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The tropical cucurbitaceous genus Zehneria as traditionally circumscribed displays much morphological diversity. Recent taxonomic revisions have resulted in its redefinition through both recognition and subsequent lumping of several additional genera. This study utilized plastid and nuclear DNA sequence data to reconstruct a molecular phylogeny of...
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... Cucurbitaceae family is well represented in the paleotropics, and includes several genera widely distributed across many environments. Species in the genus Zehneria Endl. as formerly taxonomically circumscribed are found across the paleotropics, with several species in Africa, and mainland and insular Asia (de Wilde and Duyfjes, 2006a; Schaefer and Renner, 2011a). Zehneria in that sense is species-rich in East Africa, but most diverse in South East Asia. Zehneria was first described by Endlicher based on collections made by Bauer on Norfolk Island in 1804–5 (Endlicher, 1833). Cogniaux considered Zehneria to be part of the genus Melothria L., a large assemblage with many species in the Old and New Worlds (Cogniaux, 1916, 1881). The definition of Melothria was broadly constrained to cucurbitaceous plants sharing a common anther and floral type with a pantropical distribution. Jeffrey (1962) later on recognized differences between the Old and New World species and described the Old World taxa as the genus Zehneria (excluding Mukia Arn. and Solena Lour.). Jeffrey (1990, 1980) placed Zehneria and Melothria in subtribe Cucumerinae, along with ten other genera, including Cucumis L., Cucumella Chiov., and Mukia . This group was loosely defined as sharing small, compressed seeds and anther thecae fringed with trichomes. In Jeffrey’s classification Zehneria remained a diverse genus of some 35 species, stretching from Africa to French Polynesia, of which many species were poorly collected or studied. More recent studies of Zehneria for the Flora Malesiana project (de Wilde and Duyfjes, 2010) included an extensive review of museum and field collections of Asian and African species of Zehneria , and revealed significant differences and clear divisions among the species based on flower and fruit characters. As a result, de Wilde and Duyfjes recognized six genera, Zehneria (in a restricted sense), and five new ones: Indomelothria W.J.de Wilde & Duyfjes , Neoachmandra W.J.de Wilde & Duyfjes , Papuasicyos Duyfjes, Scopellaria W.J.de Wilde & Duyfjes, and Urceodiscus W.J.de Wilde & Duyfjes (de Wilde and Duyfjes, 2006a, 2006b; Duyfjes et al. , 2003). In 2009, de Wilde and Duyfjes further split Zehneria by reinstating the old name Pilogyne Schrad. for African and remaining Asian species except Z. baueriana Endl. (de Wilde and Duyfjes, 2009a, 2009b), leaving Zehneria monotypic. The above-mentioned six genera were differentiated from Zehneria and each other based on several mainly floral characters: the presence or absence of a probract, color and aestivation of the petals, disposition of the male flowers, shape of the disc, stigma lobes, fruit and seeds and various characters of the stamens such as their place of insertion on the receptacle-tube, size and shape of the filaments, thecae and connective (de Wilde and Duyfjes, 2009a, 2006a). Molecular phylogenetic analyses of the Cucurbitaceae have redefined much of the sub-familial classification (Chung et al. , 2003; Jobst et al. , 1998; Kocyan et al. , 2007; Schaefer and Renner, 2011a; Schaefer et al. , 2009; Zhang et al. , 2006). The most recent classification based on molecular phylogenetic data by Schaefer and Renner (2011a, 2011b) accepts for the family 95 genera in 15 tribes. The genera studied here are included in tribe Benincaseae Ser., which includes over 200 species in 24 genera. Based on combined morphological and molecular evidence, Schaefer and Renner (2011a) merged the genus Neoachmandra into Zehneria , Urceodiscus into Papuasicyos , and the morphologically distinct but poorly collected monotypic genus Anangia W.J.de Wilde & Duyfjes into Zehneria . However, based on the combined evidence it would have been more conservative to maintain the status quo. A summary of the taxonomic history of Zehneria , Neoachmandra and Pilogyne is presented in Figure 1. The present study uses a molecular phylogenetic framework to test the accuracy of the genera split from Zehneria by de Wilde & Duyfjes (2009a, 2006a, 2006b; Duyfjes et al. , 2003): Indomelothria, Neoachmandra, Papuasicyos , Pilogyne , Scopellaria , and Urceodiscus and reassesses the merging of Neoachmandra into Zehneria by Schaefer and Renner (2011a) using an expanded molecular data matrix and more comprehensive taxon sampling. DNA was obtained for this study from silica-dried as well as from herbarium material of a total of 48 taxa from 13 genera with a focus on the new genera described by de Wilde and Duyfjes (2009a, 2006a, 2006b; Duyfjes et al. , 2003). The dataset includes 1 out of 2 species of Indomelothria, 15 out of c. 30 Neoachmandra, 1 out of 1 Papuasicyos , 7 out of c. 10 Pilogyne , 1 out of 2 Scopellaria , 2 out of 7 Urceodiscus , and 12 out c. 25 Zehneria . The genus Kedrostis Medik. was included as an outgroup (Schaefer and Renner, 2011a). Table 1 includes an overview of all extracted vouchers, including species, collector, country, herbarium of deposition, and NCBI GenBank accession numbers of all 159 new sequences generated for this ...
