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Nomenclatural updates of Aristolochia subgenus Pararistolochia (Aristolochiaceae)

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Aristolochia subgenus Pararistolochia is revised and 35 species, distributed in tropical Africa and Australasia, are recognised. Fourteen new combinations and two new names are presented, resulting from the transfer of all taxa of Pararistolochia to Aristolochia. Additionally, a neotype is designated for Aristolochia preussii.
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Nomenclatural updates of Aristolochia subgenus Pararistolochia
(Aristolochiaceae)
Katja Buchwalder
A
, Marie-Stéphanie Samain
B
, Garry Sankowsky
C,D
, Christoph Neinhuis
A
and Stefan Wanke
A,E
A
Institut für Botanik, Technische Universität Dresden, Zellescher Weg 20b, D-01062 Dresden, Germany.
B
Instituto de Ecología, A.C., Centro Regional del Bajío, Avenida Lázaro Cárdenas 253, 61600 Pátzcuaro,
Michoacán, Mexico.
C
Australian Tropical Herbarium, James Cook University, Cairns Campus, PO Box 6811, Cairns, Qld 4870, Australia.
D
PO Box 210, Tolga, Qld 4882, Australia.
E
Corresponding author. Email: stefan.wanke@tu-dresden.de
Abstract. Aristolochia subgenus Pararistolochia is revised and 35 species, distributed in tropical Africa and Australasia,
are recognised. Fourteen new combinations and two new names are presented, resulting from the transfer of all taxa of
Pararistolochia to Aristolochia. Additionally, a neotype is designated for Aristolochia preussii.
Additional keywords: host plant, new combination, nomenclature, Ornithoptera, taxonomy.
Received 27 September 2013, accepted 12 May 2014, published online 30 June 2014
Introduction
The genus Aristolochia L. is subdivided into three monophyletic
groups congruent with the following subgenera: Siphisia
Raf., Pararistolochia (Hutch. & Dalziel) O.C.Schmidt, and
Aristolochia (González and Stevenson 2002; Wanke et al.
2006). Hutchinson and Dalziel (1927) rst described the genus
Pararistolochia, exclusively containing African species. Both
morphological (Huber 1960, 1985; González and Stevenson
2002) and molecular phylogenetic studies (Ohi-Toma et al.
2006; Bliss et al. 2013) have shown that Pararistolochia
occurs not only in Africa, but that species from Australasia
have to be included as well. Aristolochiaceae are important
host plants for various genera of the family Papilionidae
(Lepidoptera) (Simonsen et al. 2010). In addition, Condamine
et al.(2012) reconstructed Aristolochiaceae as the ancestral host
for Papilionidae, pointing to the importance of the group for
studying co-evolution, especially the plantbuttery interaction.
Caterpillars of a dozen Papilionidae genera (Allancastria,
Archon, Battus, Cressida, Euryades, Losaria, Luehdora,
Ornithoptera, Pachliopta, Parides, Pharmacophagus, Troides)
feed on Aristolochiaceae and the majority of the tribe
Troidini
on the genus Aristolochia (Parsons 1996a, 1996b; Simonsen
et al. 2010). The most prominent species known to feed on
species of subg. Pararistolochia are the worlds largest
butteries, Ornithoptera alexandrae and O. goliath (Straatman
and Inoue 1984; Jebb 1993; Parsons 1996a). Each of the 1013
Ornithoptera taxa is popular among collectors for their
iridescent bright colours and their size (Parsons 1992).
Because of excessive collecting and habitat loss, birdwing
butteries are listed in CITES (mostly appendix II) and on the
IUCN Red List (most listed as Vulnerable or Endangered). As a
consequence, the Commonwealth Scientic and Industrial
Research Organisation (CSIRO) and many Australian NGOs
have taken initiatives to involve the public in conservation
efforts, for example, by planting native butter y host plants
such as Aristolochia praevenosa F.Muell. and eliminating
introduced species such as A. elegans Mast., A. labiata Willd.,
or A. odoratissima L. However, Papua New Guinea decided
to follow a different strategy. The government declared the
butteries a natural resource and established the Insect
Farming & Trading Agency to develop this resource
sustainably (Hutton 1985).
