Checklist of the Cucurbitaceae of Madagascar

123 Checklist of the Cucurbitaceae of Madagascar Minosoa B. Andriamiharisoa1, Edgardo M. Ortiz2, Tendro Radanielina1, Hanno Schaefer2 1 Department of Plant Biology and Ecology, Campus universitaire d’Ambohitsaina, University of Antananarivo, BP 566, 101, Antananarivo, Madagascar 2 Plant Biodiversity, Department of Life Science Systems, Technical University of Munich (TUM), Emil-Ramann-Str. 2, D-85354 Freising, Germany Corresponding author: Hanno Schaefer ([email protected]) Copyright: © Minosoa B. Andriamiharisoa et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Checklist Abstract This checklist of Malagasy Cucurbitaceae comprises 26 genera with 77 species and five varieties, including five new species to be formally described when more complete ecological and genetic data are available. Of the 82 taxa, 65 are classified as indigenous, and of those, 52 species and 5 varieties are endemic (88% of the native taxa). Six exotic species have permanent populations outside cultivation (naturalised exotics). Another eleven exotics are cultivated and occasionally found escaped from cultivation (casuals). The checklist covers taxonomic information including type specimen(s), vernacular names, DNA sequence information, photos, global and national distribution data with status (endemic, non-endemic native, naturalised, or cultivated), plus ecological and habitat information. Our analysis of IUCN conservation status suggests that half of the native species (32) are threatened, mainly due to habitat destruction. For 12 taxa we lack information to suggest IUCN status but many of them might turn out to be threatened or even extinct. Nine species are absent from protected area, 21 were found only in a single reserve. The Andohahela National Park is home to the greatest Cucurbitaceae diversity with 13 species, followed by Zombitsy-Vohibasia with 12 species. The most species-rich genera are Peponium (up to ten endemic species), Zehneria (eight species, six of them endemic), Ampelosycios and Xerosicyos (each seven endemic species). Our study reveals overlooked diversity in a plant family of global importance and highlights the need for additional fieldwork and collection in several genera, including Cayaponia, Cyclantheropsis, Peponium, and Kedrostis. Key words: Andohahela National Park, biodiversity hotspot, crop wild relatives, endemic species, Kedrostis, Peponium, Zombitsy-Vohibasia National Park Academic editor: Yasen Mutafchiev Received: 16 May 2025 Accepted: 14 November 2025 Published: 5 December 2025 Citation: Andriamiharisoa MB, Ortiz EM, Radanielina T, Schaefer H (2025) Checklist of the Cucurbitaceae of Madagascar. PhytoKeys 267: 123–222. https://doi.org/10.3897/ phytokeys.267.159218 PhytoKeys 267: 123–222 (2025) DOI: 10.3897/phytokeys.267.159218 124 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Table of contents Introduction ........................................................................................................124 Geography ......................................................................................................124 Geology of Madagascar ................................................................................125 Climate of Madagascar .................................................................................125 History of botanical exploration in Madagascar .........................................126 Vegetation ......................................................................................................127 Objectives .......................................................................................................127 Material and methods ........................................................................................128 Checklist of the species in alphabetical order .................................................129 Accepted species found outside cultivation ................................................129 Naturalised species .......................................................................................192 Cultivated species which occasionally escape ...........................................199 Doubtful species and erroneous records .....................................................206 Results ................................................................................................................208 Total species and comparison with the lists of Keraudren and Rauh ........208 Total species with DNA sequences ..............................................................209 Endemism ......................................................................................................209 Diversity ..........................................................................................................210 Analysis of IUCN classifications ...................................................................210 Discussion and conclusions .............................................................................212 Acknowledgements ...........................................................................................214 Additional information .......................................................................................214 References ..........................................................................................................215 Supplementary material 1 .................................................................................222 Introduction Geography Madagascar is located between 12°–26°S and 43°–51°E in the southwestern part of the Indian Ocean, separated from the African continent by the Mozambique Channel. From north to south, the country stretches over 1,650 km and from west to east over 560 km, with a total surface area of 587,000 km2 (Goodman 2022). The western part accounts for up to two thirds of Madagascar’s total surface area, separated from the eastern part by the High Plateau with the highest mountain Tsaratanana, 2,876 m a.s.l. (Goodman 2022). The island can be divided into six phytogeographic domains: Central, Eastern, High Mountain, Sambirano, Southern, and Western domain (Fig. 1) (Humbert 1955), and six political provinces: Antananarivo, Toamasina, Antsiranana, Mahajanga, Fianarantsoa, and Toliary which are further divided into 24 regions (Faritra): Diana, Sava, Itasy, Analamanga, Vakinankaratra, Bongolava, Sofia, Ambatosoa, Boeny, Betsiboka, Melaky, Alaotra-Mangoro, Atsinanana, Analanjirofo, Amoron’i Mania, Vatovavy, Haute Matsiatra, Fitovinany, Atsimo-Atsinanana, Ihorombe, Menabe, Atsimo-Andrefana, Androy, and Anôsy. These regions are subdivided into 119 districts and 1,693 municipalities. In terms of demographics, according to the Institut National de la Statistique, in 2018 the total population of Madagascar was 25,674,196 (INSTAT 2020). 125 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Geology of Madagascar The 3-billion-year geological history of Madagascar includes continental separation, tectonic plate convergence, and subsequent continental collisions (Goodman 2022). More than two thirds of Madagascar’s bedrock is of Cambrian age, so it is roughly 540 million years (Myr) old (Collins and Windley 2002). The island of Madagascar broke away from Gondwana in the Late Cretaceous between 170–185 million years ago (Mya) (Goodman 2022) and today’s island of Madagascar is the result of the unification of several Gondwana blocks, the Antongil Block, Antananarivo Block, Tsaratanana Sheet, and Bemarivo Belt (Collins 2006). Several regions are today dominated by Neoproterozoic metasedimentary rocks, including the Molo, Betsimisaraka, Vohibory, and Androyen domains. The Antananarivo Block is underlain by gneiss, granitoid, and gabbros with an age of c. 2.5 billion years (Collins and Windley 2002). The Itremo sheet is mainly formed from Neoproterozoic metasedimentary rocks thrust over and imbricated with the Antananarivo Block. The Tsaratanana sheet lies on Middle to Late Archean gneiss with an age of 2.7–2.5 billion years (Cox et al. 1998). The Bemarivo Belt, located in the North, lies on east to west striking meta-sediments, granites, and gneiss overlapped by meta-volcanic rocks deformed by contraction (Thomas et al. 2009). The western part of Madagascar is dominated by meta-sediments, represented in the Northwest by the Mahajanga Basin and in the Southwest by the Morondava Basin (Collins and Windley 2002), which includes Upper Cretaceous volcanic rocks (Goodman 2022). Additionally, two types of volcanic formations are present in the central part of Madagascar: the massifs of Ankaratra and Itasy, which contain basaltic rocks from the Miocene epoch, 18 Myr old (Cucciniello et al. 2017). Climate of Madagascar Due to its wide latitude, mountain ranges, and influence of the sea, Madagascar has a very diverse climate. The warm winds from the Indian Ocean significantly increase humidity on the island’s eastern side. The dynamic interaction Figure 1. Phytogeographic domains of Madagascar based on Humbert (1955). 126 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae of subtropical anticyclones, trade winds, and monsoon flow also affects the climate (Nassor and Jury 1998). Broadly speaking, the climate of Madagascar can be categorized into four different climatic zones: the arid south, the temperate highlands, and the tropical eastern and western coastlines (Goodman 2022). The highlands experience a summer rainy season, the Southwest faces a leeward and dry climate, while the Northwest exhibits a tropical climate with heavy summer rain (Nassor and Jury 1998). The eastern part of the island has high humidity and 2,400 mm of precipitation per year, which can rise to over 6,000 mm in some places due to moisture laden winds from the Indian Ocean. In the South, it is much drier with annual precipitation of less than 600 mm, in the highlands about 1,600 mm are reached (Benstead and Goodman 2003). In the south and southwest, the lowest temperatures are between 20–25 °C in the coldest months of May–August. In comparison, average temperatures during the warmer months of November through mid-April range from 25–29 °C in the East to 27–28 °C in the South and Southwest. The central plateau, located at an elevation of around 1,650 m, typically has average temperatures between 12.5–20 °C throughout the hottest and coldest months and frost can occur at the highest elevations (Goodman 2022). History of botanical exploration in Madagascar The discovery of the Malagasy flora began in the 16th and 17th centuries, when European navigators first visited the island and brought back mainly medicinal plants and spices. Among them, Étienne de Flacourt (1607–1660), French governor of Madagascar for the French East India Company, is the most renowned. He arrived in Fort Dauphin (Tolagnaro) in 1648 and stayed until 1655. In 1658, he published the “Histoire de la grande île de Madagascar”, which includes the first description of Malagasy plant species. The French naturalist Philibert Commerson (1727– 1773) came to collect for four months in 1770, while French aristocrat Louis-Marie Aubert du Petit Thouars (1758–1831) spent ten years (1792–1802) in exile in Madagascar after the French Revolution and collected about 2000 plant specimens. In 1822–1823, the Czech botanist Wenceslas Bojer (1795–1856) explored western Madagascar and visited together with the German botanist Carl Theodor Hilsenberg (1802-1824) the rainforests of the East, where Hilsenberg died of Malaria at the age of 22. French botanist Louis Hyacinthe Boivin (1808–1852) visited Madagascar between 1847–1852 and died of Malaria the day after his return to France. The second era began in 1865 with the arrival of French explorer Alfred Grandidier (1836–1921) and his son, Guillaume Grandidier (1873–1957). Supported by the Museum d’Histoire Naturelle de Paris, they published the “Histoire physique, naturelle et politique de Madagascar” (1861) and “La bibliographie de Madagascar” (1906). The British missionary and botanist Richard Baron (1847–1907) arrived in 1872 and spent several decades collecting plants in the North and Centre. In his book “Compendium des plantes malgaches” (1901– 1906), he listed 4,100 species in 970 genera. The arrival of Henri Perrier de la Bâthie (1873–1958) in 1896 marked the beginning of the third great era of botanical discovery in Madagascar. He collected more than 20,000 herbarium specimens for the herbarium of the Museum d’Histoire Naturelle de Paris, and the National Herbarium of Madagascar (TAN) housed in the Tsimbazaza Botanical Garden in Antananarivo, which he had founded in 1925 and directed for over 40 years. In 1936, Henri Humbert (1887–1967) initiated the 127 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae project “Flore de Madagascar (Plantes vasculaires)”, which later became the “Flore de Madagascar et des Comores”, published by the Museum d’Histoire Naturelle in Paris. In the same period, French botanists Raymond Decary (1891–1973), Henri Louis Poisson (1877–1963), Jacques D. Leandri (1903–1982), Jean Marie Bosser (1922–2013), and Bernard Descoings (1931–2018) contributed to the development of our knowledge of the Malagasy flora. These colleagues collected a significant part of the Malagasy Cucurbitaceae specimens available today, among them many later type specimens but they described very few of the species. The French botanist Monique Keraudren (1928–1981) wrote her PhD thesis on the Cucurbitaceae of Madagascar (Keraudren 1968) and she worked intensively with the available herbarium material in Paris and her own new collections and described 20 new taxa. Since Keraudren, hardly any research has been conducted on the Cucurbitaceae of Madagascar. The only exception was the German Werner Rauh (1913– 2000), who was broadly interested in succulent plants of all kinds and did some work on the cucurbit genera Seyrigia, Corallocarpus, and Xerosicyos (Rauh 1995). Vegetation The first phytogeographical classification of Madagascar was established by Richard Baron who divided the country into three phytogeographical zones: East, Centre, and West (Baron 1889). Henri Perrier de la Bâthie (1921) added two types of formation in relation to southeast trade wind exposure: “Flore du vent” (east) and “Flore sous le vent (west). He divided the “Flore du vent” into three distinct zones: East, Central and Sambirano, and the “Flore sous le vent” into a western and a southern zone. Madagascar is marked by an aridity that increases from east to west due to the drying out of the trade wind after its passage over the eastern flank and central part of the island. This so-called “Foehn Effect” causes the “Flore sous le vent” of the West and South with distinct xerophytes and forms of adaptation to drought. The East of Madagascar is exposed to the trade wind all year. In the central part, the remaining wind, charged with moisture, causes the formation of fog and drizzle, which reduces the aridity of the dry season. In consequence, the vegetation varies according to temperature and rainfall in the eastern part of the island, while it varies according to different soil types in the western part. In 1955, Henri Humbert integrated both vegetation and plant distribution data into a new proposal for the phytogeographic division of Madagascar (Humbert 1955), which would later become a more refined classification in his “Carte de la végétation de Madagascar”, which comprised three scales: “région”, “domaine”, and “secteur”. He divided the island into two zones: East and West (Humbert and Darne 1965), which differ mainly in their humidity levels, underlining the edaphic variations within the different districts. The Eastern zone comprises three domains: East, Sambirano, and Center plus High Mountains; the Western zone is made up of two domains: West and South. The original classification of Humbert (1955) has been refined by Faramalala et al. (1995), Du Puy and Moat (1996), and Gautier et al. (2018) taking anthropogenic transformation into account. Objectives With this checklist, we want to provide an update on taxonomy and distribution of the Cucurbitaceae of Madagascar and add, wherever possible, also ecological 128 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae information and evaluations of the threat level. While the information for most taxa is still incomplete, we see this as a first important step toward a modern flora treatment of the Cucurbitaceae of Madagascar. Several of the Malagasy cucurbit species are close relatives of important crops (crop wild relatives, CWR) and often evolved unique adaptations to drought (succulent organs) which makes their study also relevant for breeding of new crop varieties in a world with changing climates. Material and methods The basis for our work is the last comprehensive Cucurbitaceae treatment of the “Flore de Madagascar et des Comores” (Keraudren 1966), which listed 27 genera with 70 species. To evaluate and update Keraudren’s treatment, we studied and collected Cucurbitaceae in ten