Hydrangea Gronov. ex L.

Hydrangea

Synonyms
  • Adamia Wall.
  • Broussaisia Gaudich.
  • Calyptranthe (Maxim.) Nakai
  • Cardiandra Siebold & Zucc.
  • Cianitis Reinw.
  • Cornidia Ruiz & Pav.
  • Decumaria L.
  • Deinanthe Maxim.
  • Dichroa Lour.
  • × Didrangea J.M.H.Shaw
  • Heteromalla (Rehder) H.Ohba & S.Akiyama
  • Hortensia Comm.
  • Macnemaraea P.Willemet
  • Pileostegia Hook.f. & Thomson
  • Platycrater Siebold & Zucc.
  • Sarcostyles C.Presl ex DC.
  • Schizophragma Siebold & Zucc.
  • Forsythia Walter [nom. illeg. homonym. post.]
  • Hydrangia L. [orth. var.]

Treatment and Overview

Hydrangea is the large and morphologically diverse flagship genus of the Hydrangea Family (Hydrangeaceae). Tree TSAR recognizes 111 non-hybrid living species, together with several named hybrids and accepted lower-rank entities. Most hydrangeas are familiar as deciduous or evergreen shrubs, but the genus also encompasses small trees, enormous root-climbing woody vines, perennial subshrubs, and genuinely herbaceous rhizomatous plants. That breadth is not an artifact of indiscriminate lumping. It reflects a modern phylogenetic understanding of Hydrangeeae in which plants once distributed among nine genera form a single evolutionary radiation best represented as a broadly circumscribed Hydrangea with substantial internal structure (De Smet et al. 2015; Yang et al. 2024).

The current Tree TSAR treatment is more inclusive at species rank than some global backbones. Plants of the World Online currently lists 98 accepted species under Hydrangea, whereas the Tree TSAR roster recognizes 111 non-hybrid species. The discrepancy reflects continuing differences in species delimitation, synonymy, and treatment of regional taxa rather than a major disagreement over the outer boundary of the genus. Species complexes in groups such as sect. Asperae, sect. Macrophyllae, and sect. Cornidia remain active areas of taxonomic research.

One genus from a collection of very different plants

The extraordinary variation within Hydrangea once appeared to support recognition of several separate genera. Broussaisia, Cardiandra, Decumaria, Deinanthe, Dichroa, Pileostegia, Platycrater, and Schizophragma were all maintained apart from a more narrowly defined Hydrangea. Molecular analyses demonstrated, however, that these lineages were nested among traditional hydrangeas, leaving Hydrangea sensu stricto extensively paraphyletic or polyphyletic when the satellite genera were retained. De Smet et al. (2015) therefore expanded Hydrangea to encompass the entire tribe Hydrangeeae and transferred much of the useful older generic structure to sectional rank.

A competing solution was proposed by Ohba and Akiyama (2016), who emphasized the conspicuous morphological and ecological differences among these lineages and divided broad Hydrangea into several genera, including Hortensia, Platycrater, Cornidia, Calyptranthe, and Heteromallae. Tree TSAR instead retains broad Hydrangea. The broad treatment resolves the demonstrated phylogenetic defect while allowing the same morphological and evolutionary diversity to be communicated through subgenera and sections. The most recent comprehensive phylogenomic treatment recovered five major clades and recognized five subgenera and 19 sections, providing a strong infrageneric framework without requiring renewed fragmentation of the genus (Yang et al. 2024).

This treatment also satisfies an important proportionality test. Many of the former genera remain excellent morphological concepts, but recognizing each at genus rank would convey little evolutionary information that cannot be retained more economically as infrageneric groups. The result is a genus whose diversity can initially seem surprising but becomes intelligible once its principal lineages are recognized.

Growth form: from woody climbers to rhizomatous herbs

Most Hydrangea species conform to Tree TSAR's Woody Tree, Shrub, or Vine growth-form category. This category includes not only the familiar erect shrubs and occasional small trees but also the climbing hydrangeas. Species derived from the former Decumaria, Pileostegia, and Schizophragma, together with the predominantly Neotropical sect. Cornidia, may climb high into forest canopies, but their persistent woody shoot systems make them architectural variants of the same basic woody growth form rather than a separate growth-form category. Some sect. Cornidia species are massive root climbers capable of ascending tens of metres through humid montane forest.

