Table 1.
List of genera containing species capable of CAM photosynthesis; major clades are ordered following the linear classification system of APG IV (2016). Where species names have subsequently been synonymized or segregated into other genera, the original taxonomy is indicated in parentheses after the currently accepted name. Relevant phylogenetic studies are cited for clades with multiple putative origins of CAM. A fully referenced list with citations to initial and subsequent reports of CAM activity in these taxa, together with details of names reduced to synonymy, is provided as Supplementary Data (Table S1).
| Major clade | Genus | Putative CAM origins |
|---|---|---|
| Isoëtales | ||
| Isoëtaceae | Isoëtes (including Stylites) | 1 |
| Polypodiales | ||
| Polypodiaceae | ||
| Microsoroideae | Lecanopteris; Microsorum | 2; 1 each in Lecanopteris and Microsorum (Chen et al., 2020) |
| Loxogrammoideae | Dictymia | 1 |
| Polypodioideae | Niphidium | 1 |
| Platycerioideae | Platycerium; Pyrrosia (including Drymoglossum) | 1 or 2; 1 in the ancestor of Platycerium and Pyrrosia or 1 in each clade (Wei et al., 2017) |
| Pteridaceae | ||
| Vittarioideae | Haplopteris; Anetium; Vittaria | 2; 1 in the ancestor of Anetium and Vittaria and 1 in Haplopteris (Schuettpelz et al., 2016) |
| Cycadales | ||
| Zamiaceae | Dioon | 1 |
| Welwitschiales | ||
| Welwitschiaceae | Welwitschia | 1 |
| Piperales | ||
| Piperaceae | ||
| Piperoideae | Peperomia | 5–12 (Frenzke et al., 2016; Lim et al., 2019) |
| Alismatales | ||
| Alismataceae | Sagittaria | 1 |
| Araceae | Zamioculcas | 1 |
| Hydrocharitaceae | Ottelia; Vallisneria | 2; 1 each in Ottelia and Vallisneria (Chen et al., 2022) |
| Asparagales | ||
| Orchidaceae | ||
| Epidendroideae | Acianthera; Aerangis; Aeranthes; Anathallis; Angraecum; Arachnis; Aspasia; Barkeria; Bogoria; Brassavola; Brassia; Bryobium; Bulbophyllum; Campylocentrum; Cattleya (including Sophronitis); Caularthron; Caluera; Capanemia; Chiloschista; Cischweinfia; Coelogyne (including Pholidota); Comparettia (including Scelochilus); Coryanthes; Cymbidium; Cyrtopodium; Dendrobium (including Cadetia, Dockrillia, Flickingeria and Grastidium); Dendrophylax; Didymoplexis; Domingoa; Dimerandra; Echinosepala (including Brenesia); Elleanthus; Encyclia; Epidendrum (including Lanium and Oerstedella); Eriopsis; Erycina (including Psygmorchis); Eulophia (including Acrolophia, Lissochilus, Oeceoclades and Orthochilus); Gomesa; Gongora; Guarianthe; Hintonella; Ionopsis; Jacquiniella; Laelia (including Schomburgkia); Leochilus; Lockhartia; Luisia; Lycaste; Macradenia; Macroclinium; Maxillaria (including Camaridium, Heterotaxis, Ornithidium and Trigonidium); Meiracyllium; Microcoelia (including Gussonea); Micropera; Mobilabium; Mormodes; Myoxanthus; Myrmecophila; Notylia; Oberonia; Oeonia; Oncidium; Ornithocephalus; Pabstiella; Peristeria; Phalaenopsis (including Sedirea); Platyrhiza; Plectorrhiza; Plectrophora; Pleurothallis; Pomatocalpa; Prosthechea; Psychilis; Psychopsis; Pterostemma; Quekettia; Rhinerrhiza; Robiquetia; Rodriguezia; Rossioglossum (including Chelyorchis); Saccolabiopsis; Saccolabium; Sarcochilus; Scaphyglottis; Schoenorchis; Sobralia; Solenidium; Stanhopea ; Stelis; Taeniophyllum; Tetramicra; Thrixspermum; Tolumnia; Trachoma; Trichocentrum (including Cohniella and Lophiaris); Trichoglottis; Trichopilia; Trichotosia; Trizeuxis; Vanda (including Ascocentrum); Warmingia; Zygostates | 5–9 (Silvera et al., 2009) |
| Vanilloideae | Vanilla | 1 |
| Asphodelaceae | 1; in the ancestor of Alooideae and Bulbine | |
| Alooideae | Aloe; Aloidendron; Aristaloe; Astroloba (including Poellnitzia); Gasteria; Gonialoe; Haworthia; Haworthiopsis; Tulista | |
| ‘Asphodeloideae’1 | Bulbine | |