protected areas of Madagascar from December 2021 to January 2025: Andasibe-Mantadia National Park (NP), Andohahela NP, Ankarafantsika NP, Ankarana Special Reserve, Ankodida, Behara-Tranomaro, Berenty private reserve, Ranomafana NP, Montagne d’Ambre NP, Montagne des Français, and Zombitsy-Vohibasia NP. Herbarium voucher specimens have been deposited at the herbaria of Department of Botany, University of Antananarivo, DBEV, and Technical University of Munich, TUM. In addition, we examined several thousand herbarium specimens of the following herbaria: A, BR, CAS, CNARP, G, K, L, MO, P, S, TAN, US, W, and WAG, herbarium acronyms follow Index herbarorium (sweetgum.nybg.org/science/ ih/). We also examined all Cucurbitaceae records on iNaturalist (https://www. inaturalist.org) and Tropicos (legacy.tropicos.org/Project/Madagascar), the latter including records from the Missouri Botanical Garden Team (MBG), Tsimbazaza Botanical and Zoological Park Team (PBZT-TAN), and Kew-Madagascar Conservation Centre Team (KMCC). Finally, we assembled a database of geographic occurrence records for all Malagasy Cucurbitaceae from the Global Biodiversity Facility (GBIF.org 2022, 2023). Relevant protologues and diagnoses were studied on Biodiversity Heritage Library (BHL, https:// www.biodiversitylibrary.org/), JSTOR Global Plants (http://plants.jstor.org), and Botanicus (https://www.botanicus.org/). The vegetation classification used in this study follows the typology of Gautier et al. (2018). Information on vernacular names was compiled from herbarium labels. To suggest the IUCN conservation status for all native species, we determined the Extent of Occurrence (EOO, criterion B1) and Area of Occupancy (AOO, criterion B2) (IUCN 2012) to identify species or populations with a restricted distribution that are either fragmented or occupy few locations. Population declines or extreme fluctuations in distribution areas could not be considered due to the lack of temporal information. To assess AOO, grid cells of 2 × 2 km were used as recommended by IUCN (IUCN Standards and Petitions Committee 2022) and EOO computing was based on the minimum convex polygon (MCP) from the REDLISTR package (Lee et al. 2019) using R version 4.4.2 (R Core Team 2024). A species is classified as Data Deficient (DD), when there are fewer than two occurrence data points. To obtain distribution maps, occurrence data was plotted on maps showing the phytogeographic domains (Humbert 1955) and protected areas of Madagascar (https://protectedareas.mg/). From these maps, we inferred diversity hotspots at both genus and species levels and tried to identify key areas for Cucurbitaceae conservation in Madagascar. 129 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Table 1. Native Cucurbitaceae of Madagascar with status and distribution. No. Scientific name Status Phytogeographic domain 1. Ampelosycios bosseri (Keraudren) H. Schaef. & S.S. Renner Endemic Western 2. Ampelosycios humblotii (Cogn.) Jum. & H. Perrier Endemic Sambirano, Western, Central, Eastern 3. Ampelosycios leandrii (Keraudren) H. Schaef. & S.S. Renner Endemic Western 4. Ampelosycios meridionalis Keraudren Endemic Western, Southern 5. Ampelosycios scandens Thouars Endemic Central 6. Ampelosycios sp. nov. (Ankarana) Endemic Western 7. Ampelosycios sp. nov. (Montagne des Français) Endemic Western 8. Blastania lucorum (Keraudren) H. Schaef. Endemic Western, Southern 9. Cayaponia africana (Hook.f.) Exell var. madagascariensis Keraudren Endemic Western 10. Citrullus mucosospermus (Fursa) Fursa probably native but nonendemic, CWR Western, Central, Eastern, Southern 11. Corallocarpus grevei (Keraudren) Keraudren Endemic High Mountain, Western, Southern 12. Corallocarpus perrieri (Keraudren) Keraudren Endemic Southern 13. Corallocarpus poissonii Cogn. Native non-endemic Western 14. Cucumis sacleuxii Paill. & Bois Native but not endemic, CWR Western, Central, Eastern 15. Cucumis sp. nov. (aff. cinereus)Endemic, CWR Western 16. Cucumis sp. nov. (aff. hirsutus)Endemic, CWR High Mountain, Central 17. Cucumis subsericeus Hook.f. Native non-endemic, CWR Sambirano, High Mountain, Central 18. Cyclantheropsis madagascariensis Keraudren Endemic Western, Southern 19. Gerrardanthus lobatus (Cogn.) C. Jeffrey Native non-endemic Western 20. Kedrostis cogniauxii Keraudren Endemic Sambirano 21. Kedrostis dissecta Keraudren Endemic Western, Central 22. Kedrostis elongata Keraudren Endemic Sambirano, Western, Central, Eastern 23. Kedrostis lanuginosa Keraudren Endemic Western, Eastern 24. Kedrostis laxa Keraudren Endemic Western 25. Kedrostis perrieri Keraudren Endemic Western 26. Lemurosicyos variegata (Cogn.) Keraudren Endemic Western, Southern 27. Muellerargia jeffreyana Keraudren Endemic, CWR Western, Central 28. Peponium betsiliense Keraudren Endemic Western, Central 29. Peponium boivinii (Cogn.) Engl. Endemic Sambirano, Western 30. Peponium grandidieri Keraudren Endemic Sambirano, Western, Southern 31a. Peponium hirtellum Keraudren var. hirtellum Keraudren Endemic Southern 31b. Peponium hirtellum Keraudren var. longiracemosum Keraudren Endemic Central, Eastern, Southern 32. Peponium humbertii Keraudren Endemic High Mountain, Western, Central 33. Peponium laceratum Keraudren Endemic Central 34a. Peponium perrieri Keraudren var. perrieri Endemic Western, Central 34b. Peponium perrieri Keraudren var. glabrescens Keraudren Endemic Western 35. Peponium poissonii Keraudren Endemic Western, Eastern, Southern 36. Peponium racemosum Keraudren Endemic Western, Central 37a. Peponium seyrigii Keraudren var. seyrigii Endemic Southern Checklist of the species in alphabetical order Accepted species found outside cultivation A total of 66 species of Cucurbitaceae have been found in the wild in Madagascar. Of those, 60 species (plus five varieties) are classified as indigenous species (Table 1) and most of them are endemic to Madagascar (52 species plus five varieties). An additional six species are classified as naturalised exotics, which have been introduced by human activities (Table 2), some of them with invasive tendencies, others with localised but well-established populations in the wild. 130 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae No. Scientific name Status Phytogeographic domain 37b. Peponium seyrigii Keraudren var. linearilobum Keraudren Endemic Western 38. Raphidiocystis brachypoda Baker Endemic High Mountain, Western, Central 39. Seyrigia bosseri Keraudren Endemic Central, Southern 40. Seyrigia gracilis Keraudren Endemic Western, Southern 41. Seyrigia humbertii Keraudren Endemic Southern 42. Seyrigia marnieri Keraudren Endemic Western, Southern 43. Seyrigia multiflora Keraudren Endemic Central, Southern 44. Seyrigia napifera Rauh Endemic Central, Southern 45. Trochomeriopsis diversifolia Cogn. Endemic Sambirano, Western, Central, Southern 46. Xerosicyos danguyi Humbert Endemic Western, Eastern, Southern 47. Xerosicyos decaryi Guillaumin & Keraudren Endemic Western, Southern 48. Xerosicyos hirtellus (Humbert) H. Schaef. & S.S. Renner Endemic Southern 49. Xerosicyos perrieri Humbert Endemic Western, Central, Southern 50. Xerosicyos pubescens Keraudren Endemic Southern 51. Xerosicyos sp. nov. (North Madagascar) Endemic Western 52. Xerosicyos tripartitus (Humbert) H. Schaef. & S.S. Renner Endemic Central, Southern 53. Zehneria emirnensis (Baker) Keraudren Native non-endemic High Mountain, Central, Eastern, Southern 54. Zehneria madagascariensis Keraudren Endemic High Mountain, Central 55. Zehneria martinez-crovettoi Keraudren Endemic Central, Southern 56. Zehneria peneyana (Naudin) Schweinf. & Asch. Native non-endemic Western 57a. Zehneria perrieri Keraudren var. perrieri Endemic Central, Western 57b. Zehneria perrieri Keraudren var. parvula Keraudren Endemic Central 57c. Zehneria perrieri Keraudren var. tsaratananensis Keraudren Endemic High Mountain 58. Zehneria polycarpa (Cogn.) Keraudren Endemic Sambirano, Western, Central, Eastern, Southern 59. Zehneria rutenbergiana (Cogn.) Keraudren Endemic Central, Eastern 60. Zehneria tridactyla (Hook.f.) R. Fern. & A. Fern. Native non-endemic Sambirano, Western, Central, Eastern Ampelosycios Thouars, Hist. Vég. Isles Austral. Afriq.: 68 (1808). Generic type. Ampelosycios scandens Thouars. Worldwide Distribution. endemic to Madagascar (Schaefer 2020). Ampelosycios bosseri (Keraudren) H. Schaef. & S. S. Renner, Taxon 60: 133 (2011) Fig. 2 Basionym. Odosicyos bosseri Keraudren, Bull. Soc. Bot. France, Lett. Bot. 127(5): 518. 1980 [1981]. Vernacular name. Belataka antaninolo (Masikoro). Holotype. Madagascar • Forêt de Lambomakandro entre Ranohira et Sakaraha, forêt sèche, Jan. 1962, M. Keraudren 1312 P (P00464856!). Isotypes. P (P00464854!, P00464855!). GenBank information. DNA sequences on NCBI from Kocyan et al. (2007): DQ535832, DQ536710, DQ536773, DQ648183 and Filipowicz and Renner (2010): HM008593. Distribution. Endemic to Madagascar, where it is found only in the surroundings of Sakaraha (Fig. 3). Western Domain: Atsimo-Andrefana region, from Ranohira to Sakaraha, Lambomakandro, M. Keraudren 485, 1312 (P), J. Leandri 3891 (P), M.M. Debray 1914 (P), S. Hofstätter 73242 (P); Ambiamena, Zombitsy NP, H. Schaefer et al. 88, 90 (TUM). 131 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Occurrence in Protected Areas. Zombitsy-Vohibasia NP. Ecology and habitat. Remnants of the western moist semi-deciduous forest on sandy soil at an altitude of 500 m asl.; flowering Nov.–Oct. (Keraudren-Aymonin 1980, 1982a). Uses. The species is occasionally grown as an ornamental caudex plant in the Northern Hemisphere, and wild-collected material of dubious origin is sold online. Taxonomy. According to Schaefer et al. (2009), A. bosseri is sister to A. leandrii. IUCN. The AOO of the species is equal to 7 km2, which classifies the species as Critically Endangered (CR). In addition to strong pressures from forest fires and illegal grazing, its habitat is threatened by mining activities (Gautier et al. 2018). Ampelosycios humblotii (Cogn.) Jum. & H. Perrier, Ann. Fac. Sci. Marseille 23: 29 (1915) Fig. 4 Basionym. Delognaea humblotii Cogn., Bull. Mens. Soc. Linn. Paris 1: 425 (1884). Vernacular name. Vahitakaboka (Betsileo), Vahanonoka (Betsimisaraka), Voanono (Betsimisaraka). Holotype. Madagascar • 28 Oct. 1883, Humblot 203 P (P00135525!). Isotypes. P (P00135526!), K (K000310405!), MO (MO2945897!). GenBank information. DNA sequences on NCBI published by Kocyan et al. (2007) and Bellot et al. (2020): DQ501254, DQ501261, DQ521608, NC046869, MN542396. Distribution. Endemic to Madagascar where it is found mainly in the East (Fig. 5). Sambirano Domain: Diana region, Lokobe, L.H. Boivin 1851-3 (P). Central Domain: Ihorombe region, Iakora, Begogo, Bekora, Sahalava Forest, Andrianjafy 776 (MO); Ivohibe, P. De Block, J. Tosh & F. Rakotonasolo 320, 321 (BR). Eastern Domain: Vatovavy region, Ranomafana NP, R. Rakoto 46 (WAG), H. Schaefer et al. 110, 121, 126 (DBEV, TUM); Ambatosoa region, Maroantsetra, Ambinanitelo, Marovovonana, Antilahimena 2881 (MO), G. Cours 4075 (P); Alaotra-Mangoro region, Analamazaotra, Andasibe, M. Keraudren 1752 (P), G.G. Aymonin 25373 (P), M. Keraudren-Aymonin 25366, 25373 (P), H. Perrier de la Bâthie 9017 (P), J. Bosser 15952 (P); Sava region, Masoala Peninsula Vasey 117 (WAG), R. Behasy 117 (P). Occurrence in Protected Areas. Lokobe NP, Masoala NP, Mantadia NP, Ranomafana NP, Tsingy de Bemaraha NP, and Ankeniheny-Zahamena Corridor Natural Resource Reserve. Ecology and habitat. Lowland and medium altitude moist evergreen forest on alluvial deposits between 200–1,700 m asl. Uses. Locals eat the roasted seeds (Keraudren 1968). Taxonomy. According to Schaefer et al. (2009), A. humblotii is sister to A. scandens. IUCN. Based on the AOO assessment the species is Critically Endangered (CR), but the EOO indicates a status of Least Concern (LC), which might be more appropriate. 138 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 14–19. 14. Ampelosycios sp. nov. Montagne des Français. A. Male inflorescence; B. Habit; C. Male flower in full bloom (photos by MBA, Paysages Harmonieux d’Ambohitr’Antsingy, December 2023). 15. Distribution of Ampelosycios sp. nov. Montagne des Français. 16. Blastania lucorum, dry leaf (photo by HS, Sakaraha-Vohibasia, October 2022). 17. Distribution of Blastania lucorum. 18. Cayaponia africana var. madagascariensis, Holotype, Boïna, H. Perrier de la Bâthie 6776 (P). 19. Distribution of Cayaponia africana var. madagascariensis. 14 15 16 17 18 19 139 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Western Domain: Sofia region, upper Bemarivo, H. Perrier de la Bâthie 6771, 6776, 6779 (P); Betsiboka region, alluvial deposits of Betsiboka River, H. Perrier de la Bâthie 1288 (P). Occurrence in Protected Areas. Not known from protected areas. Ecology and habitat. Dry deciduous forest at c. 500 m asl., on sandy soil. Taxonomy. The species C. africana is found in West Africa and in Madagascar, where it is represented by C. africana var. madagascariensis Keraudren (Keraudren 1968). The taxonomic status of this variety remains unclear due to the lack of material for phylogenetic or population-level studies (Duchen and Renner 2010). IUCN. We suggest Data Deficient (DD), due to the limited number of occurrence data. Citrullus Schrad. ex Eckl. & Zeyh., Enum. Pl. Afric. Austral.: 279 (1836), nom. cons. Generic type. Citrullus lanatus (Thunb.) Matsum. & Nakai. Worldwide Distribution. Eastern Mediterranean region, Africa, and Western Asia (Schaefer 2020). Citrullus mucosospermus (Fursa) Fursa, Trudy Prikl. Bot. 81: 111 (1983) Fig. 20 Holotype. ghana • 5 Aug. 1957, N.P. Oltarshevskyi 3833 (WIR k-3742). Basionym. Citrullus lanatus subsp. mucosospermus Fursa, Bot. Zhurn. (Moscow & Leningrad) 57: 38 (1972). Vernacular name. Voazavo (Tanala). GenBank information. DNA sequences of this taxon published by Chomicki and Renner (2015), and Shi et al. (2017) KY430686, NC033899, KY430687, KY430693, KX773720, KX773718, KP036543, KP058579, KM281404, and KP036531; sequences of Malagasy material (unpubl. data) will be made available. Distribution. Native to Ghana, Guinea, Nigeria, Senegal, and Sudan (POWO 2023). In Madagascar, it is mainly found in the occidental part (Fig. 21), its indigenous status remains doubtful. Occurs in the Western, Central, and Southern Domains: Zombitsy-Vohibasia NP, H. Schaefer et al. 84, 85, 86, 87 (DBEV, TUM); Ambiamena, H. Schaefer et al. 92 (DBEV, TUM); Ambinany, H. Schaefer et al. 98, 99 (DBEV, TUM). Occurrence in Protected Areas. Ranobe PK32 New Protected Area, Ankarafantsika NP, and Zombitse-Vohibasia Protected Area. Note. All specimens identified by Keraudren (1966) as Citrullus colocynthis (L.) Schrad. and revised by the authors in P belong, in fact, to C. mucosospermus. Ecology and habitat. Annual plant of the seasonally dry tropical biome, where it grows in agricultural areas and on riverbanks. According to herbarium label information (J.M. Hildebrandt 3435 (P)), C. mucosospermus is spread by wild boar, dogs and humans, who eat the fruit and swallow the seeds. Uses. In West African countries, Citrullus mucosospermus, also known as ‘egusi’ watermelon, is cultivated mainly for its nutritious, oleaginous seeds, which can be used either to extract edible oil or to thicken soups (Achigan-Dako et al. 2008). This kind of usage is not known from Madagascar, where it 140 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae is rather used as a source of water for local communities in the arid zones and to feed livestock. The flesh is low in sugar and tastes more like cucumber than melon. Taxonomy. According to Guo et al. (2013) and Renner et al. (2017), C. mucosospermus is sister to C. lanatus and might better be treated as a subspecies of the latter. IUCN. We suggest Least Concern (LC), based on IUCN criterion B2. Corallocarpus Welw. ex Hook.f., G.Bentham & J.D.Hooker, Gen. Pl. 1: 831 (1867). Generic type. Corallocarpus welwitschii (Naudin) Hook.f. Worldwide Distribution. Africa, Arabia, India, Pakistan, and Madagascar (Schaefer 2020). Corallocarpus grevei (Keraudren) Keraudren, Fl. Madagasc. 