Nevertheless, all protection measures require correct
identication and accurate species names. This is especially
true for the host plants that have received considerably less
attention than the butteries. Indeed, Jebb (1993) highlighted
several incorrect identications within Aristolochia , which have
entered the literature and caused confusion about host plants.
Therefore, it is often unclear on which particular host plant
individual birdwing butteries feed, hampering conservation
measures of these prominent agship taxa. As a consequence
of co-evolution, the distribution of the butteries is strongly
linked to the distribution of the host plants (Ehrlich and Raven
1964). In parallel to a full taxonomic revision based on natural
relationships within the subgenus Pararistolochia, a molecular-
based phylogeny study is being performed, for which correct
names are also essential. Hence, an update of nomenclature
within this group is urgently required.
Journal compilation Ó CSIRO 2014 www.publish.csiro.au/journals/asb
CSIRO PUBLISHING
Australian Systematic Botany, 2014, 27 ,4855
http://dx.doi.org/10.1071/SB13042
All species belonging to the subg. Pararistolochia are treated
here, providing new combinations and new names. The
taxonomic afliation according to different authors has been
critically revised and morphological characters used by these
authors for their respective classications are discussed, serving
as basis for a molecular phylogeny of subg. Pararistolochia and
its relationships within Aristolochiaceae.
Taxonomic history
Nomenclature of Aristolochia subg. Pararistolochia is rather
intricate (Table 1). Hooker (1865) was the rst to mention that
his newly described African species Aristolochia goldieana
Hook.f., A. triactina Hook.f. and A. mannii Hook.f. were
clearly distinguishable from other previously described
Aristolochia species by the three-lobed perianth and the
structure of the gynostemium, with 12 lobes in A. goldieana
and 10 in A. triactina and A. mannii, whereas ve or six were
characteristic for other species. Masters (1879) suggested that
his newly described A. promissa Mast. had close afnities to
the species described by Hooker (1865). Shortly afterwards,
Bentham and Hooker (1880) described sect. Polyanthera
Benth. & Hook.f. in which they included the species described
by Hooker (1865) and Masters (1879). Six decades after
Hooker, Hutchinson and Dalziel (1927, 1928) placed the
African Aristolochia species possessing a three-lobed perianth
and a cucumber-shaped indehiscent fruit in the new genus
Pararistolochia Hutch. & Dalziel and mentioned that this
genus is evidently an ancestral type (Hutchinson and Dalziel
1928, p. 22). Schmidt (1935) followed Bentham and Hooker
(1880) in recognising the group at infrageneric rank, but instead
of a section, a subgeneric rank was assigned, and he adopted
the name Pararistolochia from Hutchinson and Dalziel (1927).
In contrast to Schmidt (1935), Keay (1952), Poncy (1978
) and
Huber (1960, 1985, 1993) re-established the genus on the basis of
the following morphological characters: growth as tropical liana,
attened stem, cauliorous, indehiscent fruit (cleistocarp), seed
anatomy, as well as the occurrence in dense humid habitats. In
contrast, Ma (1992 ) again adopted subgeneric rank, whereas
Parsons (1996a) described several species in the genus
Pararistolochia. Furthermore, Ma (1992) created three new
sections within the subgenus, separating these mainly on the
basis of the gynostemium segment number.
