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Micromorphology of Sacoglottis perryi. A Marginal glandular seta B basilaminar gland, adaxial C laminar gland near margin, abaxial D sepal tip, inner side with terminal gland E paired glandular stipules and petiole scar F pollen inside sporangium G stigma with ephemeral lobes intact and showing secretion H stigma lobes shredded showing thin walls I gynoecium with diagnostic hirsute ovary J glandular disc with erose margin. Sources: A–CGillespie 2810D–JTripp 2984 (all US).
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Humiriaceae
have had little recent comparative morphological study except for their distinctive fruits. We surveyed the diversity of stamen structures in the family with consideration of dehiscence patterns and the evolutionary transitions between tetra- and disporangiate anthers. Novel interpretations of floral morphology support new combinations...
Citations
... For extrafloral nectaries, only those arranged in vegetative organs were considered, according to Weber et al. [5]. Note that calycine nectaries are not described for clade 2. Superscript numbers after the genera whose flowers are illustrated indicate the data source: 1 Farias et al. [30]; 2 Almeida [29]; 3 Narayana and Rao [31]; 4 Knapp and Mallet [32]; 5 Wurdack and Zartman [36]; 6 Prance and White [37]; 7 Niedenzu [42]; 8 Bonifácio et al. [39]; 9 Castro et al. [24]; 10 This work; 11 Oliveira [40]. Abbreviations: cn, calycine nectary; cn*, calycine nectary not confirmed by histochemical tests; el, elaiophore. ...
... When considering the occurrence of nectaries in vegetative organs (EFNs) and calycine nectaries within a phylogenetic framework of Malpighiales, it becomes apparent that, for parsimony, the most proximal relationship likely exists between elaiophores and sepal nectaries. The presence of calycine glands on the abaxial surface of the sepals is reported for the following families of Malpighiales: Euphorbiaceae [29,30,57], Linaceae [31], Passifloraceae [32][33][34], Humiriaceae [35,36], Chrysobalanaceae [37,38], Caryocaraceae [4,40,41,56], and Elatinaceae [39,42]. Additionally, evidence of secretory structures was also observed on the margins of the sepals of Bergia perennis and the apex of the sepals of Elatine gratioloides [39]. ...
Discussing homology relationships among secretory structures remains a relatively underexplored area in botanical research. These structures are widely dispersed within Malpighiales, one of the largest orders of eudicots. Within Malpighiales, both extranuptial and nuptial nectaries are present, and they do not seem homoplastic or share evolutionary connections. Particularly in Malpighiaceae, extensive research has focused on the ecological interactions mediated by glands. Botanists largely agree that elaiophores in sepals of Neotropical Malpighiaceae have evolved from extrafloral nectaries on leaves. However, the evolutionary origin of elaiophores has yet to be thoroughly examined, particularly in comparison to outgroups. This study provides empirical evidence on the ontogeny of elaiophores and investigates their evolutionary origins and homology relationships across different lineages of Malpighiales using comparative anatomy. Our findings suggest that elaiophores are likely homologous to extranuptial nectaries found in sepals of other Malpighiales lineages, originating from nectaries on leaves. This discussion is a starting point for future studies exploring the evolution of nectaries found in flowers, whether extranuptial or nuptial, and their potential origins from nectaries in vegetative organs such as leaves. Understanding these relationships could shed light on the selective pressures influencing floral morphologies.
... 12 Amaral et al. (2017). 13 Wurdack and Zartman (2019). 14 Crockett (2010). ...
... Finally, the Salicoids had nectar connective glands recovered as a synapomorphy in our analysis. Nectar connective glands have recently been suggested as a new morphological synapomorphy for Humiriaceae (Wurdack and Zartman 2019). However, this exudate is also recorded in the filament or connective glands of Violaceae within the Salicoids (Feng 2005;Wurdack and Zartman 2019). ...
... Nectar connective glands have recently been suggested as a new morphological synapomorphy for Humiriaceae (Wurdack and Zartman 2019). However, this exudate is also recorded in the filament or connective glands of Violaceae within the Salicoids (Feng 2005;Wurdack and Zartman 2019). In fact, only persistent endosperm has ever been proposed as a morphological homoplasy to circumscribe the Salicoids, making the nectar connective glands reported in this study a prominent character to further explore in this group (Stevens 2001, onwards). ...