185: 72 (1966) Fig. 22 Basionym. Calyptrosicyos grevei Keraudren, Compt. Rend. Hebd. Séances Acad. Sci. 248: 3593 (1959). Holotype. Madagascar • Mahajunga, sable-Majunga, Ouest, bois, Jan. 1919, H. Perrier de la Bâthie 12348 P (P00135484!). Isotypes. P (P00135485!, P00135486!). GenBank information. Has been sequenced by the authors (unpubl. data). Distribution. Endemic to Madagascar where it has been found mainly in the South and in a few spots in the West and Center (Fig. 23). Western Domain: Boeny region, Mahajanga, H. Perrier de la Bâthie 12348 (P); Amboanio, H. Perrier de la Bâthie 1401 (P); Ankarafantsika NP, H. Schaefer et al. 41 (TUM, DBEV). High Mountain Domain: Haute Matsiatra, Andringitra, Jean De Dieu Ratoto 1379 RN (P). Southern Domain: Atsimo-Andrefana region, Toliara, J. Bosser 13572 (P), T. B. Croat 30927 (P), M. Keraudren 738 (P); Andatabo, R. Randrianaivo 334 (WAG); N Fiherenana River, J. Léandri 3798 (P); on the road from Andranovory to Vatolatsaka, L. Allorge 2264 (P); Anjamala, M. Keraudren 677 (P); Mahafaly limestone plateau, near Ankilirano, M. Keraudren 824 (P, BR), 849, 850, 866 (P); Zombitsy NP, P. Morat 3798 (P), H. Schaefer et al. 80, 81 (DBEV, TUM); Antsepoka, Ramon 468 (P, TAN); Betioky, M. Keraudren 771 (P); along the road from Ampanihy to Ampotaka (Menarandra Downstream), M. Keraudren 903 (P); Befamata, H. Poisson 422 (P); Ambatry, J. Bosser 13564 (P); Ampandrandava, A. Seyrig 711, 711 B (P), Jardin Botanique Tananarive team 6453 (P); Ampanihy, A.M. Homolle 1662 (P); Antanimora, H. Humbert 28799bis (P); Belalanda, Ranobe PK32, Ranaivojaona 1667 (US), M. Andrianjafy 1667 (P); Beomby, J. Bosser 13557 (P), M. Keraudren 794, 797, 817 (P); from Ejeda to Fofadrevo, M. Keraudren 1437 (P); Fiherenana pass about 40 km from Miary, M. Keraudren 768 (P); Androy region, Ambovombe, R. Decary 8600 (P); Ambatomika, M. Keraudren 1081 (P); Behara, Luckow, M 4123 (WAG); Tranomaro New Protected Area, H. Schaefer et al. 5, 6A, 6B (DBEV, TUM); Tsihombe, M. Peltier 2851 (P). 141 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Anosy region, Anarafaly, J. Bosser 3760 (P), M. Keraudren 1000 (P); on the road from Amboasary to Fort-Dauphin, kilometer point n°434, M. Keraudren 1027 (P); Andohahela NP, Randriambololona 63 (MO), H. Schaefer et al. 60, 64 (DBEV, TUM); Menabe region, Morondava, Grevé 125 (P). Occurrence in Protected Areas. Andringitra NP, Ranobe PK32 New Protected Area, Andohahela NP, Ankodida Harmonious Protected Landscape, Tsinjoriake Harmonious Protected Landscape, Ankarafantsika NP, and Zombitse-Vohibasia NP. Ecology and habitat: Riverine and coastal forest on brown, red, or white sands, in dry spiny thicket from 10–700 m asl., characterised by low and irregular rainfall (< 600 mm/year), dry season of over 10 months, high to very high temperatures, significant annual temperature ranges, and high daily temperature ranges (Gautier et al. 2018); flowering throughout the year (Keraudren 1966). Taxonomy. Unresolved. IUCN. We suggest Least Concern (LC) based on IUCN criterion B2. Corallocarpus perrieri (Keraudren) Keraudren, Fl. Madagasc. 185: 68 (1966) Fig. 24. Basionym. Calyptrosicyos perrieri Keraudren, Compt. Rend. Hebd. Séances Acad. Sci. 248: 3593 (1959). Holotype. Madagascar • Toliara, Tsimanampetsotsa, Manampetsa, sables, Apr. 1933, H. Perrier de la Bâthie 19118 P (P00462267!). Isotypes. P (P00462268!, P00462269!). GenBank information. Has been sequenced by the authors (unpubl. data). Distribution. Endemic to Madagascar, where it occurs only in the South (Fig. 25). Southern Domain: Atsimo-Andrefana region, Toliara, M. Keraudren 722, 724 (P), J. Bosser 10151 (P), TB Croat 30828 (WAG, P), L. Allorge 2794 (P), Ramon 388 (MO, TAN), M. Keraudren 601 (BR); Anakao, Ramon 364 (MO, TAN); from Toliara to Manombo, P. Morat 3514 (P); Manombo, R. Decary 18696 (P); North of Toliara, J. Bosser 13563 (P); Ankilibe, M. Keraudren 601 (P); Ranobe PK32, Andrianjafy 1675 (BR, P, US, MO), M. Keraudren 682; Toliara II, Tsivonoakely, Ranaivojaona 187 (US); Belemboka, J. Dequaire 27399 (P); Efoetsy, M. Keraudren 883 (P); Itampolo, M. Keraudren 890 (P); Morombe, R. Decary 18744, 18745 (BR, P); on the roadside of the National Road n°10, R.N. Ravelonanahary 3651 (P); from Toliara to Manombo, P. Morat 3514 (P); Sarodrano, P. Morat 2502 (P); Tokilisy, M. Peltier 5490 (P); from Tulear to Morombe, M. Keraudren 707 (P); Tsimanampetsotsa, H. Perrier de la Bâthie 6790, 19118 (P), M. Peltier 3148, 3153, 3154 (P), H. Humbert 5373 (P); Ifotaka, Anarafaly, Z.S. Rogers 484 (P). Occurrence in Protected Areas. Ranobe PK32 New Protected Area, Tsimanampetsotsa NP, Tsinjoriake Harmonious Protected Landscape, Harmonious Protected Landscape of the Mangoky–Ihotry Wetland Complex. Ecology and habitat. Dry spiny thicket, with Adansonia rubrostipa and Delonix floribunda from 5–170 m asl.; flowering throughout the year (Keraudren 1966). Taxonomy. Unresolved. IUCN. We suggest Least Concern (LC), based on IUCN criterion B2. 142 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 20–25. 20. Citrullus mucosospermus. A. Shoot with leaves and male flower; B. Fruit (photos by MBA, Zombitsy-Vohibasia, October 2022). 21. Distribution of Citrullus mucosospermus. 22. Corallocarpus grevei. A. Habit and mature fruit; B. Young fruit (photos by L. Ramon, Toliara, Antsepoka, February 2017 (A) and by MBA, Andohahela NP, October 2022 (B)). 23. Distribution of Corallocarpus grevei. 24. Corallocarpus perrieri. A. Shoot with mature fruit; B. Male flower (photos A, B by S.E. Rakotoarisoa, Toliara, Ampanihy, Ankazoabo, January 2019). 25. Distribution of Corallocarpus perrieri. 20 21 22 23 24 25 Corallocarpus poissonii Cogn. A.L.P.P.de Candolle & A.C.P.de Candolle, Monogr. Phan. 3: 651 (1881) Fig. 26 Syntypes. coMores • Mayotte, Bords de la mer à Pammanzi, côte ouest parmi les broussailles; Feb. 1850; L.H. Boivin. s.n. P (P00135487!, P00135488!, P00135489!, P00135490!). Madagascar • Port Leven, M. Vesco P (P00475342!); sine loc., Baron 6457 (K), n.s. 143 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae GenBank information. Malagasy material has been sequenced by the authors (unpubl. data). Distribution. Endemic to Madagascar (where it is only known from the extreme North, Fig. 27) and Comoros, where the type specimens originated. The full distribution of C. poissonii is unclear due to confusion with C. bainesii (Hook.f.) A. Meeuse, which is a genetically distinct continental African species originally described from Botswana (Andriamiharisoa et al., in prep.). Western Domain: Diana Region, Andranovondronina, Antsisikala, Ambatoara, Analamangidy, R. Guittou 99 (P); Vovo Village, T.G, Razafindrabaeza, et al. 1702 (K); Port Leven, M. Vesco 1850 (P); Montagne des Français, H. Schaefer & M.B. Andriamiharisoa 1A, 1B, 2, 5, 6 (TUM, DBEV). Occurrence inside Protected Areas. Protected Harmonious Landscape Ambohitr’Antsingy – Montagne des Francais and Protected Harmonious Landscape of Andrafiamena Andavakoera. Ecology and habitat. Dry deciduous forest on limestone and littoral thicket on brown and red sandy soils, from sea level to 450 m asl. Taxonomy. Unresolved. IUCN. We suggest Endangered (EN) based on IUCN criterion B2. Cucumis L., Sp. Pl. 1011. 1753. Generic type. Cucumis sativus L. Worldwide Distribution. Native to Africa, Asia, Australia, and Madagascar (Schaefer 2020). Cucumis sacleuxii Paill. & Bois, Bull. Soc. Natl. Acclim. France 37: 371 (1890) Fig. 28 Neotype. Tanzania • Zanzibar, Massazine, cliff area, grassland or bush, 15th Jul. 1961, H. Faulkner 2865 BR (BR0000008887313!), designated by Kirkbride, Biosyst. Monogr. Cucumis 41 (1993). Synonyms. Cucumis sativus var. usambarensis Zimm. Cucurbitac. 2: 179 (1922); Oreosyce aspera Cogn., Pflanzenr., IV, 275 I: 268 (1916). GenBank information. DNA sequences of the species (but not from Malagasy material!) published by Renner et al. (2007), Ghebretinsae et al. (2007), and Sebastian et al. (2010): EF595895, EF595945, DQ785838, DQ785852, DQ785880, EF093520, DQ785866, HM597099; Malagasy material has been sequenced by the authors (unpubl. data). Distribution. Native to East Africa (POWO 2023) and possibly Madagascar, where it has often been confused with C. sativus, so probably more widespread than indicated in the map (Fig. 29). Western Domain: Diana region, Joffreville, H. Schaefer & M.B. Andriamiharisoa 40 (January 2024); Sofia region, Befandriana Nord, Jardin Botanique d’Antananarivo team 5098 (P); Boeny region, Bevazaha, N. Ramamonjisoa 1627 (P); Ankarafantsika forest, near Ampijoroa, M. Keraudren 1255 (P); Ihorombe region, Isalo, M. Keraudren 462 (P). Central Domain: Sofia region, Betainkankana, Ankaizina, on the forest border, J. Bosser 2633 (P). 144 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Eastern Domain: Ilaka-Est, J. Bosser 17024 (P); Fort-Dauphin, R. Decary 9944 (P); Andohahela NP, H. Schaefer et al. 46 (DBEV, TUM); outside Andohahela NP, H. Schaefer et al. 58, 72 (DBEV, TUM). Occurrence in Protected Areas. Andohahela NP, Ankarafantsika NP, Montagne d’Ambre NP, and Montagne des Français Protected Harmonious Landscape. Ecology and habitat. In seasonal zones with moderate annual rainfall, in clearings around human settlements, in wetlands, wasteland, and the forest understory, mainly on clay soils, from sea level to 700 m asl. Uses. The fruits are edible but no information on uses in Madagascar is available. Taxonomy. Cucumis sacleuxii is sister to C. rostratus and C. metuliferus (Sebastian et al. 2010). IUCN. We suggest Least Concern (LC) based on IUCN criterion B2. Cucumis subsericeus Hook. f, Fl. Trop. Afr. 2: 545 (1871) Fig. 30 Holotype. angola • Cuanza Norte, Pungo Andongo, F.M.J. Welwitsch 838 (BM). Synonyms. Oreosyce africana Hook. F., Fl. Trop. Afr. 2: 545 (1871); Cucumis oreosyce H. Schaef., Blumea 51: 171 (2007) nom. superfl.; Cucumis cecilii N.E.Br., Bull. Misc. Inform. Kew 1906: 104 (1906); Cucumis parvifolius Cogn. ex Scott Elliot, J. Linn. Soc., Bot. 29: 19 (1891); Hymenosicyos subsericeus (Hook.f.) Harms, Notizbl. Bot. Gart. Berlin-Dahlem 8: 487 (1923); Oreosyce subsericea (Hook.f.) A. Meeuse, Bothalia 8: 22 (1962). Vernacular name. Voatangondolo (Merina, Betsileo). GenBank information. DNA sequences of this species (but not from Malagasy material!) published by Ghebretinsae et al. (2007): EF595956, EF595957; by Kocyan et al. (2007): DQ535833, DQ536576; and by Sebastian et al. (2010): HM597090. Distribution: Native throughout Tropical Africa from Bioko to Ethiopia, South Africa, and Madagascar (POWO 2023), where it is widespread mainly in the Centre and higher mountains (Fig. 31). High Mountain Domain: Sava, Andapa, Ambohimirahavavy, C. Rakotovao 2520 (MO, P); Beampoko Forest, C. Randrianarivelo 380 (P); East peak of Ambohimirahavavy, C. Rakotovao 2402 (P); forest edge, H. Humbert 25663 (P); Tsaratanana massif, camp no. 2, peak, Mahavavy Valley, P. Morat 2345, 2290 (P); Tsaratanana, H. Perrier de la Bâthie 16175 (P); southern flank of Antsianongatalata, H. Humbert 18396 (P); Ankaizinana, R. Decary 1796, 1829, 1993 (P); Region of Melaky, Ambohimirahavavy, H. Andriamiarinoro 14 (P); upper Bemafo, H. Humbert 25157 (P); rocks on the banks of Bemafo, Buerki 91 (MO, P, TAN). Sambirano Domain: Diana region, Ambilobe, Marivorahona, massif of Tsaratanana, H. Humbert et al. 25792 (P). Central Domain: Analamanga region, Antananarivo, C. d’ Alleizette s.n. (P); Ilafy, R. Decary s.n. (P), Peltier 3822 (P); Ambohimanga, Waterlot 56 (P); Vakinankaratra region, Ankaratra, Scott Elliot G.F. 1947 (BR); Ankaratra, Académie Malgache s.n. (P); around Tritriva Lake, H. Perrier de la Bâthie 3763, 13069 (P); Ihorombe region, Fianarantsoa, Ivohibe, R. Decary 5642 (P). Eastern Domain: Analamanga region, Mandraka, M. Keraudren-Aymonin 25401 (P); kilometer point n°23 of National Road from Antananarivo to Toamasina, J. Bosser 13401 (P); Alaotra-Mangoro Region, Imerimandroso, R. Decary 3957 145 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 26–31. 26. Corallocarpus poissonii. A. Habit; B. Fruits (photos by MBA, Montagne des Français, December 2023). 27. Distribution of Corallocarpus poissonii. 28. Cucumis sacleuxii. A. Young fruit; B. Female flower; C. Habit (photos by MBA, Andohahela NP (A, B), October 2022 and Montagne d’Ambre NP (C), December 2023). 29. Distribution of Cucumis sacleuxii. 30. Cucumis subsericeus. A. Habit; B. Young fruit (photos by MBA, Ambatofitorahana, March 2023). 31. Distribution of Cucumis subsericeus. 26 27 28 29 30 31 146 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae (P); Zahamena, Andrangivalo massif southwest of Alaotra Lake, Natural Reserve no. 3 of Zahamena, Onibe watershed, H. Humbert 17756 (P); wet undergrowth, Antsihanaka forest, Jardin Botanique Tananarive team 2258 (P); Ihorombe region, Sakamalio, Sakamalio valley, H. Humbert 13397 (P); Anosy region, Taolagnaro, Trafonaomby, Mountain, B. Randriamampionona 712 (P); Andohahela massif, H. Humbert 13620 bis (P); Betroka, Kalambatritra Protected Area, R. Razakamalala 2002 (P); Androy region, upper Mandrare Watershed, H. Humbert 6522 (P). Occurrence inside Protected Areas: New Protected Area of Ranobe PK32, Andohahela NP, Protected Harmonious Landscape of Lac Alaotra, Protected Harmonious Landscape of the Mangoky Ihotry Wetland Complex, Natural Resource Reserve of the Marojejy-Anjanaharibe South-Tsaratanàna Corridor (Northern Part), Tsaratanàna Strict Nature Reserve, and Kalambatritra Special Reserve. Ecology and habitat. Forest edges and clearings, burnt montane ericoïd thicket, wetlands, and grasslands, on gneissic laterite and rocky outcrop areas, climbing on trees or trailing along the ground, 1,000–2,600 m asl. The species’ habitat is characterised by high rainfall (>1,200 mm/year), no or only a brief dry season (<4 months), high air humidity (mist), an average temperature with low seasonal variation, and increasing daily temperature range and solar radiation with altitude (Gautier et al. 2018); flowering throughout the year (Keraudren 1966). Uses. On the African mainland, C. subsericeus filtrate is injected to treat gonorrhea (Yineger and Yewhalaw 2007). No such use has been documented in Madagascar. Taxonomy. Cucumis subsericeus is sister species to C. bryoniifolius (Merxm.) Ghebret. & Thulin (Renner et al. 2007). IUCN. We suggest Least Concern (LC) based on IUCN criterion B2. Cucumis sp. nov. (aff. cinereus) Fig. 32 GenBank information. Has been sequenced by the authors (unpubl. data). Distribution. Endemic to the South-western of Madagascar, where it is known only from a few collections from the Zombitsy area (Fig. 33). Western Domain: Atsimo-Andrefana region, Sakaraha, Zombitsy NP, M. Keraudren 480 (P), J. Bosser 19394 (P); Plateau Mahafaly, western Betioky, H. Humbert & R. Capuron 29458 (P). Occurrence in Protected Areas. Zombitsy-Vohibasia NP. Ecology and habitat. Moist semi-deciduous forest, on limestone soil (Keraudren 1966). Taxonomy. Unresolved. IUCN. We suggest Critically Endangered (CR) following IUCN criterion B1 and B2. Since the last collection dates to the 1960s, the species might have become extinct, but a detailed exploration of its former range is needed. Cucumis sp. nov. (aff. hirsutus) Fig. 34 GenBank information. Has been sequenced by the authors (unpubl. data). Distribution: Endemic to Madagascar, where only a few collections from the Centre and the North are known (Fig. 35). 147 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae High Mountain Domain: Haute Matsiatra region, Andringitra, H. Perrier de la Bâthie 14403 (P); Between Ivato and la Mania, H. Perrier de la Bâthie 12367 (P). Central Domain: Diana region, Montagne d’Ambre, Les Rousettes, M. Keraudren 1652 (P). Occurrence in Protected Areas. Montagne d’Ambre and Andringitra NP. Ecology and habitat. Montane ericoïd thicket, 1200–1800 m asl., characterised by important rainfall, no very pronounced dry season (potential for frost), and low annual temperature ranges, high daily temperature ranges and strong solar radiation (Gautier et al. 2018); flowering Feb.–March. Taxonomy. Unresolved. IUCN. We suggest Critically Endangered (CR) based on IUCN criterion B1 and B2. Since the last collection dates to the 1960s, the species might have become extinct, but a detailed exploration of its former range is needed. Cyclantheropsis Harms, Bot. Jahrb. Syst. 23: 167. 1896. Generic type. Cyclantheropsis parviflora (Cogn.) Harms. Worldwide Distribution. East and South tropical Africa and Madagascar (Schaefer 2020). Cyclantheropsis madagascariensis Keraudren, Fl. Madagasc. 185: 164 (1966) Fig. 36 Holotype. Madagascar • Anadabolava (moyen Mandrare), vestige de forêt sèche sur rocailles, 1 Feb. 1962, J. Bosser 15723 P (P00135356!). Isotype. P (P00135357!) GenBank information. No DNA sequences available for this taxon. Distribution. Endemic to Madagascar, where it was found in the West, North and South (Fig. 37). Western Domain: Diana region, Antsiranana, G. Rahajasoa et al. (MO, P, WAG); Melaky, Tsingy de Bemaraha, C.C.H. Jongkind 3512 (BR, WAG); Atsimo-Andrefana region, Anadabolava, J. Bosser 15723 (P), M. Keraudren 1089 (P); Atsimo-Andrefana region, Analavelona forest, H. Humbert 19726 (P). Occurrence in Protected Areas. Natural Monument of the Sacred Forest of Alandraza Analavelo, Tsingy de Bemaraha NP, and Ankarana Special Reserve. Ecology and habitat. Dry deciduous forests on limestone formation (tsingy), c. 180 m asl. Taxonomy. Unresolved. IUCN. Based on IUCN criterion B1, this species would be classified as Least Concern (LC). However, since we failed to confirm the presence of the species during our expeditions, we suggest classification based on criterion B2 as Critically Endangered (CR). Gerrardanthus Harv. ex Benth. & Hook.f., Gen. Pl. 1: 840 (1867) Generic type. Gerrardanthus macrorhizus Harv. ex Benth. & Hook.f. Worldwide Distribution. Tropical Africa and South Africa (Schaefer 2020), and Madagascar. 