Primulaceae

Life-forms

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Cladus: Asterids
Ordo: Ericales

Familia: Primulaceae
Subfamiliae: Maesoideae – Myrsinoideae – Primuloideae – Theophrastoideae
Genera

Aegiceras – Amblyanthopsis – Amblyanthus – Androsace – Antistrophe – Ardisia – Ardisiandra – Badula – Bonellia – Bryocarpum – Clavija – Conandrium – Coris – Ctenardisia – Cybianthus – Cyclamen – Deherainia – Dionysia – Discocalyx – Dodecatheon – Elingamita – Embelia – Emblemantha – Fittingia – Geissanthus – Grenacheria – Heberdenia – Hottonia – Hymenandra – Jacquinia – Labisia – Loheria – Lysimachia – Lysimachiopsis – Maesa – Marantodes – Monoporus – Myrsine – Neomezia – Omphalogramma – Oncostemum – Parathesis – Pelletiera – Pleiomeris – Pomatosace – Primula – Rapanea – Sadiria – Samolus – Soldanella – Steironema – Stimpsonia – Stylogyne – Systellantha – Tapeinosperma – Tetrardisia – Theophrasta – Trientalis – Vitaliana – Votschia – Wallenia – Walleniella
Name

Primulaceae Batsch ex Borkh., Bot. Wörterb. 2: 240 (1797), nom. cons.

Type genus: Primula L. Sp. Pl. 1: 142 (1753).

Synonyms

Heterotypic
Maesaceae Anderb., B.Ståhl & Källersjö, Taxon 49: 185. 2000.
Type genus: Maesa Forssk. (1775).
Myrsinaceae R.Br., Prodr.: 532. 1810 (as Myrsineae), nom. cons.
Type genus: Myrsine L. (1753).
Theophrastaceae G.Don, Edwards’s Bot. Reg. 21: ad t. 1764. 1835 (as Theophrasteae), nom. cons.
Type genus: Theophrasta L. (1753).

References
Primary references

Borkhausen, M.B. 1797. Botanisches Wörterbuch 2: 240.

Additional references

Anderberg, A.A. & Ståhl, B. 1994. Phylogenetic interrelationships in the order Primulales, with special emphasis on the family circumscriptions. Canadian Journal of Botany 73(11): 1699–1730. DOI: 10.1139/b95-184 Paywall Reference page.
Anderberg, A.A., Ståhl, B. & Källersjö, M. 1998. Phylogenetic interrelationships in the Primulales inferred from rbcL sequence data. Plant Systematics and Evolution 211(1–2): 93–102. DOI: 10.1007/BF00984914Paywall Reference page.
Angiosperm Phylogeny Group. 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161(2): 105–121. DOI: 10.1111/j.1095-8339.2009.00996.x Open access Reference page.
Källersjö, M., Bergqvist, G. & Anderberg, A.A. 2000. Generic realignment in primuloid families of the Ericales s.l.: A phylogenetic analysis based on DNA sequences from three chloroplast genes and morphology. American Journal of Botany 87(9): 1325–1341. DOI: 10.2307/2656725 Open access Reference page.
Källersjö, M. & Ståhl, B. 2003. Phylogeny of Theophrastaceae (Ericales s. lat.). International Journal of Plant Sciences 164(4): 579–591. DOI: 10.1086/375317Paywall JSTORHybrid open access journal Reference page.
Martins, L., Oberprieler, C. & Hellwig, F.H. 2003. A phylogenetic analysis of Primulaceae sl based on internal transcribed spacer (ITS) DNA sequence data. Plant Systematics and Evolution 237(1–2): 75–85. DOI: 10.1007/s00606-002-0258-1 Paywall ResearchGate Reference page.
Reveal, J.L. 2010. A checklist of familial and suprafamilial names for extant vascular plants. Phytotaxa 6: 1–402. DOI: 10.11646/phytotaxa.6.1.1 Paywall Reference page.

Links

Stevens, P.F. 2001 onwards. Angiosperm Phylogeny Website. Version 14, July 2017 [and more or less continuously updated since]. Online. Reference page.
Tropicos.org 2014. Primulaceae. Missouri Botanical Garden. Published on the internet. Accessed: 2014 Feb. 1.
The Plant List 2013. Primulaceae in The Plant List Version 1.1. Published on the internet. Accessed: 2014 Feb. 1.
USDA, ARS, Germplasm Resources Information Network. Primulaceae Batsch ex Borkh., nom. cons. in the Germplasm Resources Information Network (GRIN), U.S. Department of Agriculture Agricultural Research Service. Accessed: 2014-03-15.
Primulaceae – Taxon details on Integrated Taxonomic Information System (ITIS).
Primulaceae – Taxon details on National Center for Biotechnology Information (NCBI).
EOL: Primulaceae