Huber (1960) was the rst to associate the Australasian
Pararistolochia species with those of Africa. He pointed out
that the New Guinean A. momandul K.Schum. also develops a
cleistocarp fruit, as do the African species. Jebb (1993) described
new Aristolochia species from New Guinea and assigned
them to the Aristolochia momandul group. Parsons (1996a)
also recognised the close afnities of the African and
Australasian species and pointed out the morphological
similarities between the African Pararistolochia triactina and
the New Guinean P. meridionaliana Mich. J.Parsons in bud,
ower, fruit and seed. When González and Stevenson (2002)
published the rst cladistic analysis of the Aristolochioideae on
the basis of morphology, they followed the classication of
Schmidt (1935), with a subgeneric rank for the monophyletic
Pararistolochia. Later, the monophyly of Pararistolochia
was conrmed by molecular-based phylogenetic analyses
(Neinhuis et al. 2005, Wanke et al. 2006, 2007). Ohi-Toma
et al.(2006) and Bliss et al.(2013) showed that the
Australasian and African species belonging to Pararistolochia
are sister groups. On the basis of morphological synapomorphies,
both subgeneric and generic rank would be acceptable for
Pararistolochia
. However, recognising Pararistolochia as
genus would cause paraphyly of Aristolochia in its current
circumscription (Wanke et al. 2006) and would thus require
additional taxonomic and nomenclatural changes within the
subfamily Aristolochioideae, especially the acceptance of
Siphisia as an additional genus. As a result, we suggest
transferring all Pararistolochia species to Aristolochia,
maintaining Aristolochia in its broadest sense because this
genus is easily recognisable.
Table 1. Historical and current classications of Aristolochia subgenus Pararistolochia
Author and year Genus Subgenus Section Note
von Mueller 18601861, A. praevenosa Aristolochia
Bentham and Hooker 1880 Aristolochia Polyanthera
Hutchinson and Dalziel 1927 Pararistolochia
Schmidt 1935 Aristolochia Pararistolochia
Poncy 1978 Pararistolochia
Huber 1960 Pararistolochia A. momandul shows
indehiscent fruits like
Pararistolochia
Ma 1992 Aristolochia Pararistolochia 1. Pararistolochioides
2. Pararistolochia
3. Aristolochioides
Jebb 1993 Aristolochia Australasian momandul
group should be
included in genus
Pararistolochia
Parsons 1996a Pararistolochia
González and Stevenson 2002 Aristolochia Pararistolochia
Wanke et al. 2006 Aristolochia Pararistolochia Suggested on the basis of
molecular phylogeny
Leal and Nguema 2011 Pararistolochia
Aristolochia subgenus Pararistolochia Australian Systematic Botany 49
Taxonomic synopsis
We reviewed the literature about the species belonging to the
subg. Pararistolochia and examined and revised ~500 herbarium
specimens from 19 herbaria (A, B, BO, BRI, BRLU, CNS, DR,
G, K, KEP, L, MO, NHM, NY, PRE, S, SING, WAG, WRSL).
Approximately 35% of the known species are being cultivated in
the Botanical Garden Dresden and phenotypic variability was
observed during multiple eld trips performed by the authors.
However, it would be premature to provide a full revision before
molecular phylogenetic data have shown the monophyly of
individual species.
Aristolochia subgenus Pararistolochia (Hutch. & Dalziel)
O.C.Schmidt in H.G.A.Engler (Ed.), Nat. Panzenfam.
2nd edn, 16b: 241 1935
Pararistolochia Hutch. & Dalziel, Fl. W. Trop. Afr. 1 : 75 (1927).
Lectotype: Pararistolochia triactina Hook.f. (designated by
Parsons 1996a).
Although the genus Pararistolochia was validly published
by Hutchinson and Dalziel in 1927, a full description was not
provided until 1928 (Hutchinson and Dalziel 1928).
This taxon is mainly characterised by having a attened stem,
an actinomorphic, three-lobed perianth, and a longitudinally more
or less ribbed berry with a eshy endocarp (Hutchinson and
Dalziel 1927, 1928; Schmidt 1935; Huber 1985). The majority
of the species are cauliorous, except the Australasian species
A. deltantha, A. ornithopterae, A. laheyana, A. sparusifolia,
A. praevenosa, A. momandul and the African A. goldieana,
which also ower on young stems. In comparison to species of
subg. Siphisia and subg. Aristolochia that have six anthers, except
for Aristolochia subsect. Pentandrae Duch. displaying ve
anthers (Schmidt 1935), subg. Pararistolochia shows a
minimum of six anthers in the Australian species and up to 24
in the African A. goldieana. Also the number of stigmatic lobes
varies, in at least some species, from 6 to 12, even within a
single individual (González and Stevenson 2000). An additional
morphological synapomorphy is the presence of a massive exine-
ridge formation on the pollen grain (González and Stevenson
2002), although this character has not been conrmed for all
species. All species are lianas or slender vines distributed in
tropical rainforest, with the exception of A. ornithopterae, which
grows in woodland with a grassy or heathy understorey (Ross
and Halford 2007).
