Primulaceae

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).{{sfn|Anderberg|2004}}{{sfn|Xu|Chang|2017}}

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.{{sfn|Anderberg|2004}}{{sfn|Xu|Chang|2017}}

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.{{sfn|Anderberg et al|2002}}

The calyx has 4 to 9 lobes and persists after flowering.{{sfn|Xu|Chang|2017}} They are grouped in unbranched, indeterminate clusters such as racemes, spikes, corymbs or umbels.{{sfn|Anderberg|2004}}

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).{{sfn|Anderberg et al|2002}}

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.{{sfn|Anderberg|2004}}

The taxonomic history of Primulaceae has been long and complex. The botanical authority for the family name is given to August Batsch (1794),{{sfn|Batsch|1793–1794|loc=ii 395}} as Batsch ex Borkh, using the term Primulae with six genera, the valid description being subsequently given by Borkhausen (1797).{{efn|"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)"{{sfn|APG IV|2016}}}}{{sfn|Borkhausen|1797|p=240}} Some earlier authors{{clarification needed|reason=so earlier authors attributed the name to a later authority??|date=November 2021}} attributed the name to Ventenat (1799), as Primulaceae Vent.,{{sfn|Takhtajan| 1997|p=200}} who described the Primulacées,{{sfn|Ventenat|1799|loc=vol. ii pp. 289–291}} but Batsch had precedence.{{sfn|APG IV |2016}}

Linnaeus (1753) placed Primula and related primuloid genera in the Hexandria Monogynia (six stamens one pistil) in his sexual classification based on reproductive characteristics.{{sfn|Linnaeus|1753}} 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).{{sfn|Jussieu|1789|p=93}} Jussieu's families were the Lysimachiae, including Primula and Theophrasta{{sfn|Jussieu|1789|p=95}} and the Sapotae, including Myrsine, these being the three main lineages in modern understanding.{{sfn|Jussieu|1789|p=151}}

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.{{sfn|Pax|Knuth|1905}}{{sfn|Martins et al|2003}} 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.{{sfn|Schneeweiss et al|2004}}{{sfn|Martins et al|2003}}

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.{{sfn|Mast et al|2001}} His circumscription of Primulaceae included about 800 species.{{sfn|Cronquist|1988|pp=357–359}}

File:Cyclamen graecum1.jpg]]

These three families were referred to as the primuloid families.{{sfn|Kallersjo et al|2000}} 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.{{sfn|Anderberg et al|2002}} 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.{{sfn|Anderberg|Ståhl|1995}}{{sfn|Anderberg et al|1998}}{{sfn|Kallersjo et al|2000}}

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).{{sfn|Anderberg et al|2002}}{{sfn|APG I|1998}} Maesa was formally segregated in 2000.{{sfn|Anderberg et al|2000}} 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 that time Primulaceae was considered to consist of nine tribes (Primuleae, Androsaceae, Ardisiandreae, Lysimachieae, Glauceae, Anagallideae, Corideae, Cyclamineae, and Samoleae).{{sfn|Takhtajan| 1997|p=200}} 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).{{sfn|Martins et al|2003}} Some authors{{sfn|Stahl|2004a}}{{sfn|Ståhl|2010}} 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.{{sfn|APG III|2009}} 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.{{sfn|Kallersjo et al|2000}}{{sfn|Mast et al|2001}}{{sfn|Anderberg et al|2002}} 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.{{sfn|APG II|2003}} 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.{{sfn|Schneeweiss et al|2004}}

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 family 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.{{sfn|APG III|2009}} Consequently, the four primuloid families were reduced to the rank of subfamilies within Primulaceae s.l.{{sfn|Byng|2014}}

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{{efn|Ericales has 21 or 22 families, depending on whether Sladeniaceae is recognised as separate from, or submerged in Pentaphylacaceae{{sfn|Rose et al|2018}}}} families as per APG IV. Ericales is one of four major clades within the asterids, where it is sister to the euasterids.{{sfn|Schonenberger et al|2005}} 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.{{sfn|Rose et al|2018}}

File:EricalesRose2018.png

The fossil record of Primulaceae s.l. is sparse, but the crown group has been estimated as c. 46-61 million years old.{{sfn|Christenhusz et al|2017|p=494}} 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.{{sfn|Rose et al|2018}}

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 the Palearctic, persisted till the last 16 mya, when it started to shift into the Nearctic.{{sfn|Rose et al|2018}}

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.{{sfn|APG IV|2016}}{{sfn|Soltis et al|2018}}{{sfn|APG III|2009}} 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.{{sfn|Soltis et al|2018}}{{sfn|Anderberg et al|2002}}{{sfn|Byng|2014}}{{sfn|Rose et al|2018}}

{{clade| style=font-size:100%;line-height:100%;

|label1= Primulaceae s.l.

