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Blumea Supplement 7. Flowers of Annonaceae: Morphology, Classification, and Evolution
... The rarity of infrastylar EGC in the Annonaceae may be attributed to the uncommon occurrence of basal placentation and/or openings on the ovaries while the prevalence of suprastylar EGC may be linked to the ubiquity of wet-type stigmas in the family. Among the 110 genera in Annonaceae (Couvreur et al., 2019), basal placentation has primarily been recorded in Annona, Artabotrys, Duguetia, Goniothalamus p.p., Guatteria, and Miliusa p.p. (van Heusden, 1992). Among early divergent angiosperms, infrastylar EGC has previously only been reported from Illiciaceae (Illicium floridanum: Williams et al., 1993) and Schisandraceae (Kadsura longipedunculata: Lyew et al., 2007; Schisandra sphenanthera . ...
... Even though the new evidence provided here does not support the interpretation of the Cyathocalyx gynoecium as syncarpous, it is noted that the solitary Cyathocalyx carpel is likely to have undergone an evolutionary increase in the number of ovules: Cyathocalyx species have 8-38 ovules per carpel, compared to only 1-10 in Drepananthus and fewer than seven in Cananga (Wang & Saunders, 2006;Surveswaran et al., 2010). As the Cyathocalyx-Drepananthus-Cananga clade is collectively sister to the genus Lettowianthus, which only has approximately two ovules per carpel (van Heusden, 1992;Surveswaran et al., 2010), an increased number of ovules per carpel is likely an apomorphic trait for Cyathocalyx. The closely related genera Cyathocalyx and Drepananthus might have undergone evolutionary adaptations in opposing directions, with Cyathocalyx experiencing a reduction in carpel number but an increase in ovule number per carpel, while some Drepananthus lineages increased the number of carpels per flower (Wang, 2004). ...
... 4.5 Gynoecial diversity and adaptations that overcome the limitations of apocarpy in Annonaceae: potential reproductive significance Annonaceae flowers are largely apocarpous with 6-20 carpels (van Heusden, 1992). Apocarpy is generally considered deleterious, restricting pollen-tube growth between different carpels and hence causing imbalanced fertilization when carpels are subject to uneven pollen deposition (Carr & Carr, 1961;Armbruster et al., 2002). ...
Most species in the early divergent angiosperm family Annonaceae are apocarpous, with very diverse gynoecial morphologies. Although several Annonaceae genera with apocarpous flowers are known to possess an extragynoecial compitum (EGC) that enables intercarpellary pollen-tube growth to enhance fertilization success, the occurrence of EGC across the whole family remains obscure. Twenty Annonaceae species from all four subfamilies (Anaxagoreoideae, Ambavioideae, Annonoideae, and Malmeoideae) are examined here, with anatomical evidence revealing the occurrence of stigmatic exudate-mediated suprastylar EGC in all four subfamilies. Histological and ontogenetic studies furthermore indicate that trichomes in Cananga and Drepananthus form a confluent zone that connects adjacent stigmas, providing a structural premise for suprastylar EGC. Infrastylar EGC are reported in the Annonaceae for the first time in several genera, including Artabotrys, Annona, and Miliusa, associated with the opening on the ovary ventral groove and/or basal placentation. In addition to the sister genera Isolona and Monodora that are clearly syncarpous, flowers of the distantly related genus Cyathocalyx with a unicarpellate gynoecium have also been hypothesized to be syncarpous. Evidence of carpel vasculature and primordium development does not support that the solitary Cyathocalyx carpel is originated from carpel fusion, however, although the increased number of ovules renders it functionally similar to syncarpy. Gynoecial features, including the extensive occurrence of EGC and the increased number of ovules per carpel (consonant with reduction to a solitary carpel) in Cyathocalyx, may have evolved to overcome limitations associated with apocarpy and possibly contribute to the reproductive success and diversification of the family.
... Long considered a "natural family" (Hutchinson, 1923), the considerable diversity of Annonaceae has been the subject of various formal and informal subfamilial classifications. Such works were based on different morphological characters (flowers, fruits, pollen) and resulted in different groupings of genera (Bentham, 1862;Baillon, 1868;Engler & Diels, 1901;Hutchinson, 1923;Fries, 1959;Walker, 1971;Van Heusden, 1992;Van Setten & Koek-Noorman, 1992). A review of historical classification systems is presented in Chatrou & al. (2012a). ...
... Finally, the phylogenomic work presented here and from previous phylogenetic studies have relied on foundational taxonomic work on this diverse pantropical plant family (Fries, 1959;Maas, 1983;Chatrou & al., 2012a). A dynamic systematic community focusing on Annonaceae have increased our understanding on the morphological diversity within the family (Le Thomas, 1980Thomas, , 1981Van Setten & Koek-Noorman, 1986, 1992Van Heusden, 1992), allow-ing for the identification of key characters in delimiting infrafamilial ranks congruent with phylogenomic molecular data. Encouragingly, this community is active and collaborative, with new species continuously being described even in recent times, reflecting ongoing taxonomic work (2022 onwards, e.g., Jaikhamseub & al., 2022;Ortiz-Rodriguez, 2022;Bazante & al., 2023;Dagallier & al., 2023;Damthongdee & al., 2023a,b;Leeratiwong & al., 2023;Ortiz-Rodriguez & Linares, 2023;Page, 2023;Yang & al., 2023;Ferreira & al., 2024;Satthaphorn & al., 2024;Damthongdee & al., in press). ...
... Finally, the phylogenomic work presented here and from previous phylogenetic studies have relied on foundational taxonomic work on this diverse pantropical plant family (Fries, 1959;Maas, 1983;Chatrou & al., 2012a). A dynamic systematic community focusing on Annonaceae have increased our understanding on the morphological diversity within the family (Le Thomas, 1980Thomas, , 1981Van Setten & Koek-Noorman, 1986, 1992Van Heusden, 1992), allow-ing for the identification of key characters in delimiting infrafamilial ranks congruent with phylogenomic molecular data. Encouragingly, this community is active and collaborative, with new species continuously being described even in recent times, reflecting ongoing taxonomic work (2022 onwards, e.g., Jaikhamseub & al., 2022;Ortiz-Rodriguez, 2022;Bazante & al., 2023;Dagallier & al., 2023;Damthongdee & al., 2023a,b;Leeratiwong & al., 2023;Ortiz-Rodriguez & Linares, 2023;Page, 2023;Yang & al., 2023;Ferreira & al., 2024;Satthaphorn & al., 2024;Damthongdee & al., in press). ...
Annonaceae is a major tropical plant family particularly diverse in tropical rain forests of the world. Although the classification and systematics of the family has significantly improved over the past decade, the most recent classification was based on a reduced set of plastid markers and incomplete genus‐level taxon sampling. This classification recognised 4 subfamilies and 20 tribes. Yet, several important problems persisted, especially the phylogenetic placement of the African genus Meiocarpidium , resolution of intertribal relationships within subfamily Malmeoideae, resolution and classification within the diverse tribe Miliuseae with 23 genera, and the contrasting placement of the liana genus Artabotrys when using nuclear versus plastid data. Here, using a previously published Annonaceae‐specific nuclear bait kit, we generated for the first time a complete genus‐level (108 taxa) phylogenomic tree of the family based on 373 loci. We show that Meiocarpidium is sister to Ambavioideae and should be considered as a tribe and not a separate subfamily. Artabotrys is recovered as belonging to tribe Duguetieae, and not Xylopieae as previously inferred based on plastid data, and is sister to two other African liana genera, Letestudoxa and Pseudartabotrys . Finally, we were able to resolve intertribal relationships within subfamily Malmeoideae and most of the relationships within tribe Miliuseae. Nevertheless, we recovered strong gene conflict mainly at the backbone of the tribe, probably linked to a rapid diversification at its origin, leading to substantial incomplete lineage sorting. We suggest that this conflict will be hard to resolve. Using this novel phylogenomic framework we recognize 25 subtribes, 21 as new, to improve the infrafamilial classification of Annonaceae.
... The classification of Desmos and Dasymaschalon have been carried out previously based on their morphology (Heusden 1992;Keßler 1993;Setten and Koek-Noorman 1992), pollen (Walker 1971), and molecular data (Wang et al. 2012;Guo et al. 2017). However, no information using leaf anatomical data is provided. ...
... Large size of druse crystals is also found in Friesodielsia (Nikmah 2020, unpublished data). This result supports Heusden (1992) that Desmos is more similar to Friesodielsia than Dasymaschalon. ...
... Desmos, Dasymaschalon, and Friesodielsia are closely related genera according to the phylogenetic study (Wang et al. 2012;Xue et al. 2019), but their floral structures are different (Chiu 2012;Guo et al. 2018). The epidermal anatomy supports a classification based on the floral morphology by Heusden (1992) who divided Annonaceae into 20 informal groups, of which Desmos, Dasymaschalon, and Friesodielsia belong to the Friesodielsia Group. The epidermal anatomy of three Friesodielsia species (Nikmah 2020, unpublished data) found that the anatomical characters of these three genera were overlapped. ...
... The classification of Desmos and Dasymaschalon have been carried out previously based on their morphology (Heusden 1992;Keßler 1993;Setten and Koek-Noorman 1992), pollen (Walker 1971), and molecular data (Wang et al. 2012;Guo et al. 2017). However, no information using leaf anatomical data is provided. ...
... Large size of druse crystals is also found in Friesodielsia (Nikmah 2020, unpublished data). This result supports Heusden (1992) that Desmos is more similar to Friesodielsia than Dasymaschalon. ...
... Desmos, Dasymaschalon, and Friesodielsia are closely related genera according to the phylogenetic study (Wang et al. 2012;Xue et al. 2019), but their floral structures are different (Chiu 2012;Guo et al. 2018). The epidermal anatomy supports a classification based on the floral morphology by Heusden (1992) who divided Annonaceae into 20 informal groups, of which Desmos, Dasymaschalon, and Friesodielsia belong to the Friesodielsia Group. The epidermal anatomy of three Friesodielsia species (Nikmah 2020, unpublished data) found that the anatomical characters of these three genera were overlapped. ...