The present treatment includes all 35 species of Aristolochia
subg. Pararistolochia, with 14 African and 21 Australasian
species (Fig. 1).
Aristolochia alexandriana (Mich.J.Parsons) Buchwalder &
Wanke, comb. nov.
Pararistolochia alexandriana Mich.J.Parsons, Bot. J. Linn. Soc.
120(3): 218 (1996).
Type: Papua New Guinea: Northern Province, Ahora Village,
north of Popondetta town, supporting trees forming secondary
forest up to 10 m tall, ~100 m, 14 Mar. 1988, M. Parsons 1 (holo:
LAE (69994)).
Distribution: Papua New Guinea (Northern Province, Oro
Province).
Aristolochia australopithecurus (Mich.J.Parsons)
Buchwalder & Wanke, comb. nov.
Pararistolochia australopithecurus Mich.J.Parsons, Bot. J. Linn.
Soc. 120(3): 227 (1996).
Type: Australia: Queensland, State Forest Reserve 310,
Rainforest, 780 m, 8 Nov. 1977, 17
13
0
S 145
42
0
E, B. Gray
784 (holo: QRS (017289).
Distribution: Australia (Queensland, Wet Tropics).
A. decandra
A. alexandriana
A. biakensis
A. dictyophlebia
A. dielsiana
A. engleriana
A. gracilifolia
A. kepara
A. manokwariensis
A. meridionaliana
A. momandul
A. paradisiana
A. pithecurus
A. schlechteri
A. sepikensis
A. tithonusiana
A. australopithecurus
A. deltantha
A. laheyana
A. peninsulensis
A. praevenosa
A. ornithopterae
A. sparusifolia
A. ceropegioides
A. gabonensis
A. goldieana
A. incisiloba
A. ju-ju
A. leonensis
A. macrocarpa
A. mannii
A. preussii
A. promissa
A. triactina
A. zenkeri
Fig. 1. Map showing the distribution of Aristolochia subgenus Pararistolochia. The two main centres of distribution are central-western
Africa as well as New Guinea and eastern Australia. A single species is reported from Borneo.
50 Australian Systematic Botany K. Buchwalder et al.
Aristolochia biakensis (Mich.J.Parsons) Buchwalder &
Wanke, comb. nov.
Pararistolochia biakensis Mich.J.Parsons, Bot. J. Linn. Soc. 120(3):
232 (1996).
Type: Indonesia: New Guinea, Jappen-Biak Island, Saroerai
bij Seroei, 27 Jul. 1939, Aet & Injan 173 (Exp. Ir L.J. van Dijk)
(holo: BO (BO13302332)).
Distribution: Indonesia (Yapen and Biak Island).
Aristolochia ceropegioides S.Moore, J. Bot. 58: 269 (1920)
Pararistolochia ceropegioides (S.Moore) Hutch. & Dalziel, Bull.
Misc. Inform, Kew. 1928: 24 (1928).
Type: Cameroon: Bilye River, 1919, G.L. Bates 1235 (holo:
BM (BM000528389); iso: MO).
Distribution: Cameroon, Gabon.
Aristolochia decandra Ding Hou, Blumea 28: 343 (1983)
Pararistolochia decandra (Ding Hou) Mich.J.Parsons, Bot. J. Linn.
Soc. 120(3): 206 (1996).
Type: Indonesia: western Borneo, Kalimantan, H. Winkler
1256 (holo: L (L0360036 !)).
Distribution: Indonesia (Borneo, Kalimantan).
Aristolochia deltantha F.Muell., Fragm. 6 (46): 179 (1868)
Aristolochia deltantha F.Muell. var. deltantha , Queensland Agric.
J. 28: 199 (1912);
Pararistolochia deltantha (F. Muell.) Mich.J.Parsons, Bot. J. Linn.
Soc. 120(3): 206 (1996).
