Diversity of flowers in Apocynaceae. (A) Aspidosperma pyrifolium, Aspidospermateae; (B) Hancornia speciosa, Willughbeieae; (C) Couma rigida, Willughbeieae; (D) Tabernaemontana solanifolia, Tabernaemontaneae; (E) Condylocarpon isthmicum, Alyxieae; (F) Thevetia peruviana, Plumerieae; (G) Allamanda puberula, Plumerieae, (H) Wrightia coccinea, Wrightieae; (I) Adenium obesum, Nerieae; (J) Mandevilla atroviolacea, Mesechiteae; (K) Pentopetia grevei, Periplocoideae; (L) Secamone parviflora, Secamonoideae; (M) Ceropegia tihamana, Ceropegieae; (N) Orbea lutea, Ceropegieae; (O) Calotropis procera, Asclepiadinae; (P) Oxypetalum rusticum, Oxypetalinae [Photos of panels (A–G,I,J,O,P) by Rapini; panels (H,K–N) by Meve].

Diversity of flowers in Apocynaceae. (A) Aspidosperma pyrifolium, Aspidospermateae; (B) Hancornia speciosa, Willughbeieae; (C) Couma rigida, Willughbeieae; (D) Tabernaemontana solanifolia, Tabernaemontaneae; (E) Condylocarpon isthmicum, Alyxieae; (F) Thevetia peruviana, Plumerieae; (G) Allamanda puberula, Plumerieae, (H) Wrightia coccinea, Wrightieae; (I) Adenium obesum, Nerieae; (J) Mandevilla atroviolacea, Mesechiteae; (K) Pentopetia grevei, Periplocoideae; (L) Secamone parviflora, Secamonoideae; (M) Ceropegia tihamana, Ceropegieae; (N) Orbea lutea, Ceropegieae; (O) Calotropis procera, Asclepiadinae; (P) Oxypetalum rusticum, Oxypetalinae [Photos of panels (A–G,I,J,O,P) by Rapini; panels (H,K–N) by Meve].

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Apocynaceae (the dogbane and milkweed family) is one of the ten largest flowering plant families, with approximately 5,350 species and diverse morphology and ecology, ranging from large trees and lianas that are emblematic of tropical rainforests, to herbs in temperate grasslands, to succulents in dry, open landscapes, and to vines in a wide variet...

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... 12,[15][16][17][18][19] This particular case reflects an evolution directly related to the plasticity of the protodermis which, together with the emergence of the pollinia, seems to have been mainly responsible for the high 14,20 giving rise to thousands of species distributed almost worldwide. 21,22 Among the different parts of the flower having secretory tissues, the style head stands out for its fundamental importance for pollination in Apocynaceae. 15,[22][23][24] Style head ...
... 21,22 Among the different parts of the flower having secretory tissues, the style head stands out for its fundamental importance for pollination in Apocynaceae. 15,[22][23][24] Style head ...
... The style head is ubiquitous in Apocynaceae, 12,22,24 and is one of the main morphological synapomorphies of the family. It forms from a postgenital fusion of the apices of the two carpel primordia during flower development. ...
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Nuptial glands are very diverse and associated with different pollination mechanisms. The greater the specifici-ty in the pollen transfer mechanism from anther to stigma, the greater the morphological elaboration of flowers and functional complexity of the nuptial glands. In Apocynaceae, pollination mechanisms reached an extreme specificity, a fact that was only possible due to an extreme morphological synorganization and a profusion of floral glands. Although these glands are of different types, the vast majority have secretory cells only in the epi-dermis. In general, these epidermal cells produce many different compounds at the same time, and previous studies have demonstrated that in the style head, the functional complexity of epidermis has become even greater. Four types of style head are found in the family, which have different degrees of functional complexity in relation to the secretion produced and pollen dispersal mechanism. The secretion is fluid in types I, II and III, and the pollen is dispersed and adhered to the pollinator by the secretion produced by the style head. In type IV, the secretion hardens and acquires a specific shape, moulded by the spatial constraints of the adjacent floral organs. This evolutionary alteration is accompanied by changes in the structure and arrangement of the secre-tory cells, as well as in pollen aggregation and position of stigma. Histochemical analysis has shown that the secretion is mixed and highly complex, especially in the style head type IV, where the secretion, called translator , is formed by a rigid central portion, which adheres to the pollinator, and two caudicles that attach to two pollinia. The translator has a distinct composition in its different parts. Further studies are needed to answer the new questions that have arisen from the discovery of this highly functional complexity of the secretory tissue.
