List of Ericaceae genera

This is a list of genera in the plant family Ericaceae, which includes the heaths, heathers, epacrids, and blueberries. As currently circumscribed, the family contains about 4000 species into more than 120 genera classified into 9 subfamilies.[1]

Bell heather, Erica cinerea, is the type species of the family.

Current understanding

The list shown here follows the phylogenetic classification of Kron et al. (2002),[2] with modifications as per Stevens et al. (2004),[3] Quinn et al. (2005),[4] Albrecht et al. (2010),[5] Gillespie & Kron (2010),[6] and Craven (2011).[7]

Nonetheless, much of the taxonomy within the Vaccinioideae is in flux. Multiple studies of the Gaultherieae have now shown Pernettya, Diplycosia (already including Pernettyopsis), and Tepuia to be nested within Gaultheria,[8][9] although publication formalizing the transfers are yet to be published. Similarly, most of Vaccinieae has not been investigated using molecular techniques, and many genera are likely paraphyletic or polyphyletic. Neotropical and Old World Vaccinioideae are particularly poorly known.[10] Within the Epacrideae preliminary evidence supports the lumping of Budawangia and Rupicola with Epacris,[2] but these genera still stand for the time being.

Relevant recent synonymy include Bruckenthalia with Erica,[11] Leiophyllum and Loiseleuria with Kalmia,[12] and Diplarche and Menziesia with Rhododendron.[7]