Nikmah IA, Rugayah R, Chikmawati T. 2020. Leaf anatomical variation in Desmos Lour. and Dasymaschalon (Hook. f. & Thomson) Dalla Torre & Harms species (Annonaceae). Biodiversitas 21: 3317-3330. The relationships between Desmos and Dasymaschalon are debated for along time. Those two genera have high morphological similarities, especially in their generative character (moniliform monocarps). Therefore, sterile specimens of Desmos are difficult to be distinguished from Dasymaschalon. Leaf anatomy in paradermal section of 20 taxa (12 species of Desmos, eight species of Dasymaschalon) have been carried out. The data were used to support the interspecific and intergeneric delimitation of Desmos and Dasymaschalon. Desmos and Dasymaschalon are two distinct genera mainly distinguished based on the anticlinal wall undulation of epidermal cells and supported by variation of the crystal type, and size. The anticlinal wall undulation of Desmos is almost straight to slightly wavy, and never sinuous, meanwhile, Dasymaschalon varies from almost straight to deeply sinusoid. The crystals of Desmos consist of rhombohedric, druse type A, and druse type B crystals, whereas Dasymaschalon has prism, druse type A, druse type B, and drue type C crystals.
... The Annonaceae genus Meiogyne possesses elaborate inner petal corrugations (Fig. 1I) that have often been referred to as "glands" (e.g., van Heusden, 1992), although without confirmation of glandular function. The corrugations (strumae) on Meiogyne hainanensis petals (Shao & Xu, 2015, as "Oncodostigma") have been shown to be rich in polysaccharides and to be associated with thrips larvae. ...
... This might be correlated with a reduction in carpel number per flower and/or flower number per plant to avoid exceeding the optimal fruit-carrying capacity of the plant. Increased ovule numbers are observed in some species of Fissistigma and Uvaria (Annonaceae), with up to 20 and 30 ovules, respectively (van Heusden, 1992). The functional significance of this in relation to seedset has never been evaluated, however. ...
... Partial syncarpy has been reported in the Annonaceae genera Annona, Anonidium, Cananga, Fusaea, and Pseudartabotrys (Deroin, 1988(Deroin, , 1997van Heusden, 1992;Deroin & Bidault, 2017). Although the hypothesis remains untested, it is possible that this incomplete syncarpy endows the same benefits as full syncarpyallowing pollen grains deposited on any stigma to fertilize any ovule in the flowerwhile enabling the development of separate fruit monocarps (fruit units derived from individual carpels) that can be dispersed individually, and hence avoiding the dispersal constraints imposed by large fruits (discussed further in Section 8). ...
Potential key functional floral traits are assessed in the species‐rich early divergent angiosperm family Annonaceae. Pollinators (generally beetles) are attracted by various cues (particularly visual, olfactory and thermogenic), with pollinators rewarded by nectar (generally as stigmatic exudate), heat and protection within the partially enclosed floral chamber. Petals sometimes function as pollinator brood sites, although this may be deceptive. Annonaceae species are self‐compatible, with outcrossing promoted by a combination of protogyny, herkogamy, floral synchrony and dicliny. Pollination efficiency is enhanced by pollen aggregation, changes in anthesis duration, and pollinator trapping involving a close alignment between petal movements and the circadian rhythms of pollinators. Most Annonaceae flowers are apocarpous, with syncarpy restricted to very few lineages; fertilization is therefore optimized by intercarpellary growth of pollen tubes, either via stigmatic exudate (suprastylar extragynoecial compitum, EGC) or possibly the floral receptacle (infrastylar EGC). Although Annonaceae lack a distinct style, the stigmas in several lineages are elongated to form ‘pseudostyles’ that are hypothesized to function as sites for pollen competition. Flowers can be regarded as immature fruits in which the ovules are yet to be fertilized, with floral traits that may have little selective advantage during anthesis theoretically promoting fruit and seed dispersal. The plesiomorphic apocarpous trait may have been perpetuated in Annonaceae flowers since it promotes the independent dispersal of fruit monocarps (derived from separate carpels), thereby maximizing the spatial/temporal distance between seedlings. This might compensate for the lack of genetic diversity among seeds within fruits arising from the limited diversity of pollen donors.
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... Scheffer (1885) did not accept that these morphological differences warranted recognition as distinct genera, although he maintained the two as separate sections within a more broadly defined Cyathocalyx. Subsequent taxonomic opinion has been split between these two approaches, with some adopting Hooker & Thomson's (1872) narrower generic delimitation (e.g., Hutchinson, 1923Hutchinson, , 1964Fries, 1959;Keßler, 1993) whilst others have adopted Scheffer's (1885) broader delimitation (e.g., Prantl, 1891;Boerlage, 1899;Engler & Diels, 1900;Engler, 1908;Sinclair, 1955;Van Heusden, 1992;Van Setten & Koek-Noorman, 1992;Bakker, 2000;Wang & Saunders, 2006a,b). ...
... Cananga possesses several characteristics that are likely to be synapomorphic, including: apiculate staminal connectives (Van Heusden, 1992); inaperturate pollen with a reduced proximal exine and without nexine foliations (Doyle & Le Thomas, in press) that is released in loose tetrahedral to tetragonal tetrads (Walker, 1971); and arillate seeds (Van Setten & Koek-Noorman, 1992), although arils are notably also present in 'Cyathocalyx pruniferus' in the Drepananthus clade. Cyathocalyx s.str. ...
... The adaxial prominence of the leaf midrib is a potential candidate (character 1, as numbered in Table 3), since the leaf midribs of other members of the ambavioid clade (e.g., Ambavia, Cleistopholis, Lettowianthus, Mezzettia) are generally impressed or flush with the lamina surface adaxially. The absence of an indument on the carpels (character 15) of Cananga and Cyathocalyx s.str. is also possibly synapomorphic given the presence of such hairs in Lettowianthus (Van Heusden, 1992). ...
Generic delimitation of Cyathocalyx and Drepananthus has been controversial, with some authors recognizing them as distinct genera, and others recognizing a more broadly defined Cyathocalyx, inclusive of Drepananthus. Some doubt also exists regarding the relationships between these taxa and Cananga. Molecular phylogenetic analyses are presented based on combined psbA‐trnH spacer, trnL‐F, matK and rbcL sequences. Results indicate that Cananga, Cyathocalyx s.str. and Drepananthus form three generally well‐supported clades, although with inadequate resolution of relationships among the three clades. Morphological variation is re‐evaluated, and the narrower delimitation of Cyathocalyx proposed, necessitating 21 new nomenclatural combinations following the recognition of Drepananthus as a distinct genus. Divergence times are estimated using an uncorrelated lognormal distributed (UCLD) relaxed molecular clock. Historical biogeographical analysis suggests that the ambavioid lineage originated in Africa, with subsequent dispersal into Asia. Alternative hypotheses for this dispersal, involving rafting on the Indian tectonic plate versus migration via the extensive boreotropical forests associated with the Eocene thermal maximum, are evaluated, and the latter route identified as the most consistent with the divergence age estimates and the geological and palaeoclimatic data.
... However, in the tribe Annoneae, a haploid chromosome number of seven has only been reported in the genus Annona; in this tribe, x = 8 has been reported in the genus Asimina (Tanaka and Okada, 1972), Disepalum (Johnson, 1989), Goniothalamus (Sauer and Ehrendorfer, 1984), and Neostenanthera (Walker, 1972). Moreover, x = 8 has also been reported in sister lineages such as the tribes Monodoreae (genera Isolona, Monodora, and Uvariopsis) (Walker, 1972) and Uvarieae (genera Dasymaschalon, Desmos, Fissistigma, Friesodielsia, Melodorum, Mitrella, Sphaerocoryne, and Uvaria) (Walker, 1972;Okada and Ueda, 1984;Sauer and Ehrendorfer, 1984;Van Heusden, 1992) and in the more basal tribes Duguetieae (genera Duguetia and Fusaea) and Xylopieae (genera Artabotrys and Xylopia) (Morawetz, 1984;Van Heusden, 1992). This could indicate that x = 7 in Annona may have arisen by aneuploid chromosome loss from ancestors with x = 8. ...
... However, in the tribe Annoneae, a haploid chromosome number of seven has only been reported in the genus Annona; in this tribe, x = 8 has been reported in the genus Asimina (Tanaka and Okada, 1972), Disepalum (Johnson, 1989), Goniothalamus (Sauer and Ehrendorfer, 1984), and Neostenanthera (Walker, 1972). Moreover, x = 8 has also been reported in sister lineages such as the tribes Monodoreae (genera Isolona, Monodora, and Uvariopsis) (Walker, 1972) and Uvarieae (genera Dasymaschalon, Desmos, Fissistigma, Friesodielsia, Melodorum, Mitrella, Sphaerocoryne, and Uvaria) (Walker, 1972;Okada and Ueda, 1984;Sauer and Ehrendorfer, 1984;Van Heusden, 1992) and in the more basal tribes Duguetieae (genera Duguetia and Fusaea) and Xylopieae (genera Artabotrys and Xylopia) (Morawetz, 1984;Van Heusden, 1992). This could indicate that x = 7 in Annona may have arisen by aneuploid chromosome loss from ancestors with x = 8. ...
... 1986a) and Duguetia (Morawetz, 1984). Hexaploid and octoploid ploidy levels were also observed in the former genus Rollinia (Walker, 1972;Van Heusden, 1992) that is now included in the genus Annona (Rainer, 2007). In this work, this hexaploidy was also observed in Annona neosalicifolia previously named Rollinia neosalicifolia. ...