Connective glands are important morphological characters for the taxonomy of some genera of Malpighiaceae, with few recent studies having just elucidated these glands' anatomical and ecological functions. In order to test the systematic relevance of connective glands to the currently accepted phylogenetic informal clades of Malpighiaceae, we characterised the anatomy and/or histochemistry of two-thirds of Malpighiaceae genera and ten species from nine families of Malpighiales to test: 1. Do connective glands occur in the flowers of all informal clades of Malpighiaceae?; and 2. Are they taxonomically relevant to characterise those clades? We sampled 25 genera and 26 species of Malpighiaceae, processing their anthers using traditional anatomical methods and characterising their glands using light microscopy and SEM imaging. Selected species were subjected to histochemical tests, and an additional 21 genera and 33 species of Malpighiaceae and nine families (ten species) of Malpighiales were included in our sampling from the literature. Three anatomical characters were scored, coded and mapped using Maximum Likelihood methods onto the molecular phylogeny of Malpighiaceae. All sampled species of Malpighiaceae showed connective glands characterised as epidermal or trichomal elaiophores. Our character-mapping analyses recovered connective elaiophores as a new synapomorphy for Malpighiaceae. Different types of epidermal or trichomal elaiophores were recovered as homoplasies for the Christianella and Banisteriopsis clades and the genera Byrsonima, Camarea and Cottsia. Our analyses also recovered the glands' place of insertion in the stamen and the exudate type as potential new synapomorphies or homoplasies for the families of Malpighiales sampled. Our results propose the connective elaiophores as a new synapomorphy for Malpighiaceae and hypothesise the role that different staminal glands might play in the systematics of Malpighiales. Further comprehensive anatomical studies are still needed for the staminal glands of most families of this order to shed new light on the patterns recovered in our study. Citation: de Almeida RF, Arévalo-Rodrigues G, de Morais IL, Cardoso-Gustavson P (2023) Evolution of connective glands reveals a new synapomorphy for Malpighiaceae and the hidden potential of staminal glands for Malpighiales systematics. PhytoKeys 232: 109-131. https://doi.
... These glands can be helpful to understand plant systematics and evolution. They not only constitute valuable characters for plant classification, species delimitation, and evolutionary studies Marazzi et al. 2019;Wurdack and Zartman 2019;Afzal et al. 2021;Mehltreter et al. 2021) but also aid to explain environmental interactions associated to herbivory (Lange and Del-Claro 2014), plant fitness (Vesprini et al. 2003;Mesquita-Neto et al. 2020), and niche distribution (Lange et al. 2021). ...
Nectar plays important roles in the relationship between plants and other organisms, both within pollination systems and as a defense mechanism. In the latter case, extranuptial nectaries (ENNs) usually attract patrolling arthropods that reduce herbivory. ENNs have been frequently reported within the “xeric clade” of Bromeliaceae, but their occurrence in other groups of bromeliads is largely unexplored, especially considering their position, secretory activity and structure. After observing the presence of ants constantly patrolling the inflorescences of Pitcairnia burchellii Mez, we searched for the presence, secretory activity, and structure of ENNs in this species. We also provide a brief review of the occurrence ENNs in Bromeliaceae. The distribution of nectaries was assessed using ant-exclusion experiments, while structural analysis was performed using standard methods for light and scanning electron microscopy. The presence of sugars in the secretion was assessed by thin-layer chromatography and glucose strip tests. Nectaries in P. burchelli are non-structured glands on the adaxial surface of floral bracts and sepals. Bracts and sepals are distinct spatial units that act over time in the same strategy of floral bud protection. Literature data reveals that ENNs might be more common within Bromeliaceae than previously considered, comprising a homoplastic feature in the family. Future perspectives and evolutionary and taxonomic implications are discussed.
... Humiriaceae belongs to Malpighiales (APG IV 2016) and has 65 species in eight genera, of which Endopleura and Hylocarpa are represented by only one species (Wurdack & Zartman 2019). The family is almost exclusively Neotropical, occurring mainly in the tropical forests from southern Brazil to Central America (Cuatrecasas 1961), except for Sacoglottis gabonenses (Baill.) ...