Genera

Genus Authority Year Subfamily / Tribe Type species # of species Distribution
Enkianthus Lour.[13]1791EnkianthoideaeEnkianthus quinqueflorus Lour.16Southeast Asia
Chimaphila Pursh[14]1813 ("1814")PyroloideaeChimaphila maculata (L.) Pursh5North and Central America, Eurasia
Moneses Salisb. ex S.F.Gray[15]1821PyroloideaeMoneses grandiflora[N 1] Salisb. ex S.F.Gray1Circumboreal
Orthilia Raf.[16]1840PyroloideaeOrthilia parvifolia[N 1] Raf.1Canada to Guatemala, Eurasia
Pyrola L.[17]1753PyroloideaePyrola rotundifolia L.ca. 30North and Central America, Eurasia
Allotropa Torr. & A.Gray[18]1858 ("1857")Monotropoideae / MonotropeaeAllotropa virgata Torr. & A.Gray1Pacific Northwest
Cheilotheca Hook.f.[19]1876Monotropoideae / MonotropeaeCheilotheca khasiana Hook.f.3
Hemitomes A.Gray[18]1858 ("1857")Monotropoideae / MonotropeaeHemitomes congestum A.Gray1West Coast
Monotropa L.[17]1753Monotropoideae / MonotropeaeMonotropa uniflora L.2America, Europe, Asia
Monotropastrum H.Andres[20]1936Monotropoideae / MonotropeaeMonotropastrum macrocarpum H.Andres2 or 3Himalaya to Sumatra and Japan
Monotropsis Schwein.[21]1817Monotropoideae / MonotropeaeMonotropsis odorata Schwein.1Southeastern United States
Pityopus Small[22]1914Monotropoideae / MonotropeaePityopus oregonus[N 2] Small1West Coast of the United States
Pleuricospora A.Gray[23]1868Monotropoideae / MonotropeaePleuricospora fimbriolata A.Gray1California to British Columbia
Pterospora Nutt.[24]1919Monotropoideae / PterosporeaePterospora andromedea Nutt.1United States and Mexico
Sarcodes Torr.[25]1851Monotropoideae / PterosporeaeSarcodes sanguinea Torr.1West Coast of United States and Mexico
Arbutus L.[17]1753ArbutoideaeArbutus unedo L.10West Coast of Canada to Central America, Mediterranean area
Arctostaphylos Adans.[26]1763ArbutoideaeArctostaphylos uva-ursi (L.) Spreng.66North and Central America, Eurasia
Comarostaphylis Zucc.[27]1837ArbutoideaeComarostaphylis arguta[N 3] Zucc.10California to Central America
Ornithostaphylos Small[22]1914ArbutoideaeOrnithostaphylos oppositifolia (Parry) Small1California, Mexico
Xylococcus Nutt.[28]1842 ("1843")ArbutoideaeXylococcus bicolor Nutt.1California
Cassiope D.Don[29]1834CassiopeoideaeCassiope tetragona (L.) D.Don18Circumboreal
Bryanthus J.G.Gmel.[30][N 4]1769Ericoideae / BryantheaeBryanthus gmelinii[N 5] D.Don
Ledothamnus Meisn.[31]1863Ericoideae / BryantheaeLedothamnus guyanensis Meisn.
Ceratiola Michx.[32]1803Ericoideae / EmpetreaeCeratiola ericoides Michx.1United States
Corema D.Don[33]1826Ericoideae / EmpetreaeCorema album (L.) D.Don ex Steud.2North America, Iberian Peninsula
Empetrum L.[17]1753Ericoideae / EmpetreaeEmpetrum nigrum L.3–18[N 6]Circumboreal and southern South America
Calluna Salisb.[36]1802Ericoideae / EriceaeCalluna vulgaris (L.) Hull1European
Daboecia D.Don[29]1834Ericoideae / EriceaeDaboecia polifolia[N 7] D.Don2Europe
Erica L.[17]1753Ericoideae / EriceaeErica cinerea L.850+Afro-Eurasia
Bejaria[N 8] Mutis[37]1771Ericoideae / PhyllodoceaeBejaria aestuans Mutis ex L.15South America to United States
Elliottia Muhl. ex Elliott[21]1817Ericoideae / PhyllodoceaeElliottia racemosa Muhl. ex Elliott4North America, Japan