|1={{clade

|1= Maesoideae

|sublabel1=24.1mya

|2={{clade

|1= Theophrastoideae

|sublabel1=70mya

|2= {{clade

|1=Primuloideae

|sublabel1=46.4mya

|2= Myrsinoideae

|sublabel2=53.2mya

}}

}}

}}

}}

Christenhusz et al. (2016, 2017) list 53 genera and 2,790 species, varying from 1 in Maesoideae to 38 in Myrsinoideae, with 8 in Theophrastoideae and the remaining 6 in Primuloideae.{{sfn|Christenhusz|Byng|2016}}{{sfn|Christenhusz et al|2017|p=494}} Byng (2014){{sfn|Byng|2014}} and Plants of the World Online list 55 accepted genera.{{sfn|POWO|2021}}{{sfn|Soltis et al|2018}}{{sfn|Anderberg et al|2002}}{{sfn|Byng|2014}}{{sfn|Rose et al|2018}} The generic limits of Myrsinoideae are not fully resolved and the status of a number of genera is under revision.{{sfn|Yan et al|2019}}

{|

|- style="vertical-align:top;"

| File:Maesa lanceolata 1.jpg|130px|left|alt=Flowers and leaves of Maesa lanceolata]]

|

{{main|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.{{sfn|Kallersjo et al|2000}}

Once included in the Primulaceae tribe Samoleae, it has most commonly been considered as part of family Myrsinaceae, till segregated into its own family{{sfn|Anderberg et al|2000}} 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.{{sfn|Kallersjo et al|2000}}

|- style="vertical-align:top;"

| File:Myrsine africana.jpg|130px|left|alt=Flowers and leaves of Myrsine africana]]

|

{{main|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.{{sfn|Rose et al|2018}} Within the Myrsinoideae, the genera represented by the restricted Myrsinaceae s.s., prior to the transfers from the then Primulaceae, form a distinct clade.{{sfn|Yan et al|2019}} 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.{{sfn|Kallersjo et al|2000}}

{{div col|colwidth=20em}}

• 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
• Monoporus A.DC.
• Myrsine L.
• Oncostemum A. de Jussieu
• Paralysimachia F.Du, J.Wang & S.Y.Yang
• 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.
• Vegaea Urban
• Wallenia Sw.

{{div col end}}

|- style="vertical-align:top;"

| File:Cowslip - Primula veris - geograph.org.uk - 1274759.jpg|130px|left|alt=Flowers of Primula veris]]

|

{{main|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.{{sfn|Kallersjo et al|2000}}

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.{{sfn|Takhtajan|1997|p=200}}

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).{{sfn|Kallersjo et al|2000}}

• Androsace L. (synonym: Douglasia, Vitaliana)
• Bryocarpum Hook. f. & Thomson
• Dionysia Fenzl
• Evotrochis {{small|Raf.}}
• Hottonia Boehr. ex L.
• Kaufmannia Regel
• Omphalogramma Franch.
• Pomatosace Maxim.
• Primula L. (synonym: Dodecatheon)
• Soldanella L.

|- style="vertical-align:top;"

| File:Theophrasta jussieui - Naples Botanical Garden - Naples, Florida - DSC09930.jpg|130px|left|alt=Specimen of Theophrasta jussieui]]

|

{{main|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.{{sfn|Kallersjo et al|2000}}

• Bonellia {{small|Bertero ex Colla}}
• Clavija {{small|Ruiz & Pav.}}
• Deherainia {{small|Decne.}}
• Jacquinia {{small|L.}}
• Neomezia {{small|Votsch}}
• Samolus {{small|L.}}
• Theophrasta {{small|L.}}
• Votschia {{small|B.Ståhl}}

|}

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 ({{literal translation|little first of spring}}), primula (feminine diminutive primus, first + veris (genitive ver, spring).{{sfn|Ventenat|1799|loc=vol. ii pp. 289–291}}

Distribution is cosmopolitan.{{sfn|Rose et al|2018}}

The British National Collection of Double Primroses is held at Glebe Garden, at North Petherwin, in North Cornwall.{{sfn|Byfield|2015}}{{sfn|Stone|2021}}{{sfn|Cornwall Gardens Trust|2021}}

