Dryandra ser. Aphragma
Encyclopedia
Dryandra ser. Aphragma is an obsolete series within the former genus Dryandra (now Banksia ser. Dryandra). It was first published at sectional rank by Robert Brown
in 1830, and was retained at that rank until 1999, when Alex George
demoted it to a series. It was discarded in 2007 when Austin Mast
and Kevin Thiele
sank Dryandra into Banksia
.
. Brown's arrangement
divided Dryandra into three groups according to what Brown perceived to be variations in the number of seed separators. He allowed for these groups to be treated at subgenus or section rank, but they are now treated as having been published as sections.
D. sect. Aphragma was defined as containing four species which Brown thought lacked seed separators. The epithet Aphragma is from the Greek
a- ("without") and phragma ("barrier"). The placement and circumscription of D. sect. Aphragma in Brown's 1830 arrangement may be summarised as follows:
published a revision of Dryandra. He retained all three of Brown's sections, and the circumscription given to D. sect. Aphragma was much the same, the only differences being the demotion of D. blechnifolia to a variety of D. pteridifolia, and the inclusion of D. drummondii (now B. drummondii
), which had been published in 1848. The placement and circumscription of D. sect. Aphragma in Meissner's arrangement
thus looks like this:
in his 1870 arrangement
, making it the only one of Meissner's infrageneric taxa not to be discarded. However, Bentham reasoned that the single seed separator in D. sect. Eudryandra was formed by the joining together of the outer seed coat of each seed; and therefore both D. bipinnatifida, in which there appears to be two plates, and D. sect. Aphragma, in which there appears to be none, represent failure of the seed coats to join. On these grounds he transferred D. bipinnatifida into D. sect. Aphragma. Thus D. sect. Aphragma was redefined as containing those species in which "Outer integuments of the two seeds not connate or readily separable from each other." He also noted that the members of this section have large involucre
s with many broad bracts, giving them "a different aspect from all others of the genus".
The placement and circumscription of D. sect. Aphragma in Bentham's arrangement
is as follows:
Bentham's arrangement would stand for well over a hundred years, with one minor exception. In 1903 Otto Kuntze
challenged Dryandra R.Br. on the grounds that the name Josephia Knight
had preceded it. In the process of transferring Dryandra into Josephia, Kuntze published the name Josephia sect. Aphragma (R.Br.) Kuntze. This was rejected, however, and J. sect. Aphragma is now considered a nomenclatural synonym of D. ser. Aphragma.
demoted D. sect. Aphragma to a series within D. subg. Dryandra
. He did not accept the previous definitions of the series, but nonetheless accepted the basic circumscription as sound; he instead used a range of flower, seed and leaf characters. The placement and circumscription of D. ser. Aphragma in George's arrangement, as amended in 1999 and 2005, may be summarised as follows:
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has been publishing results of ongoing cladistic analyses of DNA sequence
data for the subtribe Banksiinae. His analyses have provided compelling evidence of the paraphyly
of Banksia
with respect to Dryandra
; that is, it seems that Dryandra arose from within the ranks of Banksia. Early in 2007, Mast and Kevin Thiele
initiated a rearrangement of Banksia by sinking Dryandra into it as B. ser. Dryandra. This transfer necessitated the setting aside of George's infrageneric arrangement of Dryandra; thus D. ser. Aphragma is no longer current. Mast and Thiele have foreshadowed publishing a full arrangement once DNA sampling of Dryandra is complete.