Epigaea L.[17]1753Ericoideae / PhyllodoceaeEpigaea repens L.3North America, Asia
Kalmia L.[17]1753Ericoideae / PhyllodoceaeKalmia latifolia L.10Northern hemisphere
Kalmiopsis Rehder[38]1932Ericoideae / PhyllodoceaeKalmiopsis leachiana (L.F.Hend.) Rehder2Oregon
Phyllodoce Salisb.[39]1806Ericoideae / PhyllodoceaePhyllodoce taxifolia[N 9] (Pall.) Salisb.8North America, Eurasia
Rhodothamnus Rchb.[40]1827Ericoideae / PhyllodoceaeRhodothamnus chamaecistus (L.) Rchb.
Rhododendron L.[17]1753Ericoideae / RhodoreaeRhododendron ferrugineum L.1000+North America, Eurasia, Australia
Therorhodion[N 10] (Maxim.) Small[43]1914Ericoideae / RhodoreaeTherorhodion camtschaticum (Pall. Small)3Alaska, Northeast Asia
Harrimanella Coville[44]1901HarrimanelloideaeHarrimanella stelleriana (Pall.) Coville2Circumboreal
Archeria Hook.f.[45]1857Styphelioideae / ArcherieaeNone designated6Tasmania, New Zealand
Andersonia R.Br.[46]1810Styphelioideae / CosmelieaeNone designatedca. 50Western Australia
Cosmelia R.Br.[46]1810Styphelioideae / CosmelieaeCosmelia rubra R.Br.2Western Australia
Sprengelia Sm.[47]1794Styphelioideae / CosmelieaeSprengelia incarnata4Australia
Budawangia Telford[48]1992Styphelioideae / EpacrideaeBudawangia gnidioides (Summerh.) Telford1Australia
Epacris Cav.[49]1797Styphelioideae / EpacrideaeEpacris longiflora Cav.30+Oceania
Lysinema R.Br.[46]1810Styphelioideae / EpacrideaeNone designated6Australia
Rupicola Maiden & Betche[50]1899 ("1898")Styphelioideae / EpacrideaeRupicola sprengelioides Maiden & Betche4Australia
Woollsia F.Muell.[51]1873Styphelioideae / EpacrideaeWoollsia pungens (Cav.) F.Muell.1Australia
Dielsiodoxa Albr.[5]2010Styphelioideae / OligarrheneaeDielsiodoxa tamariscina (F.Muell.) Albr.5Australia
Needhamiella[N 11] L.Watson[53]1965Styphelioideae / OligarrheneaeNeedhamiella pumilio (R.Br.) L.Watson1Western Australia
Oligarrhena R.Br.[46]1810Styphelioideae / OligarrheneaeOligarrhena micrantha R.Br.1Australia
Lebetanthus[N 12] Endl.[54]1841Styphelioideae / PrionoteaeLebetanthus americana[N 13] (Hook.f.) Hook.f.1South America
Prionotes R.Br.[46]1810Styphelioideae / PrionoteaePrionotes cerinthoides (Labill. R.Br.)1Tasmania
Dracophyllum Labill.[56]1800Styphelioideae / RicheeaeDracophyllum verticillatum Labill.ca. 100Oceania
Richea[N 14] R.Br.[46]1810Styphelioideae / RicheeaeRichea dracophylla R.Br.11Australia, Tasmania
Sphenotoma (R.Br.) Sweet[N 15][57]1828Styphelioideae / RicheeaeSphenotoma gracilis (R.Br.) Sweet6
Acrothamnus Quinn[4]2005Styphelioideae / StyphelieaeAcrothamnus maccraei (F.Muell.) Quinn5+Oceania
Acrotriche R.Br.[46]1810Styphelioideae / StyphelieaeAcrotriche divaricata[N 16] R.Br.16Australia
Agiortia Quinn[4]2005Styphelioideae / StyphelieaeAgiortia cicatricata (J.M.Powell) Quinn3Australia
Androstoma Hook.f.[45]1844Styphelioideae / StyphelieaeAndrostoma empetrifolia Hook.f.2Australia, New Zealand
Astroloma R.Br.[46]1810Styphelioideae / Styphelieaenone designatedca. 10Australia
Brachyloma Sond.[59]1845Styphelioideae / StyphelieaeBrachyloma preissii Sond.7Australia
Coleanthera Stschegl.[60]1859Styphelioideae / StyphelieaeNone designated
Conostephium Benth.[61]1837Styphelioideae / StyphelieaeConostephium pendulum Benth.ca. 10Australia