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... based on the combined evidence it would have been more conservative to maintain the status quo. A summary of the taxonomic history of Zehneria, Neoachmandra and Pilogyne is presented in Figure 1. The present study uses a molecular phylogenetic framework to test the accuracy of the genera split from Zehneria by de Wilde & Duyfjes (2009a, 2006a, 2006bDuyfjes et al., 2003): Indomelothria, Neoachmandra, Papuasicyos, Pilogyne, Scopellaria, and Urceodiscus and reassesses the merging of Neoachmandra into Zehneria by Schaefer and Renner (2011a) using an expanded molecular data matrix and more comprehensive taxon sampling. ...
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... of the new genera proposed by de Wilde and Duyfjes (2009a, 2009b, 2006a, 2006bDuyfjes et al., 2003) are supported by molecular phylogenetic studies, Indomelothria, Neoachmandra, Papuasicyos, Scopellaria, Urceodiscus (cf. this study and Fig. S1 and S2 in Schaefer and Renner, 2011b). The fifth genus, Pilogyne as reinstated by Duyfjes (2009a, 2009b), is not supported by molecular phylogenetic data. The type Pilogyne suavis Schrad. is not included in the sampling, but the South Asian taxa included in this study that de Wilde and Duyfjes combined in the genus are polyphyletic in ...
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... Taxonomically, the genus is belongs to the tribe Benincaseae Ser. (Schaefer and Renner 2011;de Boer et al. 2015). The tribe comprises dioecious or monoecious climbers with simple or 3-7-lobed leaves, yellow, orange, salmon, or white flowers, three stamens (two anthers 2-thecous, one 1-thecous, less often all 2-thecous or all 1-thecous), and indehiscent fruits (Schaefer and Renner 2011 Schaefer and Renner 2011;de Boer et al. 2015). ...
... (Schaefer and Renner 2011;de Boer et al. 2015). The tribe comprises dioecious or monoecious climbers with simple or 3-7-lobed leaves, yellow, orange, salmon, or white flowers, three stamens (two anthers 2-thecous, one 1-thecous, less often all 2-thecous or all 1-thecous), and indehiscent fruits (Schaefer and Renner 2011 Schaefer and Renner 2011;de Boer et al. 2015). As a result of our discovery, the number of species of Benincaseae in Java has increased to 12 genera. ...
Melothria L. (Cucurbitaceae) is a Neotropical genus of wild cucumber comprising 12 species. The only species that occurs in Malesia is M. pendula L., which has been recorded in Peninsular Malaysia, Borneo, mainland Sumatra, the Philippines, Sulawesi, and Bali. During our botanical excursions in Bangka and West Java from 2020 to 2022, we discovered naturalized populations of M. pendula. Our findings reveal that M. pendula has an extensive distribution in Sumatra and is a newly recorded wild cucumber in Java. We present a morphological description, photographs, and a brief discussion.
... The circumscription of Zehneria Endl. has changed considerably over the last centuries and particularly in the last two decades (see Dwivedi et al. 2018 for details) but the consensus of all comprehensive molecular studies (Kocyan et al. 2007;Schaefer and Renner 2011;De Boer et al. 2015;Dwivedi et al. 2018) is that a monophyletic Zehneria must include Anangia W.J.deWilde & Duyfjes, Neoachmandra W.J.deWilde & Duyfjes, and Pilogyne Schrad. ...