| Asparagaceae | ||
| Agavoideae | Agave (including Manfreda and Polianthes); Beschorneria; Furcraea; Hesperaloe; Yucca | 3; 1 in the ancestor of Agave, Beschorneria and Furcraea, 1 in Yucca sect. sarcocarpa, and 1 in Hesperaloe (Heyduk et al., 2022) |
| Nolinoideae | Beaucarnea, Sansevieria2 | 2; 1 each in Beaucarnea and Sansevieria (Meng et al., 2021) |
| Commelinales | ||
| Commelinaceae | ||
| Commelinoideae | Callisia; Cyanotis; Tradescantia; Tripogandra | 2; 1 in the ancestor of Callisia, Tradescantia and Tripogandra; and 1 in Cyanotis (Lee et al., 2021) |
| Poales | ||
| Bromeliaceae | ||
| Bromelioideae | Acanthostachys; Aechmea (including Streptocaylx); Ananas; Androlepis; Araeococcus; Billbergia; Bromelia; Canistropsis; Canistrum; Cryptanthus; Deinacanthon; Disteganthus; Edmundoa; Eduandrea; Forzzaea; Hohenbergia; Hohenbergiopsis; Hylaeaicum; Karawata; Lymania; Neoglaziovia; Neoregelia; Nidularium; Ochagavia; Orthophytum; Portea; Pseudananas; Pseudaraeococcus; Quesnelia; Ronnbergia; Sincoraea; Ursulaea; Wittrockia | 2–5 in Bromelioideae (Givnish et al., 2014) |
| Hechtioideae | Hechtia | 1 |
| Pitcairnioideae | Deuterocohnia; Dyckia; Encholirium | 1 |
| Puyoideae | Puya | 1–3 within Puya (Givnish et al., 2014) |
| Tillandsioideae | Guzmania; Josemania; Lemeltonia; Tillandsia; Werauhia | 1–5 (Barfuss et al., 2016) |
| Saxifragales | ||
| Crassulaceae | 1; CAM assumed ancestral to the Crassulaceae | |
| Crassuloideae | Crassula (including Rochea) | |
| Kalanchoideae | Adromischus; Cotyledon; Kalanchoë; Tylecodon | |
| Sempervivoideae | Aeonium (including Greenovia); Aichryson; Cremnophila; Dudleya; Echeveria; Graptopetalum; Hylotelephium; Lenophyllum; Monanthes; Orostachys; Pachyphytum; Rosularia; Sedum (including Diamorpha); Sempervivum; Umbilicus; Villadia | |
| Vitales | ||
| Vitaceae | ||
| Vitoideae | Cissus; Cyphostemma | 2; 1 each in Cissus and Cyphostemma (Wen et al., 2018) |
| Zygophyllales | ||
| Zygophyllaceae | ||
| Larreoideae | Bulnesia | 1 |
| Cucurbitales | ||
| Cucurbitaceae | Seyrigia; Xerosicyos | 2; 1 each in Seyrigia and Xerosicyos (Guo et al., 2020) |
| Rosales | ||
| Urticaceae | Pilea | 1 |
| Malphighiales | ||
| Clusiaceae | Clusia | 1–4 within Clusia (Luján et al., 2022) |
| Passifloraceae | Adenia | 1 |
| Euphorbiaceae | ||
| Euphorbiodeae | Euphorbia (including Monadenium, Pedilanthus and Synadenium) | 1–13 within Euphorbia (Horn et al., 2014) |
| Crotonoideae | Jatropha | 1 |
| Geraniales | ||
| Geraniaceae | Monsonia (including some members of Sarcocaulon); Pelargonium | 2–8; 1 in Monsonia and 1–7 in Pelargonium (Jones et al., 2003; García-Aloy et al., 2017; van de Kerke et al., 2019) |
| Caryophyllales | ||
| Aizoaceae | 1–4; CAM might be ancestral to Aizoaceae or has evolved independently in Aizooideae, Mesembryanthemoideae, Ruschioideae and Sesuvioideae, perhaps twice in Sesuvioideae (Klak et al., 2004, 2017a; Valente et al., 2014) | |
| Aizooideae | Tetragonia | |
| Mesembryanthemoideae | Mesembryanthemum (including Aptenia, Aridaria, Aspazoma, Brownanthus, Opophytum, Phyllobolus, Prenia, Psilocaulon, Sceletium, Sphalmanthus and Synaptophyllum) | |
| Ruschioideae | Antimima; Argyroderma; Astridia; Bergeranthus; Carpobrotus; Carruanthus; Cephalophyllum; Chasmatophyllum; Cheiridopsis; Conophytum; Delosperma; Disphyma; Dracophilus; Drosanthemopsis (including Anisocalyx); Drosanthemum Eberlanzia; Erepsia; Faucaria; Fenestraria; Glottiphyllum; Hartmanthus; Hereroa; Jacobsenia; Jordaaniella; Lampranthus; Lithops; Malephora; Meyerophytum; Mitrophyllum; Monilaria; Pleiospilos; Prepodesma; Psammophora; Rabiea; Rhinephyllum; Ruschia; Sarcozona; Schlechteranthus; Stoeberia; Titanopsis; Trichodiadema; Vanheerdea | |
| Sesuvioideae | Sesuvium; Trianthema | |