Subgenus Cardiandra marks a much deeper departure from that woody framework. Yang et al. (2024) recovered growth habit as one of the useful characters defining major infrageneric groups, with subshrub habit characteristic of sect. Cardiandra and herbaceous habit characteristic of sect. Deinanthe. Tree TSAR refines this distinction using perennial plant architecture. Members of sect. Cardiandra are treated as Perennial Subshrubs From Woody Bases: their enduring framework is woody or suffrutescent at or below the base, while the aerial shoots can behave much like those of herbaceous woodland perennials. This intermediate architecture helps explain why the same lineage has often been described as herbaceous in floristic and horticultural literature but as subshrubby in other treatments. The difference is substantially one of emphasis: seasonal appearance can resemble an herb, while the persistent perennial axis retains the woody character that separates the group from genuinely herbaceous hydrangeas.

Section Deinanthe carries that architectural reduction farther. Its two species, H. bifida and H. caerulea, are fundamentally herbaceous rhizomatous perennials whose aerial growth is renewed from persistent underground rhizomes rather than from a woody basal shoot framework. Tree TSAR therefore treats them as Perennial Root-Hardy Geophytes from Rhizomes. The contrast between sect. Cardiandra and sect. Deinanthe is consequently more precise than a simple division between “shrubs” and “herbs”: one lineage retains a reduced suffrutescent perennial framework, whereas the other has crossed into genuinely herbaceous rhizomatous geophytism.

Broad Hydrangea consequently spans three Tree TSAR growth-form categories: woody trees, shrubs, and vines; perennial subshrubs from woody bases; and rhizomatous perennial geophytes. The transition from a persistent woody framework through suffrutescent architecture to complete herbaceous shoot replacement is one of the clearest expressions of morphological diversification within the genus. It also provides a useful reminder that growth form can strongly diagnose internal lineages without necessarily defining generic boundaries.

Geography and evolutionary history

Hydrangea has a strikingly disjunct native distribution. Its greatest diversity lies in eastern and southeastern Asia, but native lineages also occur in Malesia, Hawaii, eastern and central North America, Mexico and Central America, and along the mountains of western South America. Plants of the World Online records a native range extending from China and the Kuril Islands through tropical Asia and the Hawaiian Islands, as well as from eastern North America southward into Central and South America.

Phylogenomic dating places the stem of Hydrangeeae at approximately 48.4 million years ago and the crown radiation at about 23 million years ago, near the Oligocene-Miocene transition. Its modern distribution developed through repeated episodes of dispersal, vicariance, climatic change, and regional isolation. Fossil and biogeographic evidence indicates that hydrangeas formerly occupied parts of the Northern Hemisphere from which they are now absent, helping explain the modern separation of major Asian and American centers of diversity (Yang et al. 2023).

This geographic history is reflected at much finer scales as well. Population and phylogenetic work in Japan, China, and the Neotropics has repeatedly uncovered lineage structure that was hidden beneath broadly interpreted morphological species. In sect. Asperae, molecular species-delimitation analyses have revealed both cryptic lineages and cases in which traditional morphotypes do not correspond cleanly to independently evolving units. Similar work in sect. Cornidia continues to uncover previously overlooked climbing species. The higher-level circumscription of Hydrangea is therefore much more stable than the boundaries of many of its constituent species (De Smet et al. 2017; Granados Mendoza et al. 2021).

Horticultural and Agricultural Uses

Few temperate woody genera have achieved the horticultural prominence of Hydrangea. Garden culture is dominated by a relatively small portion of the genus, especially H. macrophylla and its relatives, H. paniculata, H. arborescens, H. quercifolia, and a growing assortment of Asian species. Mophead and lacecap cultivars derived principally from the H. macrophylla complex established the popular image of a hydrangea, while breeding in H. paniculata and H. arborescens has greatly expanded the range of flower forms, plant sizes, and climatic tolerances available to gardeners.