{{notelist}}

{{Reflist|20em}}

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• {{cite book|last1=Borkhausen|first1=Moritz Balthasar|title=Botanisches Wörterbuch oder Versuch einer Erklärung der vornehmsten Begriffe und Kunstwörter in der Botanik, 2 Volumes|trans-title=Botanical dictionary|date=1797|publisher=Georg Friedrich Heyer|location=Giessen|url=https://books.google.com/books?id=OxMXAAAAYAAJ|language=de}}
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• {{cite book |last=Linnaeus |first=Carl |author-link=Carl Linnaeus|title=Species Plantarum: exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas|url=http://www.biodiversitylibrary.org/bibliography/669#/summary|year=1753 |publisher=Impensis Laurentii Salvii|location=Stockholm|edition=1st.}}
• {{cite book|last1=Pax|first1=F|last2=Knuth|first2=R|author-link1=Ferdinand Albin Pax|author-link2=Reinhard Gustav Paul Knuth|title=Das Pflanzenreich: regni vegetablilis conspectus|publisher=Engelmann|location=Leipzig|url=https://bibdigital.rjb.csic.es/records/item/11071-redirection|editor=Adolf Engler|editor-link=Adolf Engler|volume=4 (237) heft 22|chapter=Primulaceae|year=1905|language=German}}
• {{cite book|last=Thomé|first=Otto Wilhelm|author-link=Otto Wilhelm Thomé |title=Flora von Deutschland, Österreich und der Schweiz. 4 vols.|url=https://books.google.com/books?id=pLjvAAAAMAAJ|year=1903|volume=IV|publisher=H. Bermühler|location=Gera|language=de|chapter=Primulaceae Plate 472|pages=16–38|chapter-url=https://archive.org/details/floravondeutschl04thom/page/n541/mode/2up}}
• {{cite book |last1=Ventenat |first1=Étienne-Pierre |author-link=E P Ventenat|title=Tableau du règne végétal, selon la méthode de Jussieu. 3 vols. |date=1799 |publisher=J. Drisonnier |location=Paris |url=https://www.biodiversitylibrary.org/creator/234152#/titles|language=fr}}

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• {{cite journal |last1=Källersjö |first1=Mari |last2=Ståhl |first2=Bertil |title=Phylogeny of Theophrastaceae (Ericales s. lat.) |journal=International Journal of Plant Sciences |date=2003|volume=164 |issue=4 |pages=579–591 |doi=10.1086/375317|s2cid=83790795|jstor=10.1086/375317 }}
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• {{cite journal |last1=Wanntorp |first1=Livia |last2=Anderberg |first2=Arne A. |title=Evolution And Diversification Of Brook Weeds (Samolus, Samolaceae, Ericales) |journal=International Journal of Plant Sciences |date=2011 |volume=172 |issue=2 |pages=250–266 |doi=10.1086/657647|jstor=10.1086/657647 |s2cid=84493389 }}

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• {{cite web |last1=Utteridge |first1=Timothy MA |title=More than cowslips: Primulaceae goes tropical |url=https://www.kew.org/read-and-watch/more-than-cowslips |publisher=Royal Botanic Gardens, Kew |access-date=10 June 2021 }}
• {{cite web|last1= Cholewa |first1=Anita F|last2=Kelso|first2=Sylvia|title=Primulaceae|url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10728|work=Flora of North America|year=2009|volume=8|access-date=11 June 2021|publisher=Oxford University Press|location=New York}}
• {{cite web |last=POWO|author-link=Plants of the World Online|title=Primulaceae Batsch ex Borkh.|url= https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:77126675-1|website=Plants of the World Online |publisher=Royal Botanic Gardens, Kew| date=2021|access-date=14 June 2021 }}
• {{cite web |last1=Elpel |first1=Thomas J. |title=Primulaceae: Primrose Family. Identify plants and flowers. |url=https://www.wildflowers-and-weeds.com/Plant_Families/Primulaceae.htm |website=Wildflowers and Weeds |access-date=16 June 2021 |date=2021}}
• {{cite web |last1=Byfield |first1=Andy |title=Double primroses: pretty, difficult |url=https://www.theguardian.com/lifeandstyle/gardening-blog/2015/jun/02/double-primroses |website=Gardening blog |publisher=The Guardian |access-date=26 June 2021 |date=2 June 2015}}
• {{cite web |last1=Stone |first1=Caroline |title=Glebe Garden: National Collection Holder for Double Primroses: Primula vulgaris and hybrid cvs. |url=https://glebegarden.co.uk/ |access-date=29 June 2021 |date=2021}}
• {{cite web |last=Cornwall Gardens Trust|title=Double Primroses (National Primrose Collection) Launceston, PL15 8JX |url=https://www.cornwallgardenstrust.org.uk/double-primroses-national-primrose-collection-launceston-pl15-8jx/ |date=2021|access-date=29 June 2021}}
• {{cite web|url=http://montana.plant-life.org/cgi-bin/family03.cgi?Primulaceae|title=Primulaceae - Primrose Family|website=Montana Plant Life|date=2018|access-date=18 June 2021}}

{{refend}}

{{wikispecies}}

• {{Commons category-inline}}

{{Angiosperm families}}

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Category:Ericales families

Category:Taxa named by August Batsch