| Portulacineae | 1; CAM likely to be ancestral to the Portulacineae (Goolsby et al., 2018) | |
| Montiaceae | Australian Calandrinia3; Calyptridium; Cistanthe; Claytonia; Lewisia; Phemeranthus | |
| Didiereaceae | ||
| Didiereoideae | Alluaudia; Alluaudiopsis; Decarya; Didierea | |
| Portulacarioideae | Portulacaria (including Ceraria) | |
| Basellaceae | Anredera; Basella | |
| Halophytaceae | Halophytum | |
| Talinaceae | Talinum (including Talinella) | |
| Portulacaceae | Portulaca | |
| Anacampserotaceae | Anacampseros; Grahamia; Talinopsis | |
| Cactaceae 4 | ||
| Cactoideae | Acanthocereus (including some members of Peniocereus); Bergerocactus; Carnegiea; Cephalocereus (including Neobuxbaumia) Cereus (including Subpilocereus); Chamaecereus; Cleistocactus; Cochemiea (including some members of Mammillaria); Consolea; Copiapoa (including Pilocopiapoa); Disocactus; Echinocactus; Echinocereus; Echinopsis; Epiphyllum; Eriosyce; Eulychnia; Ferocactus; Haageocereus; Hatiora; Leucostele; Lobivia; Lophocereus; Lophophora; Mammillaria; Melocactus; Myrtillocactus; Oreocereus; Oroya; Pachycereus; Parodia; Pelecyphora; Pilosocereus; Polaskia; Rhipsalis; Schlumbergera (including Zygocactus); Sclerocactus; Selenicereus (including Hylocereus); Stenocereus (including Ritterocereus); Stetsonia; Trichocereus; Turbinicarpus | |
| Opuntioideae | Austrocylindropuntia; Cylindropuntia; Grusonia; Maihueniopsis; Opuntia (including Nopalea); Pereskiopsis; Pterocactus; Quiabentia; Tephrocactus | |
| Pereskioideae | Maihuenia; Pereskia5 | |
| Gentianales | ||
| Rubiaceae | ||
| Rubioideae | Hydnophytum; Myrmecodia; Squamellaria | 1–3; either 1 in the ancestor of Hydnophytineae or 1 in each genus (Chomicki and Renner, 2016) |
| Apocynaceae | ||
| Apocynoideae | Pachypodium | 1 |
| Asclepiadoideae | Apteranthes; Boucerosia (including Frerea); Caralluma; Caudanthera; Ceropegia6; Cynanchum (including Folotsia and Sarcostemma); Desmidorchis; Dischidia; Duvalia; Hoodia (including Trichocaulon); Hoya; Huernia; Orbea; Quaqua; Stapelia | 3; 1 each in Marsdenieae, Asclepiadeae and Ceropegieae (Wanntorp et al., 2014; Bruyns et al., 2017; Liede-Schumann et al., 2022) |
| Lamiales | ||
| Plantaginaceae | Littorella | 1 |
| Gesneriaceae | ||
| Didymocarpoideae | Haberlea; Ramonda | 1 |
| Gesnerioideae | Codonanthopsis | 1 |
| Lamiaceae | ||
| Lamioideae | Marrubium | 1 |
| Nepetoideae | Coleus (including some members of Plectranthus) | 1 |
| Asterales | ||
| Asteraceae | ||
| Asteroideae | Baculellum; Caputia; Crassothonna; Curio; Kleinia; Othonna; Senecio | 2 or 3; 1 in the Gynuroid clade and 1 or 2 in the Faujasia–Bethencourtia clade (Pelser et al., 2007; Ozerova et al., 2017) |
| Total known origins: 66–114+ |
1Asphodeloideae is polyphyletic, with Bulbine generally sister to Alooideae, as summarized by Smith and Figueiredo (2020).
2 Sansevieria has been proposed to be subsumed within Dracaena (Takawira-Nyenya et al., 2018).
3 Calandrinia is not monophyletic (Hancock et al., 2019), and CAM has been observed only in the monophyletic clade inclusive of all Australian members of Calandrinia sensu lato.
4All cacti are assumed CAM; the genera presented here are those with published data confirming CAM activity.
5We do not include Leuenbergeria but recognize that Pereskia is non-monophyletic.
6 Ceropegia has recently been shown to be polyphyletic, with Brachystelma and the stem-succulent stapeliads (including Apteranthes, Caralluma, Caudanthera, Boucerosia, Duvalia, Hoodia, Huernia, Orbea, Quaqua and Stapelia) nested within it. These genera have been proposed to be subsumed within an expanded and recircumscribed Ceropegia (Bruyns et al., 2017) but are still recognized by Kew (POWO, 2023) and other authorities (Endress et al., 2018),.