Genome duplication or polyploidy is one of the main factors of speciation in plants. It is especially frequent in hybrids and very valuable in many crops. The genus Annona belongs to the Annonaceae, a family that includes several fruit tree crops, such as cherimoya (Annona cherimola), sugar apple (Annona squamosa), their hybrid atemoya (A. cherimola × A. squamosa) or pawpaw (Asimina triloba). In this work, genome content was evaluated in several Annona species, A. triloba and atemoya. Surprisingly, while the hybrid atemoya has been reported as diploid, flow cytometry analysis of a progeny obtained from an interspecific cross between A. cherimola and A. squamosa showed an unusual ploidy variability that was also confirmed karyotype analysis. While the progeny from intraspecific crosses of A. cherimola showed polyploid genotypes that ranged from 2.5 to 33%, the hybrid atemoyas from the interspecific cross showed 35% of triploids from a total of 186 genotypes analyzed. With the aim of understanding the possible implications of the production of non-reduced gametes, pollen performance, pollen size and frequency distribution of pollen grains was quantified in the progeny of this cross and the parents. A large polymorphism in pollen grain size was found within the interspecific progeny with higher production of unreduced pollen in triploids (38%) than in diploids (29%). Moreover, using PCR amplification of selected microsatellite loci, while 13.7% of the pollen grains from the diploids showed two alleles, 41.28% of the grains from the triploids amplified two alleles and 5.63% showed up to three alleles. This suggests that the larger pollen grains could correspond to diploid and, in a lower frequency, to triploid pollen. Pollen performance was also affected with lower pollen germination in the hybrid triploids than in both diploid parents. The results confirm a higher percentage of polyploids in the interspecific cross, affecting pollen grain size and pollen performance. The occurrence of unreduced gametes in A. cherimola, A. squamosa and their interspecific progeny that may result in abnormalities of ploidy such as the triploids and tetraploids observed in this study, opens an interesting opportunity to study polyploidy in Annonaceae.
... Boutiquea was already included in the tribe Annoneae but only based on pollen and morphological characters 2 . This sister relationship is consistent with floral and pollen morphology: both genera have an elevated floral torus, very short sepals, elongated petals, three apically connivent inner petals that form a mitriform dome over the reproductive organs 16 , and pollen grains with a granular infratectum that are released as tetrads 17,18 . In addition, both genera have septate anthers in which the sporogenous cells are partitioned by transverse or longitudinal walls comprising sterile tissue 19 Table 2. Descriptive statistics and best-fitting substitution model for each of the eight chloroplast regions and the concatenated datasets. ...
... The two genera are also palynologically similar, with large pollen tetrads with a minutely granular exine structure 18,23,24 . Fusaea differs from Duckeanthus, however, in its fused carpels, a conspicuous ring of staminodes, and the fused calyx 16 . ...
... The genus has been placed in tribe Miliuseae in previous classifications 2, 26, 27 because of their 'miliusoid' stamens (in which the connective does not extend over the thecae) with an obtuse apex. Miliusoid stamens have been shown to be morphologically highly variable, however, and likely homoplasious in both the Miliuseae and the wider family 16,28 . The phylogenetic results presented here, however, indicate that Phoenicanthus is the next-divergent branch subsequent to the tribe Fenerivieae, and strongly supported as sister to a clade comprising the tribes Dendrokingstonieae, Monocarpieae and Miliuseae (BS = 95; JK = 83; PP = 1, Fig. 2; Supplementary Fig. S7): the tribe Miliuseae is shown as sister to the Monocarpieae, with these two tribes collectively sister to the Dendrokingstonieae. ...
The Annonaceae, the largest family in the early-divergent order Magnoliales, comprises 107 genera and c. 2,400 species. Previous molecular phylogenetic studies targeting different taxa have generated large quantities of partially overlapping DNA sequence data for many species, although a large-scale phylogeny based on the maximum number of representatives has never been reconstructed. We use a supermatrix of eight chloroplast markers (rbcL, matK, ndhF, psbA-trnH, trnL-F, atpB-rbcL, trnS-G and ycf1) to reconstruct the most comprehensive tree to date, including 705 species (29%) from 105 genera (98%). This provides novel insights into the relationships of five enigmatic genera (Bocagea, Boutiquea, Cardiopetalum, Duckeanthus and Phoenicanthus). Fifteen main clades are retrieved in subfamilies Annonoideae and Malmeoideae collectively, 14 of which correspond with currently recognised tribes. Phoenicanthus cannot be accommodated in any existing tribe, however: it is retrieved as sister to a clade comprising the tribes Dendrokingstonieae, Monocarpieae and Miliuseae, and we therefore validate a new tribe, Phoenicantheae. Our results provide strong support for many previously recognised groups, but also indicate non-monophyly of several genera (Desmopsis, Friesodielsia, Klarobelia, Oxandra, Piptostigma and Stenanona). The relationships of these non-monophyletic genera—and two other genera (Froesiodendron and Melodorum) not yet sampled—are discussed, with recommendations for future research.
... Sinclair 1955Maxwell 1989;Li 1993). Many taxonomists such as Finet & Gagnepain (1906), Merrill (1915), Hutchinson (1923), Fries (1959), Bân (1975), Klucking (1986), Van Heusden (1992), Kebler (1993), Koek-Noorman et al. (1997), Sun et al. (2002) disagreed with the taxonomic treatment of Safford and supported Dalla Torre & Harms' decision to raise Dasymaschalon to the generic rank in 1901. ...
... According to Guo et al. (2018), the genus Dasymaschalon is closely related to Friesodielsia Steenis s. str. (1948:458) and is morphologically distinct with small trees and shrubs (rarely climbers), shallow conical torus, 2-3-cohering petals arranged in one whorl, echinate ornamentation and pollen with thin exine (Walker 1971;Le Thomas 1980, 1981Van Heusden 1992;Kebler 1993;Doyle & Le Thomas 2012). ...
A new Annonaceae species, Dasymaschalon leilamericanum is described from Mount Lantoy Key Biodiversity Area (KBA) in the Municipality of Argao, Cebu, Philippines. D. leilamericanum is distinguished from closely related species by morphological characters including laminar size, leaf areolation, and seed/fruit shape and size. Significant differences observed in a multivariate analysis of morphological data for D. leilamericanum, D. clusiflorum, D. filipes, D. ellipticum, and D. blumei indicate D. leilamericanum is a distinct species. A phylogenetic tree analysis performed using maturase K (matK) and ribulose 1,5-biphosphate carboxylase (rbcL) as molecular markers failed to match D. leilamericanum with other Dasymaschalon species. Taken together, the results of morphological and molecular analysis indicate a unique evolutionary pathway for D. leilamericanum with its genus.
... Friesodielsia (Annonaceae; Uvarieae) was erected by Van Steenis (1948: 458) to replace the illegitimate name Oxymitra (Blume 1830: 71) Hooker & Thomson (1855: 145), which was a later homonym of the liverwort Oxymitra Bischoff in Lindenberg (1829: 124). Although Van Steenis recognized 52 species in Friesodielsia from Africa and Asia, many taxonomists questioned whether the taxa in these two geographical regions were truly congeneric (e.g., Verdcourt 1971, Van Heusden 1992. The African and Asian species are morphologically distinct with regard to overall flower shape (broad vs elongate), inner petal arrangement (loosely coherent vs apically connivent), monocarp shape (moniliform vs subglobose), seed number per monocarp (up to five vs one or two) and pollen exine (coarsely verrucate vs echinate) (Verdcourt 1971, Walker 1971, Van Heusden 1992, Guo et al. 2017a. ...
... Although Van Steenis recognized 52 species in Friesodielsia from Africa and Asia, many taxonomists questioned whether the taxa in these two geographical regions were truly congeneric (e.g., Verdcourt 1971, Van Heusden 1992. The African and Asian species are morphologically distinct with regard to overall flower shape (broad vs elongate), inner petal arrangement (loosely coherent vs apically connivent), monocarp shape (moniliform vs subglobose), seed number per monocarp (up to five vs one or two) and pollen exine (coarsely verrucate vs echinate) (Verdcourt 1971, Walker 1971, Van Heusden 1992, Guo et al. 2017a. ...
Friesodielsia (Annonaceae; Uvarieae) was erected by Van Steenis (1948: 458) to replace the illegitimate name Oxymitra (Blume 1830: 71) Hooker & Thomson (1855: 145), which was a later homonym of the liverwort Oxymitra Bischoff in Lindenberg (1829: 124). Although Van Steenis recognized 52 species in Friesodielsia from Africa and Asia, many taxonomists questioned whether the taxa in these two geographical regions were truly congeneric (e.g., Verdcourt 1971, Van Heusden 1992). The African and Asian species are morphologically distinct with regard to overall flower shape (broad vs elongate), inner petal arrangement (loosely coherent vs apically connivent), monocarp shape (moniliform vs subglobose), seed number per monocarp (up to five vs one or two) and pollen exine (coarsely verrucate vs echinate) (Verdcourt 1971, Walker 1971, Van Heusden 1992, Guo et al. 2017a).
... Annonaceae (custard apple family) is one of the most primitive families of the Angiosperm (Hutchinson 1969) that comprises about 120-130 genera and 2500 species (van Heusden 1992). Based on APG system (Angiosperm Phylogeny Group system)) (1998), APG II system (2003) and APG III system (2009), Annonaceae was placed most closely related to the small Magnoliid family Eupomatiaceae. ...
... This problem effects the identification of unidentified species. Whereas, species of Annonaceae can be classified based on morphological characters (Maas and Westra 1984;Westra 1985;Morawetz and Le Thomas 1988;van Heusden 1992;van Setten and Koek-Noorman 1992;Johnson and Murray 1995;Doyle and Le Thomas 1996;Svoma 1998;Johnson 2003;Maas et al. 2003;Tsou and Johnson 2003;Doyle 2005, 2006;Su and Saunders 2006;Maas et al. 2007;Couvreur 2009;Huysmans et al. 2010;Surveswaran et al. 2010;Weerasooriya and Saunders 2010a). So that, the identity of species become unclear, and the taxonomical position of this species is doubtful, misidentified and misnamed (Kwon et al. 2016). ...
Lestari DA, Azrianingsih R. 2019. Species identity and taxonomical position of selected species of Annonaceae based on trnL molecular marker. Biodiversitas 20: 1012-1019. Identification based on morphological characters could be difficult when some characters are not visible. The absence of some morphological characters could effect for identification of species identity and taxonomical position of the species. Confirmation by the DNA data is needed to support species identification. The sequence was used in this research is trnL intron as non-coding sequence DNA, based on chloroplast DNA. The research aimed to estimate of species identity and determine of taxonomical position of species of Annonaceae based on trnL sequences. Methods were used through steps of DNA extraction, DNA amplification, DNA sequencing, and data analysis to selected species of Annonaceae from Purwodadi Botanic Garden (PBG), the trnL intron sequences of 10 Annonaceae species from GenBank database and two species of Magnoliaceae as out-group. Results showed that trnL sequence as non-coding gene explains the different groupings with the previous groupings in Annonaceae to this observed species. trnL sequence can estimate of species identity as much as 30%, caused by changes of nucleotide bases from mutation and missing data. Polymorphism of DNA sequences showed that 61.18% sites as conserved region, 24.05% sites as polymorphic variation and 14.76% sites as alignment gaps. Oxymitra sp. is in group (monophyletic) with Mitrephora javanica because they are genetically in close relationship (Uvariae tribes and Annonoideae sub-family), Popowia sp. is in group with Orophea enterocarpa, because they are genetically in close relationship (Miliuseae tribes and Malmeoideae sub-family).