Broad Hydrangea also reveals how incomplete that familiar horticultural image is. The former Schizophragma, Pileostegia, and Decumaria contribute spectacular woody climbers capable of covering trunks and large walls; less familiar Asian shrubs add foliage, indumentum, bark, and floral characters unlike those of common garden cultivars; and Subgenus Cardiandra extends the genus into the herbaceous perennial garden. The same phylogenetic breadth that once encouraged generic segregation thus provides horticulturists with growth forms ranging from large canopy vines to low rhizomatous woodland plants.

The famous relationship between soil conditions and flower color in H. macrophylla is also more chemically precise than the common maxim that acidic soil simply “turns hydrangeas blue.” Blue coloration depends on complexes involving delphinidin-derived anthocyanin pigments, copigments, and aluminum ions within the sepals. Soil acidity is important largely because it influences aluminum availability and uptake. Red, purple, and blue sepals are therefore alternative expressions of an interacting chemical system rather than direct color responses to soil pH alone (Yoshida et al. 2021).

Species diversity within the genus has become increasingly important to breeding. Wild taxa provide potential sources of cold hardiness, heat tolerance, disease resistance, reblooming behavior, plant architecture, and novel floral traits. At the same time, reproductive barriers, ploidy differences, and uneven cross-compatibility mean that horticultural relationships cannot be inferred simply from the ease with which species can be placed in the same genus. Broad Hydrangea is an evolutionary classification, not an assertion that all of its species are readily interfertile.

Conservation Issues

The extraordinary abundance of cultivated hydrangeas can give a misleading impression of security. Conservation concern is concentrated unevenly across the wild genus, particularly in narrowly endemic species, insular lineages, specialized montane habitats, and regions experiencing rapid forest loss.

Section Cornidia is especially instructive. Its species are predominantly root-climbing lianas of humid tropical and subtropical montane forests from Mexico through Central and South America, with one Asian lineage. Intensive fieldwork in Mexico and elsewhere has revealed substantial previously unrecognized diversity, including large climbing species that had escaped formal recognition despite their size. Mexican members studied by Samain and colleagues were threatened primarily by deforestation and land-use change, and later phylogenetic work demonstrated that the section contains considerably more evolutionary structure than older broad species concepts suggested (Samain et al. 2019; Granados Mendoza et al. 2021).

Taxonomic resolution is therefore directly relevant to conservation. A supposedly widespread species may prove to consist of several geographically restricted evolutionary units, radically changing the conservation significance of individual populations. Continued revision of sect. Cornidia illustrates how botanical exploration, species delimitation, and conservation assessment can become inseparable.

The herbaceous hydrangeas present a different conservation problem. Population-genomic work on the Deinanthe lineage found strong geographic structuring associated with climatic refugia, historical range contraction, and postglacial population history. Individual populations can therefore preserve distinct portions of the evolutionary history of these rhizomatous woodland plants. Conservation of the group requires attention not only to species survival but also to geographically structured genetic diversity (Sakaguchi et al. 2021).

Similar concerns recur elsewhere in Hydrangea: restricted island endemics, narrowly distributed Asian species, montane forest specialists, and newly separated members of difficult species complexes may each require conservation assessments that differ substantially from those of their widespread cultivated relatives. For a genus so familiar in gardens, substantial portions of its wild evolutionary diversity remain poorly represented in cultivation and incompletely studied in nature.

Infrageneric Groups

The current Tree TSAR classification follows the five-subgenus framework supported by Yang et al. (2024). These five subgenera contain 19 sections, preserving much of the morphological information once expressed through separate genera. Tree TSAR also standardizes common names for many infrageneric groups, particularly sections corresponding to lineages historically treated as genera, where established vernacular usage can be retained without obscuring the broader classification. This framework emphasizes the distinctiveness of the major lineages while keeping them within a phylogenetically coherent broad concept of Hydrangea.

Subgenus Decumaria contains sections Decumaria, Pileostegia, Schizophragma, and Sinenses. It contains several of the genus's best-developed woody climbing lineages, including plants historically familiar under the generic names Decumaria, Pileostegia, and Schizophragma.