Type: Australia: Queensland, Rockinghams Bay, Dallachy s.
n. (lecto (designated by Parsons (1996a)): MEL (MEL1553306)).
Distribution: Australia (Queensland, Cape York Peninsula,
Wet Tropics).
Aristolochia dictyophlebia Merr. & L.M.Perry, J. Arnold
Arbor. 29(2): 152 (1948)
Pararistolochia dictyophlebia (Merr. & L.M.Perry) Mich.J.Parsons,
Bot. J. Linn. Soc. 120(3): 206 (1996).
Type: Papua New Guinea: Ogeramnang, forest hill, 1650 m,
Jan. 1937, Clemens 4901 (holo: A; iso: BRI (AQ0216259 !)).
Distribution: Papua New Guinea (Morobe Province).
Aristolochia dielsiana O.C.Schmidt , Bot. Jahrb. Syst. 58: 490
(1923)
Pararistolochia dielsiana (O.C.Schmidt) Mich.J.Parsons, Bot.
J. Linn. Soc. 120(3): 206 (1996).
Type: Papua New Guinea: East Sepik Province, Etappenberg
(Base Mountain), Ledermann 9169 (holo: B, destroyed?).
Distribution: Papua New Guinea (East Sepik Province).
Aristolochia engleriana O.C.Schmidt, Repert. Spec. Nov.
Regni Veg. 23: 288 (1927)
Pararistolochia engleriana (O.C.Schmidt) Mich.J.Parsons, Bot.
J. Linn. Soc. 120(3): 206 (1996); Aristolochia ledermannii
O.C.Schmidt, Bot. Jahrb. Syst. 58: 489 (1923), nom. illeg., non
Engler (1912 (1911)).
Type: Papua New Guinea: East Sepik Province, Schraderberg
(Schrader Mountain), Ledermann 12055 (holo: B, destroyed?).
Distribution: Papua New Guinea (East Sepik Province).
Aristolochia gabonensis Buchwalder & Wanke, nom. nov.
Pararistolochia mbriata M.L.Leal & D.Nguema, Fl. Gabon 42:
9 (2011).
Type: Mondah forest, 10 km N of Libreville, Gabon, 2009,
D. Nguema et al. 1050 (holo: MO; iso: LBV, WAG) (non
Aristolochia mbriata Cham. & Schltdl., Linnaea 7: 210 (1832)
Distribution: Gabon.
Etymology: named after the West African country of Gabon,
to which the species is endemic.
Aristolochia goldieana Hook. f., Trans. Linn. Soc. London 25:
185, t. 14 (1865)
Pararistolochia goldieana (Hook. f.) Hutch. & Dalziel, Fl. W. Trop.
Afr. 1: 77 (1927).
Type: Nigeria, Thompson s.n. (holo: K).
Distribution: Cameroon, Guinea, Nigeria, Sierra Leone.
Aristolochia gracilifolia O.C.Schmidt, Bot. Jahrb. Syst. 58:
490 (1923)
Pararistolochia gracilifolia (O. C. Schmidt) Mich. J.Parsons, Bot.
J. Linn. Soc. 120(3): 206 (1996).
Type: Papua New Guinea: West Sepik, Relsspitze (Rocky
Peak), Ledermann 12458 (holo: B destroyed?).
Distribution: Papua New Guinea (West Sepik Province,
Sandaun Province).
Aristolochia incisiloba Jongkind, Bull. Jard. Bot. Nat. Belg.
60: 147 (1990)
Pararistolochia incisiloba (Jongkind) Leal & Nguema, Fl.Gabon.
42: 9 (2011).
Type: Gabon: Chaillu Mts., Songou Mt, between Dibandi and
Mouyanama, ~20 km E of Mimongo, 1
37
0
S, 11
46
0
E, primary
forest, 120-m altitude, Nov. 1983, A.M. Louis, F.J. Breteler, J. de
Bruijn 975 (holo: WAG (0132108 !); iso: LBV, MO).
Distribution: Gabon.
Aristolochia ju-ju S.Moore, J. Bot. 58: 269 (1920)
Pararistolochia ju-ju (S. Moore) Hutch. & Dalziel, Fl. W. Trop. Afr.