Croninia J.M.Powell[62]1993Styphelioideae / StyphelieaeCroninia kingiana (F.Muell.) J.M.Powell1Australia
Cyathodes Labill.[63]1805 (1804)Styphelioideae / StyphelieaeCyathodes glauca[N 17] Labill.5(?)Australia
Cyathopsis Brongn. & Gris[65][66][N 18]1864Styphelioideae / StyphelieaeCyathopsis floribunda Brongn. & Gris3New Caledonia
Decatoca F.Muell.[68]1889Styphelioideae / StyphelieaeDecatoca spencerii1New Guinea
Leptecophylla C.M.Weiller[69]1999Styphelioideae / StyphelieaeLeptecophylla juniperina (J.R.Forst. & G.Forst.) C.M.Weiller12Oceania
Leucopogon R.Br.[46]1810Styphelioideae / StyphelieaeLeucopogon lanceolatus[N 19]150+Oceania
Lissanthe R.Br.[46]1810Styphelioideae / StyphelieaeLissanthe strigosa[N 20] (Sm.) R.Br.7Australia
Melichrus R.Br.[46]1810Styphelioideae / StyphelieaeMelichrus procumbens (Cav.) Druceca. 10Australia
Monotoca R.Br.[46]1810Styphelioideae / StyphelieaeMonotoca scoparia (Sm.) R.Br.17Australia
Montitega C.M.Weiller[5]1993Styphelioideae / StyphelieaeMontitega dealbata (R.Br.) C.M.Weiller1Australia
Pentachondra R.Br.[46]1810Styphelioideae / StyphelieaeNone designated[N 21]4–5Australia, New Zealand
Planocarpa C.M.Weiller[71]1996Styphelioideae / StyphelieaePlanocarpa petiolaris (DC.) C.M.Weiller3Tasmania
Styphelia Sm.[72]1795 ("1793")Styphelioideae / StyphelieaeStyphelia tubuliflora Sm.Oceania
Trochocarpa R.Br.[46]1810Styphelioideae / StyphelieaeTrochocarpa laurina (Rudge R.Br.)10?Australia, Malesia
Andromeda L.[17]1753Vaccinioideae / AndromedeaeAndromeda polifolia L.1 or 2Circumboreal
Zenobia D.Don[29]1834Vaccinioideae / AndromedeaeZenobia speciosa[N 22] (Michx.) D.Don 1Southeastern United States
Chamaedaphne Moench[73]1794Vaccinioideae / GaultherieaeChamaedaphne calyculata (L.) Moench1Circumboreal
Diplycosia Blume[74]1826Vaccinioideae / GaultherieaeNone designated[N 23]ca. 110Southeast Asia
Eubotrys Nutt.[28]1842 ("1843")Vaccinioideae / GaultherieaeEubotrys racemosa (L.) Nutt.2Eastern United States
Gaultheria L.[17]1753Vaccinioideae / GaultherieaeGaultheria procumbens L.ca. 115Circumpacific, Americas
Leucothoe D.Don[29]1834Vaccinioideae / GaultherieaeLeucothoe axillaria (Lam.) D.Don5United States, East Asia
Pernettya[N 24] Gaudich.[75]1825Vaccinioideae / GaultherieaePernettya empetrifolia[N 25] ([[[Jean-Baptiste Lamarck|Lam.]]) Gaudich.Southern America, Oceania
Tepuia Camp[76]1939Vaccinioideae / GaultherieaeTepuia tatei[N 26] Camp7South America
Agarista D.Don ex G.Don[78]1834Vaccinioideae / LyonieaeAgarista nummularia (Cham. & Schltdl.) G.Donca. 30Americas, Africa, Madagascar
Craibiodendron W.W.Sm.[79]1911Vaccinioideae / LyonieaeCraibiodendron shanicum[N 27] W.W.Sm.5India, China, Indochina
Lyonia Nutt.[24]1919Vaccinioideae / LyonieaeLyonia ferruginea (Walter) Nutt.35United States to Mexico and Caribbeans, East Asia
Pieris D.Don[29]1834Vaccinioideae / LyonieaePieris formosa (Wall.) D.Don7United States, Cuba, East Asia
Oxydendrum DC.[80]1839Vaccinioideae / OxydendreaeOxydendrum arboreum (L.) DC.1United States
Agapetes D.Don ex G.Don[78]1834Vaccinioideae / VaccinieaeAgapetes setigera (Wall.) D.Don ex G.Don80+Himalaya
Anthopteropsis A.C.Sm.[81]1941Vaccinioideae / VaccinieaeAnthopteropsis insignis A.C.Sm.1Panama