Subgenus Heteromallae contains sect. Heteromallae. It is dominated by Asian woody shrubs and small trees and includes the lineage represented horticulturally by H. paniculata and its relatives.

Subgenus Cardiandra contains sections Cardiandra and Deinanthe. It represents the genus's most pronounced departure from ordinary woody architecture. Sect. Cardiandra consists of suffrutescent, rhizomatous plants treated by Tree TSAR as perennial subshrubs from woody bases, whereas sect. Deinanthe consists of genuinely herbaceous rhizomatous perennials. The contrast between these closely related growth strategies makes subg. Cardiandra one of the clearest demonstrations of architectural evolution within broad Hydrangea.

Subgenus Hydrangea contains sections Asperae, Calyptranthe, Cornidia, and Hydrangea. Its diversity ranges from familiar erect shrubs and North American species to temperate Asian climbers and the remarkable Neotropical canopy lianas of sect. Cornidia.

Subgenus Dichroa contains sections Broussaisia, Chinenses, Dichroa, Hirtae, Kawagoeanae, Kwangsienses, Macrophyllae, and Stylosae. It includes some of the genus's most familiar cultivated species, particularly the H. macrophylla complex, while also encompassing lineages once considered sufficiently distinctive to warrant separate generic recognition.

Together these groups reveal why modern Hydrangea is simultaneously broad and biologically coherent. Its major lineages are strongly differentiated in growth form, morphology, geography, and ecology, but those differences are better understood as the products of diversification within a single ancient radiation than as evidence that every distinctive branch requires a separate genus.