Vernacular names
العربية: ربيعية
azərbaycanca: Novruzçiçəyikimilər
беларуская: Першакветныя
български: Игликови
བོད་ཡིག: ཤང་དྲིལ།
català: Primulàcia
čeština: Prvosenkovité
dansk: Kodriver-familien
Deutsch: Primelgewächse
Ελληνικά: Ηρανθοειδή
English: Primrose family
Esperanto: Primolacoj
español: Primuláceas
eesti: Nurmenukulised
euskara: Primulazeo
فارسی: پامچالیان
suomi: Esikkokasvit
Nordfriisk: Priimeln
עברית: רקפתיים
hrvatski: Jaglačevke
hornjoserbsce: Kropačkowe rostliny
magyar: Kankalinfélék
հայերեն: Գնարբուկազգիներ
日本語: サクラソウ科
한국어: 앵초과
lietuvių: Raktažoliniai
македонски: Јаглики
Nederlands: Sleutelbloemfamilie
norsk nynorsk: Nøkleblomfamilien
norsk: Nøkleblomfamilien
polski: Pierwiosnkowate
русский: Первоцветные
slovenčina: prvosienkovité
slovenščina: Jegličevke
svenska: Viveväxter
Tagalog: Prima rosa
Türkçe: Çuha çiçeğigiller
українська: Первоцвітові
oʻzbekcha/ўзбекча: Navroʻzguldoshlar
Tiếng Việt: Họ Anh thảo
中文: 报春花科

The Primulaceae /prɪmjəˈleɪʃiː/, commonly known as the primrose family (but not related to the evening primrose family), are a family of herbaceous and woody flowering plants including some favourite garden plants and wildflowers. Most are perennial though some species, such as scarlet pimpernel, are annuals.

Previously one of three families in the order Primulales, it underwent considerable generic re-alignment once molecular phylogenetic methods were used for taxonomic classification. The order was then submerged in a much enlarged order Ericales and became a greatly enlarged Primulaceae sensu lato (s.l). In this new classification of the Angiosperm Phylogeny Group, each of the Prumulales families was reduced to the rank of subfamily of Primulaceae s.l.. The original Primulaceae (Primulaceae sensu stricto or s.s.) then became subfamily Primuloideae, and one genus (Maesa) was raised to the rank of a separate subfamily, making four in all.

Description
Botanical drawing of Primula officinalis Jacquin. Legend: (A) the whole plant; (2) and (3) vertical cross-sections of the flower; (4) stamen; (5) horizontal cross-section of flower showing the calyx around the flower crown and stigma; (6) the stigma; (7) cross section through the ovary; (8) calyx; (9) seed; (10) cross section of seed[4]

The family share a number of characteristics, including haplostemonous flowers with sympetalous corolla, stamens opposite the petals, free central placentation, bitegmic (two layered) tenuinucellate ovules, and nuclear endosperm formation.[5]
Stems

Primulaceae are mostly herbaceous, having no woody stem, except that some form cushions (spreading mats a few inches high) and their stems are stiffened by lignin. The stems can grow upright (erect) or spread out horizontally and then turn upright (decumbent).[6][7]
Leaves

Leaves are simple, being directly attached to the stem by a petiole (stalk), but unlike the leaves of most flowering plants they have no stipules. The petiole is short or the leaf tapers gradually towards the base. Leaf arrangement is typically alternate but some are opposite or whorled, and there is generally a rosette at the base of the stem. The edges are toothed (dentate) or sawtoothed. New leaves in the bud are usually involute (rolled towards the upper surface) or conduplicate (folded upwards), but a few species roll downwards.[6][7]
Flowers

Each flower is bisexual, having both stamens and carpels. They have radial symmetry; the petals can be separate or partially or fully fused together to form a tube-shaped corolla that opens up at the mouth to form a bell-like shape (as in item 8 in the figure) or a flat-faced flower. In most of the families of Ericales, stamens alternate with lobes, but in Primulaceae there is a stamen opposite each petal.[8]

The calyx has 4 to 9 lobes and persists after flowering.[7] They are grouped in unbranched, indeterminate clusters such as racemes, spikes, corymbs or umbels.[6]
Reproductive anatomy

The fruit of Primulaceae begins as an ovary and inside it are the future seeds (ovules). These are attached to a central axis without any partitions between them (an arrangement called free central placentation; see item 7 in the figure), and they are bitegmic (having a double protective layer around each ovule). Unlike in most other families of Ericales, both layers form the opening at the top (the micropyle).[8]
Seeds and fruit

As seeds develop, an endosperm grows around the embryo through free division of nuclei without forming walls (nuclear endosperm formation). The embryo forms a pair of short, narrow cotyledons (item 10 in the figure). Usually multiple seeds are in a capsule that is carried on a straight stalk (pedicel or scape). After it matures, it splits apart, releasing the seeds ballistically.[6]
Taxonomy
History