... Delimitation of the Uvaria group within the Annonaceae has focused on the following four main characters: a scrambling or climbing growth habit, stellate hairs over all organs, valvate sepals and imbricate petals. There has been broad taxonomic agreement about the core composition of this grouping within the family (Hooker and Thomson 1855; Finet and Gagnepain 1906;Sinclair 1955;Van Heusden 1992;Meade 2001); however, for example Fries (1959) included many additional genera of less obvious affinity. Disagreement has been greater regarding taxonomic differentiation of core genera within the group, primarily owing to varying emphasis on a variety of petal characters (Fries 1959;Nguyẽn 1974Nguyẽn , 2000 (often cited by his first name(s) as Bân or Tiẽn Bân); Van Heusden 1992; Utteridge 2000). ...
... Despite these recent origins within the Long Branch Clade (LBC), Uvaria is the third most species-rich genus in the Annoaceae, and on the basis of a sensu lato interpretation of Zhou et al. (2009), displays an elevated speciation rate compared with genera in the family as a whole (Couvreur et al. 2011). While this elevated speciation rate is partly attributable to the biogeographic radiation of Uvaria from Africa into Asia, it also reflects the sensu lato interpretation of Zhou et al. (2009), wherein all the Uvaria group genera of Van Heusden (1992) and Meade (2001) are subsumed into Uvaria on the principle of generic monophyly. In many branches of the Annonaceae, discrete differences in taxonomy and morphology readily distinguish adjoining monophyletic clades, and allow the maintenance of multiple smaller genera with much more recent origins than Uvaria (Couvreur et al. 2011;Chatrou et al. 2012). ...
This paper presents a revision of Uvaria L. (Annonaceae) in continental Asia and outlying islands, north of the Thailand–Malaysia border, on the basis of a combination of new morphological analyses and recent phylogenetic data. Two new taxonomic groupings are defined within the genus in Asia on the basis of detailed morphological character analysis in ~1800 specimens. Stamen and carpel structure are shown to be significantly more informative than calyx and corolla characters for subgeneric differentiation, and reliably discriminate basal and derived radiations when mapped onto recent molecular phylogenies for the genus in Asia. Thirty-three species and species varieties are recognised within Uvaria in continental Asia, incorporating taxa formerly assigned to Anomianthus Zoll., Cyathostemma Griff., Ellipeia Hook.f. & Thomson, Ellipeiopsis R.E.Fr. and Dasoclema J.Sinclair. Four new combinations are proposed, and outstanding taxonomic, nomenclatural and typification issues are resolved for included taxa. Keys for both flowering and fruiting material are included, and distribution data are provided for all taxa.
... Dasymaschalon by Hooker and Thomson (1885), then was raised as the generic rank as Dasymaschamon by Dalla Torre & Harms (1901) which followed by the many toaxonomists (e.g. Finet & Gagnepain 1906, Merrill 1915, Hutchinson 1923, Fries 1959, Bân 1975, Klucking 1986, Heusden 1992, Keßler 1993, Koek-Noorman et al. 1997, Sun et al. 2002, but Safford (1912) was considered it as a section of Desmos Lour. (1790: 352), Desmos sect. ...
... (1948: 458) (Gou et al. 2012. As currently delimited, Dasymaschalon is characterized by small trees and shrubs (rarely climbers), the 2-3-cohering petals arranged in one whorl, a shallowly conical torus, and pollen has a thin exine and echinate ornamentation (Walker 1971, Le Thomas 1980, 1981, Van Heusden 1992, Keßler 1993, Doyle & Le Thomas 2012. The genus has been well studied Indochina and surrounding areas (e.g. ...
A new species of Dasymaschalon (Annonaceae), D. bachmaensis, from Bạch Mã NP, Thừa Thiên-Huế Province from North Central Coast region, Vietnam, is described and illustrated here. It shares several characteristics with D. glaucum, D. sootepense and D. wallichii (the leaves are glaucous, the petals are somewhat ovate and spirally twisted when mature, the stamen connectives are truncate and glabrous, the ovaries are hairy, and the stigmas are glabrous) but differs from them in having an obtuse leaf base, larger triangular-ovate sepals that are hairy on both surfaces, pale greenish-yellow petals, fewer stamens, smaller fruits that are not constricted between seeds, and smaller subglobose seeds. Details on distribution, ecology, phenology preliminary conservation assessment of D. bachmaensis, a key to Dasymaschalon species in Vietnam and a distribution map are provided.
... The macromorphology of the unknown Annonaceae from Vietnam was studied from six herbarium specimens (dried and pickled material: HUAF collectors 2009-03-19-ND;Chaowasku 129, 130, 131, 165, 166; see below for more details). The relevant morphological information of related taxa were taken from literature (Huber, 1985;van Heusden, 1992;Kessler, 1993;Mols & Kessler, 2000a, 2000b, 2003Mols et al., 2004a;Chaowasku et al., 2012bChaowasku et al., , 2013bChaowasku et al., , 2014Turner, 2012) or derived from herbarium specimens for Phoenicanthus obliquus (Hook. f. & Thomson) Alston [Huber 515,540,565 (E); Huber 518, 577 (L)]. ...
... Further, the monocarps of the unknown taxon from Vietnam are subsessile with a stout stipe (Fig. 2D), while those of Monocarpia are completely sessile ( Fig. 2B; Mols & Kessler, 2000b;Turner, 2012). Other features of the Vietnamese new genus fit well with the diagnostic traits of Monocarpieae (van Heusden, 1992;Mols & Kessler, 2000b;Chaowasku et al., 2012b;Chatrou et al., 2012): enlarged stigmas (more or less peltate and lobed; Fig. 2H, 5J), a percurrent tertiary venation of the leaves (Fig. 2C, 6), a reduced carpel number to 3 -4 per flower (Fig. 5A), multiple ovules per ovary (and hence seeds per monocarp) arranged in two rows, relatively large monocarps with a thick and hardened wall when dry (Fig. 2D, 5K), and spiniform endosperm ruminations (Fig. 5N). The tribe Monocarpieae is, therefore, enlarged to include the new genus from Vietnam. ...
Recent botanical expeditions in central Vietnam yielded an unknown species of Annonaceae that could not be confidently identified to subfamily, tribe, and genus. Preliminary BLAST® searches based on plastid data have suggested that this taxon is genetically closely-related to the following tribes of subfamily Malmeoideae: Malmeeae, Fenerivieae, Maasieae, Phoenicantheae, Dendrokingstonieae, Monocarpieae, and Miliuseae. Using representatives of Piptostigmateae, another tribe of Malmeoideae, as outgroups and including representatives of all other tribes of Malmeoideae, molecular phylogenetic analyses of seven combined plastid markers (rbcL, matK, ndhF, ycf1 exons; trnL intron; trnL-trnF, psbA-trnH intergenic spacers) inferred the enigmatic Vietnamese taxon as belonging to the monotypic tribe Monocarpieae. Detailed morphological comparisons between this taxon and its sister group, Monocarpia Miq., warranted the recognition of a second genus of Monocarpieae to accommodate our unknown taxon: Leoheo Chaowasku with a single species, Leoheo domatiophorus Chaowasku, D.T. Ngo & H.T. Le. The morphology of the new genus agrees well with the diagnostic traits of Monocarpieae, e.g., a percurrent tertiary venation of the leaves, a highly reduced number of carpels per flower, enlarged and lobed stigmas, multiple ovules/seeds per ovary/monocarp, considerably large monocarps with a hardened pericarp when dry, and spiniform ruminations of the endosperm. However, the new genus does not exhibit two characteristic features of Monocarpia: Terminal inflorescences and generally distinct intramarginal leaf veins. In addition, the new genus possesses three autapomorphic characters: Hairy domatia on the lower leaf surface, longitudinal ridges on the monocarp surface, and subsessile monocarps with a stout stipe. The tribe Monocarpieae is consequently enlarged to include the genus Leoheo. The enlarged Monocarpieae, along with the recently established monotypic tribe Phoenicantheae and two other related tribes, Dendrokingstonieae and Miliuseae, are discussed.
... Characters found only in (Malmea (Cremastosperma, Pseudoxandra)) are a lobed stigma (7), an autapomorphy of Cresmastosperma (Pirie, 2005, Figs. 3 and 5A) and the presence of staminodes (8) in a single species of Pseudoxandra (Figs. 3 and 5D). Staminodes are present in few genera of Annonaceae, notably in Anaxagorea, the sister genus to the rest of the family (Chatrou et al., 2012;van Heusden, 1992). Staminodes are also present in Eupomatiaceae, the sister family of Annonaceae (The Angiosperm Phylogeny Group, 2016;Endress, 1984). ...
... Thus, the evolution of a convex midrib is a synapomorphy of the oxandra euneura group. Another traditionally important feature in previous classifications of Annonaceae, and in the recognition of Oxandra, is the stamen shape (Prantl, 1891;van Heusden, 1992;Junikka et al., 2016). A narrow stamen with a tongue-like connective, found in Oxandra and other Malmeeae, is ancestral for the tribe. ...