Robert Brown (botanist)
Robert Brown was a Scottish botanist and palaeobotanist who made important contributions to botany largely through his pioneering use of the microscope...
in 1830, and was retained at that rank until 1999, when Alex George
Alex George
Alexander Segger George is a Western Australian botanist. He is the authority on the plant genera Banksia and Dryandra...
demoted it to a series. It was discarded in 2007 when Austin Mast
Austin Mast
Austin R. Mast is a research botanist. Born in 1972, he obtained a Ph.D. from the University of Wisconsin–Madison in 2000. He is currently an associate professor within the Department of Biological Science at Florida State University , and has been director of FSU's since August 2003.One of his...
and Kevin Thiele
Kevin Thiele
Kevin R. Thiele is curator of the Western Australian Herbarium. His research interests include the systematics of the plant families Proteaceae, Rhamnaceae and Violaceae, and the conservation ecology of grassy woodland ecosystems...
sank Dryandra into Banksia
Banksia
Banksia is a genus of around 170 species in the plant family Proteaceae. These Australian wildflowers and popular garden plants are easily recognised by their characteristic flower spikes and fruiting "cones" and heads. When it comes to size, banksias range from prostrate woody shrubs to trees up...
.
According to Brown
Aphragma was first published by Brown in his 1830 Supplementum primum prodromi florae Novae HollandiaeSupplementum Primum Prodromi Florae Novae Hollandiae
Supplementum primum Prodromi florae Novae Hollandiae is an 1830 supplement to Robert Brown's Prodromus florae Novae Hollandiae et Insulae Van Diemen. It may be referred to by its standard botanical abbreviation Suppl. Prodr. Fl. Nov...
. Brown's arrangement
Brown's taxonomic arrangement of Dryandra
Robert Brown's taxonomic arrangement of Dryandra was the first arrangement of what is now Banksia ser. Dryandra. His initial arrangement was published in 1810, and a further arrangement, including an infrageneric classification, followed in 1830...
divided Dryandra into three groups according to what Brown perceived to be variations in the number of seed separators. He allowed for these groups to be treated at subgenus or section rank, but they are now treated as having been published as sections.
D. sect. Aphragma was defined as containing four species which Brown thought lacked seed separators. The epithet Aphragma is from the Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
a- ("without") and phragma ("barrier"). The placement and circumscription of D. sect. Aphragma in Brown's 1830 arrangement may be summarised as follows:
- Dryandra (now B. ser. Dryandra)
- Dryandra verae (18 species)
- D. sect. Diplophragma (1 species)
- D. sect. Aphragma
- D. nervosa (now B. alliaceaBanksia alliaceaBanksia alliacea is a shrub endemic to Western Australia.It was known as Dryandra nervosa until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...
) - D. callophylla (now B. calophyllaBanksia calophyllaBanksia calophylla is a shrub endemic to Western Australia. It was known as Dryandra calophylla until 2007, when all Dryandra species were transferred to Banksia by research botanists Austin Mast and Kevin Thiele.-External links:...
) - D. pteridifolia (now B. pteridifoliaBanksia pteridifoliaBanksia pteridifolia, commonly known as Tangled Honeypot, is a shrub endemic to Western Australia. It was known as Dryandra pteridifolia until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-External links:...
) - D. blechnifolia (now B. pellaeifoliaBanksia pellaeifoliaBanksia pellaeifolia is a shrub endemic to Western Australia.It was known as Dryandra blechnifolia until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...
)
- D. nervosa (now B. alliacea
- Hemiclidia (1 species)
According to Meissner
In 1856, Carl MeissnerCarl Meissner
Carl Daniel Friedrich Meissner was a Swiss botanist.Born in Bern, Switzerland on 1 November 1800, he was christened Meisner but later changed the spelling of his name to Meissner. For most of his 40 year career he was Professor of Botany at University of Basel...
published a revision of Dryandra. He retained all three of Brown's sections, and the circumscription given to D. sect. Aphragma was much the same, the only differences being the demotion of D. blechnifolia to a variety of D. pteridifolia, and the inclusion of D. drummondii (now B. drummondii
Banksia drummondii
Banksia drummondii, commonly known as Drummond's Dryandra, is a shrub endemic to Western Australia. Depending on the subspecies, flowering occurs either mid-year, or at year's end.-Description:...