The taxonomic history of Primulaceae has been long and complex. The botanical authority for the family name is given to August Batsch (1794),[1] as Batsch ex Borkh, using the term Primulae with six genera, the valid description being subsequently given by Borkhausen (1797).[a][9] Some earlier authors attributed the name to Ventenat (1799), as Primulaceae Vent.,[10] who described the Primulacées,[11] but Batsch had precedence.[2]

Linnaeus (1753) placed Primula and related primuloid genera in the Hexandria Monogynia (six stamens one pistil) in his sexual classification based on reproductive characteristics.[12] Jussieu arranged Linnaeus' genera in a hierarchical system of ranks based on the relative value of a much wider range of characteristics. In his Genera plantarum (1789) he organised the primuloid genera into two Ordo (families), within a class (VIII) he called Dicotyledones Monopetalae Corolla Hypogyna, based on the cotyledons (two), form of the petals (fused), and position of the corolla with respect to the ovary (below).[13] Jussieu's families were the Lysimachiae, including Primula and Theophrasta[14] and the Sapotae, including Myrsine, these being the three main lineages in modern understanding.[15]

The most complete treatment of the Primulaceae family, with nearly 1,000 species arranged into 22 genera, was by Pax and Knuth in 1905 in the Engler system. They divided the family into five tribes (and several subtribes); Androsaceae, Cyclamineae, Lysimachieae, Samoleae and Corideae.[16][17] Many systems since have lacked consistency, but generally recognised two major groups as either tribes or subfamilies, the Lysimachieae and Primuleae (the Androsaceae of Pax and Knuth), with the largest genera being Primula, Lysimachia and Androsace.[18][17] In the Cronquist system (1988), Cronquist included the three closely related families, Primulaceae, Myrsinaceae and Theophrastaceae in the order Primulales, of subclass Dilleniidae, based on morphological characteristics, in particular, ovaries with free-central placentation, a feature considered synapomorphic.[19] His circumscription of Primulaceae included about 800 species.[20]
Molecular phylogenetics

These three families were referred to as the primuloid families.[5] With the later development of molecular phylogenetic methods, the Primulales were found to be more closely related to other families within the Ericales, and the three primuloid families were subsequently absorbed into an expanded Ericales (Ericales sensu lato or s.l.), making 24 families within that order, where the primuloids formed a monophyletic clade.[8] It was also apparent that Myrsinaceae were paraphyletic, unless the genus Maesa was segregated and elevated to become a new monogeneric family, Maesaceae, but also that Primulaceae were probably paraphyletic.[21][22][5]

In the first consensus taxonomic classification, the Angiosperm Phylogeny Group (APG 1998), these proposals were recognised by including Primulaceae within Ericales, as Eudicots, forming one of three clades in the Asterids (Asteridae).[8][23] Maesa was formally segregated in 2000.[24] Further changes came from analysis of DNA sequence data. This led to the move of genera (primarily terrestrial non-basal-rosette) from Primulaceae to Mysinaceae and Theophrastaceae. At this time Primulaceae was considered to consist of nine tribes (Primuleae, Androsaceae, Ardisiandreae, Lysimachieae, Glauceae, Anagallideae, Corideae, Cyclamineae, and Samoleae).[10] Notably, Lysimachieae and three smaller tribes, Corideae, Cyclamineae and Ardisiandreae, were transferred to Myrsinaceae, and Samoleae to Theophrastaceae. This enlarged Myrsinaceae is distinguished as Myrsinaceae s.l. in comparison to the previous smaller family, Myrsinaceae s.s. (less Maesa).[17] Some authors[25][26] preferred to raise Samoleae to its own family, Samolaceae, but this has not been accepted by subsequent authors, placing it within Theophrastaceae, while recognising its distinct position within that grouping.[27] These transfers, to preserve monophyly at the family level essentially left two tribes remaining in Primulaceae, the Primuleae and Androsaceae, with about 15 genera sharing a number of common characteristics.[5][19][8] These additional changes were reflected in the 2003 revision of the APG system (APG II), where the now four primuloid families were among 23 in Ericales.[28] This restricted Primulaceae sensu stricto (s.s.) consisted of three groups: The Primulae, including Primula, the largest genus; the Androsaceae, including Androsace, the second largest genus; together with a small third group containing Soldanella, Hottonia, Omphalogramma and Bryocarpum.[18]

The APG third classification system (APG III, 2009) discussed all the taxonomic challenges arising from the phylogenetic studies, and placed all primuloid genera into one large Primulaceae s.l., corresponding to Cronquist's Primulales. They stated that "The biggest problem for APG III was the question of how to treat Primulaceae and their immediate relatives, a closely related group that in the past has often been recognized as a separate order". The decision to treat all genera as a single famioly was based on the observation that the new circumscriptions had little in the way of apomorphies, but the entire group had numerous synapomorphies and were easy to recognise. This resulted in an Ericales with 22 families.[27] Consequently, the four primuloid families were reduced to the rank of subfamilies within Primulaceae s.l..[29]
Phylogeny