154 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae IUCN. Based on IUCN criterion B1 the species would be classified as Vulnerable (VU). However, based on our field observations we suggest assigning the status Endangered (EN), which better reflects its current condition. Kedrostis perrieri Keraudren, Bull. Soc. Bot. France 108: 242 (1961). Holotype. Madagascar • Mahajanga, Ouest: partie Nord-Ouest du domaine, Tsitampiky, 28 Sep. 1930, R. Decary 8157 P (P00135481!), our Fig. 50. Isotypes. P (P00135482!, P00135483!). GenBank information. No DNA sequences available. Distribution: Endemic to Madagascar where it is only known from the West (Fig. 51). Western Domain: Boeny region, Mahavavy-Kinkony Complexe, Ambongo, A. Pervillé 589 (P); Tsitampiky, R. Decary 8157 (HOLOTYPE, P); Sofia region, Bongolava between Port Bergé and Antsijomitondraka, H. Perrier de la Bâthie 6782 (P). Occurrence inside Protected Areas. Harmonious Protected Landscape Mahavavy Kinkony Wetland Complex. Ecology and habitat. Dry deciduous forest on sandy soil at c. 200 m asl. Taxonomy. Unresolved. IUCN. Based on IUCN criteria B1 and B2 we suggest classifying the species as Critically Endangered (CR) reflecting its highly restricted distribution and the few known occurrence records. Lemurosicyos Keraudren, Bull. Soc. Bot. France 110: 405. 1963 (publ. 1964). Generic type. Lemurosicyos variegata (Cogn.) Keraudren. Worldwide Distribution. Madagascar endemic (Schaefer 2020). Lemurosicyos variegata (Cogn.) Keraudren, Bull. Soc. Bot. France 110: 405 (1964) Fig. 52 Basionym. Luffa variegata Cogn., Abh. Naturwiss. Vereins Bremen 7: 250 (1882). Holotype. Madagascar • Nossy-Bé, Rutenberg s.n. (B, destroyed). Neotype. Madagascar • Toliara, Ambatomika, le long du terrain d’aviation d’Ambatomika, route Amboasary à Tsivory, 8 Apr. 1960, M. Keraudren 1080 P (P00475174!), designated by Keraudren (1963b: 405). GenBank information. DNA sequences of this taxon are available on NCBI published by Kocyan et al. (2007) and Schaefer and Renner (2011): DQ491011, DQ491017, DQ501257, DQ501266, HQ201984. Distribution: Endemic to Madagascar, where it is widespread across the country (Fig. 53). Western Domain: Diana Region, Ankarana Special Reserve, M. Keraudren 1688 (P), M. Bardot-Vaucoulon 1518 (P), H. Schaefer & M. B. Andriamiharisoa 11, 25 (TUM, DBEV); Nosy Be, Anjiabe, Analabe forest, R. Rabevohitra et al. 4559 (P); Sava Region, Bekaraoka forest, L. Gautier 4323 (P); Nosy Sata, Leven, Sata isle, L. H. Boivin 2572 (P); Betsiboka Region, Morataitra, H. Perrier de la Bâthie 519 (P); Maevatanana, R. Decary 19213 (P); Firingalava, H. Perrier de la Bâthie 519 (P); Melaky, Antsalova, on dry tsingy, J.F. Villiers 4907 (P); Antsingy, Ambodiriana ford, J. Léandri 155 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae 2680 (P); Tsiandro, Antsingy, J. Léandri 3012, 3058 (P); Antsalova, P. Morat 4858 (P); Tsingy of Bemaraha, D.J. Du, J.-N. Labat & A.M. Couté 891 (P, WAG), J. Léandri 1027, 1058 (P), C.C.H. Jongkind, J.L. Andriantiana & H.E. Razanatsoa 3233 (WAG); Tsingy of Marovato, S.G. Razafimandimbison 65 (P); Upper Bemarivo, H. Perrier de la Bâthie 6773 (P); Boeny Region, Tsaramandroso, M. Peltier 5216 (P); Bevazaha, RN 33, 1883 (P); Madirovalo dry forest, H. Perrier de la Bâthie 6786 (P); iNaturalist observation: https://www.inaturalist.org/observations/1967893). Southern Domain: Androy region, in Ampandrandava, A. Seyrig 615, 615B (P); Atsimo-Andrefana region, Zombitsy NP, Lambomakandro Forest, M. Keraudren 1314 (P), P. Morat 3795bis (P); Mahaboboka, M. Peltier 5823 (P), Randrianarivony, F. Rakotoarivony & Rehary 275 (MO, P), B. Tefy Andriamihajarivo & F. Rakotoarivony 1868 (P), Andriamihajarivo 1868 (MO); Anjoho forest, P. Morat 2489 (P); Mahafaly Plateau, H. Humbert 29455, 29508 (P), B. Descoings 761 (P); Beomby, J. Bosser 13558 (P), M. Keraudren 792 (P), H. Humbert 20288 (P); from Sakamena to Sakoa Valley, H. Humbert 29432 (P); Tanandava, J. Bosser 16089, 16091, 16112 (MO, P); Ankazoabo, P. Morat 2548, 2517 (P), J. Bosser 17273, 17704 (P); Andriambe, P. Morat 3810 (P); Beroroha, Androtsy forest, R. Razakamalala 5946 (P); Androy region, Ifotaka, J. Bosser 13562 (P); Bekiria, M. Keraudren 1012 (BR, P); Ambovombe, clearing area, R. Decary 8597 (P); Ambatomika, M. Keraudren 1080 (BR), M. Keraudren 1080 (P); Menabe Region, 20 km SW of Besely on the road to Tongobory, M. A. Luckow 4194 (US, WAG); Ihorombe region, on the Ranotsara plain boundaries, M. Keraudren 328 (P). Occurrences inside Protected Areas. Ankarafantsika NP, Tsingy de Bemaraha NP, Zombitse-Vohibasia NP, Loky Manambato Harmonious Protected Landscape, Nord-Ifotaka Harmonious Protected Landscape, and Ankarana Special Reserve. Ecology and habitat. Moist semi deciduous and dry deciduous forest, on riverbanks, forest edges, roadsides, clearings, and in savanna on red sand, limestone, and alluvial deposits, from 100–1,100 m asl.; flowering Dec.–Apr. Taxonomy. Lemurosicyos variegata is sister to Solena heterophylla and Borneosicyos simplex (Kocyan et al. 2007). IUCN. Based on IUCN criterion B1, the species would qualify as Endangered (EN). However, based on B2, we suggest Least Concern (LC). Muellerargia Cogn., Monogr. Phan. 3: 630. 1881. Generic type. Muellerargia timorensis Cogn. Worldwide Distribution. Northeastern Australia, New Guinea, and Madagascar (Schaefer 2020). Muellerargia jeffreyana Keraudren, Adansonia, n.s., 5: 423 (1965) Fig. 54 Holotype. Madagascar • District de Diégo-Suarez, canton d’Anivorano Nord, forêt d’Antenampandrana, km 89,500 de la route d’Ambilobe, calcaires de l’Ankarana, 1 Feb. 1960, G. Cours 5586 P (P00135452!). GenBank information. DNA sequences published by Schaefer et al. (2009): EU436337, EU436361, EU436387, EU436411. Distribution. Endemic to Madagascar, where it is found in the North, Centre, and West (Fig. 55). 156 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 50–55. 50. Kedrostis perrieri, Holotype, Tsitampiky, R. Decary 8157 (P). 51. Distribution of Kedrostis perrieri. 52. Lemurosicyos variegata. A. Habit; B. Female flower; C. Young fruit (photos A by MBA, Ankarana Special Reserve, December 2023, B and C by F. Rakotoarivony, Sakaraha-Mahaboboka, February 2011 (fruit), and March 2013 (flower)). 53. Distribution of Lemurosicyos variegata. 54. Muellerargia jeffreyana. A. Habit; B. Fruit (photos A, B by F. Rakotonasolo, Antananarivo, Itasy, February 2024). 55. Distribution of Muellerargia jeffreyana. 51 52 53 54 50 55 157 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Western Domain: Diana, Montagne des Francais, M. Bardot-Vaucoulon 1648 (MO, P); Ankarana Special Reserve, G. Cours 5586 (P), H. Humbert 32537 (P), M. Bardot-Vaucoulon 1690 (P); Sofia Region, Betainkankana, Ankaizina forest edge, J. Bosser 2626 (P); Melaky Region, Beanka (Tsingy), R. F. Bolliger 165 (P); Atsimo-Andrefana region, Mikoboka, Milenaky, Ambotakebo, Randrianarivony 1029 (MO, TAN); undergrowth of Anjoho, P. Morat 2559 (P). Central Domain: Itasy Region, primary vegetation remnants, trachyte scree, Itasy Lake, J. Bosser 19189 (P). Occurrence inside Protected Areas. Natural Monument of the Sacred Forest of Alandraza Analavelo, Montagne des Français Protected Harmonious Landscape, Harmonious Protected Landscape of Beanka, and Ankarana Special Reserve. Ecology and habitat. Tropical Forest on limestone hills and plateaus, dry, deciduous forest on clay soil on Eocene limestone, in canyons, at forest edges, remnants of primary forest on trachyte scree, c. 100–1,300 m asl.; flowering Feb.–May (Keraudren 1966). Taxonomy. Muellerargia jeffreyana is sister to M. timorensis from Australia and New Guinea (Schaefer et al. 2009; Telford et al. 2011). IUCN. Based on IUCN criterion B1 we suggest classifying the species as Endangered (EN). Peponium Engl., H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam., Nachtr. 1: 318 (1897). Generic type. Peponium mackenii (Naud.) Engl. Worldwide Distribution. Tropical and Southern Africa, Madagascar, and Seychelles (Schaefer 2020). Peponium betsiliense Keraudren, Notul. Syst. (Paris) 16: 148 (1960) Fig. 56 Holotype. Madagascar • Fianarantsoa, environs d’Ambatofitorahana (Betsileo), au km 300 de la route Tananarive-Fianarantsoa, restes de forêt ombrophile sur argiles latéritiques, 1575 m, Mar. 1955, H. Humbert 30210 P (P00135362!). Isotype. P (P00135363!, P00135364!). GenBank information. Has been sequenced by the authors (unpubl. data). Distribution. Endemic to Madagascar where it is mainly found in the Southeast (Fig. 57). Central Domain: Amoron’i Mania region, Ambatofitorahana, J. Bosser 14903 (MO, P), M. Keraudren 269, 1167, 1554 (P), H. Humbert 30210 (P). Western Domain: Diana region, Sahafary forest, between Diego and Anivorano, M. Keraudren 1678 (P); Atsimo-Andrefana region, Zombitsy NP, J. Bosser 19373, 19894, M. Keraudren 457 (P). Occurrence in Protected Areas. Zombitse-Vohibasia NP. Ecology and habitat. Medium altitude moist evergreen and dry deciduous forest on lateritic clay, characterised by high rainfall (>1,200 mm/year), no or brief dry season (< 4 months), and high air humidity (Gautier et al. 2018), flowering Jan.–Apr. (Keraudren 1966). Taxonomy. Unresolved. 158 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae IUCN. Based on IUCN criterion B1 the species is classified as Critically Endangered (CR). Peponium boivinii (Cogn.) Engl., Nat. Pflanzenfam., Nachtr. 1: 318 (1897) Fig. 58 Basionym. Peponia boivinii Cogn. Monogr. Phan. 3: 408 (1881). Holotype. Madagascar • Bords du ruisseau d’Andradroite, Nossi-be; Mar. 1851; L.H. Boivin s.n. P (P00135366!). GenBank information. No DNA sequences available. Distribution. Endemic to Madagascar where it is found only in the extreme North (Fig. 59). Western Domain: Sava Region, Daraina, Solaniampilana-Maroadabo forest, L. Gautier 4519 (G). Sambirano Domain: Diana region, Nosy-Be, on the streamside of Andradroite, L.H. Boivin s.n. (P), Djabalabe, L.H. Boivin s n. (P). Occurrence inside Protected Areas. Loky Manambato Harmonious Protected Landscape. Ecology and habitat. Littoral thicket and forest by the sea on sandy soil up to 100 m asl. Taxonomy. Unresolved. IUCN. Based on IUCN criterion B1 the species is classified as Critically Endangered (CR). Peponium grandidieri Keraudren, Notul. Syst. (Paris) 16: 148 (1960) Fig. 60 Holotype. Madagascar • Mouroundava [Morondava], Mar. 1869, A. Grandidier s.n. P (P00135368!). Syntype. P (P00135369!). GenBank information. Has been sequenced by the authors (unpubl. data). Distribution. Endemic to Madagascar, where it was found mainly in the Southwest (Fig. 61). Western Domain: Boeny region, Ankarafantsika, near Ampijoroa forest station, M. Keraudren 1203 (P); Atsimo-Andrefana region, Sakaraha, Lambomakandro forest, J. Bosser 15757 (P), M. Keraudren 1313 (P); Zombitsy, P. Morat 3799 (P); Tsaramasao 20 km South of Sakaraha, J. Bosser 19906 (MO), P. Morat 3501 (P); undergrowth of Anjohobe, P. Morat 2600 (P); Menabe, Morondava, A. Grandidier s.n. (P). Sambirano Domain: Diana region, Nosy-Komba, J. Bosser 14758 (P), M. Keraudren 1601 (P). Southern Domain: Anosy region, Ankodida New Protected Area, H. Schaefer et al. 23A,B (DBEV, TUM). Occurrence inside Protected Areas. Ankarafantsika NP, Zombitse-Vohibasia NP, and Ankodida Protected Harmonious Landscape. Ecology and habitat. Undergrowth of dry deciduous forests and in remnants of primary forest, c. 200 m asl. Taxonomy. Unresolved. IUCN. We suggest Least Concern (LC) based on IUCN criterion B2. 159 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 56–61. 56. Peponium betsiliense. A. Fruit, longitudinal section; B. Habit; C. Young pubescent fruit; D. Young leaf with inflorescence (photos by MBA, Ambatofitorahana, November 2023). 57. Distribution of Peponium betsiliense. 58. Peponium boivinii, plant with female flower (photo by F. Ratianarivo, Tsaratanana Nature Reserve, May 2023). 59. Distribution of Peponium boivinii. 60. Peponium grandidieri, habit (photo by S. Cable, Ankarafantsika NP, February 2018). 61. Distribution of Peponium grandidieri. 56 57 58 59 60 61 160 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Peponium hirtellum Keraudren, Notul. Syst. (Paris) 16: 147 (1960). Fig. 62 Peponium hirtellum Keraudren var. hirtellum, Notul. Syst. (Paris) 16: 147 (1960) Holotype. Madagascar • Toliara, environs de Fort Dauphin, près de Bevilany, 250 m, 14 Sep. 1928, H. Humbert & C.F. Swingle 5697bis P (P00135370!). Isotypes. P (P00135371!, P00135372!). GenBank information. No DNA sequences available. Peponium hirtellum var. longiracemosum Keraudren Fl. Madagasc. 185: 135 (1966). Holotype. Madagascar • Toliara, Nord-Ouest de Fort Dauphin, Haute Mananara, près du poste forestier d’Imonty (RN) et du village d’Ambatohabo, dans les restes de végétation sèche sur rochers, avec Alluaudia, Pachypodes et Aloe, Feb. 1962, M. Keraudren 1518 P (P00135373!), our Fig. 64. Isotypes. P (P00135374!, P00135375!, P00465122!), MO (MO102597723!). Distribution. Endemic to Madagascar where both varieties of the species have been found only in the Southeast (Figs 63, 65). Southern Domain: Atsimo-Andrefana region, on the road to Manombo in the North of Toliara, P. Morat 3841 (P); Anosy region, on the road from Amboasary to Fort Dauphin, the kilometric point no. 434, J. Bosser 15832 (P), M. Keraudren 1023, 1480 (P), M. Keraudren-Aymonin 25052 (P); near Bevilany, H. Humbert 5697bis (P); Androy region, between Lavanono and Marovato, Croat 31574 (MO). Eastern Domain: Anosy Region, on the mountain between Andohahela and Elakelake, H. Humbert 13845 (P); Upper Manampanihy Valley, between the Saindro pass and Eminiminy, H. Humbert 14024 (P); near Imonty Forestry station, a few kilometers from Antotohabo, M. Keraudren 1518 (MO, P). Occurrence inside Protected Areas. Peponium hirtellum var. hirtellum: Ankodida Harmonious Protected Landscape. Peponium hirtellum var. longiracemosum: Andohahela NP. Ecology and habitat. Peponium hirtellum var. hirtellum grows in dry spiny thicket on sandy soils at the edge of the forest, c. 250 m asl.; flowering Sept.–Feb. (Keraudren 1966). Peponium hirtellum var. longiracemosum grows in montane ericoid thicket with Alluaudia, Pachypodium, and Aloe, in transition of dry spiny thicket and lowland evergreen moist forest, and in medium altitude moist evergreen forest on lateritic gneiss, c. 600–1,200 m asl. Taxonomy. Unresolved. IUCN. Peponium hirtellum var. hirtellum: We suggest Critically Endangered (CR) based on IUCN criterion B1 for a species with a very restricted area of occurrences. Peponium hirtellum var. longiracemosum: We suggest Critically Endangered (CR) based on IUCN criterion B1 for a species with a very restricted area of occurrences. Peponium humbertii Keraudren, Notul. Syst. (Paris) 16: 142 (1960). Holotype. Madagascar • Massif de Marivorahona, au Sud-Ouest de Manambato (Haute Mahavavy du Nord, district d’Ambilobe), vallons supérieurs, 161 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae parties un peu claires, 2,100 m, 1 Mar. 1951, H. Humbert & R. Capuron 25808 P (P00135376!), our Fig. 66. Isotypes. P (P00135377!, P00135378!, P00465144!), MO (MO102332705!). GenBank information. Has been sequenced by the authors (unpubl. data). Distribution. Endemic to Madagascar where it was found mainly in the North (Fig. 67). High Mountain Domain: Diana region, Marivorahona Massif, H. Humbert 25808 (P, MO). Western Domain: Sofia region, Bealanana, northeastern Mangidirano, C. Rakotovao 2361 (P, MO). Central Domain: Andasibe, Mantadia NP, H. Schaefer et al. 11, 21 (TUM, DBEV). Occurrence inside Protected Areas. Natural Resources Reserve of the Marojejy-Anjanaharibe Sud-Tsaratanàna Corridor, northern part. Ecology and habitat. Medium altitude moist evergreen forest, and montane ericoid thicket, climbing on bamboo and Pandanus, on roadsides, lake shores, ridge forests, and clearings on gneiss or sandstone, c. 800–2,200 m asl.; flowering in March (Keraudren 1966). Taxonomy. Unresolved. IUCN. We suggest Critically Endangered (CR) based on IUCN criterion B1 for a species with a very restricted area of occurrences. Peponium laceratum Keraudren, Notul. Syst. (Paris) 16: 146 (1960). Holotype. Madagascar • Alaotra, Bord d’un chemin en forêt d’Antsihanaka, 1 Nov. 1936, Jardin Botanique Tananarive 2254 P (P00135379!), our Fig. 68. GenBank information. No DNA sequences available. Distribution: Endemic to Madagascar, where it is known only from the Alaotra region and Nosy Mangabe (Fig. 69). Central Domain: Alaotra Mangoro region, Alaotra, Antsihanaka forest, Jardin Botanique Tananarive 2254 (P); Ambatosoa region, Maroantsetra, Nosy Mangabe, M. Keraudren-Aymonin 24512 (P). Occurrence inside Protected Areas. Nosy Mangabe NP. Ecology and habitat. Lowland and medium altitude moist evergreen forest; flowering in March. Taxonomy. Unresolved. IUCN. Due to the small number of collections known, we suggest Data Deficient (DD). Peponium perrieri Keraudren, Notul. Syst. (Paris) 16: 144 (1960). Fig. 70 Peponium perrieri Keraudren var. perrieri, Notul. Syst. (Paris) 16: 144 (1960) Holotype. Madagascar • Ouest: haut Bemarivo, sables humides et boisées au bord des rivières, 200 m, Sep. 1907, H. Perrier de la Bâthie 6770 P (P00135380!). Isotype. P (P00135381!, P00135382!, P00135383!, P00135384!, P00465534!). Paratype. Madagascar • Antananarivo, Manerinerina, Centre: bois des pentes occidentales, à Manerinerina, sur le Tampoketsa, entre l’Ikopa et la Betsiboka, 1,500 m, 1 Dec. 1924, H. Perrier de la Bâthie 16750 P (P00135386!). 