Anthopterus Hook.[82]1839Vaccinioideae / VaccinieaeAnthopterus racemosus Hook.ca. 20Neotropical
Cavendishia Lindl.[83]1835 ("1836")Vaccinioideae / VaccinieaeCavendishia nobilis Lindl.ca. 100Neotropical
Ceratostema Juss.[84]1789Vaccinioideae / VaccinieaeCeratostema peruvianum J.F.Gmel.35Andes
Costera J.J.Sm.[85]1910Vaccinioideae / VaccinieaeCostera ovalifolia J.J.Sm.10
Demosthenesia A.C.Sm.[86]1936Vaccinioideae / VaccinieaeDemosthenesia mandonii (Britton) A.C.Sm.12Neotropical
Didonica Luteyn & Wilbur[87]1977Vaccinioideae / VaccinieaeDidonica pendula Luteyn & Wilbur4Neotropical
Dimorphanthera[N 28]N/AVaccinioideae / VaccinieaeNone designated80+Malesia to Australia
Diogenesia Sleumer[88]1934Vaccinioideae / VaccinieaeDiogenesia octandra Sleumer13Eastern Venezuela, Northern Andes
Disterigma (Klotzsch) Nied.[89]1889 ("1890")Vaccinioideae / VaccinieaeDisterigma empetrifolium (Kunth) Drudeca. 40Mexico to Northern South America
Gaylussacia Kunth[90]1819Vaccinioideae / VaccinieaeGaylussacia buxifolia Kunth50Americas
Gonocalyx Planch. & Linden[91]1856Vaccinioideae / VaccinieaeGonocalyx pulcher Planch. & Linden11Southern America
Lateropora A.C.Sm.[92]1932[N 29]Vaccinioideae / VaccinieaeLateropora ovata A.C.Sm.3
Macleania Hook.[82]1837Vaccinioideae / VaccinieaeMacleania floribunda Hook.ca. 50Southern Mexico to Peru
Mycerinus A.C.Sm.[93]1931Vaccinioideae / VaccinieaeMycerinus sclerophyllus A.C.Sm.3Southern America
Notopora Hook.f.[94]1873Vaccinioideae / VaccinieaeNotopora schomburgkii[N 30] Hook.f.5
Oreanthes Benth.[95]1844Vaccinioideae / VaccinieaeOreanthes buxifolius Benth.7Neotropical
Orthaea Klotzsch[96]1851Vaccinioideae / VaccinieaeOrthaea secundiflora (Poepp. & Endl.) Klotzsch34Neotropical
Paphia Seem.[97]1864Vaccinioideae / VaccinieaePaphia vitiensis Seem.20+Australasia
Pellegrinia Sleumer[98]1935Vaccinioideae / VaccinieaePellegrinia grandiflora (Ruiz & Pav. ex G.Don) Sleumer6
Plutarchia A.C.Sm.[86]1936Vaccinioideae / VaccinieaePlutarchia rigida (Benth.) A.C.Sm.11Ecuador, Colombia
Polyclita A.C.Sm.[86]1936Vaccinioideae / VaccinieaePolyclita turbinata (Kuntze) A.C.Sm.1
Psammisia Klotzsch[96]1851Vaccinioideae / VaccinieaePsammisia falcata Klotzsch70Neotropical
Rusbya Britton[99]1893Vaccinioideae / VaccinieaeNone designated[N 31]1Bolivia
Satyria Klotzsch[96]1851Vaccinioideae / VaccinieaeSatyria warszewiczii Klotzsch
Semiramisia Klotzsch[96]1851Vaccinioideae / VaccinieaeSemiramisia speciosa (Benth.) Klotzsch4Ecuador, Venezuela to Peru
Siphonandra[N 32] Klotzsch[96]1851Vaccinioideae / VaccinieaeSiphonandra elliptica (Ruiz & Pav. ex G.Don) Klotzsch3Neotropical
Sphyrospermum Poepp. & Endl.[102]1835Vaccinioideae / VaccinieaeSphyrospermum buxifolium Poepp. & Endl.22Neotropical
Symphysia C.Presl[103]1827Vaccinioideae / VaccinieaeSymphysia martinicensis[N 33] C.Presl
Themistoclesia Klotzsch[96]1851Vaccinioideae / VaccinieaeThemistoclesia pendula[N 34]22Costa Rica to Peru
Thibaudia Ruiz & Pav. ex J.St.-Hil.[104][N 35]1805Vaccinioideae / VaccinieaeThibaudia mellifera[N 36] Ruiz & Pav. ex J.St.-Hil.60Neotropical
Utleya Wilbur & Luteyn[87]1977Vaccinioideae / VaccinieaeUtleya costaricensis Wilbur & Luteyn1
Vaccinium L.[17]1753Vaccinioideae / VaccinieaeVaccinium uliginosum L.500+Near cosmopolitan

