e34.1-35.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. calyptratum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003ecal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e87.5-93.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e91.5\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e90.8-92.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e55.0-61.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e57.4\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e56.6-58.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. canastrense\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003ecan\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e50.0-60.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e56.5\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e55.5-57.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e42.5-52.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e47.6\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e46.7-48.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. cipoanum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003ecip\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e65.0-71.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e68.5\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e67.9-69.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e43.7-50.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e47.5\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e46.7-48.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. elegans\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003eele\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e62.5-67.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e64.2\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e63.6-64.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e37.5-41.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e39.0\u0026plusmn;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e38.5-39.5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. glauscesens\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003egla\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e72.5-77.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e74.3\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e73.5-75.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e45.0-50.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e47.4\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e46.6-48.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. morelianum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003emor\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e60.0-70.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e65.0\u0026plusmn;0.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e64.0-66.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e45.0-52.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e48.4\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e47.6-49.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. papilio\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003epap\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e78.7-82.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e81.0\u0026plusmn;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e80.4-81.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e52.5-47.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e49.6\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e49.0-50.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. puniceum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003epun\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e75.0-82.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e79.3\u0026plusmn;0.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e78.3-80.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e32.5-42.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e36.9\u0026plusmn;0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e35.7-38.1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. reginae\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003ereg\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e55.0-60.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e58.1\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e57.5-58.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e37.5-43.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e40.5\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e39.7-41.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. reticulatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003eret\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e62.5-70.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e65.5\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e64.7-66.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e40.0-46.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e42.2\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e41.4-43.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. santacatarina\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003esan\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e50.0-57.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e55.9\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e55.1-56.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e45.0-50.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e46.5\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e47.2-45.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. stapfianum\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003esta\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e75.0-82.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e77.3\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e77.1-79.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e45.0-55.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e48.3\u0026plusmn;0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e46.9-49.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. striatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003estr\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e57.5-62.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e60.6\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e60.0-61.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e40.0-43.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e41.8\u0026plusmn;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e41.2-42.4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. stylosum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003esty\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e62.5-68.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e64.9\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e64.1-65.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e37.5-45.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e40.7\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e39.8-41.5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. vittatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003evit\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e62.5-70.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e67.2\u0026plusmn;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e66.3-68.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e42.5-50.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 10px;\"\u003e\n \u003cp\u003e46.3\u0026plusmn;0.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e45.3-47.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003ex- arithmetic mean; sx \u0026ndash; standard deviation of the mean; IC- confidence interval.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 2\u003c/strong\u003e Measurements (\u0026micro;m) (\u003cem\u003en\u003c/em\u003e = 10) of pollen grains in polar view of\u003cem\u003e\u0026nbsp;Hippeastrum\u0026nbsp;\u003c/em\u003especies. \u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"560\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003eSpecie \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 219px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; Polar \u0026nbsp;Diameter \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 219px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Equatorial Diameter\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003eRange\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003ex\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003eRange\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003ex\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. angustifolium\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e30.0-37.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e34.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e60.0-62.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e60.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. aulicum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e50.0-57.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e54.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e105.0-110.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e106.7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. blossfeldiae\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e37.5-43.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e41.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e65.0-70.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e67.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. brasilianum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e42.5-47.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e45.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e67.5-72.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e70.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. breviflorum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e33.8-37.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e35.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e67.5-72.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e70.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. calyptratum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e47.5-55.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e52.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e87.5-95.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e92.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. canastrense\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e42.5-46.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e43.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e55.0-58.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e56.5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. cipoanum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e45.0-47.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e46.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e66.2-70.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e68.1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. elegans\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e37.5-40.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e39.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e65.0-70.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e67.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. glauscesens\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e42.5-47.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e45.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e70.0-75.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e72.7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. morelianum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e40.0-50.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e44.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e65.0-70.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e67.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. papilio\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e45.0-55.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e48.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e77.5-80.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e79.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. puniceum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e32.5-37.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e35.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e70.0-80.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e73.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. reginae\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e37.5-42.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e39.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e55.0-58.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e56.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. reticulatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e37.5-42.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e40.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e65.0-70.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e67.4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. santacatarina\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e45.0-47.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e46.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e50.0-55.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e52.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. stapfianum\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e33.8-40.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e36.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e72.5-82.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e80.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. striatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e32.5-40.