162 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 62–67. 62. Peponium hirtellum var. hirtellum, Holotype, Bevilany, H. Humbert & C.F. Swingle 5697bis (P). 63. Distribution of Peponium hirtellum var. hirtellum. 64. Peponium hirtellum var. longiracemosum, Holotype, Ambatohabo, M. Keraudren 1518 (P). 65. Distribution of Peponium hirtellum var. longiracemosum. 66. Peponium humbertii, Holotype, Manambato, H. Humbert & R. Capuron 25808 (P). 67. Distribution of Peponium humbertii. 62 63 64 65 66 67 163 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Vernacular name. Takotakokely (Merina). GenBank information. No DNA sequences available. Peponium perrieri var. glabrescens Keraudren Holotype. Madagascar • Centre: bois des pentes occidentales, Tampoketsa d’Antongodrahoza, entre la Betsiboka et la Mahazamba, 1,300 m, 1 Sep. 1922, H. Perrier de la Bâthie 14841 P (P00135387!), our Fig. 71. Isotype. P (P00135388!). Distribution. Endemic to Madagascar, where it is found mainly in the Centre and West (Figs 72, 73). Peponium perrieri var. perrieri Western Domain: Diana region, 45 km southern Diégo, Sahafary forest, M. Keraudren 1678 (P). Betsiboka region, between Betsiboka and Mahajamba, Antongondrahoza, H. Perrier de la Bâthie 14841 (P). Central Domain: Bongolava region, Belobaka, J. Bosser 17661 (P); Analamanga, Ankazobe, J. Bosser 15976 (P); Manerinerina, H. Perrier de la Bâthie 16750 (P); Alaotra-Mangoro region, Andasibe, Analamazaotra forest, M. Keraudren-Aymonin 25373 bis (P); Andranobe forest, road from Ambatondrazaka to Andriamena, J. Bosser 19838 (P). Peponium perrieri var. glabrescens Western Domain: Sofia, region, Upper Bemarivo, riverbank, H. Perrier de la Bâthie 6770 (P). Occurrence inside Protected Areas. Not known from protected areas. Ecology and habitat. Peponium perrieri var. perrieri has been found in medium altitude moist evergreen forest, on riverbanks and at the forest edge, c. 200–1,600 m asl.; flowering Sept.–Mar. (Keraudren 1966). Peponium perrieri var. glabrescens has been collected in medium altitude moist evergreen forest on a riverbank; flowering in July (Keraudren 1966). Taxonomy. Unresolved. IUCN. Peponium perrieri var. perrieri: Critically Endangered (CR), based on IUCN criterion B1 for a species with a very restricted area of occurrences. Peponium perrieri var. glabrescens: Since there is only a single collection known, we suggest Data Deficient (DD). Peponium poissonii Keraudren, Notul. Syst. (Paris) 16: 146 (1960) Fig. 74 Holotype. Madagascar • Toliara, Ambovombe, forêt claire, 3 Feb. 1931, R. Decary 8473 P (P00135389!). Isotypes. P (P00135390!, P00135391!). GenBank information. No DNA sequences available. Distribution. Endemic to Madagascar where it has been found only in the South and Southwest (Fig. 75). Western Domain: Atsimo-Andrefana region, Toliara, H. Poisson 387 (P); National Road from Isalo-Sakaraha, Zombitsy, L. Allorge 2098 (P). Southern Domain: Anosy region, Taolagnaro Prefecture, Fort-Dauphin, Magnambaro Township, Petriky Forest, R. Gereau 3346 (US); Androy region, Ambovombe, clear forest, R. Decary 8473 (P). 170 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 81–86. 81. Raphidiocystis brachypoda. A. Female flower; B. Mature fruit; C. Habit; D. Dehiscent fruit with seeds in red pulp (photos by MBA, Ranomafana NP, October 2022 (A–C), and Andasibe NP, December 2021 (D)). 82. Distribution of Raphidiocystis brachypoda. 83. Seyrigia bosseri. A. Mature and immature fruit; B. Shoot with ripe fruit; C. Male inflorescence (photos by MBA, Andohahela NP, October 2022). 84. Distribution of Seyrigia bosseri. 85. Seyrigia gracilis. A. Fruit with seeds; B. Female flower; C. Fruit; D. Habit (photos by MBA, Tranomaro New Protected Area, December 2024). 86. Distribution of Seyrigia gracilis. 81 82 83 84 85 86 171 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Isotype. P (P00135503!). GenBank information. DNA sequences published by Ayre et al. (2003) and Zhang et al. (2006): AY379781, AY968421, AY968443, AY968457, AY968510, AY968526. Distribution. Endemic to Madagascar, where it is known only from the Southwest (Fig. 88). Southern Domain: Atsimo-Andrefana region, Ankaloha-IOT, Ramon 484 (MO); Belalanda, Ramon 583 (MO); Gorges of the Fiherenana river, H. Humbert 19947 (P); Fiherenana valley, W. Rauh 7377 (P), Humbert 5148 (P), J. Bosser 13569 (P), M. Keraudren 766 (P); Miary, M. Keraudren 640, 671 (P). Occurrence inside Protected Areas. Ranobe New Protected Area. Ecology and habitat. Dry spiny thicket on limestone rocks, from sea level up to c. 200 m asl. Taxonomy. Seyrigia humbertii forms a clade with Trochomeriopsis diversifolia (Kocyan et al. 2007). IUCN. We suggest the category Endangered (EN) based on IUCN criterion B2 for a species with a very restricted distribution. Seyrigia marnieri Keraudren, Bull. Soc. Bot. France 114: 446 (1968). Holotype. Madagascar • Saint Jean Cap Ferrat, jardin botanique “Les Cèdres”, M. Keraudren 5 P (P00135504!). GenBank information. Has been sequenced by the authors (unpubl. data). Pictures. colour photographs in Rauh (1995, p. 195, Figs 698–696, 701–702). Distribution. Endemic to Madagascar, only known from the Southwest (Fig. 89). Southern Domain: Atsimo-Andrefana region, on limestone near Betioky (Rauh 1995); bord du Lac Hanomay, près de lklioky, fourrés xérophiles sur calcaires, Bosser 191816 (P); 20 km de Sakaraha, route de Ankazoabo, J. Bosser 19971 (P). Occurrence inside Protected Areas. Not known from protected areas. Ecology and habitat. Dry spiny thicket on limestone. Taxonomy. Unresolved. IUCN. We suggest the category Data Deficient (DD) since the species is poorly known and has not been seen for decades. It might have become extinct. Seyrigia multiflora Keraudren, Bull. Soc. Bot. France 107: 299 (1961) Fig. 90 Holotype. Madagascar • Bas mandrare, piste d’Amboasary a Ambatomika, dans bush dégradé le long du terrain d’aviation d’Ambatomika, 1 Apr. 1960, M. Keraudren 1083 P (P00135506!). Isotypes. K (K000242621!), P (P00135507!, P00135508!). GenBank information. Has been sequenced by the authors (unpubl. data). Distribution. Endemic to Madagascar, where it is only found in the South (Fig. 91). Southern Domain: Atsimo-Andrefana region, Beza Mahafaly Reserve, P.B. Phillipson 1651 (P); Around Tsimanampetsotsa lake, J. Léandri 3995 (P); from Ampanihy to Ejeda, L. Allorge 2319 (P); Bevoalava, M. Keraudren 1467 (P); from Ampanihy to Ampotaka, M. Keraudren 917 (P); from Androka to Ampanihy, M. Keraudren 897 (P), J. Bosser 13560 (P); Anadabolava, M. Keraudren 1547 (P), 172 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae H. Humbert 12553 (P); Betioky, H. Humbert 29436 (P); Sakoa, R. Decary 15953 (P); Anosy region, Ambatomika, M. Keraudren 1083 (K, P); Isomony, H. Humbert 12978 (P); Vohipolaka, H. Humbert 11619 (P); Tranomaro New Protected Area, H. Schaefer et al. 17 (DBEV, TUM), Tanambao Morafeno, H. Schaefer et al. 2A,B, 8, 12A,B (DBEV, TUM); Androy Region, Sihanadahy, from Beloha to Ambovombe, M. Keraudren 941 (P). Central Domain: Androy region, East ridge, between Bekily to Tsivory, rocks, half-shade, A. Seyrig 344B (P). Occurrence inside Protected Areas. Tranomaro New Protected Area, Andohahela NP, Tsimanampesotse NP, and Bezà-Mahafaly Special Reserve. Ecology and habitat. Dry spiny thicket and dry deciduous and transitional forest with Euphorbia and Adansonia fony in sandy, calcareous, and rocky soils with gneiss, c. 50–1,100 m asl.; flowering Oct.-Mar. (Keraudren 1966). Taxonomy. Unresolved. IUCN. We suggest the category Least Concern (LC) based on IUCN criterion B2 for a species with a wide distribution. Seyrigia napifera Rauh, Succ. & Xeroph. Pl. Madagascar 2: 193 (1998). Holotype. Madagascar • 60 km west of Tolanaro, Sept. 1987, Rauh 68582 (HEID!). GenBank information. No DNA sequences are available for this taxon. Pictures. colour photographs in Rauh (1995, p. 195, figs 703-705). Distribution. Endemic of Madagascar where it is known only from the Southeast (Fig. 92). Southern Domain: near Tolagnaro (Fort Dauphin) in the Southeast, Rauh 68582 (HEID). Central Domain: Anosy region, Integral Natural Reserve, plot no. 2, on the track to Ambatohabo, bush xerophytic, M. Randriambololona et al. 82 (BR, WAG, P). Occurrence inside Protected Areas. Andohahela NP. Ecology and habitat. In Alluaudia bush on sandy soil. Taxonomy. Perhaps closely related to S. humbertii with which it shares densely white hairy shoots (Rauh 1995). IUCN. Critically Endangered (CR) according to Rauh (1995). Trochomeriopsis Cogn, Monogr. Phan. 3: 661. 1881. Generic type. Trochomeriopsis diversifolia Cogn. Worldwide Distribution. Madagascar endemic (Schaefer 2020). Trochomeriopsis diversifolia Cogn., Monogr. Phan. 3: 661 (1881) Fig. 93 Vernacular name. Voatangonala (South). Holotype. Madagascar • Port Leven, 1849, L.H. Boivin 2573 P (P00135491!). GenBank information. DNA sequences published by Kocyan et al. (2007): DQ535859, DQ536607, DQ536746, DQ536878. Distribution. Endemic to Madagascar where it is widespread in the lower regions of the West and South (Fig. 94). 173 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 87–92. 87. Seyrigia humbertii, shoot covered in whitish pubescence (photo by HS, cultivated in Freising, Germany, September 2024). 88. Distribution of Seyrigia humbertii. 89. Distribution of Seyrigia marnieri. 90. Seyrigia multiflora, shoot with male inflorescence (photo by MBA, Andohahela NP, October 2022). 91. Distribution of Seyrigia multiflora. 92. Distribution of Seyrigia napifera. 87 88 89 90 91 92 174 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Western Domain: Diana region, Vovo village, D. K. Harder et al. 1699 (P); Montagne des Français, A.M. Homolle 319 (P), M. Bardot-Vaucoulon 1627 (P), H. Perrier de la Bâthie 16199 (P), H. Schaefer & M.B. Andriamiharisoa 3 (TUM, DBEV); Mangakoa, G. Aymonin 25614 (P); Ramena, J.B. Leopold 14 (P); Analamera, H. Humbert 19120 (P); Ambodivahibe, Ampasindava, Ratovoson 985 (CNARP, TAN, MO, P), R. Ramananjanahary 310 (P); Ankarana Special Reserve, C.C.H. Jongkind & S. Rapanarivo 980 (BR, P, WAG, US), H. Humbert 19075 bis (P), M. Bardot Vaucoulon 45, 444, 474, 714 (P), H.Schaefer & M.B. Andriamiharisoa 10, 14,14B (TUM, DBEV), L. Allorge 2758, 2759 (P); Antsisikala, J.B. Leopold 80 (P); Orangea, G. Aymonin 25485, 25474 (P); Beaomby, J. Bosser 20190 (P); Irodo, C. Hongwa 243243 (P); Sava Region, Port Leven, J. Vesco s.n. (P), L.H. Boivin 2573 (P). Sambirano Domain: Diana region, Integral Natural Reserve (RNI) no. 6 Lokobe, C.R. Birkinshaw 181 (P); Nosy-Be, L.H. Boivin s. n. (P); Boeny Region, near the sea, M. Keraudren 1710 (K, P); Ampijoroa, M. Keraudren 1250, 1251 (P); Ankarafantsika NP, H. Schaefer et al. 34, 44 (TUM, DBEV), H. Perrier de la Bâthie 6736, 13835, 14691, 15869 (P), A.H. Gentry 62119 (P); Maevarano, H. Perrier de la Bâthie 12101 (P); around Marovoay, H. Perrier de la Bâthie 17961 (P); Betsiboka region, Antsiafabositra, N.M. Andrianjafy 1051 (P); Mojeramanga, H. Perrier de la Bâthie 6732 (P); Vilanandro, H. Perrier de la Bâthie 6735 (P); Sofia region, Mampikony, R. Decary 14427 (P). INaturalist observations: https://www.inaturalist.org/observations/8292193 and https://www.inaturalist.org/observations/66955435. Central Domain: Ramena, Ankorikakely, F. Ratovoson 981 (P). Southern Domain: Atsimo-Andrafana region, along Onilahy River, M. Keraudren 727 (P); Anadabolava, M. Keraudren 1085, 1535 (P); Anakao, L. Allorge 2277 (P); Andoharano, P. Montagnac 47 (P); Bevovo, J. Léandri 4040 (P); surroundings of Toliara, M. Keraudren 546 (P); Eastern Mahabo, G. Aymonin 25563, 25563bis (P); Fiherenana, M. Keraudren 767 (P); Miary, M. Keraudren 639 (BR, P); Itambono Corridor, P.B. Phillipson & S. Rabesihanaka 3137 (BR, MO, P, US, WAG); Lambomakandro, M. Keraudren 1310, 1317 (P); Tsefanoka, A. Randrianasolo & R. Ranaivojaona R. 1465 (BR, P); Ankilimalinike, P.B. Phillipson 5946 (MO, P); Mahafaly Plateau, M. Keraudren 874 (P); Belalanda, Ranobe, M. Andrianjafy 1831 (P); Eastern Andrevo, R. Ranaivojaona et al. 1447 (BR, P); Tongobory, T.B. Croat 31186 (P); Tsivonoa N., M. Peltier 1373 (P), J. Bosser 10126 (P); Onilahy Valley, J. Bosser 13566 (P); Fiherenana Valley, J. Bosser 13579 (P); Androy region, Ampandrandava, A. Seyrig 712 B (P); Betioky, H. Humbert 29428 (P); Masiaboay, Beara, R. Randrianaivo 1025 (P); between Tranoroa and Beloha, J. Bosser 13576 (P); southern Tranoroa, M. Keraudren 924 (P); Ejeda, Service Forestier Madagascar 373 r16 (P); Itampolo, M. Keraudren 876 (P); 218 km to Toliara, L. Allorge 2167 (P); Basibasy, Ihotry, H.N. Manjakahery & J. Razanantsoa 193 (P); Anosy Region, Ampihamy, H. Humbert 11498 (P); Ifotaka, J. Bosser 13582 (P); Lower stream Mandrare River, M. Keraudren 1007 (P); Tranomaro New Protected Area, H. Schaefer et al. 3A,B (DBEV, TUM). Menabe region, Ankilizato, G. Aymonin 25849 (P); from Morondava to Belo, G. Aymonin 25884 (P). iNaturalist observations: https://www.inaturalist.org/observations/12948635, https://www.inaturalist.org/observations/14070295, https:// www.inaturalist.org/observations/70548360. 175 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Occurrence inside Protected Areas. Ambohitr’Antsingy-Montagne des Français Harmonious Protected Landscape, Amoron’i Onilahy Harmonious Protected Landscape, Analamerana Special Reserve, Andrafiamena Andavakoera Harmonious Protected Landscape, Ankarafantsika NP, Ankarana Special Reserve, Mangoky Ihotry Wetlands Complex Harmonious Protected Landscape, Menabe Antimena Harmonious Protected Landscape, Nord-Ifotaka Harmonious Protected Landscape, and Ranobe PK32 New Protected Area. Ecology and habitat. Open dry deciduous forest, dry spiny thickets, moist semi-deciduous, gallery forest, woodland and dunes by the sea on red, white, or brown sands, limestone, alluvial deposits or clay soil and outcropping boulders, from sea level up to c. 700 m asl., flowering Nov.-Jan. Uses. Not known, but leaves eaten by goats in the dry South. Taxonomy. According to molecular data, T. diversifolia forms a clade with Seyrigia humbertii (Kocyan et al. 2007). IUCN. We suggest the category Least Concern (LC) based on IUCN criterion B2 for a species with a wide distribution. Xerosicyos Humbert, Compt. Rend. Hebd. Séances Acad. Sci. 208: 220. 