Androdioecy is the rarest sexual system among plants. The majority of androdioecious species are herbaceous plants that have evolved from dioecious ancestors. Nevertheless, some woody and androdioecious plants have hermaphrodite ancestors, as in the Annonaceae, where androdioecious genera have arisen several times in different lineages. The majority of androdioecious species of Annonaceae belong to the Neotropical tribe Malmeeae. In addition to these species, Pseudoxandra spiritus-sancti was recently confirmed to be androdioecious. Here, we describe the morphology of male and bisexual flowers of Pseudoxandra spiritus-sancti, and investigate the evolution of androdioecy in Malmeeae. The phylogeny of tribe Malmeeae was reconstructed using Bayesian inference, maximum parsimony and maximum likelihood of 32 taxa, using DNA sequences of 66 molecular markers of the chloroplast genome, sequenced by next generation sequencing. The reconstruction of ancestral states was performed for characters associated with sexual systems and floral morphology. Sex ratio was analysed for Ephedranthus, Klarobelia, Pseudomalmea and Pseudoxandra spiritus-sancti. The phylogenetic analyses reconstructed three main groups in Malmeeae, ((Malmea (Cremastosperma, Pseudoxandra)) sister to the rest of the tribe, and ((Unonopsis (Bocageopsis, Onychopetalum)) sister to (Mosannona, Ephedranthus, Klarobelia, Oxandra, Pseudephedranthus fragrans, Pseudomalmea, Ruizodendron ovale). Hermaphroditism is plesiomorphic in the tribe, with four independent evolutions of androdieocy, which represents a synapomorphy of two groups, one that includes three genera and 14 species, the other with a single genus of seven species. Male flowers are unisexual from inception and bisexual flowers possess staminodes and functional stamens. Pseudoxandra spiritus-sancti is structurally androdioecious and unlikely to be functionally dioecious.
... The exine seems not essential in A. montana, that presents a special and rare reproductive characteristic in angiosperms. As the largest family within Magnoliales, Annonaceae exhibits an extraordinary variety of morphological and structural features over a long period of evolution [19,20]. Probably, more species with exine shedding might exist in basal angiosperms, especially in Annonaceae, since four Annona species (A. ...
Background
It is generally considered that exine is vital for interactions with stigmas, and pollen exine shedding during pollination has never been reported in angiosperms. However, Annona montana, located at Annonaceae family in basal angiosperms, was observed spontaneously shedding exine in stigma exudates and any watery solution.
Results
We demonstrated the entire process of A. montana pollen exine shedding in vivo and in vitro, verified that water is the only necessary medium for A. montana pollen shedding, and exine shedding is an unavoidable step before pollen germination. Since pollen volume of A. montana did not change remarkably before and after hydration, it is undoubtedly that exine shedding in this species might be related with exine, rather than the intine-absorbing water.
Conclusions
Pollen shedding exine in A. montana exhibits a special and rare reproductive characteristic in angiosperms, which will update what we have previously recognized that exine is essential in reproductive processes.
... Uvariopsis (Annonaceae, subfamily Annonoideae, tribe Monodoreae) (Chatrou et al., 2012;Guo et al., 2017) is a highly distinctive and easily recognised genus, since most of its species have unisexual flowers, a calyx with two basally connate sepals, and the petals in a single whorl of four (very rarely three, see below). Annonaceae are otherwise characterised by bisexual flowers with trimerous perianths (van Heusden, 1992). Most species of the genus Uvariopsis are cauliflorous small trees, the flowers being produced from the trunk, although some species are ramiflorous or bear axillary flowers (Kenfack et al., 2003;Couvreur et al., in press). ...
Background
The Ebo Forest area is a highly threatened centre of diversity in the Littoral Region of Cameroon, globally important for conservation with many threatened species including 68 threatened species of plant, yet not formally protected. The tropical African evergreen forest tree genus Uvariopsis Engl. & Diels (Annonaceae) is characterised by unisexual, usually cauliflorous flowers with a uniseriate corolla of four petals, and two sepals. Cameroon is the centre of diversity of the genus with 14 of the 19 known species.
Methods
The herbarium collection MacKinnon 51 from Ebo is hypothesized to represent a new species to science of Uvariopsis . This hypothesis is tested by the study of herbarium specimens from a number of herbaria known to hold important collections from Cameroon and surrounding countries.
Results
We test the hypothesis that MacKinnon 51 represents a new species to science, using the most recent dichotomous identification key, and comparing it morphologically with reference material of all known species of the genus. We make a detailed comparative morphological study focussing on three other Cameroonian species, Uvariopsis solheidii, U. korupensis and the sympatric U. submontana . In the context of a review of the pollination biology of Uvariopsis , we speculate that in a genus otherwise with species with dull, flesh-coloured (pink, red to brown) flowers pollinated (where known) by diptera, orthoptera and blattodea (flies, crickets and cockroaches), the glossy, pale yellow-green flowers of Uvariopsis dicaprio , with additional traits unique in the genus, may be pollinated by nocturnal moths. Based on MacKinnon 51, we formally name Uvariopsis dicaprio Cheek & Gosline (Annonaceae) as new to science, and we describe, and illustrate, and map it. Restricted so far to a single site in evergreen forest in the Ebo Forest, Littoral Region, Cameroon, Uvariopsis dicaprio is provisionally assessed as Critically Endangered using the IUCN, 2012 standard because the forest habitat of this species remains unprotected, and there exist imminent threats of logging and conversion to plantations.
Discussion
We show that the highest density of species of the genus (12), and of narrow endemics (5), is found in the Cross-Sanaga Interval of SE Nigeria and Western Cameroon. A revised key to the 14 Cameroonian species of Uvariopsis is presented. We review the other seven narrowly endemic and threatened species unique to the Ebo forest of Cameroon and discuss the phytogeographic affinities of the area.
Conclusions
Uvariopsis dicaprio adds to the growing list of species threatened with extinction at Ebo Forest due to current anthropogenic pressures.
... We analyzed 30 collections of Xylopia maasiana and around 40 of X. nitida. The study followed the standard methodology used in plant taxonomy, and morphological terminology follows Radford et al. (1974), Van Heusden (1992, and Van Setten and . In the description, the term stamens includes both fertile stamens and staminodes. ...
... Meiocarpidium becomes the Toward a phylogenetic reclassification of the subfamily Ambavioideae (Annonaceae): establishment of a new subfamily and a new tribe second diverging lineage of Annonaceae after Anaxagorea. The genus is easily identifiable because of the possession of lepidote indumentum (e.g., Heusden 1992;Keßler 1993), which is rarely found elsewhere in the family (i.e., in Duguetia; Maas et al. 2003). Several palynological features, especially the obvious intine extrusion (Thomas 1980;1981;Hesse et al. 1985), have suggested that Meiocarpidium may be placed phylogenetically somewhere near Anaxagorea. ...
A molecular phylogeny of the subfamily Ambavioideae (Annonaceae) was reconstructed using up to eight plastid DNA regions (matK, ndhF, and rbcL exons; trnL intron; atpB-rbcL, psbA-trnH, trnL-trnF, and trnS-trnG intergenic spacers). The results indicate that the subfamily is not monophyletic, with the monotypic genus Meiocarpidium resolved as the second diverging lineage of Annonaceae after Anaxagorea (the only genus of Anaxagoreoideae) and as the sister group of a large clade consisting of the rest of Annonaceae. Consequently, a new subfamily, Meiocarpidioideae, is established to accommodate the enigmatic African genus Meiocarpidium. In addition, the subfamily Ambavioideae is redefined to contain two major clades formally recognized as two tribes. The tribe Tetramerantheae consisting of only Tetrameranthus is enlarged to include Ambavia, Cleistopholis, and Mezzettia; and Canangeae, a new tribe comprising Cananga, Cyathocalyx, Drepananthus, and Lettowianthus, are erected. The two tribes are principally distinguishable from each other by differences in monoploid chromosome number, branching architecture, and average pollen size (monads). New relationships were retrieved within Tetramerantheae, with Mezzettia as the sister group of a clade containing Ambavia and Cleistopholis.
... Annonaceae is one of the primitive families from Angiosperms that plays an ecological role in species diversity, especially in tropical lowland rainforest ecosystems, namely Paleotropic and Neotropic. Annonaceae consists of 120-130 genera and 2100-2300 species with habitus of tree, shrub and woody climber (Van Heusden, 1992;Couvreur et al., 2012). Plant species from Annonaceae require ex-situ conservation to avoid threat of extinction. ...
Plant species of Annonaceae family were collected in Purwodadi Botanic Garden (PBG) needs ex-situ conservation efforts to prevent threat of extinction. Annonaceae plants collection in PBG have experienced dynamics over the past of 12 years which affect to diversity and conservation efforts. This study aimed to determine the dynamics or development of Annonaceae plants collection in PBG for the past of 12 years and how their conservation efforts. The methods were used are primary and secondary data collections from Registration Unit, measurement of stem diameter, plant height, branch-free stem height (BFS) of Annonaceae plants collection and microclimatic data (temperature, humidity, soil pH and light intensity) in each block of Annonaceae plants collection. Data were analyzed descriptively and by Principal Component Analysis using the statistical program PAST 4.0. The Annonaceae plant collections in PBG have fluctuated in the number of plants, genera, species, specimens and unidentified species with the addition of 28, 10, 6, 58, and 7 plants respectively. There were also 81 times of species identity changes. Annonaceae plant collections in PBG was able to well adapt and develop based on the average of stem diameter and BFS, with humidity and light intensity became the most influential environmental factors for plant growth. The management of Annonaceae plant collections in PBG consisted of the maintenance activities and studies to support the conservation. By regulating and manipulating of humidity and light intensity in Annonaceae plant collections, plant growth will increase and well maintenance. Management efforts can be planned and developed properly by knowing the dynamics of plant collections.
... Polyalthia Blume (1830: 68) was long considered one of the largest genera in the family Annonaceae (Sinclair, 1955), at one point thought to comprise 155 to 160 species distributed across the tropical areas of Asia, New Guinea, Australia, Solomon Islands, Fiji, Madagascar, and East Africa (Van Heusden 1992). Recently, phylogenetic analyses based on several molecular markers (Mols et al. 2004, 2012), have shown the genus to be highly polyphyletic, and in recent years many species have been transferred to other described proposed genera, such as Fenerivia Diels (1925: 355;Saunders et al. 2011), Marsypopetalum Scheffer (1870Xue et al. 2011), Monoon Miquel (1865Xue et al. 2012), Goniothalamus Hooker & Thomson (1855: 105;Tang et al. 2013), andMeiogyne Miquel (1865: 12;Xue et al. 2014). ...
A new species, Polyalthia khaoyaiensis, is proposed based on materials from central Thailand. The species appears most similar to Polyalthia meghalayensis and Polyalthia evecta, differing from these taxa in having shorter pedicels and usually two (rather than one) ovules per carpel. From P. meghalayensis it differs further in absence of a claw from the inner petals. The new species is described and illustrated. The conservation status of the species is considered and it is given an assessment of Least Concern (LC), as it is locally common in the hill evergreen forest of Khao Yai National Park.