Accepted Taxa

Accepted taxa of Hydrangea
Hydrangea acuminata Siebold & Zucc. Sour hydrangea
Hydrangea acuminata var. australis (T.Yamaz.) S.Fujii, Sh.Murak. & M.Maki Southern sour hydrangea
Hydrangea acuminata var. yakushimensis (Yahara & Tagane) S.Fujii, Sh.Murak. & M.Maki Yakushima sour hydrangea
Hydrangea × agricola (J.M.H.Shaw) J.M.H.Shaw Crûg Farm herb hydrangea
Hydrangea albostellata Samain, Najarro & E.Martínez Downy sprawling hydrangea
Hydrangea alternifolia Siebold Japanese herb hydrangea
Hydrangea × amagiana Makino Izu hydrangea
Hydrangea amamiohsimensis (Koidz.) Y.De Smet & C.Granados Nansei herb hydrangea
Hydrangea ampla (Chun) Y.De Smet & C.Granados Windmill hydrangea vine
Hydrangea ampla Choufenianum Form Choufen hydrangea vine
Hydrangea ampla Megalocarpa Form Giant-bract hydrangea vine
Hydrangea anomala D.Don Chinese climbing hydrangea
Hydrangea arborescens L. Smooth hydrangea
Hydrangea arguta (Gaudich.) Y.De Smet & C.Granados Kanawao hydrangea
Hydrangea aspera Buch.-Ham. ex D.Don Roughleaf hydrangea
Hydrangea aspera ssp. robusta (Hook.f. & Thomson) E.M.McClint. Tall hydrangea
Hydrangea asterolasia Diels Costa Rican sprawling hydrangea
Hydrangea barbara (L.) Bernd Schulz Woodvamp hydrangea
Hydrangea bifida (Maxim.) Y.De Smet & C.Granados Two-lobed herb hydrangea
Hydrangea breedlovei Samain, Najarro & E.Martínez Cloudforest sprawling hydrangea
Hydrangea bretschneideri Dippel Mongolian hydrangea
Hydrangea caerulea (Stapf) Y.De Smet & C.Granados Blue herb hydrangea
Hydrangea carroniae Samain & E.Martínez Veracruz sprawling hydrangea
Hydrangea caucana Engl. Gland-tooth sprawling hydrangea
Hydrangea chungii Rehder Fujian hydrangea
Hydrangea cinerea Small Ashy hydrangea
Hydrangea coenobialis Chun Monastery hydrangea
Hydrangea corylifolia (Chun) Y.De Smet & C.Granados Hazel-leaf hydrangea vine
Hydrangea crassa (Hand.-Mazz.) Y.De Smet & C.Granados Leatherleaf hydrangea vine
Hydrangea crassa var. elliptica J.Anthony [TSAR]* Elliptic-leaf hydrangea vine
Hydrangea daimingshanensis (Y.C.Wu) Y.De Smet & C.Granados Daimingshan bluebead hydrangea
Hydrangea davidii Franch. David hydrangea
Hydrangea densifolia (C.F.Wei) Y.De Smet & C.Granados Taiwanese herb hydrangea
Hydrangea diplostemona (Donn.Sm.) Standl. Shrubby sprawling hydrangea
Hydrangea durifolia Briq. Santander sprawling hydrangea
Hydrangea fauriei (Hayata) Y.De Smet & C.Granados Taiwanese hydrangea vine
Hydrangea febrifuga (Lour.) Y.De Smet & C.Granados Common bluebead hydrangea
Hydrangea fistulosa (G.H.Huang & G.Hao) U.B.Deshmukh & M.B.Shende Hollowstem bluebead hydrangea
Hydrangea glaucescens (Rehder) Y.De Smet & C.Granados Pale hydrangea vine
Hydrangea goudotii Briq. Columbian sprawling hydrangea
Hydrangea gracilis W.T.Wang & M.X.Nie Slender hydrangea
Hydrangea grosseserrata Engl. Yakushima hydrangea
Hydrangea heteromalla D.Don Woolly hydrangea
Hydrangea hirsuta (Gagnep.) Y.De Smet & C.Granados Downy bluebead hydrangea
Hydrangea hirta (Thunb.) Siebold Nettleleaf hydrangea
Hydrangea hwangii J.M.H.Shaw Yunnan bluebead hydrangea
Hydrangea hydrangeoides (Siebold & Zucc.) Bernd Schulz Japanese hydrangea vine
Hydrangea hydrangeoides Concolor Form Greenback hydrangea vine
Hydrangea hydrangeoides Ulleungdo Form Ulleung hydrangea vine
Hydrangea hydrangeoides var. taquetii (H.Lév.) J.M.H.Shaw Jeju hydrangea vine
Hydrangea hydrangeoides var. yakushimensis (J.M.H.Shaw) J.M.H.Shaw Yakushima hydrangea vine
Hydrangea hypoglauca Rehder Whiteback hydrangea
Hydrangea integrifolia Hayata Philippine sprawling hydrangea
Hydrangea involucrata Siebold Bracted hydrangea
Hydrangea involucrata var. tokarensis M.Hotta & Shiuchi Nansei bracted hydrangea
Hydrangea jelskii Szyszyl. Andean sprawling hydrangea
Hydrangea kawagoeana Koidz. Tokara hydrangea
Hydrangea kawakamii Hayata Taiwan roughleaf hydrangea
Hydrangea kwangsiensis Hu Guangxi hydrangea
Hydrangea kwangtungensis Merr. Guangdong hydrangea
Hydrangea lalashanensis S.S.Ying Lalashan hydrangea
Hydrangea lingii G.Hoo Narrowleaf hydrangea