), which had been published in 1848. The placement and circumscription of D. sect. Aphragma in Meissner's arrangement
Meissner's taxonomic arrangement of Dryandra
Carl Meissner's taxonomic arrangement of Dryandra, now Banksia ser. Dryandra, was published in 1856 as part of his chapter on the Proteaceae in A. P. de Candolle's Prodromus systematis naturalis regni vegetabilis...
thus looks like this:
- Dryandra (now Banksia ser. Dryandra)
- D. sect. Eudryandra (8 series, 47 species, 7 varieties)
- D. sect. Diplophragma (2 species)
- D. sect. Aphragma
- D. pteridifolia (now B. pteridifoliaBanksia pteridifoliaBanksia pteridifolia, commonly known as Tangled Honeypot, is a shrub endemic to Western Australia. It was known as Dryandra pteridifolia until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-External links:...
)- D. pteridifolia var. blechnifolia (now B. pellaeifoliaBanksia pellaeifoliaBanksia pellaeifolia is a shrub endemic to Western Australia.It was known as Dryandra blechnifolia until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...
)
- D. pteridifolia var. blechnifolia (now B. pellaeifolia
- D. calophylla (now B. calophyllaBanksia calophyllaBanksia calophylla is a shrub endemic to Western Australia. It was known as Dryandra calophylla until 2007, when all Dryandra species were transferred to Banksia by research botanists Austin Mast and Kevin Thiele.-External links:...
)- D. calophylla var. acaulis (now B. drummondii subsp. drummondii)
- D. nervosa (now Banksia alliaceaBanksia alliaceaBanksia alliacea is a shrub endemic to Western Australia.It was known as Dryandra nervosa until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...
) - D. drummondii (now Banksia drummondiiBanksia drummondiiBanksia drummondii, commonly known as Drummond's Dryandra, is a shrub endemic to Western Australia. Depending on the subspecies, flowering occurs either mid-year, or at year's end.-Description:...
)
- D. pteridifolia (now B. pteridifolia
- Hemiclidia (1 species)
According to Bentham
D. sect. Aphragma was retained by George BenthamGeorge Bentham
George Bentham CMG FRS was an English botanist, characterized by Duane Isely as "the premier systematic botanist of the nineteenth century".- Formative years :...
in his 1870 arrangement
Bentham's taxonomic arrangement of Dryandra
George Bentham's taxonomic arrangement of Dryandra was published in 1870, in Volume 5 of Bentham's Flora Australiensis...
, making it the only one of Meissner's infrageneric taxa not to be discarded. However, Bentham reasoned that the single seed separator in D. sect. Eudryandra was formed by the joining together of the outer seed coat of each seed; and therefore both D. bipinnatifida, in which there appears to be two plates, and D. sect. Aphragma, in which there appears to be none, represent failure of the seed coats to join. On these grounds he transferred D. bipinnatifida into D. sect. Aphragma. Thus D. sect. Aphragma was redefined as containing those species in which "Outer integuments of the two seeds not connate or readily separable from each other." He also noted that the members of this section have large involucre
Involucre
Involucre may refer to* involucral bract, a bract, bract pair, or whorl of bracts surrounding a flower or inflorescence* a term sometimes misused for the cupule surrounding developing nuts in the Fagaceae...
s with many broad bracts, giving them "a different aspect from all others of the genus".
The placement and circumscription of D. sect. Aphragma in Bentham's arrangement
Bentham's taxonomic arrangement of Dryandra
George Bentham's taxonomic arrangement of Dryandra was published in 1870, in Volume 5 of Bentham's Flora Australiensis...
is as follows:
- Dryandra (now Banksia ser. Dryandra)
- D. sect. Eudryandra
- D. sect. Aphragma
- D. tenuifolia (now B. tenuisBanksia tenuisBanksia tenuis is a shrub endemic to Western Australia. It was known as Dryandra tenuifolia until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...