... Humiriaceae is composed mostly of shrubs or treelets, with alternate leaves having serrate (e.g., Humiriastrum and Sacoglottis) or entire (e.g., Humiria and Vantanea) leaf margins. The most important reproductive structure for genera and/or species delimitation is the morphological variation of the androecium (Wurdack & Zartman 2019). ...
... Inflorescences corymbose, terminal and axillary; peduncle 3-4 cm long, cylindrical, glabrescent. Flowers with pedicels 1-2 cm long, cylindrical, hirsute; sepals 1-1.5 × 1 mm, ovate, ciliate; petals 3-4 × 1-1.5 mm, deciduous, lanceolate, hirsute; stamens 20, (Wurdack & Zartman 2019). The species has a greater morphological similarity with V. bahiaensis, whose morphological delimitation was presented in the comments of that species. ...
Belonging to Malpighiales, Humiriaceae has about 65 species in eight genera, distributed almost exclusively in the Neotropics, occurring from southern Brazil to Central America. In Brazil, 35 species have been recorded (ca. 54% of the family’s diversity), wherein the greatest diversity is found in the Amazon (27 spp.), Atlantic Forest (9 spp.) and Cerrado (6 spp.) domains. This work is part of the “Flora do Espírito Santo” project, which aims to contribute to the knowledge of the state’s floristic diversity. Field expeditions were carried out from January/2019 to March/2020. Humiriaceae, in the state of Espírito Santo, is represented by six species distributed in four genera: Humiria, Humiriastrum, Sacoglottis and Vantanea. Among the registered genera, Vantanea is the most diversified, with three species. In the dense rain forest, all Humiriaceae species recorded here were found, followed by pioneer formations of restingas, with four species and seasonal semideciduous forest, with one species. The main morphological characters that assist in the delimitation of Humiriaceae species in Espírito Santo, are androecium characteristics, such as length (homodynamous or heterodynamous), number of stamens and the number of their series, as well as anther morphology.
... The Humiriaceae is a relatively small monophyletic flowering plant family within the order Malpighiales. The Humiriaceae contains eight genera (Sacoglottis, Schistostemon, Humiriastrum, Humiria, Hylocarpa, Endopleura, Duckesia and Vantanea), and circa 65 extant species (Cuatrecasas, 1961;Herrera et al., 2010Herrera et al., , 2014Wurdack and Zartman, 2019). ...
... Although there are numerous records of stranded drift endocarps of S. amazonica from the coasts of northern South America, Central America, Caribbean Region, Gulf of Mexico, and North America as far north as North Carolina (L'Ecluse, 1605;Sloane, 1696aSloane, , b, 1725Hemsley, 1897;Morris, 1889Morris, , 1895Cuatrecasas, 1961;Gunn, 1968;Gunn et al., 1984Gunn et al., , 1999Gunn and Dennis, 1972, 1973, Zies, 1997Dennis, 2000;Katz, 2001;Boykin, 2003;Sullivan, 2003Sullivan, , 2004Foreman, 2004;Norton, 2007Norton, , 2008Perry and Dennis, 2010;Witherington, 2011, 2017;Anon., 2021b), it is strange that the species has not become naturally established northwards of Costa Rica or any of the offshore Caribbean Islands other than Trinidad and Tobago (Guppy, 1917;Ridley, 1930;Acevedo-Rodriguez and Strong, 2012;Wurdack and Zartman, 2019). Guppy (1917) suggested that the seeds within drifting S. amazonica endocarps probably lose their germination capacity after six months. ...
During early December 2020, LMN discovered a drift endocarp of the Blister Pod or Cojon de Burro Scaoglottis amazonica Mart. measuring 40mm in length and 30mm in width stranded on Fanore Beach, Fanore Mor, County Clare (53.1196 o N, 9.2882 o W), on the Atlantic coast of western Ireland. The specimen represents the second record of S. amazonica from Irish maritime shores and the fourth from the NW European Atlantic.
There many families and orders where arborescent traits are basal while herbaceous traits are derived. But the reverse situation is quite rare. To explain this asymmetrical distribution, a hypothesis about plant evolution is advanced.
Information concerning typical features of families whose names starting with the letter ‘H’ is provided. Besides, important economic plants with notes on the larger genera are provided and, also, photographic pictures and line drawings.