Primulaceae s.l. sensu APG III form part of the speciose (species rich) Asterid order Ericales s.l., with about 12,000 species and 22[b] families as per APG IV. Ericalees is one of four major clades within the asterids, where it is sister to the euasterids.[31] The phylogenetic structure of Ericales, as shown in the following cladogram, consists of seven major suprafamilial clades (e.g. balsaminoids, styracoids) and a group of "core" Ericales. Within the eracalean families, Primulaceae s.l. is shown as a sister group to Ebenaceae, and both are sister to Sapotaceae. These three families make up the primuloid clade.[30]
Evolution and biogeography

The fossil record of Primulaceae s.l. is sparse, but the crown group has been estimated as c. 46-61 million years old.[32] The crown primuloids have been dated to c. 102 mya, with Primulaceae/Ebenaceae divergence at 80 mya. Crown ages for the Primulaceae subfamilies vary from 24 mya for the Maesoideae, the basal group, to 70 mya for the Theophrastoideae.[30]

The primuloids probably originated in a shared Neotropical/Indo-Malaysian ancestral range, with the Primulaceae/Ebenaceae clade occupying the neotropics. Theophrastoideae is nearly all neotropical with a more recent migration out of the realm found in the aquatic Samolus genus. The divergence between Theophrastoideae and Primuloideae-Myrsinoideae at 70 mya represents a vicariant event between the Neotropics and the Palearctic in the case of the latter. The Primuloideae originating in thr Palearctic, persisted till the last 16 mya, when it started to shift into thr Nearctic.[30]
Subdivision

The three former families of the Primulales, together with the segregated Maesaceae, have been re-circumscribed into the broadly defined Primulaceae sensu lato (s.l.) The two uniting features of this family are a free central placenta and one stamen opposite each of the corolla lobes.[2][33][27] The cladogram below shows the infrafamilial phylogenetic relationships, together with the subfamilial crown ages. Maesoideae forms the basal group, while Primuloideae and Myrsinoideae are in a sister group relationship.[33][8][29][30]

Primulaceae s.l.	Maesoideae 24.1mya     Theophrastoideae 70mya     Primuloideae 46.4mya   Myrsinoideae 53.2mya

 

Christenhusz et al. (2016, 2017) list 2,790 species and 53 genera, varying from 1 in Maesoideae to 38 in Myrsinoideae, with 8 in Theophrastoideae and the remaining 6 in Primuloideae.[34][32] Byng (2014)[29] and Plants of the World Online list 55 accepted genera.[3][33][8][29][30] The generic limits of Myrsinoideae are not fully resolved and the status of a number of genera is under revision.[35]
Subfamilies
Flowers and leaves of Maesa lanceolata
Maesa lanceolata

Maesoideae (A. DC.) A. DC.
Main article: Maesoideae

Maesoideae is a monogeneric subfamily, whose single genus is Maesa Forssk.. It is a genus of trees. The flowers are small, with short tubules and semi-inferior ovary, arranged in axillary or terminal racemes, or in panicles. Maesa has about 100 species, and is distributed in both tropical and subtropical regions of the Old World, ranging from East Africa to Japan.[5]

Once included in the Primulaceae tribe Samoleae, it has most commonly been considered as part of family Myrsinaceae, till segregated into its own family[24] and then submerged as a Primulaceae s.l. subfamily. It has characteristics that distinguish it from Myrsinaceae and forms the basal group of the clade as sister to all other subfamilies.[5]
Flowers and leaves of Myrsine africana
Myrsine africana

Myrsinoideae Burnett
Main article: Myrsinoideae

The relatively large Myrsinoideae, has been treated as a number of tribes, e.g. Ardisieae and Myrsineae, and more recently Lysimachiaeae, a transfer from Primulaceae s.s.. Ardisieae and Myrsineae represent the woody clades within the subfamily.[30] Within the Myrsinoideae, the genera represented by the restricted Myrsinaceae s.s., prior to the transfers from the then Primulaceae, form a distinct clade.[35] Genera in Lysimachiaeae are Trientalis, Anagallis, Glaux , Lysimachia, Asterolinon and Pelletiera. Coris had its own tribe within Primulaceae s.s., Corideae, but its molecular affinities led to its transfer to Myrsinoideae. Similarly, Ardisiandra was the tribe Ardisiandreae and Cyclamen the tribe Cyclamineae.[5]