Zehneria is one of the most diverse genera in Cucurbitaceae with 75 accepted species mainly in Southeast Asia and tropical Africa. Here, we describe Zehneria neorensis, a new species from Neora Valley in the mountains of West Bengal, India, which has 7–10 cm long twisted fruiting pedicels, the longest pedicels reported in the genus so far. Based on morphological data, we also suggest the transfer of Melothria morobensis to the genus Zehneria. With a molecular phylogenetic and global biogeographic analysis based on 3856 nucleotides of plastid and nuclear ribosomal DNA, we demonstrate that the genus Zehneria most likely originated on the African continent 24 (30–19) million years ago and spread from there at least five times to Madagascar and three times to Asia. Zehneria neorensis represents an independent colonization event from Africa to India about 11 (15–7) million years ago. Three lineages reached New Guinea/Australia and finally moved into Polynesia. We infer a rate of at least 20 long-distance dispersal (LDD) events per 10 million years in the genus. This high LDD frequency is most likely a result of the small berry fruits and small flattened seeds of Zehneria, which seem perfectly adapted to long-distance bird dispersal. Field observations are needed to investigate a potential effect of the newly discovered extended and coiling pedicels in Zehneria neorensis on seed dispersal efficiency. The new species adds to a growing list of rather old Cucurbitaceae lineages in the Himalayan foothills, supporting the hypothesis of long climatic stability in the region.
... Over the past two decades, multiple molecular phylogenetic studies have strongly supported that the remaining segregate genera Neoachmandra W.J. de Wilde & Duyfjes (2006a: 12) and Pilogyne Schrad. in Schrader (1835: 5) are paraphyletic, and together with the monotypic Anangia W.J.De Wilde & Duyfjes (2006b: 219) are deeply nested in Zehneria s.s. (Kocyan et al. 2007;Schaefer et al. 2009;Schaefer & Renner 2011;De Boer et al. 2015, Dwivedi et al. 2018, Wei et al. 2020. From the perspective of nomenclature, Zehneria, a validly described genus, has priority over all the segregated generic names, including Anangia, Neoachmandra, and Pilogyne. ...
... concept was applied in one broadly circumscribed genus by Dwivedi et al. (2018). Schaefer & Renner (2011), De Boer et al. (2015, Dwivedi et al. (2018), andPelser et al. (2021) formally reincorporated all species of the segregate genera Anangia, Neoachmandra, and Pilogyne into Zehneria, except for one species distributed in Papua New Guinea. Given that there is no validated name for that Zehneria species (Z. ...
The segregated genera, Neoachmandra and Pilogyne were merged into Zehneria, based on recent molecular phylogenetic studies. Accordingly, Z. morobensis (Merr. & L.M.Perry) N.Wei, G.W.Hu & Q.F.Wang, comb. nov., a new combination in Cucurbitaceae, is proposed herein.
... monocarpa, which was separated from Z. scabra recently (Ngumbau et al. 2020). Furthermore, we also found that species of Neoachmandra in the sense of De Wilde and Duyfjes (2006a) and De Boer et al. (2015), are paraphyly. In line with the conclusion made by Dwivedi et al. (2018), the whole genus tended to be separated into two major clades (clade 1 and clade 2), with African taxa being the basal lineages. ...
Zehneria grandibracteata, a new species of Cucurbitaceae from western Kenya, is described here, based on morphological and molecular data. It has long been misidentified as the widely-distributed species Z. scabra. However, it differs by its ovate leafy probract at the base of the inflorescences, subglabrous condition of the entire plant, shorter receptacle-tube and filaments, as well as denser and sessile inflorescences. Furthermore, the molecular phylogenetic analysis of Zehneria, based on nrITS sequences, further supports the argument that Z. grandibracteata should be segregated from Z. scabra.
... Its members are small to medium sized herbaceous creepers and climbers, sometimes with a woody base when old. They are distributed throughout the Old-World tropics and subtropics with their center of diversity found in Southeast Asia (De Boer et al. 2015, Dwivedi et al. 2018. The genus consists of about 94 species in its broad circumscription, whereby ca. 15 species are found in East Africa and eight species in Kenya (Jeffrey 1967, Dwivedi et al. 2018. ...
... With more new species from this genus being discovered in East Africa, especially in Kenya recently (Zhou et al. 2016, Watuma et al. 2019, there is a need for a revision of the genus Zehneria, as East Africa has been neglected in the collection and studying of this genus since 1967 (De Boer et al. 2015). A phylogeny of African Zehneria, including the well-resolved Asian Zehneria, will help scientists to comprehensively understand the taxonomic relationships and evolutionary history of the genus. ...