Small shrubs, treelets or large trees; wood hard, aromatic, often with balsamic sap, heartwood reddish, alburnum yellow or yellowish; bark smooth, striate or fissured. Leaves alternate, simple, distichous, coriaceous or subcoriaceous, penninerved, brochidodromous, from small to large, margins entire, crenulate, dentate or slightly serrate, petiolate or rarely sessile, sometimes decurrent along branches, often punctate-glandulose (nectariferous glands) near margins (or basal) on underside; stipules very small, geminate, often deciduous. Inflorescences (synflorescences) axillary, pseudo terminal or rarely terminal, cymose-paniculate (or thysoid) or often corymbiform, of dichasial type and trichotomous, but through reduction with dichotomous or alternate (cincinnate) branching; branchlets often with incrassate ends, articulate; pedicels short, articulate; bracts and bracteoles persistent or deciduous, small, amplectant. Flowers hermaphroditic, actinomorphic, slightly aromatic; sepals 5, persistent, thick and carnose at base, thinner toward margins, suborbicular or triangular, more or less connate in tube or cupule in varying degrees, glabrous, pubescent or tomentose outside, margins always ciliate, sometimes with marginal or dorsal glands; aestivation quincuncial or imbricate; petals 5, deciduous or sometimes persistent, free, thick or membranaceous, usually 3–5-veined, oblong, linear or oblong-lanceolate, acute to obtuse, 1.5–16 mm long, exceptionally 30–40 mm long, rarely with gland at top, margins smooth, sometimes with tooth at one side near apex, above glabrous, below glabrous or pilose, white, greenish white, or yellowish white, rarely red or purple; aestivation contorted, cochlear or quincuncial; stamens monadelphous, numerous and pluriseriate or of variable number, 1–2-seriate; filaments filiform (when numerous), slender and flexuose, or thick, complanate, linear, acute at apex, straight and glabrous or papillose; connate at base in a more or less long tube, alternating in different lengths, sometimes five alternating with petals are trifurcate at apex and triantheriferous; sometimes with additional staminodial filaments; anthers dorsifixed or subbasifixed; thecae 2, bilocular, laterally attached, ellipsoid-oblong and each cell dehiscing by longitudinal slit, or 4 unilocular, rounded or ellipsoid disjunct thecae (2 lateral and 2 basal), dehiscing by detachment, or two unilocular, disjunct, basal, dehiscing by detachment; connective thick, fleshy, ovoid or lanceolate, obtuse at apex or most commonly produced in apiculum or linguiform appendage; pollen shed as isopolar monads; intrastaminal free disc surrounding ovary, membranaceous or subcoriaceous, nectariferous, annular, tubular or cupular, dentate, lobate, laciniate or composed of 10–20 free scales; style single, entire, columnar, erect, as long as stamens (1.2–12 mm long, exceptionally 30 mm long or shorter (0.3–0.9 mm long), rarely very short and rather thick or longer; stigma narrowly or broadly capitate, 5-lobate or 5-radiate; ovary superior, ovoid or ellipsoid, sessile, syncarpous, (4-)-5(−7) septate with axile placentation, locules uniovulate or biovulate; ovules anatropous, epitropous with 2 integuments, pendant at inner angle of ovary cells, micropyle pointing upward, raphe ventral; when 2 ovules present in each cell, superposite and lower one hanging from longer funiculus. Fruit a drupe (drupoid), from small (not exceeding 16 mm) to large, black, blackish, reddish, yellow, or orange when mature, usually aromatic; exocarp with smooth surface, glabrous, or pilose; mesocarp hard-fleshy varying from pulpy to fibrous, subcoriaceous texture, often aromatic and edible; endocarp woody, usually very hard, compact or with many resin-filled, round cavities, rarely spongy-woody, 5 septate, commonly with only 1–2(−5) seeds developed; surface smooth, bullate, rugose, or tuberculate, slightly striate or strongly costate; with dehiscence germinal, provided with as many longitudinal opercula or valves as carpels, which may open or be pushed away by emerging embryo at germination of seed inside fruit; often subapical foramina present in Duckesia, Endopleura, Humiria, and Humiriastrum. Seeds oblong, with double testa, exterior often adherent to endocarp, inner membranaceous, thin; one or two per locule; embryo straight or slightly curved, cotyledons oblong or ovate, often subcordate at base, radicle half as long, endosperm fleshy and oily.