Aegiceras Gaertn.
Amblyanthopsis Mez
Amblyanthus A.DC.
Antistrophe A.DC.
Ardisia Sw.
Ardisiandra Hook. f.
Badula Juss.
Conandrium (K.Schumann) Mez
Coris L.
Ctenardisia
Cybianthus Mart.
Cyclamen L.
Discocalyx Mez
Elingamita G.T.S. Baylis
Embelia Burm.f.
Emblemantha B.C.Stone
Fittingia Mez (synonym Abromeitia)
Geissanthus Bentham & Hooker
Gentlea – Glaux – Graphardisia – Grenacheria
Heberdenia Banks ex A.DC.
Hymenandra A.DC.
Labisia Lindley
Loheria Merrill
Lysimachia Tourn. ex L.
Mangenotiella M.Schmid
Microconomorpha (Mez) Lundell
Monoporus A.DC.
Myrsine L.
Oncostemum A. de Jussieu
Parathesis J.D.Hooker f.
Pleiomeris A.DC.
Sadiria Mez
Solonia Urban (synonym Walleniella)
Stimpsonia C.Wright ex A.Gray
Stylogyne A.DC.
Systellantha B.C.Stone
Tapeinosperma Hook.f.
Trientalis L.
Vegaea Urban
Wallenia Sw.

Flowers of Primula veris
Primula veris

Primuloideae Burnett
Main article: Primuloideae

The subfamily is characterised by scapose inflorescences, distinctly tubular flowers having campanulate or hypocrateriform corolla, imbricate corolla aestivation, isodiametric corolla epidermal cells, leaves almost always forming a basal rosette, and ovules rarely immersed in the placenta. In addition they often have syncolpate or sometimes polycolpate pollen, without margo.[5]

Subdivision has included the tribes Androsaceae and Primulae. Takhtajan (1997) gives the four genera of Androsaceae as Androsace (including Douglasia), Vitaliana, Stimpsonia and Pomatosace, and the ten genera of Primulae as Omphalogramma, Bryocarpum, Primula, Dionysia, Cortusa, Kauffmannia, Hottonia, Srediskya, Dodecatheon and Soldanella.[10]

In contrast Kallersjo et al. (2000) place all genera in Primulae, and list thirteen in all: Androsace, Douglasia, Omphalogramma, Soldanella, Dodecatheon, Cortusa, Primula, Dionysia, Vitaliana, Hottonia, Bryocarpum, and Pomatosace. These two (or one) tribes represent the remaining genera in Primuloideae (Primulaceae s.s.) following redistribution among the Primulales on molecular grounds. The 600 species mainly belong to Androsace and Primula and belong to the mountainous regions of Europe and Asia (mainly China).[5]

Androsace L. (synonym: Vitaliana)
Bryocarpum Hook. f. & Thomson
Dionysia Fenzl
Hottonia Boehr. ex L.
Kaufmannia Regel
Omphalogramma Franch.
Pomatosace Maxim.
Primula L. (synonym: Dodecatheon)
Soldanella L.

Specimen of Theophrasta jussieui
Theophrasta jussieui

Theophrastoideae A. DC.
Main article: Theophrastoideae

Theophrastoideae consist of a relatively small subfamily, whose flowers are staminodial and bear berries. The ovary is superior, and the corolla is often tubular. The eight genera are confined to South and Central America. Samolus is a distinct genus, being sister to all other Theophrastoideae. Previously it formed its own tribe, Samoleae within Primulaceae s.s., and in some systems, its own family, Samolaceae, but was subsequently transferred to Theophrastoideae.[5]
Etymology

The Primulaceae are named for their nominative and type genus, Primula. Linnaeus used this name to reflect its place among the first flowers of spring, given the primrose's vernacular Latin name of primula veris (lit. 'little first of spring'), primula (feminine diminutive primus, first + veris (genitive ver, spring).[11]
Distribution and habitat

Distribution is cosmopolitan.[30]
Cultivation

The British National Collection of Double Primroses is held at Glebe Garden, at North Petherwin, in North Cornwall.[36][37][38]
Notes

"More additions affectingconserved familial names are from Batsch (1794),which is to be considered as the place for valid publication of Melanthiaceae and Primulaceae, both accepted and conserved with the authorship of‘Batsch ex Borkh. 1797’ but validly published in 1794 by a reference in the introduction of that book to the corresponding descriptions in Batsch (1786)"[2]

Ericales has 21 or 22 families, depending on whether Sladeniaceae is recognised as separate from, or submerged in Pentaphylacaceae[30]