Zehneria monocarpa (Cucurbitaceae), a new species from the fragmented lowland coastal forests of Kenya is described here and illustrated with photographs. It resembles Z. oligosperma and Z. longiflora, but can be distinguished by its solitary female flowers and fruits. Furthermore, the new species has 1–23 conspicuous dark green glands on the apex of the leaf and the inner part of its receptacle tube is entirely hairy.
... It is only in the past two decades that several studies on this genus have been published (Thulin 1996, Verdcourt 1997, Zhou et al. 2016, Wei et al. 2017. Although South East Asia remains the diversity hotspot for this genus (Dwivedi et al. 2018), it has been suggested that East Africa is a neglected centre of diversity (De Boer et al. 2015). Following this current contribution, there are 16 species known from the region. ...
A new plant species in the genus Zehneria (Cucurbitaceae) is here described and illustrated. The species was collected at Taita Hills tropical montane forests in Kenya, a fragment of the larger Eastern Arc Mountains biodiversity hotspot. The new species, Zehneria tuberifera, is similar to Z. oligosperma and Z. longiflora, but distinguished by its tuberous rhizome and subcoriaceous leaf blades. Furthermore, it has scattered bristles along the midrib on the adaxial surface of its young leaf blades, a shorter receptacle tube, a trilobed disk in the male flower, solitary or 2-clustered female flowers, and up-to-20-flowered and subumbelliform male inflorescences.
... This is not supported by the molecular phylogeny, which is why Schaefer et al. (2009, Schaefer & Renner (2011a, 2011b) argued against over-splitting of the group. In regard to the new species we discuss here, we agree with the latter view (Schaefer et al. 2009, Schaefer & Renner 2011b, De Boer et al. 2015, and put it into Zehneria in the broad sense. ...
... As a center of distribution for Zehneria, East Africa has been neglected in collecting and studying this genus since 1967 (De Boer et al. 2015), when Charles Jeffrey finished editing the Cucurbitaceae part of Flora of Tropical East Africa (FTEA; Jeffrey 1967). However, a few papers about Zehneria from East Africa have been published in the last five decades (Verdcourt 1997). ...
Zehneria longiflora, a new species of Cucurbitaceae from Nandi County, Kenya, is described and illustrated here. It is most similar to Z. oligosperma, but differs mainly by its longer receptacle-tube, stamens that are inserted at the middle of the receptacle-tube and with longer filaments, as well as characters of the male and female inflorescence.
... The Old World species of the group have either been accommodated in various genera (De Wilde & Duyfjes 2006 or united under one large genus Zehneria Endlicher (1833: 69, Jeffrey 1962, Schaefer & Renner 2011. The latter concept has been supported by pollen exine ornamentation (Van der Ham & Pruesapan 2006) and more recently by molecular phylogenetic analyses (Schaefer & Renner 2011, De Boer et al. 2015. One of the species, Zehneria thwaitesii (Schweinfurth) Jeffrey (1962: 15), is subject to molecular phylogenetic investigations at a larger geographic scale (Dwivedi et al., in prep.). ...
In Zehneria thwaitesii long standing uncertainties persisted with respect to taxonomic delimitation. This study is a first step in clarifying the case using both morphological and molecular phylogenetic analyses. We here provide a complete overview on the nomenclatural history of Z. thwaitesii. For the African plants hitherto included in the broadly circumscribed species the name Z. tridactyla is reinstated and a lectotype is designated. Diagnostic features distinguishing both taxa are summarised. Z. thwaitesii is now considered restricted to Asia.
... Zehneria Endl. (1833: 69) is a genus of Cucurbitaceae, which is widely distributed in the Old World tropics and contains 30-60 species worldwide (Jeffrey 1967, Verdcourt 1997, Lu & Jeffrey 2011, Schaefer & Renner 2011, De Boer 2015. There are about 11 species recorded in tropical East Africa (Uganda, Kenya and Tanzania), most of which could be found in Kenya (Jeffrey 1967). ...
Zehneria subcoriacea Y.D. Zhou and Q.F. Wang, a new species of Cucurbitaceae is described and illustrated here, which is recorded as Zehneria sp. A in Flora of Tropical East Africa. It is easily recognized by its sub-coriaceous leaves and hairy stamens and staminodes.