A more detailed species-by-species taxonomic history of the family is given in Cuatrecasas (1961) and only the major contributions before that date are highlighted here. The first genera of Humiriaceae were described and illustrated by Aublet (1775), Houmiri Aubl., with the single species H. balsamifera (later Latinized to Humiria) and Vantanea guianensis Aubl. Aublet placed these genera in the class Polyandria and order Monogynia of the Linnaean system. Nees and Martius (1824) described the genus Helleria, which was later placed in synonymy with Vantanea Aubl. (Bentham 1853). Martius (1827) described the genus Sacoglottis with a single species S. amazonica and also Humirium floribundum (now a variety of Humiria balsamifera). He considered these genera to be related to Meliaceae, Symplocaceae, and Styracaceae, an opinion supported by Endlicher (1840). Jussieu (1829) was the first to treat the group at the family level including two genera: Humirium with two species and Helleria of Nees and Martius (1824) also with two species that are now in Vantanea. He also recognized Sacoglottis as belonging in this family, so in effect he brought together all three genera of the family that had been described by 1829. Planchon (1848) compared the Humiriaceae with Erythroxylaceae and the Linaceae due to morphological affinities to Erythroxylum P. Browne and Roucheria Planch, respectively. Bentham (1853) published a concise summary of the family treating the three genera Humiria, Sacoglottis, and Vantanea. The separation of these genera was mainly based on the number of stamens, Vantanea with numerous stamens, Humiria with 20 stamens, and Sacoglottis with 10 stamens. He also noted the number of ovules, two in Vantanea and one in Sacoglottis, but Humiria with one or two ovules. Baillon (1873) united all known Humiriaceae into the single genus Houmiri, which included 15 species placed in five sections: Humirium (20 stamens), Aubrya (10 free stamens), Saccoglottis (10 stamens + 10 staminodia), Vantanea (20–60 stamens), and Vantaneoides (never formally published), and the genus was placed in his Linaceae. Baillon (1874) repeated the same division of Humiriaceae and placed his series Humiriées in the Linaceae. The next major contribution to the family was by Urban (1877) in Martius’ Flora brasiliensis. Urban recognized three genera: Humiria with three species, Saccoglottis with nine species, and Vantanea with four species. Saccoglottis was grouped into three subgenera: Eusaccoglottis with two Brazilian species but recognizing that the African Aubrya gabonensis of Baillon (1862) belonged here; subgenus Humiriastrum with three species; and subgenus Schistostemon with four species. Urban divided the family into two groups distiguished by the number of stamens and number of cells in the thecae of the anthers: Vantanea with thecae with two locules and many stamens and Humiria and Saccoglottis with unilocular thecae and fewer stamens. The subgenera were divided by staminal characters: subg. Eusaccoglottis with 10 stamens, subg. Humiriastrum with 20 undivided stamens, and subg. Schistostemon with 20 stamens, the latter with 5 tridentate ones bearing 3 anthers. Urban (1893) later described Sacoglottis glaziovii Urb. based on material sent to him from Brazil, and this was transferred to Humiriastrum by Cuatrecasas (1961). Hallier (1921) followed Baillon in keeping the Humiriaceae and Linaceae united into a single family. Ducke (1922–1933; 1935–1937) in a series of important papers about Amazon plants published four new species of Vantanea and five of Sacoglottis, as well as several new varieties. Two of Ducke’s new species of Sacoglottis were later given generic status by Cuatrecasas (1961) one named appropriately Duckesia (after Adolpho Ducke whose work was a major contribution to the taxonomy of the Amazonian species of Humiriaceae) and the other Hylocarpa. Ducke (1938) summarized much of his work on Humiriaceae and provided a key to the genera Sacoglottis and Vantanea, placing them in Linaceae. Winkler (1931) in the second edition of Pflanzenfamilien followed Baillon and Hallier in including the family in Linaceae, but as the subfamily Humirioideae. Otherwise Winkler (1931) followed Urban’s system without any changes.
The Humiriaceae consists exclusively of woody tropical plants that are mainly large trees and a few smaller shrubs or sprawling shrubs in rocky places. In some of the larger trees, buttresses may occur.