References

Batsch 1793–1794, ii 395.
APG IV 2016.
POWO 2021.
Thomé 1903.
Kallersjo et al 2000.
Anderberg 2004.
Xu & Chang 2017.
Anderberg et al 2002.
Borkhausen 1797, p. 240.
Takhtajan 1997, p. 200.
Ventenat 1799, vol. ii pp. 289–291.
Linnaeus 1753.
Jussieu 1789, p. 93.
Jussieu 1789, p. 95.
Jussieu 1789, p. 151.
Pax & Knuth 1905.
Martins et al 2003.
Schneeweiss et al 2004.
Mast et al 2001.
Cronquist 1988, pp. 357–359.
Anderberg & Ståhl 1995.
Anderberg et al 1998.
APG I 1998.
Anderberg et al 2000.
Stahl 2004a.
Ståhl 2010.
APG III 2009.
APG II 2003.
Byng 2014.
Rose et al 2018.
Schonenberger et al 2005.
Christenhusz et al 2017, p. 494.
Soltis et al 2018.
Christenhusz & Byng 2016.
Yan et al 2019.
Byfield 2015.
Stone 2021.

Cornwall Gardens Trust 2021.

Bibliography
Books

Byng, James W. (2014). "Primulaceae". The Flowering Plants Handbook: A practical guide to families and genera of the world. Plant Gateway Ltd. pp. 399ff. ISBN 978-0-9929993-1-5.
Christenhusz, Maarten J. M.; Fay, Michael F.; Chase, Mark W. (2017). "Primulaceae". Plants of the World: An Illustrated Encyclopedia of Vascular Plants. University of Chicago Press. p. 494. ISBN 978-0-226-52292-0.
Kubitzki, K., ed. (2004). Celastrales, Oxalidales, Rosales, Cornales, Ericales. The Families and Genera of Vascular Plants. VI. Berlin: Springer-Verlag. doi:10.1007/978-3-662-07257-8. ISBN 978-3-662-07257-8. S2CID 12809916.
Bhattacharyya, Bharati (2005). Systematic botany. Alpha Science International. pp. 216–218. ISBN 9781842652510.
Cronquist, Arthur (1988) [1968]. The evolution and classification of flowering plants (2nd ed.). Bronx, N.Y., USA: New York Botanical Garden. ISBN 9780893273323. (Available here at Internet Archive)
Datta, Subhash C. (1988). "Primulaceae". Systematic botany. New Age Intl. pp. 387–388. ISBN 9788122400137.
Judd, W.S.; Campbell, C.S.; Kellogg, E.A.; Stevens, P.F.; Donoghue, M.J. (2002). "Ericales". Plant Systematics: A Phylogenetic Approach (2nd ed.). Sinauer Associates. pp. 425–436. ISBN 978-0-87893-403-4.
Soltis, Douglas; Soltis, Pamela; Endress, Peter; Chase, M.W.; Manchester, Steven; Judd, Walter; Majure, Lucas; Mavrodiev, Evgeny (2018) [2005]. Phylogeny and Evolution of the Angiosperms: Revised and Updated Edition. University of Chicago Press. pp. 266, 269–271. ISBN 978-0-226-44175-7.

Takhtajan, Armen Leonovich (1997). Diversity and Classification of Flowering Plants. Columbia University Press. ISBN 978-0-231-10098-4.(Available here at Internet Archive)
Tomlinson, P. Barry (2016) [1986]. "Primulaceae (Myrinoideae: Myrsinaceae)". The Botany of Mangroves (2nd ed.). Cambridge University Press. pp. 302–308. ISBN 9781316790656.
Xu, Zhenghao; Chang, Le (2017). "2. Primulaceae". Identification and Control of Common Weeds. 3 vols. 3 (3rd ed.). Springer. pp. 51–82. ISBN 978-981-10-5403-7.

Chapters

Stahl, B; Anderberg, A. A. (2004). Maesaceae. pp. 255–257., in Kubitzki (2004)
Stahl, B; Anderberg, A. A. (2004). Myrsinaceae. pp. 266–281., in Kubitzki (2004)
Anderberg, A. A. (2004). Primulaceae. pp. 313–319., in Kubitzki (2004)
Stahl, B (2004a). Samolaceae. pp. 387–389., in Kubitzki (2004)
Stahl, B (2004). Theophrastaceae. pp. 472–478., in Kubitzki (2004)