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e37.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e60.0-65.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e62.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. stylosum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e35.0-40.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e38.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e62.5-70.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e66.7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 122px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. vittatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 140px;\"\u003e\n \u003cp\u003e45.0-50.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 79px;\"\u003e\n \u003cp\u003e47.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 135px;\"\u003e\n \u003cp\u003e65.0-70.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e67.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cstrong\u003eTable 3\u0026nbsp;\u003c/strong\u003eMeasurements (\u0026micro;m) (n = 10) of apertures and exine layers of pollen grains of \u003cem\u003eHippeastrum\u0026nbsp;\u003c/em\u003especies.\u0026nbsp;\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"840\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003eSpecies\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 189px;\"\u003e\n \u003cp\u003eSulcus\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"3\" valign=\"top\" style=\"width: 274px;\"\u003e\n \u003cp\u003eExine thickness\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003eLumen\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eMuri\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003eLength\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003eWidth\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003eExine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003eSexine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003eNexine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003eDiameter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eWidth\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. angustifolium\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e56.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e9.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e4.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. aulicum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e101.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e8.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e3.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. blossfeldiae\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e59.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e9.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e3.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. brasilianum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e66.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e7.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e2.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. breviflorum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e64.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e6.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e3.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. calyptratum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e85.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e7.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e6.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. canastrense\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e48.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e8.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. cipoanum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e64.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e9.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e3.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. elegans\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e\u0026nbsp;57.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e10.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e3.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e____\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. glauscesens\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e70.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e7.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e4.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. morelianum\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e61.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e11.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e4.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. papilio\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e75.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e9.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e4.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e0.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. puniceum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e70.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e10.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e5.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e___\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. reginae\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e52.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e7.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e3.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e____\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. reticulatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e41.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e9.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e2.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e0.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. santacatarina\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e44.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e9.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. stapfianum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e74.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e14.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e____\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e____\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. striatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e51.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e20.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e4.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. stylosum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e61.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e7.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e2.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 166px;\"\u003e\n \u003cp\u003e\u003cem\u003eH. vittatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e62.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e13.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 85px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e4.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cstrong\u003eTable 4\u0026nbsp;\u003c/strong\u003eCumulative variance and vector values of principal component analysis (PCA).\u0026nbsp;\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 192px;\"\u003e\n \u003cp\u003ePrincipal Axis\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 120px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e1\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 142px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e2\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 192px;\"\u003e\n \u003cp\u003eVARIANCE (%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 120px;\"\u003e\n \u003cp\u003e63.06\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 142px;\"\u003e\n \u003cp\u003e12.93\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 192px;\"\u003e\n \u003cp\u003eCharacters\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 262px;\"\u003e\n \u003cp\u003eAxis\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 192px;\"\u003e\n \u003cp\u003eLargest diameter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 120px;\"\u003e\n \u003cp\u003e0.5160\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 142px;\"\u003e\n \u003cp\u003e-0.0884\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 192px;\"\u003e\n \u003cp\u003eSmallest diameter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 120px;\"\u003e\n \u003cp\u003e0.1900\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 142px;\"\u003e\n \u003cp\u003e0.1281\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 192px;\"\u003e\n \u003cp\u003ePolar diameter\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 120px;\"\u003e\n \u003cp\u003e0.1934\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 142px;\"\u003e\n \u003cp\u003e0.3489\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 192px;\"\u003e\n \u003cp\u003eEquatorial diameter\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 120px;\"\u003e\n \u003cp\u003e0.5114\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 142px;\"\u003e\n \u003cp\u003e-0.1433\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 192px;\"\u003e\n \u003cp\u003eSulcus length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 120px;\"\u003e\n \u003cp\u003e0.6225\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 142px;\"\u003e\n \u003cp\u003e-0.0875\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 192px;\"\u003e\n \u003cp\u003eSulcus width\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 120px;\"\u003e\n \u003cp\u003e0.0746\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 142px;\"\u003e\n \u003cp\u003e-0.3179\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 192px;\"\u003e\n \u003cp\u003eMuri width\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 120px;\"\u003e\n \u003cp\u003e0.0183\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 142px;\"\u003e\n \u003cp\u003e0.5369\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 192px;\"\u003e\n \u003cp\u003eLumen diameter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 120px;\"\u003e\n \u003cp\u003e0.0727\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 142px;\"\u003e\n \u003cp\u003e0.6607\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"brazilian-journal-of-botany","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"brjb","sideBox":"Learn more about [Brazilian Journal of Botany](https://www.springer.com/journal/40415)","snPcode":"40415","submissionUrl":"https://www.editorialmanager.com/brjb/default2.aspx","title":"Brazilian Journal of Botany","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Asparagales, Morphology, Pollen, Taxonomy","lastPublishedDoi":"10.21203/rs.3.rs-6785943/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6785943/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eNative to the American continent, the genus \u003cem\u003eHippeastrum\u003c/em\u003e Herb. is distributed from Mexico to Argentina. In Brazil, it is represented by approximately 30 species. This study examined the pollen grains of 20 taxa within the genus to determine whether differences in pollen morphology could support a clearer delimitation and characterization of subgenera and subordinate species. Pollen grains were treated with 40% lactic acid, measured, described, and photomicrographed. Non-acetolyzed pollen grains were examined using scanning electron microscopy. The analyzed taxa have large to very large pollen grains, shed as monads, monosulcate, with an elliptical shape in polar view and a reticulate or retipilate sexine. Species with a retipilate sexine could be distinguished based on the characteristics and organization of pila, whereas those with a reticulate sexine could be differentiated by murus, lumen, and columella characters. Thus, it was possible to develop a palynological key. Qualitative and quantitative analyses revealed differences in pollen morphology among species. However, pollen characters were generally uniform among subgenera. Overall, pollen characters proved informative for delimiting and describing species within the genus \u003cem\u003eHippeastrum\u003c/em\u003e.\u003c/p\u003e","manuscriptTitle":"Palynological analysis of representatives of Hippeastrum Herb. (Amaryllidaceae: Amaryllidoideae), with implications for systematics","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-11 11:03:16","doi":"10.21203/rs.3.rs-6785943/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Minor revisions","date":"2025-07-04T06:44:22+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2025-06-09T18:27:54+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-06-09T17:58:56+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-06-05T14:24:42+00:00","index":"","fulltext":""},{"type":"submitted","content":"Brazilian Journal of Botany","date":"2025-05-31T10:29:02+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"brazilian-journal-of-botany","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"brjb","sideBox":"Learn more about [Brazilian Journal of Botany](https://www.springer.com/journal/40415)","snPcode":"40415","submissionUrl":"https://www.editorialmanager.com/brjb/default2.aspx","title":"Brazilian Journal of Botany","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"be3a45c9-1a03-4d7f-b32c-63399e0a60e5","owner":[],"postedDate":"June 11th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2025-07-16T17:00:50+00:00","versionOfRecord":[],"versionCreatedAt":"2025-06-11 11:03:16","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6785943","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6785943","identity":"rs-6785943","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}