1939. Generic type. Xerosicyos danguyi Humbert. Worldwide Distribution. Madagascar endemic (Schaefer 2020). Xerosicyos danguyi Humbert, Compt. Rend. Hebd. Séances Acad. Sci. 208: 221 (1939) Fig. 95 Holotype. Madagascar • Toliara, Belalanda, environs de Tuléar-Delta du Fiherenana, lieux sablonneux, dunes, également sur les plateaux calcaires rocailleux, 6 m, 14 Sep. 1924, H. Humbert & H. Perrier de la Bâthie 2440 P (P00135509!). Isotypes. K (K000272553!, K000272554!) GenBank information. DNA sequences published e.g., by Zhang et al. (2006): AJ235648, AY968388, AY968423, AY968459, AY968479, AY968512. Distribution. Endemic to Madagascar where it is mostly found in the South (Fig. 96). Western Domain: Ihorombe region, along the road to Ivohibe, L. Bernardi 11200 (P); Ihosy, M. Keraudren 335 (P); Analavoky, J. Léandri 345 (BR), 3443 (P), 3484 (P). Eastern Domain: Anosy region, Ankodida New Protected Area, Mahavelo Forest, H. Schaefer et al. 25A,B (DBEV, TUM). Southern Domain: Atsimo-Andrefana region, Toliara, P. Morat 1476 (P), A.M. Homolle 1587 (P), R. Decary 16159 (P), AC d’Alleizette s.n. (L); Onilahy river, H. Perrier de la Bâthie 12753 (P); Natural Reserve of Tsimanampetsotsa, H. Humbert 5289 (P, US), M. Keraudren 1426 (P); Belalanda, H. Humbert 2420 (K, P), H. Manjakahery 111 (MO, P, TAN); Fiherenana Valley, H. Humbert 5156 ter (P), J. Léandri 3696 (P); Belemboka, J. Dequaire 27341 (P); Songeritelo, B. Koechlin s.n. (P); Bevato, B. Koechlin s.n. (P); Mangoky, H. Perrier de la Bâthie 4381 (P); Morombe, Ihosy, J. & M. Peltier 2662 (P); Betioky, near Analafaly, L. Sussman 331 (P), Rapanarivo 214 (MO, BR, P, US, WAG); Ambatry, M. Peltier 2554 (P); between Anakao and Betioky, R. Decary 16084 (P); Tongobory, J. Léandri 3740 176 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae (P); Mahaleotse, S.E. Rakotoarisoa 303 (P); 40 km from Toliara, M. Keraudren 545 (P); La Table, M. Keraudren 568 (P); Andranovory, M. Peltier 2519 (P); on the roadside to Ifaty, P. Morat 7897 (P); Ambohimahavelona, Rabarivola M. L. 594 (P); on the road to Sarodrano, F. Chauvet 345 (BR, P), G. Grandidier s.n. (P), M. Keraudren-Aymonin 24704 (P); St. Augustin Township, G. Cours 3144 (P); Ampanihy, H. Humbert & C. Swingle 5551 (K, P, US); from Ampanihy to Beloha, M. Keraudren 926 (P); Itampolo, J. Léandri 4104 (P), P.B. Phillipson et al. 3731 (P); Anosiary, limestone plateau, J. Peltier 2492 (P); Ambovombe, J. Bosser 10616 (P); Imanombo-Androy, M. Decorse s.n. (P); Ambatomika, M. Keraudren 1082 (P), Ambatomika, B. Koechlin s.n. (P); Behara, Réserves Naturelles Madagascar-Rakotoson 6608 RN (P); Tanandava, M. Keraudren 734 (P); Tranomaro, H. Schaefer et al. 14, 15 (DBEV, TUM); Anosy region, Andohahela NP, H. Schaefer et al. 68 (DBEV, TUM), M.B.A. Randriambololona 82 (WAG); Andehomana, H. Schaefer et al. 76 (DBEV, TUM); between Ambovombe to Tsihombe, M. Keraudren 993 (P); Northern Ambovombe on the road to Antanimora, M. Keraudren-Aymonin 25058 (P), R. Decary 2731 (P); Ambararata, R. Decary 2988, 2988bis (P); Behara, R. Decary 3234 (P); Antanimora, R. Decary 9200 (P); road to Ifotaka, M. Keraudren 1473 (P), Fort Dauphin, Vinanibe dry forest, N. Dumetz 1301 (P); Fort Dauphin, Antanimora, R. Decary 4627 (P); Andavaka, Andrahomana, C. Alluaud 12 (P). iNaturalist observations: https://www.inaturalist.org/observations/8869856; https://www.inaturalist.org/observations/1968017; https://www.inaturalist.org/observations/2742739; https://www.inaturalist.org/observations/12949398; https://www.inaturalist.org/observations/64028998; https:// www.inaturalist.org/observations/105662184; https://www.inaturalist.org/observations/72461191; https://www.inaturalist.org/observations/136852990; https://www.inaturalist.org/observations/141155576; https://www.inaturalist.org/observations/142572038; https://www.inaturalist.org/observations/143142452; https://www.inaturalist.org/observations/143557311; https://www.inaturalist.org/observations/102665883; https://www.inaturalist.org/observations/105481841; https://www.inaturalist.org/observations/30103565; https://www.inaturalist.org/observations/74353724; https:// www.inaturalist.org/observations/11069140; https://www.inaturalist.org/observations/66240542; https://www.inaturalist.org/observations/36689701. Occurrence inside Protected Areas. Amoron’i Onilahy Harmonious Protected Landscape, Ankodida Harmonious Protected Landscape, Andohahela NP, Bezà-Mahafaly Special Reserve, Cap Sainte Marie Special Reserve, Ranobe PK32 New Protected Area, Tsimanampesotse NP, and Tsinjoriake Harmonious Protected Landscape, Ankodida Harmonious Protected Landscape, and Tranomaro New Protected Area. Ecology and habitat. Dry deciduous, transitional forest and dry spiny forest, where it grows with Euphorbia, Grewei grevei, Pachypodium geayi, and Fabaceae on rocky outcrops, limestone plateaus and hillsides from sea level up to 800 m asl. Uses. Widely grown as an ornamental plant (“silver dollar plant”) and easily reproduced from cuttings. Taxonomy. According to Schaefer et al. (2009) X. danguyi is sister to X. perrieri. IUCN. We suggest the category Least Concern (LC) based on IUCN criterion B2 for a species with a wide distribution. 177 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Xerosicyos decaryi Guillaumin & Keraudren, Notul. Syst. (Paris) 16: 127 (1960) Fig. 97 Holotype. Madagascar • Mahajanga, Namoroka, District de Soalala, Namoroka, reserve n°8, 17 Sep. 1940, R. Decary 15791 P (P00135510!). Isotype. P (P00135511!). GenBank information. DNA sequence published by Schaefer and Renner (2011): HQ202007. Distribution. Endemic to Madagascar where it is restricted to the South and Southwest (Fig. 98). Western Domain: Melaky region, Bemaraha Tsingy, H. Perrier de la Bâthie 2228 (P); Bekopaka, P. Morat 81, 777 (P); Boeny Region, Namoroka, R. Decary 15791 (P); Atsimo-Andrefana region, Zombitsy, M. Keraudren 460 (P), L. Allorge 2109 (P), H. Schaefer et al. 79, 93 (DBEV, TUM); Ihorombe region, Ihosy District, from Ambalavao to Ihosy, M. Keraudren 1091 (P). Southern Domain: Atsimo-Andrefana region, Toliara, M. Keraudren 526 (P); Anjamala, M. Keraudren 760, 1353 (P); Beza Mahafaly Reserve, P.B. Phillipson 2531, 3686 (P); Androy region, Behompy, Mihary, Behompy, G. Cours 3140 (P); Anosy region, Bekiria, M. Keraudren 1021 (P); Andohahela NP, H. Schaefer et al. 65 (DBEV, TUM). iNaturalist observation: https://www.inaturalist.org/observations/38956168. Occurrence inside Protected Areas. Andohahela NP, Namoroka NP, Nord-Ifotaka Harmonious Protected Landscape, Ranobe PK32 New Protected Area, Tsimanampesotse NP, and Zombitse-Vohibasia NP. Ecology and habitat. Dry deciduous forest on both sandy soil and rocky woodland on Jurassic limestone from sea level up to c. 800 m asl. Taxonomy. Unresolved. IUCN. We suggest the category Least Concern (LC) based on IUCN criterion B2 for a species with a wide distribution. Xerosicyos hirtellus (Humbert) H. Schaef. & S.S. Renner, Taxon 60: 135 (2011) Fig. 99 Basionym. Zygosicyos hirtellus Humbert, Bull. Soc. Bot. France 111: 170 (1945). Holotype. Madagascar • Anadabolava, Vallée moyenne du Mandrare près d’Anadabolava, forêt sèche; 225 m, 1 Dec. 1933, H. Humbert 12439bis P (P00135518!). Isotype. P (P00135518!). GenBank information. Has been sequenced by the authors (unpubl. data). Distribution. Endemic to Madagascar where it is known only from the type material from the Mandrare valley (Fig. 100). Southern Domain: Androy region, Anadabolava, Middle Mandrare Valley near Anadabolava, dry forest, H. Humbert 12439bis (P). Occurrence inside Protected Areas. Not known from protected areas. Ecology and habitat. Dry deciduous forest, c. 200 m asl. Taxonomy. Morphological very close to X. tripartitus and perhaps just a variety of that species. IUCN. We suggest Critically Endangered (CR) based on criterion B1. The species is likely to face threats from illegal trade. 178 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 93–98. 93. Trochomeriopsis diversifolia, habit (photo by MBA, Ankarana Special Reserve, December 2023). 94. Distribution of Trochomeriopsis diversifolia. 95. Xerosicyos danguyi. A. Flower; B. Habit (photos by MBA, Andohahela NP, October 2022). 96. Distribution of Xerosicyos danguyi. 97. Xerosicyos decaryi, habit (photo by MBA, Zombitsy-Vohibasia NP, October 2022). 98. Distribution of Xerosicyos decaryi. 93 94 95 96 97 98 179 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Xerosicyos perrieri Humbert, Compt. Rend. Hebd. Séances Acad. Sci. 208: 221 (1939) Fig. 101 Holotype. Madagascar • Toliara, Ampanihy, Sud-Ouest: Environs d’Ampanihy, Bush à Euphorbes et Didierea, 1 Jul. 1932, H. Perrier de la Bâthie 18659 P (P00135512!). Isotypes. P (P00135514!), K (K000272576!). Syntypes. Madagascar • ♀, Toliara, Benenitra, Ouest: Environs de Benenitra (Onilahy), Broussailles à xérophytes, 600 m, 1 Jul. 1919, H. Perrier de la Bâthie 2707 P (P00135516!); Mahajanga, Ouest: bois sablonneux près de Maintirano, H. Perrier de la Bâthie 1994 P (P00135517!). GenBank information. DNA sequences of the species have been published by Schaefer et al. (2009): EU436344, EU436372, and EU436421. Distribution: Endemic to Madagascar, where it is restricted to the South (Fig. 102). Western Domain: Ihorombe region, between Ihosy and Ivohibe, Tsimirimbo, J. Léandri 3467 (P); Ihosy, B. Descoings 3664 (P); Zazafotsy, M. Keraudren 319, 1264 (P); Melaky region, near Maintirano, H. Perrier de la Bâthie 1994 (P). Southern Domain: Atsimo-Andrefana region, 40 km from Toliara, M. Keraudren 518 (P); Ampanihy, H. Perrier de la Bâthie 4380, 18659 (P, K); Bealoka, L. Allorge 691, 692 (P); Benenitra, H. Perrier de la Bâthie 12707 (P); Beroroha, H. Humbert 11286 (P); Beza Mahafaly Reserve, P.B. Phillipson 2531, 3686 (BR, WAG), M. Keraudren 899 (P); Sakoa Valley, R. Decary 15947 (P); Fiherenana, Miary, M. Keraudren 649 (P), B. Descoings 1034 (P); Zombitsy, J. Léandri 3624 (P), H. Schaefer et al. 94, 95 (DBEV, TUM); Sakaraha, H. Schaefer et al. 78 (DBEV, TUM); Androy region, Berenty C.M. Hladik 1255 (P), P.B. Phillipson 2316 (BR, WAG), M. Keraudren-Aymonin & G.G. Aymonin 24837, 24838 (BR, P); Ambovombe, J. Bosser 10384 (P), R. Decary 2890 (P), M. Keraudren 972 (BR, P); Ampandrandava, A. Seyrig 356 (P); Antanimora, R. Decary 4613 (K); Anosy region, Fort Dauphin, R. Decary 4266 (P); Imangiry, R. Decary 8938 (P); Imanombo, M. Decorse s.n. (P); Tranomaro New Protected Area, H. Schaefer et al. 10, 16A,B (DBEV, TUM); Amboasary, X. Aubriot 56 (P); Bekiria, Ifotaka, M. Keraudren 1020 (P); between Amboasary and Fort-Dauphin, M. Keraudren-Aymonin 24859 (P); Imonty, J. Léandri 4201, 4564 (P); Menabe Region, Mahabo, J. Dequaire 27145 (P), J. Bosser s.n. (P). iNaturalist observations: https://www.inaturalist.org/observations/1339915, https://www.inaturalist.org/observations/12949026. Occurrence inside Protected Areas. Mangoky Ihotry Wetlands Complex Harmonious Protected Landscape, Nord-Ifotaka Harmonious Protected Landscape, and Zombitse-Vohibasia NP. Ecology and habitat. Moist semi-deciduous and dry deciduous forest, and dry spiny thicket, and coppice associated with Euphorbia, Alluaudia procera, and Baudouinia rouxevillei on sand and sandstone, gneissic rocks, and alluvial deposits from sea level up to c. 800 m asl.; flowering in July (Keraudren 1966). Taxonomy. According to Schaefer et al. (2009), X. perrieri is sister to X. danguyi. IUCN. We suggest the category Least Concern (LC) based on IUCN criterion B2. 186 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Western Domain: Ihorombe region, Sakamalio Valley, rocks, gneiss, xerophilous bush, H. Humbert 13312 (P). Southern Domain: Androy region, Ampandrandava, between Bekily and Tsivory, A. Seyrig 260, 260b (P). Occurrence inside Protected Areas. Not known from protected areas. Ecology and habitat. Dry spiny thicket in riverbed, on sands and rocks, c. 650 m asl., flowering throughout the year (Keraudren 1966), fruits Feb.-Mar. Taxonomy. Unresolved. IUCN. Due to the lack of spatial occurrence data, we suggest Data Deficient (DD). Zehneria peneyana (Naudin) Schweinf. & Asch., Beitr. Fl. Aethiop.: 268 (1867) Fig. 115 Basionym. Pilogyne peneyana Naudin, Ann. Sci. Nat., Bot. sér. 5, 5: 38 (1866). Synonyms. Melothria peneyana (Naudin) Cogn., Monogr. Phan. 3: 592 (1881). Holotype. sudan • Nibblanc, 29 Dec. 1840, Peney s.n. (P, BR0000008889089!). GenBank information. DNA sequence published by Ranjan et al. (2022): MZ409453. Distribution. Zehneria peneyana is native to Sudan, South-Eastern Kenya to North-Eastern Tanzania. In Madagascar, it has been recorded in the West (Fig. 116). Western Boeny region, West Bekalila, Marovoay District, on rice field slope, J. Bosser 8416 (P); Sofia region, Trabonjy, J.M. Hildebrandt 3448 (P); Menabe Region, Mahabo farm, marsh edge grass, J. Dequaire 27166 (P). Occurrence inside Protected Areas. Not known from protected areas. Ecology and habitat. Rice paddies and marsh edges, c. 60–200 m asl.; flowering Sept.–May, fruits in Sept. (Keraudren 1966). Taxonomy. According to Ranjan et al. (2022), Z. peneyana is phylogenetically close to Z. pallidinervia. IUCN. Due to the lack of spatial occurrence data, we suggest Data Deficient (DD). Zehneria perrieri Keraudren, Adansonia, n.s., 4: 337 (1964) Fig. 117 Holotype. Madagascar • Analamazaotra, H.Perrier de la Bâthie 6789 P (P00135441!). Holotype of Zehneria perrieri var. parvula Keraudren: Madagascar • Ifandana, Farafangana, R. Decary 5167 P (P00135442!). Holotype of Zehneria perrieri var. tsaratananensis Keraudren: Madagascar • Mont Tsaratanana, altitude 1,700–2,000 m, 1923, H. Perrier de la Bâthie 15494 P (P00135445). GenBank information. DNA sequences published by Dwivedi et al. (2018): KY523270, KY523339, KY523378, KY523444. Distribution. Endemic to Madagascar where Z. perrieri var. perrieri is present in the Central, and Western Domain (Fig. 118). Central Domain: Diana region, Montagne d’Ambre, rainforest, J. Bosser 20338 (P), S.M. Trigui 277 (P), H. Schaefer & M.B. Andriamiharisoa 41, 41B (DBEV, TUM); Analamanga region, Angavokely, J. & M. Peltier 1903 (P); M. Keraudren 25178, 25223 (P); 30 km East of Antananarivo, Service Forestier Madagascar 96 187 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 111–116. 111. Zehneria madagascariensis, Holotype, Antsirabe, H. Humbert 7149 (P). 112. Distribution of Zehneria madagascariensis. 113. Zehneria martinez-crovettoi, Holotype, Ampandrandava, A. Seyrig 260 (P). 114. Distribution of Zehneria martinez-crovettoi. 115. Zehneria peneyana, Trabonjy, J.M. Hildebrandt 3448 (P). 116. Distribution of Zehneria peneyana. 111 112 113 114 115 116 188 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae SF (P); around Antananarivo, M. Keraudren 30 (P); Ravine cut by the new road, J. Léandri 3243 (P), J. Bosser 13624, 13625, 15302 (P); Tsinjoarivo, Onive waterfall, M. Keraudren 1560 (P); Amoron’i Mania region, Ambatofitorahana, M. Keraudren 246 (P); Ambatomboay, C. Rakotovao 9355 (P); Haute Matsiatra region, Mahazony, Ambalavao, C. Rakotovao 9191bis (P); Tsitondroina township, Herb. Jard. Bot. Tananarive 4802 (P); Vatovavy region, Ranomafana NP, H. Schaefer et al. 113, 120, 122, 124 (DBEV, TUM); Ihorombe region, Begogo, N.M. Andrianjafy 766 (P); Ivohibe, G. Cremers 1515 (P); Alaotra-Mangoro region, Forest northern Sihanaka, Herb. Jard. Bot. Tananarive 2936 (P); Mangoro, H. Perrier de la Bâthie 18072 (P); Analamazaotra, H. Perrier de la Bâthie 6789 (P), M. Keraudren 1746 (P); Anoribe, J. Bosser 16462 (P); Ambatosoratra, C. Rakotovao 11028 (P). Western Domain: Anosy region, Andohahela Massif (South-East), rainforest on the gneissic ridge, H. Humbert 13620 (P); humid forest, Andohahela NP, H. Schaefer et al. 53, 55 (DBEV, TUM). Zehneria perrieri var. parvula is only known from the type collected at Farafangana: Eastern Domain, Atsimo-Antsinana region, Farafangana, Ifandana, forest edge, R. Decary 5167 (P). Zehneria perrieri var. tsaratananensis was described from the High Mountain Domain, Diana region, in herbaceous undergrowth of Mount Tsaratanana, c. 2,000 m asl., H. Perrier de la Bâthie 15494, 16421 (P). Occurrence inside Protected Areas. Zehneria perrieri var. perrieri: Montagne d’Ambre NP, Ranomafana NP, Andohahela NP. – Zehneria perrieri var. parvula: Not known from protected areas. – Zehneria perrieri var. tsaratananensis: Tsaratanana Integral Natural Reserve. Ecology and habitat. Zehneria perrieri var. perrieri grows in undergrowth of medium altitude moist evergreen forest, near waterfalls, in forest remnants, and primary vegetation, preferably on gneiss, c. 1,000–1,800 m asl. Flowering throughout the year (Keraudren 1966), fruits found in May. Zehneria perrieri var. parvula was found at a forest edge; fruits in Sept. (Keraudren 1966). Zehneria perrieri var. tsaratananensis was found in the undergrowth of forest bordering montane grassland, c. 1,700–2,000 m asl.; fruits found in April (Keraudren 1966) Taxonomy. According to Dwivedi et al. (2018) Z. perrieri is sister to Z. emirnensis. IUCN. Zehneria perrieri var. perrieri: Based on IUCN criterion B2, we suggest Least Concern (LC) for a species with wide distribution. Zehneria perrieri var. parvula and Z. perrieri var. tsaratananensis: We suggest Data Deficient (DD), as there are insufficient occurrence records to enable a proper assessment of the taxa, which have not been seen for decades and might have become extinct already. Zehneria polycarpa (Cogn.) Keraudren, Adansonia, n.s., 4: 333 (1964) Fig. 119 Basionym. Melothria polycarpa Cogn., J. Linn. Soc., Bot. 29: 20 (1891). Holotype. Madagascar • Fort-Dauphin, G.F. Scott Elliot 2316 K (K000313362!). Isotype. P (P00135446!). GenBank information. DNA sequences published by Dwivedi et al. (2018): KY523276, KY523342, KY523381, KY523439. 189 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Distribution. Endemic to Madagascar where it is found mainly in the North and East (Fig. 120). Western Domain: Montagne des Francais, M. Keraudren 1640 (P); Ankarana, M. Keraudren 1692 (P), H. Schaefer & M.B. Andriamiharisoa 23, 23B (DBEV, TUM); Manongarivo Massif, H. Perrier de la Bâthie 6785 (P); Ihorombe region, Zazafotsy, G. Cremers 581 (P). Sambirano Domain: Ambanja, Réserves Naturelles Madagascar 7435 RN (P), Nosy Komba, L.H. Boivin 2132 bis (P); Nosy-Be, L.H. Boivin s.n. (P). Central Domain: Diana region, Montagne d’Ambre NP, O. Andrianantoanina 986 (P), J. Bosser 20320 (P); Analamaitso forest, Anivorano North, M. Keraudren 1713 (P); Sava region, Daraina, Antsahabe forest, A. Rakotondrafara 344 (P); Anhabe forest, L. Nusbaumer 1276 (P); Alaotra Mangoro region, Andasibe NP, H. Schaefer et al. 3, 4, 17, 18, 19 (TUM, DBEV); Mantadia NP, H. Schaefer et al. 8, 10 (TUM, DBEV). Eastern Domain: Vatovavy region, Ranomafana NP, Schatz G.E. 1718 (BR), H. Schaefer et al. 103, 104, 114, 115 (DBEV, TUM); Anosy region, Andohahela NP, H. Schaefer et al. 49, 51, 52; Amboahangy, Fort-Dauphin, G.F. Scott Elliot 2542 (BR, K), M. Keraudren 1030, 1490 (P), P. Morat 1336 (P), M. Decorse s.n. (P); St Louis Peak, H. Humbert 5909 (P), J. Bosser 13559 (P), G.F. Scott-Elliot 2316 (BR, K, P), N. Dumetz 559 (WAG); Mandrare Valley, C d’ Alleizette, s.n. (L). Southern Domain: Androy region, Upper basin of Mandrare, H. Humbert 6843, 6590 (P); Manambolo Valley, H. Humbert 13291 (P). Occurrence inside Protected Areas. Andohahela NP, Analamazaotra NP, Mantadia NP, Montagne d’Ambre NP, Ranomafana NP, Ambohitr’Antsingy-Montagne des Français Harmonious Protected Landscape, Loky Manambato Harmonious Protected Landscape, and Ankarana Special Reserve. Ecology and habitat. Medium altitude moist evergreen to moist semi-deciduous forest, on roadside and riverbanks, in cliff, and dunes, growing in lateritic gneiss, limestone soils, and sands, c. 10–1,200 m asl.; flowering Oct.