... Molecular phylogenetic studies of Annonaceae using plastid DNA sequence data (Richardson et al. 2004;Mols et al. 2004;Pirie et al. , 2006Couvreur et al. 2011;Chatrou et al. 2012;Guo et al. 2017) have shown strong support for a clade including all the above-mentioned taxa plus various African and a large number of Asian species, classified by Chatrou et al. (2012) under the subfamily Malmeoideae (Fig. 5). Pirie et al. (2006) identified a clade including all Malmeoideae genera with distributions centred in South America (the "SAC clade"; Pirie et al. 2006), which corresponded to the Cremastosperma group of Van Heusden (1992) plus Unonopsis, Bocageopsis and Onychopetalum. This SAC clade was classified formally as tribe Malmeeae by Chatrou et al. (2012) and sister group relationships between Cremastosperma and one or other lineage of (a not demonstrably monophyletic) Pseudoxandra and between the Cremastosperma/Pseudoxandra clade and Malmea were confirmed. ...
We present a taxonomic revision of Cremastosperma , a genus of Neotropical Annonaceae occurring in lowland to premontane wet forest, mostly in areas surrounding the Andean mountain chain. We recognise 34 species, describing five as new here: from east of the Andes, C.brachypodum Pirie & Chatrou, sp. nov. and C.dolichopodum Pirie & Maas, sp. nov. , endemic to Peru; C.confusum Pirie, sp. nov. , from southern Peru and adjacent Bolivia and Brazil; and C.alticola Pirie & Chatrou, sp. nov. , at higher elevations in northern Peru and Ecuador; and from west of the Andes, C.osicola Pirie & Chatrou, sp. nov. endemic to Costa Rica, the most northerly distributed species of the genus. We provide an identification key, document diagnostic characters and distributions and provide illustrations and extensive lists of specimens, also presenting the latter in the form of mapping data with embedded links to images available online. Of the 34 species, 22 are regional endemics. On the basis of the extent of occurrence and area of occupancy of species estimated from the distribution data, we designate IUCN threat categries for all species. Fourteen species proved to be endangered (EN) and a further one critically endangered (CR), reflecting their rarity and narrow known distributions.
... Morphological characters were observed by phenetic character includes qualitative and quantitative characters. Qualitative characters are observed fifty-five characters and quantitative characters are observed fifteen characters (van Heusden, 1992;Kessler, 1993;Priyanti, 2001;Bioversity International, 2008;Folorunso and Olorode, 2008). Each plant sample is repeated based on a number of plant specimens in PBG. ...
Morphological characters are one of the most consistent of taxonomical markers are used in Annonaceae today. One of the plants collections from Annonaceae species which came from East Java and conserved in Purwodadi Botanic Garden-Indonesian Institute of Sciences, Purwodadi, and East Java has some problems in morphological characters. It has some problems that causing the taxonomical position of these species are not clear, i.e. presence of unidentified plant until level species. The aim of this study is to classify and identify of species that have not been identified in order to taxonomical position be clearly. Material samples are used Annonaceae collection from East Java and Magnoliaceae as out-group. Samples of these plants were observed qualitatively and quantitatively through morphological characters were analyzed using cluster analysis and then synapomorphy, autapomorphy and apomorphy character analysis to identify and to determine the taxonomical position. The results showed that there are 4 tribes and two sub-families in the dendrogram. Tribe includes Miliuseae, Xylopiae, Annoneae and Uvariae, as well as sub-family of Malmeoideae and Annonoideae. Sub-family distinguished by inner petal and habit, sub-family of Malmeoideae has connate inner petals and trees habit, sub-family of Annonoideae has free inner petal and woody climber habit, some of the trees. Species are unidentified suspected as a different species based on the proximity of group formed. There are five plant species that have not been identified to the species level. These species can be recommended the taxonomical position and proposed of species name through dendrogram with the name of the specimen. Specimen 1 is predicted to have close relations with Annona muricata, specimen 2 is predicted to have close relations with Miliusa macropoda, specimen 3 is predicted to have close relations with Artabotrys uncinatus and identified as Uvaria micrantha, specimen 4 is predicted to have close relations with Fissistigma latifolium and specimen 5 is predicted to have close relations with Saccopetalum horsfieldii, because these species are in one group. © 2017, Society for Indonesian Biodiversity. All rights reserved.
... The most common pollinators are small beetles (Nitidulidae, Curculionidae, Staphylinidae and Chrysomelidae), whereas larger beetles (Scarabaeidae) pollinate a few plant species (Gottsberger, 2012). Nevertheless, the dominance of beetle pollination did not constrain the amazing floral diversification, reflecting the wide functional diversity of beetle visitors (van Heusden, 1992;Saunders, 2010). In many genera, thick petals create pollination chambers offering a warmer environment through thermogenesis, shelters from predators, sufficient food and potential mates (Gottsberger, 2012). ...
In angiosperms, the exine is essential for pollen‒pistil interactions and is difficult to separate from the intine. In Annona montana, however, the exine spontaneously separates from the intine during hydration, leaving the entire male gametophyte exposed and activated, which does not affect its reproduction, and fruit are produced every year. In the present study, we used light microscopy, transmission electron microscopy, immunocytochemistry, and FTIR spectroscopy to explore whether pectins located in the cavities of the exine in A. montana dissolved in water, resulting in exine shedding, and whether water was the only necessary medium for A. montana pollen exine shedding. Exine shedding has also been reported in gymnosperms (e.g., Cupressaceae, Taxaceae, and Taxodiaceae), but A. montana has been the only species with exine shedding in angiosperms. Pollen shedding exine in A. montana exhibits a special and rare reproductive characteristic in angiosperms, which will update what we have previously recognized that exine is essential in reproductive processes.
Premise
The family Annonaceae possesses a broad array of floral phenotypes and pollination specialisations, and are important in the plant-pollinator interactions of tropical rainforests. Although there has been considerable effort to assess their interactions with pollinators, attempts to characterise their visual and olfactory communication channels are scarce.
Methods
Here, we investigated the pollination biology of 12 Annonaceae species from five genera, viz. Meiogyne , Monoon , Polyalthia , Pseuduvaria , and Uvaria . Furthermore, their floral colour was characterised by reflectance spectroscopy and floral odour chemistry was assessed using gas chromatography-mass spectrometry. Floral scent was further compared across the whole family using non-metric dimensional scaling plots to identify specialisation in floral odour.
Results
The Meiogyne species are likely pollinated by small beetles; the Polyalthia and Pseuduvaria species are likely pollinated by beetles and flies; and the Uvaria species is likely pollinated by beetles and bees. Flowers of most species are UV non-reflective, and have various spectral reflectance profile across the remaining visible spectra. Multiple species produce floral odour resembling ripe fruits. The flowers of Meiogyne species and Polyalthia xanthocarpa emitted mostly branched-chain esters, while flowers of Uvaria released mainly straight-chain esters. The Pseuduvaria species instead emitted scent reminiscent of rotten fruits, largely consisting of 2,3-butanediol. The inner petal corrugation in Meiogyne functions as a food reward, and the inner petal growth serves as a nectary gland for Pseuduvaria .
Conclusions
Our study identifies the visual and olfactory cues of multiple Annonaceae species and provides insights into how Annonaceae flowers attract different guilds of pollinators.
Lestari DA, Ningrum LW, Nada FMH, Pradipta NN, Harsono DR. 2023. Flowering and fruiting phenology of Anaxagorea luzonensis A. Gray (Annonaceae). Biodiversitas 24: 784-792. Anaxagorea luzonensis is one of the critical plant collections from the Annonaceae family, with only one specimen in Purwodadi Botanic Garden (PBG). In protecting these plants, it is necessary to know the process of flowering and fruiting through the phenological process. This research is expected to observe plant propagation, physiology, environmental response, and increased reproductive success. The aim of this research is to know the details of flowering and fruiting phenology stages, determine reproductive success, and explain the floral biology of A. luzonensis. Research on the flowering and fruiting phenology of A. luzonensis was carried out from December 2021 – November 2022 in one season at PBG. The method is observative, including changes in color, shape, and size of flowers and fruit in each phenological stage. In addition, the method includes the count of reproductive success and observation of the A. luzonensis flower. This study's results stated that the flowering and fruiting phenology of A. luzonensis experienced a shorter flowering time than fruiting time and a low percentage (<30%) of reproductive success. Flowering stages are divided into five stages, i.e. initiation, flower bud, before bloom, bloom, and anthesis; fruiting stages are divided into three stages, i.e. immature-sized, mature-sized, and ripe fruit, which occurred in period 73-121 ± 85 days. Environmental and genetic factors cause the low value of reproductive success. This research can be useful for the conservation strategy of A. luzonensis, especially for reproduction biology and seed conservation.
Angiosperm mesofossils are described from the Lower Cretaceous Almargem Formation exposed near the village of Catefica, Portugal, and are thought to be of Aptian-early Albian age. The mesofossil assemblage from Catefica is diverse
and, in addition to the angiosperms described here, also contains a rich assemblage of non-angiosperm fossils, including leafy axes of bryophytes and lycopsids, lycopsid and salvinialean megaspores, and sporangia, sori and leaf fragments of ferns. There are also twigs, cones, cone scales, seeds and sporangia of several kinds of conifers. Other seed plants include 11 species of chlamydospermous seeds and vegetative axes related to the BEG group (Bennettiales-Erdtmanithecales-Gnetales). In terms of the number of plant fragments identified, angiosperms are most abundant in the Catefica assemblage and account for more than half of all specimens. Angiosperms also dominate in number of species, but because the non-angiosperm fossils have not been studied in detail the total number of species in the flora is not yet established. Sixty-seven species of angiosperms are recognized. Angiosperm diversity is mainly at the level of non-eudicots, including ANA-grade angiosperms, Chloranthaceae and magnoliids. Remains of chloranthoid angiosperms are especially common, both in the number of specimens and in number of species recognized. About 40% of the specimens, and more than 25% of the species are chloranthoids. Remains of magnoliid angiosperms(Magnoliales, Laurales, Canellales, Piperales) are also prominent among the angiosperms. Eudicots are subordinate:
only 3–4% of all angiosperm specimens can be assigned confidently to eudicot angiosperms. Five new genera and six new species of angiosperms are established (Canrightia foveolata sp. nov., Elasmostemon paisii gen. et sp. nov., Endressistemon
cateficensis gen. et sp. nov., Ibericarpus cuneiformis gen. et sp. nov., Proencistemon portugallicus gen. et sp. nov., Valvidistemon globiferus gen. et sp. nov.). Several other new taxa are also described, but not formally named.