Notes

  1. As unnecessary renamings of species described earlier by Carl Linnaeus, these names are illegitimate and treated respectively as Moneses uniflora (L.) A.Gray, Orthilia secunda (L.) House, and Phyllodoce caerulea (L.) Bab.
  2. Small's species, which he originally named Pityopus oregona, was later found to be the same as Alice Eastwood's Monotropa californica. It is now treated as Pityopus californius (Eastw.) H.F.Copel.
  3. Currently treated as a synonym of Comarostaphylis discolor (Hook.) Diggs.
  4. The name is often attributed to David Don.[29] Although no valid species name were published in the two last volumes of Gmelin's work (which did not use the Linnean binomial system), his genera are considered valid.
  5. This species' correct name is Bryanthus musciformis (Poir.) Nakai
  6. Murray et al. (2009)[34] note that there are no recent comprehensive reviews of this genus "where different authors rarely, if ever, arrive at the same conclusion." (Hultén, 1971)[35]
  7. The correct name of the species is Daboecia cantabrica (Huds.) K.Koch
  8. As Befaria. The corrected spelling was later conserved.
  9. John Lindley and Joseph Paxton had originally moved Pallas' Rhododendron camtschaticum to a new genus Rhodothamnus,[41] a name that Reichenbach had already coined for another Ericacea genus.[40] Small elevated Carl Maximowicz' Rhododendron sect. Therorhodion[42] to generic level to replace that name.
  10. Robert Brown's original Needhamia[46] could not be used because Needhamia Scop.,[52] a rejected synonym of Tephrosia (Fabaceae) antedates it.
  11. Endlicher substituted this name (as "Lebethanthus") for his own Allodape,[54] which he considered preoccupied by Allodape Lepeletier & Serville 1825,[55] a genus of bees. Since homonymy with zoological name is irrelevant under the ICN (Art. 54.1), preserving the usage of the later name (with corrected spelling) required its conservation.
  12. Now included within Lebetanthus myrsinites (Lam.) Dusén.
  13. Sometimes misattributed to Labillardière,[56] but that name is a rejected name against Brown's, and a synonym of Craspedia (Asteraceae:Gnaphalieae).
  14. Not "R.Br. ex Sweet" or just "Sweet". Brown's Dracophyllum [unranked] Sphenotoma[46] had no rank attached. Although such names cannot affect priority, they can still serve as basionyms so long as they are validly published (ICBN, art. 35).
  15. Not recorded in ING (Index Nominum Genericorum), but see Paterson (1960)[58]
  16. Not recorded in ING, but see Weiller (1996)[64]
  17. ING and IPNI cite both journals but the issue of the Bulletin was not published before late January 1865.[67]
  18. An illegitimate renaming of Henry Cranke Andrews Styphelia parviflora. The correct name is Leucopogon parviflorus (Andrews) Lindl.
  19. Not recorded in APNI or ING, but see Crayn et al. (2003)[70]
  20. This would have to be either Pentachondra involucrata or Pentachondra pumilla.
  21. The genus' sole species is now known as Zenobia pulverulanta (W.Bartram ex Willd.) Pollard
  22. This would be either Diplycosia pilosa, D. heterophylla or D. latifolia (≡D. heterophylla var. latifolia).
  23. Gaudichaud originally spelled the name, which honors Antoine-Joseph Pernety, as "Pernettia". The spelling was later conserved with "y".
  24. The correct name in Pernettya is Pernettya pumila (L.f.) Hook.. Although the species is known to belong in Gaultheria, it currently lacks a name in that genus (Gaultheria pumila Hook. already exists).
  25. Although Tepuia species now belong to Gaultheria, this species currently lacks a valid name in the latter genus because of the existence of Gaultheria tatei A.C.Sm..[77]
  26. Currently treated as a synonym of Craibiodendron stellatum (Pierre) W.W.Sm.
  27. Authorship, date and proper place of publication of this genus are listed in a number of different, contradictory ways.
  28. A printing error misstates the year as "1942".
  29. Hooker also listed the species with the epithet "schomburgkiana".
  30. This would have to be Rusbya taxifolia, as the other of Britton's original species is currently treated as Demosthenesia pearcei (Britton) A.C.Sm. and its designation as lectotype would replace Demosthenesia with Rusbya.
  31. Because the name is preceded by Siphonandra Turcz.,[100] it has been proposed for conservation.[101]
  32. A synonym of Symphysia racemosa (Vahl) Stearn
  33. A synonym for Themistoclesia dependens (Benth.) A.C.Sm.
  34. The fourth and fifth volumes of Ruiz and Pavón's Flora Peruviana, et Chilensis were not published during their time, but the plates that were to illustrate volume 4 were circulated and lead to the publication of some names.[105]
  35. It is not clear whether the correct spelling is "mellifera" or "melliflora".

References

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  3. Stevens, P.F.; et al. (2004). "Ericaceae". In Klaus Kubitzki (ed.). The Families and Genera of Vascular Plants. 6. Berlin: Springer. pp. 145–194. ISBN 978-3-540-06512-8.
  4. Quinn, Christopher J.; et al. (2005). "Generic concepts in Styphelieae (Ericaceae): the Cyathodes group". Australian Systematic Botany. 18 (5): 439–454. doi:10.1071/SB05005.
  5. Albrecht, D.E.; et al. (2010). "Generic concepts in Ericaceae: Styphelioideae: the Monotoca group". Australian Systematic Botany. 23 (5): 320–332. doi:10.1071/SB10009.
  6. Gillespie, Emily; Kron, Kathleen (2010). "Molecular phylogenetic relationships and a revised classification of the subfamily Ericoideae (Ericaceae)". Molecular Phylogenetics and Evolution. 56 (2): 343–354. doi:10.1016/j.ympev.2010.02.028. PMID 20193767.
  7. Craven, L.A. (2011). "Diplarche and Menziesia transferred to Rhododendron (Ericaceae)". Blumea. 56 (1): 33–35. doi:10.3767/000651911X568594.
  8. Powell, E. Ann; Kron, Kathleen A. (2001). "An Analysis of the Phylogenetic Relationships in the Wintergreen Group (Diplycosia, Gaultheria, Pernettya, Tepuia; Ericaceae)". Systematic Botany. 26 (4): 808–817. doi:10.1043/0363-6445-26.4.808 (inactive 2021-01-15). JSTOR 3093862.CS1 maint: DOI inactive as of January 2021 (link)
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