-Mar., fruits Oct.-July. Taxonomy. Ranjan et al. (2022) place Z. polycarpa close to Z. parvifolia. IUCN. Based on IUCN criterion B2 we suggest Least Concern (LC) for a species with wide distribution. Zehneria rutenbergiana (Cogn.) Keraudren, Adansonia, n.s., 4: 333 (1964) Fig. 121 Basionym. Melothria rutenbergiana Cogn., Abh. Naturwiss. Vereins Bremen 7: 251 (1882). Vernacular name. Siramboalavo (Sihanaka). Holotype. Madagascar • Ambatondrazaka, Lac Alaotra, Rutenberg s.n. (B; destroyed). Neotype. Madagascar • Central, s.d., R. Baron 2348 K (K000391257!), designated by Keraudren (1966), Cucurbitacees, Fl. Madagasc. 185: 47. GenBank information. DNA sequence published by Ranjan et al. (2022): MZ409454. Distribution. Endemic to Madagascar, where it has been found only in the Alaotra region (Fig. 122). Central Domain: Alaotra-Mangoro region, Ambatondrazaka, Ambatosoratra, Andreba Gara, Alaotra Lake, Rakotoarivelo 842 (MO); Ambatondrazaka, 190 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 117–122. 117. Zehneria perrieri var. perrieri, plant with green (immature) and red (mature) fruits (photo by MBA, Ranomafana NP, October 2022). 118. Distribution of Zehneria perrieri var. perrieri. 119. Zehneria polycarpa, habit (photo by MBA, Andohahela NP, September 2022). 120. Distribution of Zehneria polycarpa. 121. Zehneria rutenbergiana. A. Habit; B. Longitudinal section of ripe fruit (photos by N. Manjato, Andreba Gara, January 2017). 122. Distribution of Zehneria rutenbergiana. 117 118 119 120 121 122 191 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Andaingo Gara, Bembary, Randriambololomamonjy 466 (MO, TAN); Ambohinajanhary, J. Bosser 8179 (P). Eastern Domain: Zahamena, Manaka-East Township, Ambatondrazaka District, C. Rakotovao 11277, 11821 (P). Occurrence inside Protected Areas. Lac Alaotra Harmonious Protected Landscape and Zahamena NP. Ecology and habitat. Marshes and bog and island habitats associated with Cyperus latifolius and Psidium guajava, c. 800 m asl.; fruits in June. Taxonomy. According to Dwivedi et al. (2018), Z. rutenbergiana is sister to Z. capillacea. IUCN. We suggest Endangered (EN) based on IUCN criterion B2. Zehneria tridactyla (Hook.f.) R.Fern. & A.Fern., Mem. Junta Invest. Ultramar, 2 Ser. 34: 118 (1962) Fig. 123 Basionym. Melothria tridactyla Hook.f., Fl. Trop. Afr. 2: 562 (1871). Lectotype. MozaMbique • Shupanga, climbing on trees and among grass; Apr. 1862, J. Kirk, s.n. K (K000313374!), designated by Bräuchler et al. (2016), Phytotaxa 284(2): 144. Syntypes. angola • F.M.J. Welwitsch 826 (BM, G00458275!, K); congo • R. Burton s.n. K (K000313436!); MozaMbique • J. Kirk s.n. K (K000313373!). GenBank information. DNA sequences published by Dwivedi et al. (2018): KY523415, KY523414, KY523318, KY523466, KY523467, KY523350, KY523317. Distribution. Native to Southwest India, Sri Lanka, and Madagascar, where it is scattered throughout the country (Fig. 124). Western Domain: Diana region, Lohariandava, Andovoranto, R. Viguier 621 (P); Ankarana, H. Humbert 25531 (P), M. Keraudren 1690 (P); Boeny region, Marovoay, K. Afzelius s.n. (P); Ambongo-Boïna, H. Perrier de la Bâthie 6772 (P); Melaky, Tsingy de Bemaraha, C.C.H. Jongkind (WAG); Bongolava region, Babay-ville Sakay, M.W. Armand 129 (P); Kianjasoa, J. Bosser 783 (P), Jard. Bot. Tananarive 171 (P); Kinkony, R. Decary 7752 (P). Sambirano Domain: Diana region, Nosy-Be, A. Pervillé s.n. (P), L.H. Boivin 2132 (P). Central Domain: SAVA region, Marojejy massif, H. Humbert 22142bis (P); Betsiboka region, Maevatanana, H. Perrier de la Bâthie 1074 (P). Sofia region, Ankaramy, Maromandia (Ankaramy), R. Decary 1318 (P); Alaotra-Mangoro region, Ivondro river, J.P. Goudot 1833 (BR); Atsimo-Andrefana region, Sakaraha, Zombitsy, T. Mitchell & H. Schaefer (TUM). Eastern Domain: Fenoarivo-Antsiranana, Sainte Marie, L.H. Boivin s.n. (P). Alaotra-Mangoro, Ambodimanga, G. Cours 476 (P); Ambatosoa region, Mananara, R. Decary 55 (P). Occurrence inside Protected Areas. Tsingy de Bemaraha NP, Mahavavy Kinkony Wetlands Complex Harmonious Protected Landscape, and Zombitse-Vohibasia NP. Ecology and habitat. Marshes and bogs, rice fields, forest edges, and in montane ravines, c. 250–800 m asl.; flowering Sept.–May, fruits June–Aug. Taxonomy. Zehneria tridactyla is sister to Z. giletii, and Z. capillacea (Dwivedi et al. 2018). IUCN. We suggest Least Concern (LC) based on IUCN criterion B2. 192 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 123, 124. 123. Zehneria tridactyla, habit (photo by HS, Zombitsy-Vohibasia NP, March 2013). 124. Distribution of Zehneria tridactyla. 123 124 Naturalised species Six species of Cucurbitaceae are classified as naturalised species (Table 2) which have been introduced on purpose or by accident in the recent past and established permanent populations in natural habitats. Coccinia Wight & Arn., Prodr. Fl. Ind. Orient. 1: 347 (1834). Generic type. Coccinia indica Wight & Arn. Worldwide Distribution. Africa and 1 species introduced in tropical regions worldwide (Schaefer 2020). Coccinia grandis (L.) Voigt, Hort. Suburb. Calcutt. 59 (1845) Fig. 125 Basionym. Bryonia grandis L., Mant. Pl. 126 (1767). Lectotype. india • Linnaeus s.n. (LINN No. 1153.2), designated by Jeffrey (1967), Fl. Trop. E. Afr., Cucurbitac.: 68. GenBank information. Sequence data available on GenBank from Holstein and Renner (2011); Malagasy material has been sequenced by the authors (unpubl. data). Distribution. Native to Tropical Africa, the Western Arabian Peninsula, Tropical and perhaps subtropical Asia. Introduced in Madagascar, where it is found cultivated and escaped in the South (Fig. 126). Table 2. Naturalised Cucurbitaceae of Madagascar with status and distribution. No. Scientific name Status Distribution 1. Coccinia grandis (L.) Voigt Naturalised Eastern, Southern 2. Cucumis anguria L. Naturalised Southern 3. Cucumis dipsaceus Ehrenb. ex Spach Naturalised Southern 4. Cucumis melo L. Naturalised Sambirano, Western, Central, Southern 5. Diplocyclos palmatus (L.) C.Jeffrey Naturalised Western, Central, Eastern 6. Momordica charantia L. Naturalised All 193 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Southern Domain: Atsimo-Andrefana region, Andranomaitso, near Zombitsy NP, H. Schaefer et al. 83 (DBEV, TUM). Easter Domain: Anosy region, Fort Dauphin, city, H. Schaefer et al. 59 (DBEV, TUM); Cultivated, climbing fence, Nahampoana Village, H. Schaefer et al. 27 (January 2025). Occurrence inside Protected Areas. Not known from protected areas. Ecology and habitat. Disturbed ground near settlements, on sandy soil, c. 50 m asl.; flowers and fruits Oct.-Dec. Uses. Young leaves, shoots, and green fruits are used as vegetables in Asia (HS, pers. obs.). No such use has been documented from Madagascar. Taxonomy. According to Holstein and Renner (2011), C. grandis is sister species to C. ogadensis. Cucumis L., Sp. Pl. 1011. 1753. Fig. 127 Cucumis anguria L., Sp. Pl. 2:1011 (1753) Type. JaMaica (cult. in Sweden) • Linnaeus s.n. (Lectotype. LINN 1152.6), designated by Staples, in RA Howard, Fl. Less. Antill., Dicot. 3: 482 (1989). GenBank information. DNA sequences published by Endl et al. (2018): KY434397, KY458066, KY434332, HM597065, KY434505. Distribution. Native to the African continent (Angola, Botswana, KwaZulu-Natal, Malawi, Mozambique, Namibia, Northern Provinces, Swaziland, Tanzania, Zambia, Zaire, Zimbabwe), introduced and invasive in many (sub)tropical regions of the world (POWO 2023). In Madagascar, C. anguria occurs only near settlements in the extreme South (Fig. 128), where it has been misidentified as C. africanus in the past (e.g., Keraudren 1001 (P)), a species which probably does not (or no longer) exist in the country. Southern Domain: Androy region, Ambovombe, R. Decary 9540 (P); Anarafaly, J. Bosser 13568, M. Keraudren 1001 (P); Androka, J. Bosser 301 (P); Tranomaro New Protected Area, H. Schaefer et al. 7 (DBEV, TUM); Atsimo-Andrefana region, Ranobe Forest, Ankilimalinike, P.B. Phillipson 5927 (P); Anosy region, dry forest, Andohahela NP, H. Schaefer et al. 66, 69 (DBEV, TUM). Occurrence inside Protected Areas. Andohahela NP, Ankodida Harmonious Protected Landscape. Ecology and habitat. Disturbed ground in dry spiny thicket and dry deciduous clearings, in savanna with scattered bushes on sandy soils, c. 50–100 m asl. Uses. In South America and the West Indies, the fruits are grown as vegetables (Baird and Thieret 1988) but no such use has been documented in Madagascar. Taxonomy. Cucumis anguria is sister to the clade including C. pustulatus, C. prophetarum, C. dipsaceus, C. ficifolius, C. baladensis, and C. rigidus (Sebastian et al. 2010). Cucumis dipsaceus Ehrenb. ex Spach, Hist. Nat. Vég. 6: 211 (1838) Fig. 129 Holotype. saudi arabia • G. Ehrenberg s.n. (B, destroyed). 194 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 125–128. 125. Coccinia grandis. A. Mature fruit; B. Flower (photos by S. Philizot, Toliara, February 2024 (A) and by MBA, Fort Dauphin, October 2022 (B)). 126. Distribution of Coccinia grandis. 127. Cucumis anguria. A. Habit; B. Longitudinal section of the fruit (photos by MBA, Andohahela NP, October 2022). 128. Distribution of Cucumis anguria. 125 127 126 128 Neotype. saudi arabia • Wadi Kamme east of al-Qunfidha, Feb. 1825, G. Ehrenberg & Hemprich s.n. (MPU), designated by Kirkbride, Biosyst. Monogr. Cucumis 45 (1993). Synonyms. Momordica dasycarpa Hochst. ex A.Rich., Tent. Fl. Abyss. 1: 291 (1848). 195 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae GenBank information. DNA sequences for this taxon (but not from Malagasy material) published by Ghebretinsae et al. (2007), Renner et al. (2007), Endl et al. (2018): AF006800, AF006801, AF013310, DQ785828, DQ785844, EF595872, EF595922, EF093513, MZ489698, MG993578, MG993587. Distribution. Introduced to Madagascar and invasive in the South (Fig. 130). Southern Domain: Tranomaro Village, H. Schaefer et al. 4, 9A, 9B (DBEV, TUM); Anosy region, Bevilany village, roadside, Andohahela NP, H. Schaefer et al. 67 (DBEV, TUM). iNaturalist observations: https://www.inaturalist.org/observations/25662008; https://www.inaturalist.org/observations/70549193. Occurrence inside Protected Areas. Andohahela NP, Amoron’i Onilahy Harmonious Protected Landscape. Ecology and habitat. Disturbed areas of dry spiny thicket, often climbing on roadsides and in xerophytic shrub, c. 100 m asl.; flowers and fruits Oct.–Feb. Uses. In Peru, the fresh fruits of C. dipsaceus are used for dandruff, adding shine and beauty to hair, hair loss prevention, and to stop babies from breastfeeding (Bussmann and Sharon 2016). No such use has been documented in Madagascar. Taxonomy. Cucumis dispsaceus is sister species to C. ficifolius (Sebastian et al. 2010). Cucumis melo L., Sp. Pl.: 1011 (1753) Fig. 131 Holotype. cult. at Uppsala (Sweden), Linnaeus s.n. (Herb. LINN No. 1152/8), designated by Meeuse, Bothalia 8 (1962): 61. Synonyms. https://powo.science.kew.org/taxon/292238-1. Vernacular name. Voatango (Merina). GenBank information. DNA sequences of the species published e.g., by Endl. et al. (2018): KY434411, KY434412, KY458087, KY458086, KY434470, KY434472, KY434471, KY434354, KY434355. Distribution. Cultivated in Madagascar for its fruit and occasionally found in the wild (Fig. 132). Western Domain: Diana region, Manongarivo Special Reserve, L. Gautier 2972 (MO); Ankarana, H. Schaefer & M.B. Andriamiharisoa 28, 31, 31B (TUM, DBEV); Menabe, West coast, from Manombo to Morondava, A. Grandidier s.n. (P); Mahobo Farm, J. Dequaire 27010; Andranofotsy transit hut, H. Poisson 482 (P). Sambirano Domain: Diana region, Nosy Be, L.H. Boivin 2129 (P). Central Domain: Analamanga region, C. d’Alleizette (P); Alaotra-Mangoro region, East-Ilaka, J. Bosser 17024 (P). Eastern Domain: Alaotra-Mangoro region, Andasibe-Perinet village, H. Schaefer et al. 12 (TUM, DBEV). Southern Domain: Atsimo-Andrefana region, north of Toliara, A. Rakotozafy (P); Ambolonkira, M. Keraudren 664 (P), J. Bosser 13581 (P); Vohitany, M. Keraudren 1436 (P); Near Ambatry southern Betioky, M. Keraudren 790 (P); Andatabo, M. Keraudren 556 (P); Mahafaly limestone plateau, M. Keraudren 867 (P); Ambiamena, Zombitsy NP, H. Schaefer et al. 91 (DBEV, TUM), B. Descoings 2275 (P); near Andabolava, H. Humbert 12419, 27193 (P); Tanandava, Mangoky, J. Bosser 16092 (P); Ampandrandava, A. Seyrig 557 (P). 202 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Lagenaria Ser., Mém. Soc. Phys. Genève 3: 26. 1825. Generic type. Lagenaria vulgaris Ser. Worldwide Distribution. Africa, cultivated and escaped in tropical and temperate regions worldwide (Schaefer 2020). Lagenaria siceraria (Molina) Standl., Publ. Field Columb. Mus., Bot. Ser. 3: 435 (1930) Fig. 140 Basionym. Cucurbita siceraria Molina, Sag. Stor. Nat. Chili 133 (1782). Type. souTh africa • Roggefeld, Rietpoort, 1 Jan. 1875, A. Rehmann 3247 (Z-000004481). Synonyms. https://powo.science.kew.org/taxon/134809-2. Vernacular name. Tsilaniantitra (Merina). GenBank information. DNA sequences of the species from outside Madagascar have been published e.g., by N’dri et al. (2016): KF982914, KF982913. Malagasy material has been sequenced by the authors (unpubl. data). Distribution. Often cultivated and recorded as a casual from three provinces, including the Center, North, and South. Ecology and habitat. Cultivated mainly on sandy soil, from 100–1,800 m asl. Uses. The young, tender peeled fruits are eaten as vegetables, the hard, dried shells of the calabash are used as a water container or to make musical instruments. Taxonomy. Lagenaria siceraria is sister to L. breviflora (Schaefer et al. 2009). Lagenaria sphaerica (Sond.) Naudin, Ann. Sci. Nat., Bot., sér. 5, 5: 9 (1866). Basionym. Luffa sphaerica Sond., Fl. Cap. 2: 490 (1862). Syntypes. souTh africa • zwischen Omsamculo und Omcomas, in feuchten Niederungen und sumpfigem Thal, 1832, J.F. Drège s.n. HBG (HBG506395!), TUB (TUB-004706!). Synonyms. Lagenaria sphaerocarpa Arn., J. Bot. (Hooker) 3: 277 (1841); Sphaerosicyos meyeri Hook.f., Fl. Trop. Afr. 2: 532 (1871), nom. superfl.; Sphaerosicyos sphaericus (Sond.) Cogn., Monogr. Phan. 3: 466 (1881); Lagenaria mascarena Naudin, Ann. Sci. Nat., Bot., sér. 4, 18: 187 (1862). GenBank information. DNA sequences of the species (but not from Malagasy material!) published e.g., by Kress and Erickson (2007): EF590539, EF590627, EF590405, EF590779. Pictures. https://powo.science.kew.org/taxon/293050-1. Distribution. Widely cultivated and sometimes found as casual. Western Domain: Diana region, Ambahatra, midstream Ambahatra on the left side, between Anketrakabe and Antafiabe, S.-L. Stiefel 77 (P). Sambirano Domain: Diana region, Sambirano alluvial deposits, H. Perrier de la Bâthie 16268 (P); Nosy Be, J.M. Hildebrandt 2961 (P), L.H. Boivin 2128 (P); Nosy Komba, J. Bosser 14757, 14757bis, 15915 (P). Eastern Domain: Antsinanana region, Befalafa, L. Gautier 3350 (P, WAG). Ecology and habitat. Riverbanks and humid forest, mainly on clayey soil or alluvial deposits, from 400–1,000 m asl. 203 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Figures 137–140. 137. Cucumis sativus, habit (photo by MBA, Montagne des Français, January 2024). 138. Cucurbita moschata. A. Flower; B. Habit (photos by MBA, Joffreville, January 2024). 139. Cucurbita pepo. A. Fruit from cultivation; B. Habit, (photos by MBA, Antananarivo, March 2024 (A) and Betafo, Antsirabe, January 2020 (B)).140. Lagenaria siceraria. A. Habit; B. Mature fruit (photos by MBA, Antananarivo, December 2023, March 2024). 137 138 139 140 Uses. Vegetable, the immature fruit is peeled and cooked or fried or used in curries. Luffa Mill., Gard. Dict. Abridg. ed. 4. 1754. Generic type. Luffa aegyptiaca Mill. Worldwide Distribution. Tropics and Subtropics of the Old and New world (Schaefer 2020). Luffa acutangula (L.) Roxb., Fl. Ind., ed. 1832. 3: 713 (1832) Fig. 141 Basionym. Cucumis acutangulus L., Sp. Pl.: 1011 (1753). Holotype. PhiliPPines • Antipolo, Rizal Province, Luzon, Nov. 1914, Merill (Species Blancoanae No. 107), n.s. Synonyms. Cucurbita acutangula (L.) Blume, Bijdr. Fl. Ned. Ind.: 932 (1826). Vernacular name. Papangay (Betsimisaraka, Sakalava). GenBank information. DNA sequences of the species from outside Madagascar have been published e.g., by Filipowicz et al. (2014): KF487441, KF487394, KF487331, KF487330. Malagasy material has been sequenced by the authors (unpubl. data). 