Uvaria chamae P. Beauv is known as a key shrub species providing several goods and services for sustaining livelihoods in Sub-Saharan Africa. Despite its great medicinal importance for local communities, little is known about its conservation status and sustainable management strategies regarding the current overexploitation of the species’ fruits through traditional agroforestry systems. Here, we addressed a global systematic review of the current state of knowledge on several aspects of research of U. chamae for setting further breeding programmes and conservation initiatives. A total of 744 publications were identified based on the extensive bibliometric review of its sustainable management and conservation status over the last three decades (1991-2021) through existing online databases. Only 257 publications were finally included in the current review after deep scrutinization which were in line with several aspects of the conservation ecology and management of U. chamae in Africa. All retained papers came globally from the five sub-regions, and particularly 13 countries in Africa. Most of them were recorded in West Africa (n = 245) compared to the other sub-regions where few studies exist on this intensively harvested shrub species. Approximately 89% of the retained publications came from five of West African countries including Nigeria (n = 151), Benin (n = 30), Côte d'Ivoire (n = 18), Guinea (n = 16), and Togo (n = 14). In-depth bibliometric analysis revealed critical knowledge gaps on U. chamae in terms of its geographic distribution; conservation status; tree growth, productivity and propagation; morphological diversity; molecular genetic diversity; reproductive biology; ecophysiological performances; socio-economic importance; biochemical analysis; and structural characterization. The current review paves the way for developing further long-term management programs of U. chamae in Africa.
Annonaceae is a major pantropical family with 113 genera and about 2550 species. Cameroon is one of the most biodiverse countries in Africa but its flora remains incompletely known. In this volume of the Flora of Cameroon, we describe 166 native taxa representing 163 species in 28 native genera within the family Annonaceae. A total of 22 species (about 13%) are endemic to the country. We provide keys to all native genera, species, and infraspecific taxa. For each species a detailed morphological description and a map of its distribution in Cameroon are provided. Distribution maps and diversity analyses are based on a taxonomically verified database of 2073 collections. Across Africa, Cameroon is a center of diversity for Annonaceae harboring one of the highest numbers of species and genera. For example, Cameroon harbors the highest number of African species for the only pantropical genus of Annonaceae, Xylopia . Annonaceae are found across all 10 administrative regions of Cameroon but diversity is concentrated within the tropical rain forest areas situated in the south and South-West. The areas around Bipindi and Mount Cameroon show the highest levels of diversity, but this is correlated with collection effort. Line drawings and/or photographs accompany most species. One species new to science Uvariopsis etugeana Dagallier & Couvreur sp. nov. is described. We also undertake a number of nomenclatural changes such as lectotypifications, six new synonymies and two new combinations ( Uvaria anisotricha (Le Thomas) Couvreur, comb. nov. ; Uvariodendron fuscum var. giganteum (Engl.) Dagallier & Couvreur, comb. nov. ).
Societal Impact Statement
The United Nations Decade on Ecosystem Restoration underlines the importance of understanding how different taxa are affected by human induced, global changes in ecosystems. Here, we investigate if this impact can be quantified for the globally distributed tropical plant group Annonaceae (Soursop family) using distributional data and International Union for Conservation of Nature (IUCN) Red List assessments. We find that even for a taxonomically well‐studied tropical plant family such as Annonaceae, little is known about the true distribution and ecological requirements of, and threats to, species in this group. We discuss several improvements in data collection that should enable more in‐depth analyses in the future.
Summary
The Sustainable Development Goals of the United Nations (UN), formulated with the overarching aims to end poverty and protect the planet, are also aimed at implementing sustainable management of all types of forests, to stop deforestation and to restore degraded forests. This led to the declaration of the UN Decade on Ecosystem Restoration. To meaningfully restore ecosystems, it is important to increase our understanding on the distribution of taxa and obtain insight in how different taxa are affected by human induced, global changes in ecosystems.
Here, we investigate if this impact can be quantified for the globally distributed tropical plant group Annonaceae (Soursop family) using spatial data and International Union for Conservation of Nature (IUCN) Red List assessments. Insight is gained in how Annonaceae are distributed over biomes and anthropogenic biomes (anthromes) and how threatened Annonaceae are based on their distribution.
We find that even for a taxonomically well‐studied group such as Annonaceae, very little is known about the true distribution and ecological requirements of, and threats, to species.
We urge to invest in (1) the exploration of ecological requirements of species in relation to their genetic patterns, in order to understand the impact of ecosystems changes, (2) research on distributional patterns in a temporal framework as the available data collected over decades might not reflect current distributions over biomes and anthromes well and (3) high‐quality spatial data collection that should adhere to the Findability, Accessibility, Interoperability and Reuse (FAIR) data principles, so that the quality of spatial analyses as well as IUCN Red List assessments will increase.
Miliusa agasthyamalana (Annonaceae), a new species from the Western Ghats, India is described and illustrated. The new species is morphologically similar to M.paithalmalayana and M. wightiana. Detailed notes on the species along with its color photographs and illustrations are provided.
Based on morphological characteristics, Cyathocalyx tsukayae, which is endemic to Borneo is transferred
to Monoon. The Bornean endemic Monoon sympetalum is the most morphologically similar species, mainly because
of the basally connate petals it shares with M. tsukayae. The two species can be chiefly distinguished by differences
in outer petal size, monocarp size and monocarp/stipe length ratio.
The delimitation of the genera Pyramidanthe and Mitrella in the tribe Uvarieae of the family Annonaceae is assessed
by molecular phylogenetic analyses and morphological re-evaluation. Using up to six plastid DNA regions (matK, ndhF, rbcL
exons; trnL intron; psbA-trnH, trnL-trnF intergenic spacers) and including, among others, two accessions of Pyramidanthe and
five accessions of Mitrella, the resulting phylogeny shows that Pyramidanthe and Mitrella are recovered in a clade sister to the Fis-
sistigma clade. The Pyramidanthe-Mitrella clade is composed of a trichotomy: a clade consisting of Pyramidanthe accessions and
two clades containing Mitrella accessions. In combination with negligible morphological distinctions between the two genera, they
are consequently merged, with 11 new combinations under the chosen name Pyramidanthe: P. beccarii, P. clementis, P. cylindrica,
P. dielsii, P. elegans, P. kentii, P. ledermannii, P. mabiformis, P. schlechteri, P. sylvatica and P. tiwiensis. The names M. dielsii (the
basionym of P. dielsii) and P. rufa (a heterotypic synonym of P. prismatica) are lectotypified. Pyramidanthe s. lat. possesses the
following diagnostic traits: usually indistinct secondary leaf veins with a brochidodromous to brochidodromous-eucamptodromous
venation, a reticulate tertiary leaf venation, axillary inflorescences, presence of a basal excavation on an inner side of each outer
petal, and inner petals that are much smaller than the outer petals and cohering marginally at anthesis.
Annonaceae is the most species-rich family in the order Magnoliales and has a pantropical distribution, includes 109 genera and 2,440 species. It is subdivided into four subfamilies, Anaxagoreoideae, Ambavioideae, Annonoideae and Malmeoideae. The latter comprises eight tribes, including the Malmeeae, the only tribe of Malmeoideae represented in Brazil, with 13 genera and 73 species. The Amazon is the center of diversity of the tribe, with eight genera restricted to this biome, the other five genera having broader distributions. The present study sought to characterize the species of Malmeeae that occur in the state of Rondônia. Seventeen species belonging to eight genera were identified and characterized: Oxandra with five species, Bocageopsis with three, Onychopetalum, Pseudoxandra and Unonopsis with two, and Cremastosperma, Ephedranthus and Klarobelia with one species each. Bocageopsis mattogrossensis and Pseudoxandra lucida are recorded for the state for the first time. A key to the species, morphological descriptions, geographical distribution and comments are provided.
Two new species of Polyalthia from Peninsular Thailand, Polyalthia heliopetala and P. taweensis, are described and illustrated. Polyalthia heliopetala most closely resembles P. oblonga and P. motleyana. Polyalthia taweensis most closely resembles P. obliqua. Polyalthia pumila is newly reported for Thailand from a single locality; the species was previously known only from two localities in Peninsular Malaysia and has not been re-collected in over 100 years.
Meiogyne oligocarpa sp. nov. (Annonaceae) is described from Yunnan Province in Southwest China. It is easily distinguished from all previously described Meiogyne species by the possession of up to four carpels per flower, its bilobed, sparsely hairy stigma, biseriate ovules and cylindrical monocarps with a beaked apex. A phylogenetic analysis was conducted to confirm the placement of this new species within Meiogyne . Meiogyne oligocarpa represents the second species of Meiogyne in China: a key to the species of Meiogyne in China is provided to distinguish it from Meiogyne hainanensis . Paraffin sectioning was undertaken to study the anatomy of the corrugations on the inner petals of Meiogyne oligocarpa to verify whether they are glandular.
Meiogyne kwangtungensis is a rare species endemic to Hainan, China, known just from two fruiting collections made in the 1930s. Although it was published under the name Meiogyne in 1976, it was suggested that it might be better placed within Pseuduvaria or Mitrephora . For decades, this species was never collected again, thus its true generic affinity remained unresolved due to the lack of flowers. During a field exploration in Hainan, we re-discovered this species and collected a flowering specimen for the first time. The flower immediately confirmed its affinity with Pseuduvaria . Phylogenetic analyses of five chloroplast regions ( psbA-trnH , trnL-F , matK , rbcL , and atpB-rbcL ; ca. 4.2 kb, 70 accessions) also unambiguously placed Meiogyne kwangtungensis in the Pseuduvaria clade (PP = 1.00, ML BS = 99%). Morphologically, it is most similar to P. multiovulata which is endemic to Myanmar and Thailand, both with often-paired flowers, long pedicels and short peduncles, and often 1–2 monocarps. However, it differs in having smaller flowers with kidney-shaped glands on the inner petals, fewer stamens and carpels, smaller ovoid monocarps with an apicule and fewer seeds. On the basis of the combined molecular phylogenetic and morphological data, we propose a new combination, Pseuduvaria kwangtungensis (P.T.Li) Qing L.Wang & B.Xue. A full description including floral characters and a color plate are provided here for this species. A key to species in the genus Pseuduvaria in China is also provided.