) - D. proteoides (now B. proteoidesBanksia proteoidesBanksia proteoides, commonly known as King Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra proteoides until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....
)- D. proteoides var. ferruginea (now B. rufaBanksia rufaBanksia rufa is a shrub endemic to Western Australia. It was first published by Carl Meisner in 1855, where it was given the name Dryandra ferruginea by Richard Kippist. The following year, Meisner published what was purportedly a distinct species, Dryandra runcinata...
)
- D. proteoides var. ferruginea (now B. rufa
- D. runcinata (now B. rufa subsp. rufaBanksia rufa subsp. rufaBanksia rufa subsp. rufa is a subspecies of Banksia rufa. It was known as Dryandra ferruginea subsp. ferruginea until 2007, when Austin Mast and Kevin Thiele sunk all Dryandra into Banksia. Since the name Banksia ferruginea had already been used, Mast and Thiele had to choose a new specific epithet...
) - D. obtusa (now B. obtusaBanksia obtusaBanksia obtusa, commonly known as Shining Honeypot, is a shrub endemic to Western Australia. It was known as Dryandra obtusa until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....
) - D. bipinnatifida (now B. bipinnatifidaBanksia bipinnatifidaBanksia bipinnatifida is a shrub endemic to Western Australia.-Description:It grows as a prostrate shrub with an underground stem and a lignotuber. It has few leaves; these are deeply lobes, and the lobes are themselves deeply lobed, giving the impression of a bipinnate leaf structure...
) - D. pteridifolia (now B. pteridifoliaBanksia pteridifoliaBanksia pteridifolia, commonly known as Tangled Honeypot, is a shrub endemic to Western Australia. It was known as Dryandra pteridifolia until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-External links:...
) - D. calophylla (now B. calophyllaBanksia calophyllaBanksia calophylla is a shrub endemic to Western Australia. It was known as Dryandra calophylla until 2007, when all Dryandra species were transferred to Banksia by research botanists Austin Mast and Kevin Thiele.-External links:...
)
- D. tenuifolia (now B. tenuis
Bentham's arrangement would stand for well over a hundred years, with one minor exception. In 1903 Otto Kuntze
Otto Kuntze
Otto Carl Ernst Kuntze was a German botanist.-Biography:Otto Kuntze was born in Leipzig.An apothecary in his early career, he published an essay entitled Pocket Fauna of Leipzig. Between 1863 and...
challenged Dryandra R.Br. on the grounds that the name Josephia Knight
Joseph Knight (horticulturist)
Joseph Knight , gardener to George Hibbert, was one of the first people in England to successfully propagate Proteaceae...
had preceded it. In the process of transferring Dryandra into Josephia, Kuntze published the name Josephia sect. Aphragma (R.Br.) Kuntze. This was rejected, however, and J. sect. Aphragma is now considered a nomenclatural synonym of D. ser. Aphragma.
According to George
In 1996, Alex GeorgeAlex George
Alexander Segger George is a Western Australian botanist. He is the authority on the plant genera Banksia and Dryandra...
demoted D. sect. Aphragma to a series within D. subg. Dryandra
Dryandra subg. Dryandra
Dryandra subg. Dryandra is an obsolete series within the former genus Dryandra . It was first published at sectional rank as Dryandra verae in 1830, before being renamed Eudryandra in 1847, the replaced by the autonym at subgenus rank in 1996...
. He did not accept the previous definitions of the series, but nonetheless accepted the basic circumscription as sound; he instead used a range of flower, seed and leaf characters. The placement and circumscription of D. ser. Aphragma in George's arrangement, as amended in 1999 and 2005, may be summarised as follows:
- Dryandra (now Banksia ser. Dryandra)
- D. subg. DryandraDryandra subg. DryandraDryandra subg. Dryandra is an obsolete series within the former genus Dryandra . It was first published at sectional rank as Dryandra verae in 1830, before being renamed Eudryandra in 1847, the replaced by the autonym at subgenus rank in 1996...