Hydrangea linkweiensis Chun Lingui hydrangea
Hydrangea liukiuensis Nakai Ryukyu bluebead hydrangea
Hydrangea lobbii Maxim. Philippine hydrangea
Hydrangea longifolia Hayata Longleaf hydrangea
Hydrangea longipes Franch. Longstalk hydrangea
Hydrangea luteovenosa Koidz. Sweet hydrangea
Hydrangea luteovenosa var. yakushimensis (Masam.) Sugim. Yakushima sweet hydrangea
Hydrangea macrocarpa Hand.-Mazz. Silverback hydrangea
Hydrangea macrophylla (Thunb.) Ser. Bigleaf hydrangea
Hydrangea macrosepala Hayata Large-sepal hydrangea
Hydrangea mangshanensis C.F.Wei Mangshan hydrangea
Hydrangea marunoi Tagane & S.Fujii Kimotsuki herb hydrangea
Hydrangea mathewsii Briq. Rusty sprawling hydrangea
Hydrangea medogensis X.Z.Lan, W.L.Zheng & W.Q.He Medog hydrangea
Hydrangea minamitanii (H.Ohba) Yahara Hyuga hydrangea
Hydrangea × mizushimarum H.Ohba Mizushima hydrangea
Hydrangea moellendorffii Hance Chinese herb hydrangea
Hydrangea mollissima (Merr.) Y.De Smet & C.Granados Hainan bluebead hydrangea
Hydrangea nahaensis Samain & E.Martínez Lacandon sprawling hydrangea
Hydrangea nebulicola Nevling & Gómez Pompa Mountain sprawling hydrangea
Hydrangea obtusifolia (Hu) Y.De Smet & C.Granados Chinese scrambling hydrangea
Hydrangea oerstedii Briq. Ruby sprawling hydrangea
Hydrangea ofeliae Sodusta & Lumawag Philippine bluebead hydrangea
Hydrangea otontepecensis Samain & E.Martínez Otontepec sprawling hydrangea
Hydrangea panamensis Standl. Panamanian sprawling hydrangea
Hydrangea paniculata Siebold Panicle hydrangea
Hydrangea peruviana Moric. ex Ser. Ecuadorian sprawling hydrangea
Hydrangea petiolaris Siebold & Zucc. Japanese climbing hydrangea
Hydrangea pingtungensis S.S.Ying Pingtung climbing hydrangea
Hydrangea platyarguta Y.De Smet & C.Granados Japanese tea hydrangea
Hydrangea pottingeri Prain Chinese hydrangea
Hydrangea preslii Briq. Peruvian sprawling hydrangea
Hydrangea quercifolia W.Bartram Oakleaf hydrangea
Hydrangea radiata Walter Silverleaf hydrangea
Hydrangea sargentiana Rehder Sargent hydrangea
Hydrangea scandens (L.f.) Ser. Deutzia hydrangea
Hydrangea schizomollis Y.De Smet & C.Granados Chinese hydrangea vine
Hydrangea schlimii Briq. Mophead sprawling hydrangea
Hydrangea seemannii L.Riley Mexican sprawling hydrangea
Hydrangea serrata (Thunb.) Ser. Mountain hydrangea
Hydrangea serrata var. angustata (Franch. & Sav.) H.Ohba Narrowleaf mountain hydrangea
Hydrangea serratifolia (Hook. & Arn.) F.Phil. Sawtooth sprawling hydrangea
Hydrangea sikokiana Maxim. Shikoku hydrangea
Hydrangea sousae Samain, Najarro & E.Martínez Chiapas sprawling hydrangea
Hydrangea stenophylla Merr. & Chun Willowleaf hydrangea
Hydrangea steyermarkii Standl. Tacaná sprawling hydrangea
Hydrangea strigosa Rehder Late hydrangea
Hydrangea stylosa Hook.f. & Thomson Himalayan hydrangea
Hydrangea sungpanensis Hand.-Mazz. Songpan hydrangea
Hydrangea taiwaniana Y.C.Liu & F.Y.Lu Taiwan sprawling hydrangea
Hydrangea tapalapensis Samain, Najarro & E.Martínez Tapalapa sprawling hydrangea
Hydrangea tarapotensis Briq. Tarapoto sprawling hydrangea
Hydrangea tomentella (Hand.-Mazz.) Y.De Smet & C.Granados Rusty evergreen climbing hydrangea
Hydrangea trianae Briq. Leatherleaf sprawling hydrangea
Hydrangea × versicolor (Fortune) J.M.H.Shaw Sapphire hydrangea
Hydrangea viburnoides (Hook.f. & Thomson) Y.De Smet & C.Granados Chinese evergreen climbing hydrangea
Hydrangea villosa Rehder Velvetleaf hydrangea
Hydrangea vinicolor Chun Wineleaf hydrangea
Hydrangea wallichii J.M.H.Shaw Himalayan bluebead hydrangea
Hydrangea weberbaueri Engl. Matapalo sprawling hydrangea
Hydrangea xanthoneura Diels Nodding hydrangea
Hydrangea xinfeniae W.B.Ju & J.Ru Xinfen bluebead hydrangea
Hydrangea yaoshanensis (Y.C.Wu) Y.De Smet & C.Granados Luomeng bluebead hydrangea
Hydrangea yayeyamensis Koidz. Nansei hydrangea
Hydrangea yesoensis Koidz. Hime hydrangea
Hydrangea × ytiensis (J.M.H.Shaw) J.M.H.Shaw Y Tý hydrangea
Hydrangea zhewanensis P.S.Hsu & X.P.Zhang Zhejiang hydrangea