... Overall, both regions presented higher range sizes than forests. The two results follow the trend of recent radiation of the family from forests to drier environments (Bitencourt et al. 2021), with niche conservatism potentially playing an important role in the differences observed in the richness patterns of forests and savannas. Considering solely the savanna environments included in our analysis, the widespread distribution observed for some species can also be linked to the overall simpler topography of the region. ...
... Apocynaceae are hypothesized to have originally occupied wet forests in austro-and boreotropical latitudes until the Eocene-Oligocene climate transition, later developing multiple adaptations (e.g. wind-dispersed, twining, pollinia-bearing) that allowed them to disperse into open vegetation types (Bitencourt et al. 2021). For lianas, our result follows that reported for different families and other regions of the world since they are usually not as abundant in shrublands as they are in forest-type physiognomies (Harper et al. 2005). ...
... In addition, the distribution of species richness was positively associated with a latitudinal gradient (Fig. 1B), the highest richness cell values being observed in tropical latitudes between 14°S and 10°S, and the mean richness values being higher in subtropical latitudes between 22°S and 26°S. These results show that most of the diversity of the group is concentrated in tropical and subtropical forests, following previous quantifications of Apocynaceae diversity in the Neotropics (BFG 2015) and the biogeographical and evolutionary trends reported for the family (Bitencourt et al. 2021). In these environments, the complexity of the forest structure has been hypothesized to enhance the diversification of climbing modes and, consequently, to increase the species richness of lianas. ...
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... Apocynaceae, the dogbane family, are one of the ten largest families in angiosperms, comprising 378 genera and more than 5000 species (Endress et al., 2018). The majority of species in this family are found in tropical and subtropical regions of the world, and they exhibt wide diversity in terms of their habitats, growth forms and morphological characteristics (Fishbein et al., 2018;Ollerton et al., 2019;Bitencourt et al., 2021). Furthermore, many Apocynaceae species have high economic values. ...
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Apocynaceae are one of the ten species-richest angiosperm families. However, the backbone phylogeny of the family is yet less well supported, and the evolution of plastid genome structure has not been thoroughly studied for the whole family. Herein, a total of 101 complete plastomes including 35 newly sequenced, 24 reassembled from public raw data and the rest from NCBI GenBank database, representing 26 of 27 tribes of Apocynaceae, were used for comparative plastome analysis. Phylogenetic analyses were conducted using a combined plastid data matrix of 77 protein-coding genes from 162 taxa, encompassing all tribes and 41 of 49 subtribes of Apocynaceae. Plastid genome lengths ranged from 150,897 bp in Apocynum venetum to 178,616 bp in Hoya exilis. Six types of boundaries between the inverted repeat (IR) regions and single copy (SC) regions were identified. Different sizes of IR expansion were found in three lineages, including Alyxieae, Ceropegieae, and Marsdenieae, suggesting multiple expansion events of the IRs over the SC regions in Apocynaceae. The IR regions of Marsdenieae evolved in two ways: expansion towards the large single copy (LSC) region in Lygisma + Stephanotis + Ruehssia + Gymnema (Cosmopolitan Clade), and expansion towards both LSC and small single copy (SSC) region in Dischidia-Hoya alliance and Marsdenia (Asia-Pacific Clade). Six coding genes and five non-coding regions were identified as highly variable, including accD, ccsA-ndhD, clpP, matK, ndhF, ndhG-ndhI, trnG(GCC)-trnfM(CAU), trnH(GUG)-psbA, trnY(GUA)-trnE(UUC), ycf1, and ycf2. Maximum likelihood and Bayesian phylogenetic analyses resulted in nearly identical tree topologies and produced well-resolved backbone relationships. Fifteen clades in the Apocynaceae backbone were identified. The subfamily Periplocoideae were embedded in the Apocynoid grade and were sister to the Echiteae-Odontadenieae-Mesechiteae clade with high support values. Three tribes (Melodineae, Vinceae, and Willughbeieae), the subtribe Amphineuriinae, and four genera (Beaumontia, Ceropegia, Hoya, and Stephanotis) were not resolved as monophyletic. Our work sheds light on the backbone phylogenetic relationships in the family Apocynaceae and offers insights into the evolution of Apocynaceae plastomes using the most densely sampled plastome dataset to date.