1: 77 (1927).
Type: Nigeria: Degema Division, 1916, P.A.Talbot 3766
(holo: BM (BM000528396 !)).
Distribution: Nigeria.
Note: this species is possibly synonymous with Aristolochia
mannii Hook.f. However, more specimens are needed to draw a
nal conclusion.
Aristolochia subgenus Pararistolochia Australian Systematic Botany 51
Aristolochia kepara (Mich.J.Parsons) Buchwalder & Wanke,
comb. nov.
Pararistolochia kepara Mich.J.Parsons, Bot. J. Linn. Soc. 120(3):
219 (1996).
Type: Papua New Guinea: Northern Province, near Kamondo
Village near Kopkoda, 1 km UTM grid square EL 8714, on
supporting tree in primary forest, ~400-m elev., 18 May 1991,
M. Parsons 17 (holo: LAE (69995); iso: K).
Distribution: Papua New Guinea (Northern Province, Oro
Province).
Aristolochia laheyana (Mich.J.Parsons) Buchwalder &
Wanke, comb. nov.
Aristolochia deltantha var. laheyana F.M.Bailey Queensland Agric.
J. 28: 199 (1912); Pararistolochia laheyana (F.M.Bailey) Mich.
J.Parsons, Bot. J. Linn. Soc. 120(3): 206 (1996).
Type: Australia: Queensland, MacPherson Range, C.T.White
s.n. (holo: BRI (AQ0333077))
Distribution: Australia (Queensland, Border Range to New
South Wales).
Aristolochia leonensis Mast., Bot. J. Linn. Soc., 30: 95 (1895)
Pararistolochia leonensis (Mast.) Hutch. & Dalziel, Fl. W. Trop. Afr.
1: 77. 1927.
Type: Sierra Leone: env. Kassa, 30 Mar. 1892, Scott-Elliot
5062 (holo: K)
Distribution: Guinea, Ivory Coast, Liberia, Sierra Leone.
Aristolochia macrocarpa Duch., in A.L.P.P. Candolle, Prodr.
15(1): 497. 1864
Pararistolochia macrocarpa (Duch.) Poncy, Adansonia, sér. 2
17(34): 488 (1978); Pararistolochia macrocarpa (Duch) Poncy var.
macrocarpa Poncy, Adansonia, sér. 2 17(34): 488 (1978).
Type: in Africa occidentalis tropicae regione dicta Gabon,
1854, Aubry-Lecomte s.n. (holo: P (P00487096 !); iso: MPU
(MPU018708))
Aristolochia soyauxiana Oliv., Hook. Ic. Pl. 15: 8, 1410 (1884);
Pararistolochia soyauxiana (Oliv.) Hutch. & Dalziel, Kew. Bull. 23
(1928); Pararistolochia macrocarpa Duch. var. sojauxiana (Oliv.)
Poncy, Adansonia, sér. 2 17(34): 488 (1978).
Type: Gaboon river, 27 Nov. 1881, H. Soyaux 317 (iso:
P (P00487098 !)).
Aristolochia staudtii Engl., Bot. Jahrb. Syst. 24: 491 (1898);
Pararistolochia staudtii (Engl.) Hutch. & Dalziel, Kew Bull. 1928:
24 (1928).
Type: Cameroon: Lolodorf, Mar. 1896, G. Staudt 186 (holo:
B destroyed?; iso: P (P00487095 !)).
Aristolochia os-avis A. Chev., J. Bot. (Morot) 22: 129 (1909);
Pararistolochia os-avis (A.Chev.) Hutch. & Dalziel, Fl. W. Trop.
Afr. 1: 77 (1927).
Type: Côte dIvoire: 4 Apr. 1909, A. Chevalier 21145 (holo:
P? n.v.).
Aristolochia tessmannii Engl., Bot. Jahrb. Syst. 46: 413 (1911).
Type: Spanish Guinea: Bebai im Campogebiet, Dec. 1908;
Tessmann 717. Cameroon: Bipindi, im Urwald, May 1899,
Zenker 2056; in der Njabilandschaft, Mar. 1900, Zenker 2261
(syntypes: HBG).