204 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Distribution. Widely cultivated and occasionally found as casual, e.g., in the Western and Central Domains. Western Domain: Diana region, Ankarana, H. Schaefer & M.B. Andriamiharisoa 29, 30, 33, 34 (TUM, DBEV); Boeny region, Mahajanga, Poisson 98 (P); Ankarafantsika, Ampijoroa, on the forest edge, M. Keraudren 1250 (P); Melaky region, Soalala District, Namoroka, Andranomavo township, H. Randriamiera 7997 (P). Central Domain: Diana region, Montagne d’Ambre, H. Schaefer & M.B. Andriamiharisoa 43, 43B (January 2024). Ecology and habitat. Cultivated on forest edges and in shaded areas at lower altitudes. Uses. Young fruits and leaves are cooked or fried and eaten in soups and sauces. Taxonomy. Luffa acutangula is sister to all other Luffa species (Filipowicz et al. 2014). Luffa aegyptiaca Mill., Gard. Dict., ed. 8.: [s.p.] (1768) Fig. 142 Type. Unclear. Synonyms. Luffa cylindrica auct., non (L.) Roem., Syn. Mon. 2: 63 (1846). Vernacular name. Papangay (Sakalava, Merina). GenBank information. DNA sequences of the species from outside Madagascar published e.g., by Filipowicz et al. (2014) and Wu et al. (2020): CM029398 JAETGN010000000, KF487456, KF487500, KF487377, KF487344, KF487450. Malagasy material has been sequenced by the authors (unpubl. data). Distribution. Luffa aegyptiaca is widely cultivated and frequently reported as casual near settlements. Ecology and habitat. Cultivated and locally escaped growing on alluvial deposits at lower altitudes. Uses. Young fruits and leaves eaten as vegetables. Taxonomy. Luffa aegyptiaca is sister to L. graveolens and L. saccata (Filipowicz et al. 2014). Sicyos L., Sp. Pl. 2: 1013. 1753. Generic type. Sicyos angulatus L. Worldwide Distribution. South, Central and North America, Hawaii, New Zealand, Australia, invasive in Africa and Eurasia (Schaefer 2020). Sicyos edulis Jacq., Enum. Syst. Pl.: 32 (1760) Fig. 143 Holotype. JaMaica • N.J. von Jacquin BM (BM000559376!). Synonyms. Sechium edule (Jacq.) Sw., Fl. Ind. Occid. 2: 1150 (1800). Vernacular name. Sôsety (Merina). GenBank information. DNA sequences of the species from outside Madagascar published e.g., by Kocyan et al. (2007) and Sebastian et al. (2012): DQ536727, 205 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae JN560580, DQ535843, JN560209, DQ536589, DQ536861, JN560486. Malagasy material has been sequenced by the authors (unpubl. data). Distribution. Widely cultivated throughout the Tropics including Madagascar and sometimes found outside cultivation but not truly naturalised on the island. Central Domain: Analamanga region, Antananarivo, A.C d’Alleizette, s.n. (L), M. Keraudren 1556 (P); Analamazaotra, C. d’ Alleizette s. n. (P); on the Mangoro riverbank near Antandrokomby, frequent sub-spontaneous, around the village in the warmer parts of the Central East, but not naturalised, current introduction, relatively recent introduction (during the 20th century), H. Perrier de la Bâthie 17190 (P). Western Domain: Ankarafantsika NP, H. Schaefer et al. 42 (TUM, DBEV); eastern Mahabo, G. Aymonin 25961 (P). Ecology and habitat. Widely cultivated in Madagascar but does not seem to occur in the wild. Uses. Fruits, young leaves, shoots, and tuberous roots are commonly consumed as vegetables. Taxonomy. Sicyos edulis is sister to Sechium chinantlense (Sebastian et al. 2012). Telfairia Hook., Bot. Mag. 2751-52. 1827. Generic type. Telfairia pedata (Sm.) Hook. Worldwide Distribution. African Tropics (Schaefer 2020). Telfairia pedata (Sm.) Hook., Bot. Mag. 54: t. 2751 (1827). Basionym. Fevillea pedata Sm., Bot. Mag. 53: t. 2681 (1826). Type. Tanzania • Zanzibar, 1833, W. Bojer s.n. (K). Synonyms. Joliffia africana Delile, Mém. Soc. Hist. Nat. Paris 3: 314 (1827); Telfairia africana (Delile) A.Chev., Rev. Int. Bot. Appl. Agric. Trop. 29: 217 (1949). GenBank information. DNA sequences of the species (but not from Malagasy material!) published e.g., by Kocyan et al. (2007): DQ374439, DQ491021, DQ501271, DQ535853. Pictures. https://www.inaturalist.org/observations?taxon_id=1139409. Distribution. Introduced, only known from one ancient collection from the North, no recent reports. Sambirano Domain: Nosy-Be, cultivated, L.H. Boivin s.n. (P). Ecology and habitat. Was probably cultivated in humid low altitude areas of Madagascar in the past but not seen more recently. Uses. Seeds are eaten raw and used to produce vegetable oil on the African continent, but no such use has been documented in Madagascar. Taxonomy. Telfairia pedata is sister to T. occidentalis (Kocyan et al. 2007). Trichosanthes L. Sp. Pl. 2: 1008. 1753. Generic type. Trichosanthes anguina L. Worldwide Distribution. Australasia (Schaefer 2020). 206 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Trichosanthes cucumerina L., Sp. Pl.: 1008 (1753) Fig. 144 Holotype. indonesia • Java, s.d., T. Horsfield 499981A U (U0008345). Isotype. U (U0008346!). Vernacular name. Mapangay (Sakalava). GenBank information. DNA sequences of the species from outside Madagascar have been published e.g., by Schaefer et al. (2008) and De Boer et al. (2012): EU155603, EU155609, HE661486, HF586722, HF586731, HF586766, HF586768. Malagasy material has been sequenced by the authors (unpubl. data). Distribution. Introduced to Madagascar and cultivated as a vegetable and rarely found as a casual. Ecology and habitat. Cultivated at low altitudes. Uses. Fruits and leaves are consumed as vegetables. Taxonomy. Trichosanthes cucumerina is sister to T. nervifolia (De Boer et al. 2012). Doubtful species and erroneous records Cucumis africanus L. fil., Suppl. Pl.: 423 (1782). Lectotype. france • Jardin d’expérience de Collioure, de l’intérieur de l’Afrique, 1874, C. Naudin s.n. P (P00346219!), designated by Kirkbride, Biosyst. Monogr. Cucumis 31 (1993). Distribution. Native to the African continent (Angola, Botswana, South Africa, Namibia, and Zimbabwe (POWO 2023), based on our herbarium studies, all reports from Madagascar seem to be in error for C. anguria. Cucumis cinereus (Cogn.) Ghebret. & Thulin, Novon 17: 177 (2007). Basionym. Kedrostis cinerea Cogn., Bull. Herb. Boissier, sér. 2, 1: 883 (1901). Holotype. naMibia • Hereroland, An der Giftkopje, 12th Feb., 1900, M.K. Dinter 1440 Z (Z000004457!). Isotype. BR (BR0000008888297!). Synonyms. Cucumella cinerea (Cogn.) C.Jeffrey, Kew Bull. (1962); Melothria cinerea (Cogn.) A. Meeuse, Bothalia 8: 17 (1962). Distribution. Native to Namibia, Angola, Tanzania, Kenya (POWO 2023) and has been reported from Madagascar in error. Taxonomy. Genetic analyses show that Malagasy material identified as this species in fact belongs to a different undescribed species. Cucumis hirsutus Sond., W.H. Harvey Cap. 2: 497 (1862). Holotype. ziMbabwe • Marandellas, 7 Jan. 1942, G. Dehn 582 m (M0105801! / 27188! / 18229!). Distribution. Native to continental Africa (including Angola, Botswana, Burundi, Cameroon, Cape Provinces, Congo, Kenya, KwaZulu-Natal, Malawi, Mozambique, Northern Provinces, Sudan, Swaziland, Tanzania, Zambia, Zaïre, Zimbabwe) (POWO 2023) and has been reported from Madagascar in error. 207 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Taxonomy. Genetic analyses show that Malagasy material identified as this species in fact belongs to a different undescribed species. Momordica trifoliolata Hook.f., Fl. Trop. Afr. 2: 537 (1871). Holotype. Tanzania • Morogoro Dist., Ngomero, Grant s.n. K (K000310605!). GenBank information. DNA sequences of the species (but not from Malagasy material!) published e.g., by Schaefer and Renner (2010): GQ162917, GQ162962, GQ163454, GQ163091, GQ163214, GQ163336. Distribution. Only known from a single collection in Antananarivo, Mandraka, near Mandraka forest, C. d’ Alleizette 1079 (P). It is doubtful if this specimen really came from Madagascar. Taxonomy. Momordica trifoliolata is sister to M. rostrata (Schaefer and Renner 2010). Zehneria maysorensis (Wight & Arn.) Arn., J. Bot. (Hooker) 3: 275 (1841). Basionym. Bryonia maysorensis Wight & Arn., Prodr. Fl. Ind. Orient. 1: 345 (1834) Vernacular name. Vahifamakiela (Betsileo). Lectotype. india • Peninsula Ind. Orientalis [Chovei], E.P. Wright 1116 P (P00218566!). GenBank information. DNA sequences of the species (but not from Malagasy material!) published e.g., by Dwivedi et al. (2018): KY523335, KY523256, KY523425, KY523368. Figure 141–144. 141. Luffa acutangula, habit (photo by MBA, Ankazomiriotra, March 2023). 142. Luffa aegyptiaca. A. Female flower; B. Young fruit; C. Seeds (photos by MBA, Marovoay, March 2020). 143. Sicyos edulis, cultivated plant with flowers and young fruit (photo by MBA, Ranomafana village, October 2022). 144. Trichosanthes cucumerina. A. Young fruit; B. Habit (photos by MBA, Andringitra, March 2023). 141 142 143 144 208 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Distribution. Zehneria maysorensis is native to the Indian Subcontinent and has also been reported from Madagascar (POWO 2023). However, it is unlikely that the Malagasy material belongs to the same taxon found in India. The species has been reported from the Central Domain: Haute-Matsiatra region, Ambalavao District, Mahazony township, Ankidontany, C. Rakotovao 9173 (P), and s. loc., P. Commerson s.n (BR). Zehneria thwaithesii (Schweinf.) C. Jeffrey, Kew Bull. 15: 371 (1961, publ. 1962). Basionym. Melothria thwaitesii Schweinf., Reliq. Kotschy.: 44 (1868) Lectotype. sri lanka • Walker 275 K (K000742779), designated by C Jeffrey, Kew Bulletin 15(3): 371(1962). GenBank information. DNA sequences of the species (but not from Malagasy material!) published by Dwivedi et al. (2018): AM981145, KY523315, KY523349, KY523412, KY523465. Distribution. Tropical Africa, southern India, and Sri Lanka (De Wilde and Duyfjes 2006). Note. Not present in Madagascar, where it has been confused with Z. tridactyla. Results Total species and comparison with the lists of Keraudren and Rauh Following Keraudren’s study of the Cucurbitaceae family in Madagascar, which resulted in the publication of the respective volume of the Flore de Madagascar et des Comores (Keraudren 1966), very few studies of the Cucurbitaceae have been conducted, with the notable exception of Rauh’s work on the succulent taxa (1995). Monique Keraudren (1966) listed 27 genera, eight of them endemic, 66 species, and nine varieties, including six for the genus Peponium and three for Zehneria. In our revision, we accept 26 genera, five of which are endemic to Madagascar, comprising 82 taxonomic units. This total includes 77 species and five varieties: three for the genus Peponium and two for Zehneria. Among the additions in our list are new species described or reported by Keraudren after the publication of her flora volume: Seyrigia marnieri (Keraudren 1967), Odosicyos bosseri (Keraudren-Aymonin 1980, 1982a), Diplocyclos palmatus (Keraudren-Aymonin 1982b), and Seyrigia napifera, which has been described by Rauh (1995). Due to modern phylogenetic analyses, genus circumscriptions changed for many of the Malagasy cucurbits, leading to numerous name changes mainly at the genus level: Cucumella and Oreosyce were included in Cucumis, Odosicyos and Tricyclandra were included in Ampelosycios, Sechium was moved to Sicyos, Zombitsia moved to Blastania, and Zygosicyos included in Xerosicyos (Kocyan et al. 2007; Schaefer 2007; Schaefer and Renner 2011; Sebastian et al. 2012; Lindner et al. 2017). Most or all records of Cucumis africanus, Cucumis sativus, Cucurbita maxima, and Citrullus colocynthis by Keraudren (1966) seem to have been in error for Cucumis anguria, Cucumis sacleuxii, Cucurbita moschata, and Citrullus mucosospermus. Cucumis dipsaceus and Coccinia grandis seem to be recently introduced invaders that arrived in Madagascar after the 1980s. 209 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Despite considerable efforts, we were unable to locate living plants of 24 taxa. Five species remain doubtful because they have most likely been confused with species occurring in India or Africa: Cucumis cinereus, Cucumis hirsutus, Momordica trifoliolata, Zehneria thwaitesii, and Z. maysorensis. A total of five new species were identified in this study, two for the genus Ampelosycios, two for Cucumis, and one for Xerosicyos. In addition, five species from continental Africa and India are newly reported from Madagascar: Citrullus mucosospermus, Cucumis anguria, Coccinia grandis, Gerrardanthus grandiflorus, and Zehneria tridactyla. Total species with DNA sequences For the vast majority of the 82 Cucurbitaceae species and varieties, DNA sequence data from Madagascar is already available on GenBank (https://www. ncbi.nlm.nih.gov/) or has been collected during our project and will be made available publicly in the near future. The genera, where more fieldwork and collection are needed most are Cayaponia, Cyclantheropsis, Peponium, and Kedrostis, especially those taxa known only from the types in the P herbarium. Endemism Regarding the distribution status of the Cucurbitaceae family in Madagascar, we classify the species in our checklist as follows: 60 are native, representing 80% of the total diversity. Endemism level is high with 86% (52 species plus 5 varieties), eight species are probably native but not endemic (Fig. 145). We find that 17 species have been introduced, eleven of them are found in cultivation and rarely as casuals near settlements, while six exotic species have established permanent populations outside cultivated areas and some even start to become invasive in natural habitats (e.g., Cucumis dipsaceus). Figure 145. Status of Malagasy Cucurbitaceae by genus. 210 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Diversity The diversity analysis conducted across Madagascar’s five phytogeographic divisions (Humbert 1955) indicates that both at species and genus levels, Western, Southern and Central domains are most diverse (Fig. 146). Most of the native species, (47, 68%) occur in the Western domain, 33 (48%) in the Southern, and 32 (46%) in the Central domain. Only 17 species (25%) have been found in the Eastern domain, 11 (16%) in the Sambirano, and eight (12%) in the High Mountain Domain. Analysis of IUCN classifications Our preliminary analysis of extinction risks of Malagasy wild Cucurbitaceae based on IUCN criteria B1 and B2 (Table 4), revealed that the available information for 12 taxa is insufficient, so that we suggest classifying them as Data Deficient (DD), even though some of them might have become extinct already. For 24 taxa, we suggest classification as Critically Endangered (CR), and for eight taxa, the category Endangered (EN) seems most appropriate. The remaining 21 are classified Least Concern (LC) (Fig. 147). Figure 146. Number of Cucurbitaceae genera and species per phytogeographic domain. Figure 147. Distribution of Red list categories of Malagasy Cucurbitaceae based on IUCN Criterion B1, B2, and author’s attribution. 211 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. Andriamiharisoa et al.: Checklist of Malagasy Cucurbitaceae Table 4. Estimated Area of Occupancy, Extent of Occurrence, and suggested IUCN classification of the native Cucurbitaceae of Madagascar. Scientific name AOO (km2) EOO (km2) AOO Category EOO Category Final Assessment Ampelosycios bosseri 7 121.4 CR EN CR Ampelosycios humblotii 10 252074.8 EN LC LC Ampelosycios leandrii 729229.8 CR LC CR Ampelosycios meridionalis 3 13375.6 CR VU DD Ampelosycios scandens 10 31560.9 EN LC LC Ampelosycios sp. nov. Ankarana 7163.3 CR EN EN Ampelosycios sp. nov. Montagne des Français EN EN Blastania lucorum 68574.5 CR VU CR Cayaponia africana var. madagascariensis DD Citrullus mucosospermus 20 252769.8 EN LC LC Corallocarpus grevei 29 230860 EN LC LC Corallocarpus perrieri 22 40750.2 EN LC LC Corallocarpus poissonii 3 1957.8 CR EN EN Cucumis sacleuxii LC LC Cucumis sp. nov. (aff. cinereus)2 0 CR CR CR Cucumis sp. nov. (aff. hirsutus)2 0 CR CR CR Cucumis subsericeus 27 123453.3 EN LC LC Cyclantheropsis madagascariensis 5 233722.4 CR LC CR Gerrardanthus grandiflorus 2 0 CR CR CR Kedrostis cogniauxii DD Kedrostis dissecta 3 51910.4 CR LC CR Kedrostis elongata 20 230798.4 EN LC LC Kedrostis lanuginosa 3 40348.2 CR LC CR Kedrostis laxa 11 8340.4 EN VU EN Kedrostis perrieri 2 0 CR CR CR Lemurosicyos variegata 36 376031.6 EN LC LC Muellerargia jeffreyana 12 185339.2 EN LC EN Peponium betsiliense 6116897.7 CR LC CR Peponium boivinii 3 709.5 CR EN CR Peponium grandidieri 8 230445.2 CR LC LC Peponium hirtellum var. hirtellum 3 198.3 CR EN CR Peponium hirtellum var. longiracemosum 4 20 CR CR CR Peponium humbertii 4 697.6 CR EN CR Peponium laceratum DD Peponium perrieri var. glabrescens DD Peponium perrieri var. perrieri 6198370.2 CR LC CR Peponium poissonii 5 30988.5 CR LC CR Peponium racemosum 426850.8 CR LC CR Peponium seyrigii var. linearlobum DD Peponium seyrigii var. seyrigii DD Raphidiocystis brachypoda 30 126767.9 EN LC LC Seyrigia bosseri 4213.9 CR EN CR Seyrigia gracilis 28 56717.7 EN LC LC Seyrigia humbertii 5 187.2 CR EN EN Seyrigia marnieri 2 0 CR CR DD Seyrigia multiflora 17 44153.6 EN LC LC Seyrigia napifera 2 0 CR CR CR Trochomeriopsis diversifolia 61 460246.7 EN LC LC Xerosicyos danguyi 65 102590.5 EN LC LC Xerosicyos decaryi 16 147808.8 EN LC LC Xerosicyos hirtellus CR Xerosicyos perrieri 33 104911.3 EN LC LC Xerosicyos pubescens 3 36.2 CR CR CR 218 PhytoKeys 267: 123–222 (2025), DOI: 10.3897/phytokeys.267.159218 Minosoa B. 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