... With about 3400 species, 181 genera, 17 subtribes, and 5 tribes (Asclepiadeae, Ceropegieae, Eustegieae, Fockeeae, Marsdenieae), this subfamily comprises 60% of all species in the family Apocynaceae (Fishbein & al., 2018;Endress & al., 2019). In addition to its ancestral African distribution (Bitencourt & al., 2021), four independent radiation events occurred in the Americas (Rapini & al., 2003), of which onethe speciesrich MOG-clade comprising subtribes Metastelmatinae, Oxypetalinae and Gonolobinae, along with five smaller subtribes of the tribe Asclepiadeaeincludes 75% of all American Asclepiadoideae species (Rapini & al., 2007;Silva & al., 2012;Endress & al., 2019) (Fig. 1). While Secamonoideae are mostly woody with predominantly lianescent or twining growth forms and to a lesser extent also small erect shrubs (Endress & al., 2019), Asclepiadoideae are defined by a wide array of non-woody life forms. ...
... These various woody and non-woody growth forms thrive in habitats ranging from lowland rainforests in Southeast Asia to diverse, often disturbed, forests and shrublands in Africa and the Americas (Endress & al., 2019). Interestingly, Bitencourt & al. (2021) observed a family-wide evolutionary transition in habitat occupation, ranging from stable and dense tropical rainforests typical of the rauvolfioids and some apocynoids, towards highly diverse twining lineages in more open, semi-arid, pioneer vegetation in Asclepiadoideae. ...
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The Apocynaceae subfamilies Secamonoideae and Asclepiadoideae have undergone several transitions during their evolution with regard to growth form and degree of woodiness. In this study, we present a wood anatomical overview of both subfamilies that complements previous work on the remaining Apocynaceae. Detailed microscopic wood descriptions using light and scanning electron microscopy were performed on 60 species that cover most Secamonoideae genera and all major woody Asclepiadoideae lineages. Our observations are in line with subfamilial and (sometimes sub)tribal delimitations. Furthermore, we present for the first time an overview of the estimated number of shifts from herbaceousness to (phylogenetically) derived woodiness in Asclepiadoideae, along with a derived woody species list with distribution and habitat information. In total, at least 168 derived woody species resulting from at least 28 independent woodiness transitions were found, with drought possibly being one of the main drivers of most of these transitions.
... While the DIVALIKE model reconstructs tropical Asia as ancestral area for the tribe (though with a rather high probability for Africa or Africa-tropical Asia, suppl. Fig. S16), the DEC model gives widespread Africa-tropical Asia as ancestral area, a result also obtained by Bitencourt & al. (2021). In our analysis, stem age of Marsdenieae is estimated to be somewhat younger (30.2 Ma) than in with 39.5 Ma, and crown age somewhat older (28.8 Ma vs. 22.8 Ma) than in . ...
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... While the DIVALIKE model reconstructs tropical Asia as ancestral area for the tribe (though with a rather high probability for Africa or Africa-tropical Asia, suppl. Fig. S16), the DEC model gives widespread Africa-tropical Asia as ancestral area, a result also obtained by Bitencourt & al. (2021). In our analysis, stem age of Marsdenieae is estimated to be somewhat younger (30.2 Ma) than in with 39.5 Ma, and crown age somewhat older (28.8 Ma vs. 22.8 Ma) than in . ...
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Using the Apocynaceae as an example, the present paper demonstrates how the new tool Rhakhis highlights taxonomic problems in a large vascular plant family and, at the same time, invites specialists in the family to solve these problems with the aim of producing an up‐to‐date taxonomy. This taxonomy can then be used to estimate species and genus numbers, thus approaching the goal of completing our picture of the diversity surrounding us and providing a solid basis for science and conservation. The most recent estimates for genus and species number in the Apocynaceae are provided.
Preprint
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Diante da atual crise da biodiversidade e visto que os recursos para sua conservação são limitados, estabelecer áreas prioritárias tem se mostrado um recurso-chave e, ao mesmo tempo, desafiador. Os métodos tradicionais são contestáveis, já que o número de espécies e o endemismo são indicadores pobres do nível de ameaça das regiões.