Inner staminodes are widespread in Magnoliales and present in Anaxagorea and Xylopia, but were lost in the other genera of Annonaceae and have no counterparts in derived angiosperms. The coexistence of normal stamens, modified stamens and inner staminodes in Anaxagorea javanica is essential to understand the homology and pollination function of the inner staminodes. A. javanica was subjected to an anatomical study by light and scanning electron microscopy, and the chemistry of secretions was evaluated by an amino acid analyser. Inner staminodes have a secretory apex, but do not have thecae. They bend toward either tepals or carpels at different floral stages, and function as a physical barrier preventing autogamy and promoting outcrossing. At the pistillate phase, the exudates from the inner staminodes have high concentration of amino acid, and provide attraction to pollinating insects; while abundant proline was only detected in stigmas exudates, and supply for pollen germination. Modified stamens have a secretory apex and one or two thecae, which are as long as or shorter than that of the normal stamens. As transitional structures, modified stamens imply a possible degeneration progress from normal stamens to inner staminodes: generating a secretory apex first, shortening of the thecae length next, and then followed by the loss of thecae. The presence of modified stamens together with the floral vasculature and ontogeny imply that the inner staminodes are homologous with stamens.
Mischogyne (Annonaceae, tribe Monodoreae) is a genus of small- to medium-sized tropical trees and shrubs. It is characterised by a combination of: (1) stamens and carpels on a more or less extended torus; (2) carpels divergent from each other at the apex of the torus above the anthers; (3) anthers linear and anther connectives not expanded above the thecae; (4) inflorescences extra-axillary (or sometimes terminal in M. michelioides) with usually solitary flowers; (5) petals reflexed at anthesis (except M. michelioides), and (6) prominent reticulate tertiary veins. The genus is found in the lowland tropical rainforests of Africa with annual precipitation of 1000 – 4000 mm and in the dry coastal foothills of Angola. Five species and one variety of Mischogyne are recognised. One newly discovered species from the Eastern Arc Mountains, M. iddii, is described. The variety M. elliotiana var. glabra is reduced into synonymy with M. elliotiana var. elliotiana. Mischogyne elliotiana var. gabonensis is raised to species level as M. gabonensis. Specimens from the Congo are recognised as a new species, M. congensis. Preliminary conservation status assessments are provided for each species, as well as an identification key and detailed species descriptions. An unusual distribution pattern for the genus is discussed: dry coastal Angola as well as West, Central and East African wet forest.
The present study focuses on the emitted and endogenous scent profiles of Uvaria hamiltonii flowers. Among the 34 compounds identified, sesquiterpenoids were found to dominate the floral volatiles composition. Profiles from endogenous scent volatiles showed higher number of compounds than the emitted ones. The anthocyanin pigment responsible for the flower colour was also explored. It was found that a single anthocyanin compound, cyanidin-3-O-glucoside, was principally responsible for petal colour. Total phenolic content was evaluated and antioxidant capacities were studied with the help of DPPH, FRAP and ABTS assays. The total phenolic content and the antioxidant capacity were higher in methanolic extract as compared to aqueous, petroleum ether and ethyl acetate extracts of U. hamiltonii flowers.
Greenwayodendron (Annonaceae) is a tropical African genus of trees occurring mainly in rain forests. Until recently, Greenwayodendron contained only two species: Greenwayodendronoliveri from West Africa and Greenwayodendronsuaveolens from Central and East Africa.
Genetic data, using chloroplast haplotypes and nuclear microsatellites as well as morphometric analyses, provided important information on the delineation of species. Greenwayodendron now contains six species, including two new species ( Greenwayodendronglabrum Lissambou, Hardy & Couvreur, sp. nov. and Greenwayodendronlittorale Lissambou, Dauby & Couvreur, sp. nov. ). Greenwayodendronsuaveolensvar.gabonica and Greenwayodendronsuaveolenssubsp.usambaricum are recognised as distinct species: Greenwayodendrongabonicumcomb. nov. and Greenwayodendronusambaricumcomb. nov. , respectively. A key, detailed descriptions of morphology and geographic distributions, as well as notes on their ecology and uses are presented for all species. Preliminary conservation assessments following IUCN criteria are also provided. Two species are preliminarily identified as threatened, one as Endangered and one as Vulnerable.
RESUMO São apresentadas as espécies de Annonaceae da Reserva Biológica da Represa do Grama. A Reserva está localizada em Descoberto, Minas Gerais, e abrange uma área de 263,8 hectares de floresta estacional semidecidual. São encontrados cinco gêneros e sete espécies: Annona cacans, Guatteria nigrescens, G. sellowiana, Rollinia dolabripetala, Unonopsis bauxitae, Xylopia brasiliensis, X. sericea e a nova espécie Unonopsis bauxitae, aqui descrita. São apresentadas chave de identificação das espécies, descrições, ilustrações, e informações sobre floração, frutificação, distribuição geográfica e hábitats.
A revision of the 45 species of the pantropical genus Xylopia in Tropical Africa includes descriptions of six new species and a new section of the genus. The fruits and seeds of Xylopia show specializations that promote vertebrate dispersal, primarily by hornbills and monkeys. Over half of the African species have an Area of Occupancy (AOO) less than 80 km ² , suggesting that they are in need of protection. African species are classified into five sections. Section Neoxylopia , with four species, is centered in the Guineo-Congolian Region and includes X.globosasp. nov. Section Ancistropetala, with three species, occurs in the same region. Both of these sections are endemic to Africa. Section Xylopia, which extends to Madagascar and the American tropics, has only a single species in Africa, X.aethiopica . The three species of section Verdcourtia sect. nov. are restricted to the East African coast and Madagascar. The largest number of African species, (34) belong to section Stenoxylopia, in which the seeds lack the arils found in the other sections and instead have a fleshy sarcotesta. Section Stenoxylopia is divided into two informal groups, one centered in eastern and southern Africa ( X.odoratissima group) and the other centered in the wetter forests of western and central Africa ( X.acutiflora group). Five new species are described in section Stenoxylopia: Xylopianiloticasp. nov. from Sudan, South Sudan, and Uganda, Xylopiacalvasp. nov. from Nigeria and Cameroon, which is allied to X.phloiodora , and Xylopiamonticolasp. nov. from Nigeria and Cameroon, X.pirataesp. nov. from Ivory Coast and Ghana, and X.unguiculatasp. nov. from Gabon. The latter three species are segregates of the former Xylopiaacutiflora s. l. One new combination is made at the species level, X.shirensiscomb. nov. Keys, descriptions, illustrations, distribution maps, and an index to numbered collections document diversity and assist with species identification. The name Unonaoliveriana Baill. was found to pre-date the name Unonalepidota Oliv., requiring the combination Meiocarpidiumoliverianumcomb. nov.
We present revisions of the Neotropical genera Ruizodendron and Pseudephedranthus (Annonaceae). Ruizodendron includes a single species R. ovale. Pseudephedranthus now comprises two species, with the description of the new species P. enigmaticus sp. nov. extending the range of the genus beyond the Upper Rio Negro region of Brazil (Amazonas) and adjacent Venezuela (P. fragrans), to include Guyana, Suriname, and the Brazilian state of Pará. An overview is provided of current revisions of Neotropical Annonaceae genera that will aid in accessing proper species information for this frequently encountered tropical rain forest family.
Background and aims – Annonaceae are an important family in tropical rain forests of Africa. Here, we present a revision of the west-central African genus Piptostigma of the tribe Piptostigmateae. Previous molecular phylogenetic analyses of Piptostigmateae have revealed that the genus as currently circumscribed is paraphyletic. The aims of the article are: (1) to clarify the generic delimitation of the genus Piptostigma, especially in relation to its current polyphyletic delimitation; (2) to provide a new updated key to the genera of tribe Piptostigmateae; and (3) to provide for the first time in over 100 years a complete taxonomic revision of genera Brieya and Piptostigma. Material and methods – Normal practices of herbarium taxonomy were used for the morphological study of about 200 specimens from B, BR, BRLU, G, HBG, K, L, LBV, LISC, MO, P, U, US, WAG, YA and Z. We sequenced three plastid regions for 32 samples representing 26 ingroup (Piptostigmateae) and six outgroup species in order to explore the paraphyly of Piptostisma. The open source software QGis was used to generate the distribution maps of the species and their conservation status was obtained using the IUCN categories and criteria. Key results – Molecular and morphological evidence support the reinstatement of the genus Brieya De Wild. Three keys are presented, one to the genera of Piptostigmateae and the two others to the species of Brieya and Piptostigma. A total of thirteen species are described for Piptostigma and two for the genus Brieya. Four new species are presented for the genus Piptostigma: P. macrophyllum, P. mayndongtsaeanum, P. goslineanum and P. submontanum.
Key words – Annonaceae, Piptostigma, Brieya, taxonomy, tropical Africa.
Oncodostigma Diels (1912: 143) was erected by Diels (1912) based on the type species O. leptoneura Diels (1912: 143). Five additional species were subsequently added to the genus, viz.: O. wilsonii Guillaumin (1931: 224), O. monosperma (Hooker & Thomson 1872: 57) Sinclair (1951: 605), O. mindorense (Merrill 1907: 273) Bân (1974: 1779), O. hainanense (Merrill 1925: 131) Tsiang & Li (1979: 81) and O. microflorum Okada (1996: 8). Van Heusden (1992) conducted a detailed comparative study of the floral morphology of Annonaceae and noted that Oncodostigma is not clearly distinct from Meiogyne Miquel (1865: 12). She subsequently transferred O. mindorense and O. monosperma to Meiogyne and reduced O. wilsonii to synonymy with M. cylindrocarpa (Burck 1911: 433) Heusden (1994: 499). The name Oncodostigma hainanense (Merrill 1925: 131) Tsiang & Li (1979: 81) had previously been transferred to Meiogyne by Bân (1973).
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