Additional Information

References and Further Reading

De Smet Y, De Clerck O, Uemachi T, Granados Mendoza C, Wanke S, Goetghebeur P, Samain M-S (2017) Multilocus coalescent species delimitation to evaluate traditionally defined morphotypes in Hydrangea sect. Asperae (Hydrangeaceae). Molecular Phylogenetics and Evolution 114: 415–425. https://doi.org/10.1016/j.ympev.2017.05.021 (opens in a new tab)

De Smet Y, Granados Mendoza C, Wanke S, Goetghebeur P, Samain M-S (2015) Molecular phylogenetics and new (infra)generic classification to alleviate polyphyly in tribe Hydrangeeae (Cornales: Hydrangeaceae). Taxon 64(4): 741–753. https://doi.org/10.12705/644.6 (opens in a new tab)

Granados Mendoza C, Martínez Salas EM, Goetghebeur P, Wanke S, Samain M-S (2021) Molecular phylogeny, character evolution, and biogeography of Hydrangea section Cornidia, Hydrangeaceae. Frontiers in Plant Science 12: 661522. https://doi.org/10.3389/fpls.2021.661522 (opens in a new tab)

McClintock E (1957) A monograph of the genus Hydrangea. Proceedings of the California Academy of Sciences, Fourth Series 29(5): 147–256. https://www.biodiversitylibrary.org/part/51707 (opens in a new tab) https://doi.org/10.5281/zenodo.15905786 (opens in a new tab)

Ohba H, Akiyama S (2016) Generic segregation of some sections and subsections of the genus Hydrangea (Hydrangeaceae). Journal of Japanese Botany 91(6): 345–350. https://doi.org/10.51033/jjapbot.91_6_10703 (opens in a new tab)

Sakaguchi S, Asaoka Y, Takahashi D, Isagi Y, Imai R, Nagano AJ, Qiu Y-X, Li P, Lu R, Setoguchi H (2021) Inferring historical survivals of climate relicts: The effects of climate changes, geography, and population-specific factors on herbaceous hydrangeas. Heredity 126(4): 615–629. https://doi.org/10.1038/s41437-020-00396-4 (opens in a new tab)

Samain M-S, Hernández Najarro F, Martínez Salas EM (2019) The climbing hydrangeas (Hydrangeaceae) of Mexico, including description of six (critically) endangered new species. Acta Botanica Mexicana 126: e1463. https://doi.org/10.21829/abm126.2019.1463 (opens in a new tab)

Yang X, Xue T, Gao T, Liang Y, Smets EF, Gadagkar SR, Yu S (2024) An updated infrageneric classification based on phylogenomics and character evolution in Hydrangea (Hydrangeaceae). Taxon 73(2): 503–518. https://doi.org/10.1002/tax.13158 (opens in a new tab)

Yang X, Zhang X, Xue T, Zhang X, Yang F, Yu J, Janssens SB, Bussmann RW, Yu S (2023) Phylogenomics and historical biogeography of Hydrangeeae (Hydrangeaceae) elucidate the effects of geologic and climatic dynamics on diversification. Proceedings of the Royal Society B: Biological Sciences 290(2003): 20230659. https://doi.org/10.1098/rspb.2023.0659 (opens in a new tab)

Yoshida K, Oyama K, Kondo T (2021) Insight into chemical mechanisms of sepal color development and variation in hydrangea. Proceedings of the Japan Academy, Series B 97(2): 51–68. https://doi.org/10.2183/pjab.97.003 (opens in a new tab)