- D. ser. FloribundaeDryandra ser. FloribundaeDryandra ser. Floribundae is an obsolete series within the former genus Dryandra . It was first published by George Bentham in 1870, and was given a new circumscription by Alex George in 1996, but was ultimately discarded in 2007 when Austin Mast and Kevin Thiele sunk Dryandra into...
(1 species, 4 varieties) - D. ser. ArmataeDryandra ser. ArmataeDryandra ser. Armatae is an obsolete series within the former genus Dryandra . It was first published by George Bentham in 1870, and was given a new circumscription by Alex George in 1996, but was ultimately discarded in 2007 when Austin Mast and Kevin Thiele sunk Dryandra into Banksia.-According...
(21 species, 7 subspecies, 4 varieties) - D. ser. Marginatae (1 species)
- D. ser. Folliculosae (1 species, 5 varieties)
- D. ser. Acrodontae (4 species, 2 varieties)
- D. ser. CapitellataeDryandra ser. CapitellataeDryandra ser. Capitellatae is an obsolete series within the former genus Dryandra . It was published by Alex George in 1996, but discarded in 2007 when Austin Mast and Kevin Thiele sank Dryandra into Banksia.-Publication:George published the series in his 1996 "New taxa and a new infrageneric...
(2 species, 2 subspecies) - D. ser. IlicinaeDryandra ser. IlicinaeDryandra ser. Ilicinae is an obsolete series within the former genus Dryandra . It was first published by Carl Meissner in 1856, but was discarded by George Bentham in 1870...
(3 species, 2 varieties) - D. ser. Dryandra (3 species, 2 subspecies)
- D. ser. Foliosae (3 species, 2 subspecies)
- D. ser. Decurrentes (1 species)
- D. ser. Tenuifoliae (2 species, 2 varieties)
- D. ser. Runcinatae (4 species, 7 subspecies)
- D. ser. Triangulares (3 species, 3 subspecies)
- D. ser. Aphragma
- D. pteridifolia (now B. pteridifoliaBanksia pteridifoliaBanksia pteridifolia, commonly known as Tangled Honeypot, is a shrub endemic to Western Australia. It was known as Dryandra pteridifolia until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele.-External links:...
)- D. pteridifolia subsp. inretita (now B. pteridifolia subsp. inretita)
- D. pteridifolia subsp. pteridifolia (now B. pteridifolia subsp. pteridifoliaBanksia pteridifolia subsp. pteridifoliaBanksia pteridifolia subsp. pteridifolia is a subspecies of Banksia pteridifolia. As an autonym, it is defined as encompassing the type material of the species. It was known as Dryandra pteridifolia subsp. pteridifolia until 2007, when Austin Mast and Kevin Thiele sunk all Dryandra into Banksia. As...
) - D. pteridifolia subsp. vernalis (now B. pteridifolia subsp. vernalis)
- D. fililoba (now B. fililobaBanksia fililobaBanksia fililoba is a shrub endemic to Western Australia....
) - D. shanklandiorum (now B. shanklandiorumBanksia shanklandiorumBanksia shanklandiorum is a shrub endemic to Western Australia....
) - D. nervosa (now B. alliaceaBanksia alliaceaBanksia alliacea is a shrub endemic to Western Australia.It was known as Dryandra nervosa until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...
) - D. blechnifolia (now B. pellaeifoliaBanksia pellaeifoliaBanksia pellaeifolia is a shrub endemic to Western Australia.It was known as Dryandra blechnifolia until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele...
) - D. porrecta (now B. porrectaBanksia porrectaBanksia porrecta is a shrub endemic to Western Australia.An assessment of the potential impact of climate change on this species found that its range is likely to contract by between 50% and 80% by 2080, depending on the severity of the change....
) - D. aurantia (now B. aurantiaBanksia aurantiaBanksia aurantia is a shrub endemic to Western Australia....