Historical sources

Batsch, August Johann Georg Karl (1793–1794). Synopsis vniversalis analytica genervm plantarvm fere omnivm hvcvsque cognitorum qvam secvndvm methodvm sexvalem corollinam, et carpologicam adivnctis ordinibvs naturalibvs adhibitis vltra Linnaeana monitis et adavctionibvs meritissimorvm Avbletii, Lovreirii, Forskolii, Thvnbergii, Forsteri, Vahlii, Gaertneri, Hedwigii, Schreberi, Ivssievii, Swarzii, et aliorum. 2 vols (in Latin). Ienae: Svmtibvs Bibliopolii Croekeriani.
Borkhausen, Moritz Balthasar (1797). Botanisches Wörterbuch oder Versuch einer Erklärung der vornehmsten Begriffe und Kunstwörter in der Botanik, 2 Volumes [Botanical dictionary] (in German). Giessen: Georg Friedrich Heyer.
Jussieu, Antoine Laurent de (1789). Genera plantarum: secundum ordines naturales disposita, juxta methodum in Horto regio parisiensi exaratam, anno M.DCC.LXXIV [Genera of Plants Arranged According to Their Natural Orders, Based on the Method Devised in the Royal Garden in Paris in the Year 1774] (in Latin). Paris. OCLC 5161409.(translated into French, with revisions, by Ventenat (1799) as Tableau du règne végétal selon la méthode de Jussieu)
Linnaeus, Carl (1753). Species Plantarum: exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas (1st. ed.). Stockholm: Impensis Laurentii Salvii.
Pax, F; Knuth, R (1905). "Primulaceae". In Adolf Engler (ed.). Das Pflanzenreich: regni vegetablilis conspectus (in German). 4 (237) heft 22. Leipzig: Engelmann.
Thomé, Otto Wilhelm (1903). "Primulaceae Plate 472". Flora von Deutschland, Österreich und der Schweiz. 4 vols (in German). IV. Gera: H. Bermühler. pp. 16–38.
Ventenat, Étienne-Pierre (1799). Tableau du règne végétal, selon la méthode de Jussieu. 3 vols (in French). Paris: J. Drisonnier.

Articles

Anderberg, Arne A.; Ståhl, Bertil (1995). "Phylogenetic interrelationships in the order Primulales, with special emphasis on the family circumscriptions". Canadian Journal of Botany. 73 (11): 169–91730. doi:10.1139/b95-184.
Anderberg, Arne A.; Ståhl, Bertil; Källersjö, Mari (1998). "Phylogenetic relationships in the Primulales inferred from rbcL sequence data". Plant Systematics and Evolution. 211 (1–2): 93–102. doi:10.1007/BF00984914. S2CID 37418900.
Christenhusz, Maarten JM & Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3): 201–217. doi:10.11646/phytotaxa.261.3.1.
Martins, L.; Oberprieler, C.; Hellwig, F. H. (2003). "A phylogenetic analysis of Primulaceae s.l. based on internal transcribed spacer (ITS) DNA sequence data". Plant Systematics and Evolution. 237 (1–2): 75–85. doi:10.1007/s00606-002-0258-1. S2CID 43988431.

Ericales

Anderberg, A.A.; Rydin, C.; Kallersjo, M. (1 April 2002). "Phylogenetic relationships in the order Ericales s.l.: Analyses of molecular data from five genes from the plastid and mitochondrial genomes". American Journal of Botany . 89 (4): 677–687. doi:10.3732/ajb.89.4.677. PMID 21665668.
Källersjö, M.; Bergqvist, G.; Anderberg, A.A. (2000). "Generic realignment in primuloid families of the Ericales s. l.: a phylogenetic analysis based on DNA sequences from three chloroplast genes and morphology". American Journal of Botany. 87 (9): 1325–41. doi:10.2307/2656725. JSTOR 2656725. PMID 10991903.
Schönenberger, Jürg; Anderberg, Arne A.; Sytsma, Kenneth J. (2005). "Molecular Phylogenetics and Patterns of Floral Evolution in the Ericales". International Journal of Plant Sciences. 166 (2): 265–288. doi:10.1086/427198. JSTOR 10.1086/427198.
Rose, Jeffrey P.; Kleist, Thomas J.; Löfstrand, Stefan D.; Drew, Bryan T.; Schönenberger, Jürg; Sytsma, Kenneth J. (2018). "Phylogeny, historical biogeography, and diversification of angiosperm order Ericales suggest ancient Neotropical and East Asian connections" (PDF). Molecular Phylogenetics and Evolution. 122: 59–79. doi:10.1016/j.ympev.2018.01.014. PMID 29410353.

Maesoideae

Anderberg, Arne A.; Ståhl, Bertil; Källersjö, Mari (May 2000). "Maesaceae, a new primuloid family in the order Ericales s.l.". Taxon. 49 (2): 183–187. doi:10.2307/1223834. JSTOR 1223834.

Myrsinoideae

Anderberg, Arne A.; Manns, Ulrika; Källersjö, Mari (2007). "Phylogeny and Floral Evolution of the Lysimachieae (Ericales, Myrsinaceae): Evidence from ndhF Sequence Data". Willdenowia. 37 (2): 407–421. JSTOR 20371369.
Yan, Xiaokai; Liu, Tongjian; Yuan, Xun; Xu, Yuan; Yan, Haifei; Hao, Gang (2019). "Chloroplast Genomes and Comparative Analyses among Thirteen Taxa within Myrsinaceae s.str. Clade (Myrsinoideae, Primulaceae)". International Journal of Molecular Sciences. 20 (18): 4534. doi:10.3390/ijms20184534. PMC 6769889.