) - D. calophylla (now B. calophyllaBanksia calophyllaBanksia calophylla is a shrub endemic to Western Australia. It was known as Dryandra calophylla until 2007, when all Dryandra species were transferred to Banksia by research botanists Austin Mast and Kevin Thiele.-External links:...
) - D. lepidorhiza (now B. lepidorhizaBanksia lepidorhizaBanksia lepidorhiza is a shrub endemic to Western Australia....
)
- D. pteridifolia (now B. pteridifolia
- D. ser. Ionthocarpae (1 species, 2 subspecies)
- D. ser. Inusitatae (1 species)
- D. ser. Subulatae (1 species)
- D. ser. Gymnocephalae (11 species, 4 subspecies, 2 varieties)
- D. ser. Concinnae (3 species)
- D. ser. Obvallatae (7 species, 2 varieties)
- D. ser. Pectinatae (1 species)
- D. ser. Acuminatae (1 species)
- D. ser. NiveaeDryandra ser. NiveaeDryandra ser. Niveae is an obsolete series within the former genus Dryandra . It was first published by George Bentham in 1870, and was given a new circumscription by Alex George in 1996, but was ultimately discarded in 2007 when Austin Mast and Kevin Thiele sunk Dryandra into Banksia.-According to...
(7 species, 7 subspecies)
- D. ser. Floribundae
- D. subg. HemiclidiaDryandra subg. HemiclidiaDryandra subg. Hemiclidia is an obsolete plant taxon that encompassed material that is now included in Banksia. Published at genus rank as Hemiclidia by Robert Brown in 1830, it was set aside by George Bentham in 1870, but reinstated at subgenus rank by Alex George in 1996...
(2 species) - D. subg. DiplophragmaDryandra subg. DiplophragmaDryandra subg. Diplophragma is an obsolete subgenus within the former genus Dryandra . It was first published by Robert Brown in 1830, but was discarded by George Bentham in 1870...
(1 species)
- D. subg. Dryandra
Recent developments
Since 1998, Austin MastAustin Mast
Austin R. Mast is a research botanist. Born in 1972, he obtained a Ph.D. from the University of Wisconsin–Madison in 2000. He is currently an associate professor within the Department of Biological Science at Florida State University , and has been director of FSU's since August 2003.One of his...
has been publishing results of ongoing cladistic analyses of DNA sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...
data for the subtribe Banksiinae. His analyses have provided compelling evidence of the paraphyly
Paraphyly
A group of taxa is said to be paraphyletic if the group consists of all the descendants of a hypothetical closest common ancestor minus one or more monophyletic groups of descendants...
of Banksia
Banksia
Banksia is a genus of around 170 species in the plant family Proteaceae. These Australian wildflowers and popular garden plants are easily recognised by their characteristic flower spikes and fruiting "cones" and heads. When it comes to size, banksias range from prostrate woody shrubs to trees up...
with respect to Dryandra
Dryandra
Banksia ser. Dryandra is a series of 94 species of shrub to small tree in the plant genus Banksia. It was considered a separate genus named Dryandra until early 2007, when it was merged into Banksia on the basis of extensive molecular and morphological evidence that Banksia was paraphyletic with...
; that is, it seems that Dryandra arose from within the ranks of Banksia. Early in 2007, Mast and Kevin Thiele
Kevin Thiele
Kevin R. Thiele is curator of the Western Australian Herbarium. His research interests include the systematics of the plant families Proteaceae, Rhamnaceae and Violaceae, and the conservation ecology of grassy woodland ecosystems...
initiated a rearrangement of Banksia by sinking Dryandra into it as B. ser. Dryandra. This transfer necessitated the setting aside of George's infrageneric arrangement of Dryandra; thus D. ser. Aphragma is no longer current. Mast and Thiele have foreshadowed publishing a full arrangement once DNA sampling of Dryandra is complete.