Primuloideae

Mast, Austin R.; Kelso, Sylvia; Richards, A. John; Lang, Daniela J.; Feller, Danielle M. S.; Conti, Elena (2001). "Phylogenetic Relationships in Primula L. and Related Genera (Primulaceae) Based on Noncoding Chloroplast DNA". International Journal of Plant Sciences. 162 (6): 1381–1400. doi:10.1086/323444. JSTOR 10.1086/323444. S2CID 59929813.
Mast, Austin R.; Reveal, James L. (2007). "Transfer of Dodecatheon to Primula (Primulaceae)". Brittonia. 59 (1): 79. doi:10.1663/0007-196X(2007)59[79:TODTPP]2.0.CO;2.
Schneeweiss, Gerald M; Schönswetter, Peter; Kelso, Sylvia; Niklfeld, Harald (2004). "Complex Biogeographic Patterns in Androsace (Primulaceae) and Related Genera: Evidence from Phylogenetic Analyses of Nuclear Internal Transcribed Spacer and Plastid trnL-F Sequences". Systematic Biology. 53 (6): 856–876. doi:10.1080/10635150490522566. PMID 15764556.
Trift, Ida; Källersjö, Mari; Anderberg, Arne A. (2002). "The Monophyly of Primula (Primulaceae) Evaluated by Analysis of Sequences from the Chloroplast Gene rbcL". Systematic Botany. 27 (2): 396–407. ISSN 0363-6445. JSTOR 3093879.

Theophrastoideae

Källersjö, Mari; Ståhl, Bertil (2003). "Phylogeny of Theophrastaceae (Ericales s. lat.)". International Journal of Plant Sciences. 164 (4): 579–591. doi:10.1086/375317. JSTOR 10.1086/375317. S2CID 83790795.
Ståhl, Bertil (2010). "Theophrastaceae". Flora Neotropica. 105: 1–160.
Wanntorp, Livia; Anderberg, Arne A. (2011). "Evolution And Diversification Of Brook Weeds (Samolus, Samolaceae, Ericales)". International Journal of Plant Sciences. 172 (2): 250–266. doi:10.1086/657647. JSTOR 10.1086/657647.

APG

Angiosperm Phylogeny Group (1998), "An ordinal classification for the families of flowering plants", Annals of the Missouri Botanical Garden, 85 (4): 531–553, doi:10.2307/2992015, JSTOR 2992015
Angiosperm Phylogeny Group II (2003), Bremer, B., K. Bremer, M.W. Chase, J.L. Reveal, D.E. Soltis, P.S. Soltis & P.F. Stevens, "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II", Botanical Journal of the Linnean Society, 141 (4): 399–436, doi:10.1046/j.1095-8339.2003.t01-1-00158.x
Angiosperm Phylogeny Group III (October 2009), Bremer, B., K. Bremer, M.W. Chase, M.F. Fay, J.L. Reveal, D.E. Soltis, P.S. Soltis & P.F. Stevens, "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III", Botanical Journal of the Linnean Society, 161 (2): 105–121, doi:10.1111/j.1095-8339.2009.00996.x
Angiosperm Phylogeny Group IV; Chase, M. W.; Christenhusz, J. M.; Fay, M. F.; Byng, J. W.; Judd, W. S.; Soltis, D. E.; Mabberley, D. J.; Sennikov, A. N.; Soltis, P. S.; Stevens, P. R. (2016), "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV", Botanical Journal of the Linnean Society, 181 (1): 1–20, doi:10.1111/boj.12385

Websites
Utteridge, Timothy MA. "More than cowslips: Primulaceae goes tropical". Royal Botanic Gardens, Kew. Retrieved 10 June 2021.
Cholewa, Anita F; Kelso, Sylvia (2009). "Primulaceae". Flora of North America. New York: Oxford University Press. Retrieved 11 June 2021.
POWO (2021). "Primulaceae Batsch ex Borkh". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 14 June 2021.
Elpel, Thomas J. (2021). "Primulaceae: Primrose Family. Identify plants and flowers". Wildflowers and Weeds. Retrieved 16 June 2021.
Byfield, Andy (2 June 2015). "Double primroses: pretty, difficult". Gardening blog. The Guardian. Retrieved 26 June 2021.
Stone, Caroline (2021). "Glebe Garden: National Collection ® Holder for Double Primroses: Primula vulgaris and hybrid cvs". Retrieved 29 June 2021.
Cornwall Gardens Trust (2021). "Double Primroses (National Primrose Collection) Launceston, PL15 8JX". Retrieved 29 June 2021.
"Primulaceae - Primrose Family". Montana Plant Life. 2018. Retrieved 18 June 2021.