Aristolochia tribrachiata S. Moore, Cat. Pl. Oban 92 (1913);
Pararistolochia tribrachiata (S.Moore) Hutch. & Dalziel, Fl.
W. Trop. Afr. 1: 77 (1927).
Type: Nigeria, Oban, Talbot 1542 (holo: BM
(BM000528392); iso: K (K000350282)).
Distribution: Angola (Province Cabinda), Cameroon, Central
African Republic, Democratic Republic of the Congo, Gabon,
Ghana, Equatorial Guinea, Ivory Coast, Liberia, Nigeria,
Republic of the Congo.
Aristolochia mannii Hook. f., Trans. Linn. Soc. London 25:
186 (1865)
Pararistolochia mannii (Hook. f.) Keay, Kew Bull. 1952: 159 (1952).
Type: Nigeria: Old Calabar, Feb. 1883, Mann 2323 (holo:
K (K000350284 !); iso: P (P00482858 !)).
Distribution: Benin, Republic of the Congo, Ivory Coast,
Nigeria.
Aristolochia manokwariensis (Mich.J.Parsons) Buchwalder
& Wanke, comb. nov.
Pararistolochia manokwariensis Mich.J.Parsons, Bot. J. Linn. Soc.
120(3): 220 (1996).
Type: Indonesia: Irian Jaya, Wamaré to Mokwam foot track,
open primary hill forest, ~500-m elev., 22 Nov. 1992, M.Parsons
002 (holo: BO; iso: K).
Distribution: Indonesia (Irian Jaya, Manokwari).
Aristolochia meridionaliana (Mich.J.Parsons) Buchwalder &
Wanke, comb. nov.
Pararistolochia meridionaliana Mich.J.Parsons, Bot. J. Linn. Soc.
120(3): 214 (1996).
Type: Papua New Guinea: Central Province, along Hiritano
Highway in disturbed forest by the roadside, Hopkins &
M.Parsons 913 (holo: LAE (69991); iso: K, UPNG (13444)).
Distribution: Papua New Guinea (Central Province).
Aristolochia meridionaliana subsp. milnensis
(Mich.J.Parsons) Buchwalder & Wanke, comb. nov.
Pararistolochia meridionaliana subsp. milnensis Mich.J.Parsons,
Bot. J. Linn. Soc. 120(3): 218 (1996).
Type: Papua New Guinea: Milne Bay Province, Pini Range,
near Alotau, Milne Bay, ~150-m elev., 1 Jun. 1991, M.Parsons 23
(holo: LAE (69993)).
Distribution: Papua New Guinea (Milne Bay Province).
Aristolochia meridionaliana subsp. popondettensis
(Mich.J.Parsons) Buchwalder & Wanke, comb. nov.
Pararistolochia meridionaliana subsp. popondettensis Mich.
J.Parsons, Bot. J. Linn. Soc. 120(3): 216 (1996).
Type: Papua New Guinea: Northern Province, Hegata Village,
immediately out of Popondetta, 100-m elev., 16 Mar. 1988,
Parsons 2 (holo: LAE (69992)).
52 Australian Systematic Botany K. Buchwalder et al.
Distribution: Papua New Guinea (Northern Province, Oro
Province).
Aristolochia momandul K.Schum, in K.Schumann &
M.Hollrung, Fl. Kais. Wilh. Land.: 105 (1889)
Pararistolochia momandul (K.Schum.) Mich.J.Parsons, Bot. J. Linn.
Soc. 120(3): 207 (1996).
Type: Papua New Guinea: Madang, Astrolabe Bay,
Konstantinhafen, Hollrung 520 (holo: B, destroyed?; iso: BO
(BO108760 !), WRSL).
Distribution: Papua New Guinea (Madang Province, Morobe
Province).
Aristolochia ornithopterae Buchwalder & Wanke, nom. nov.
Pararistolochia linearifolia Mich.J.Parsons, Bot. J. Linn. Soc.
120(3): 235 (1996).
Type: Australia: Queensland, Moa Island, 14 Feb. 1989,
B.Gray 5006 (holo: QRS 091513 !) (non Aristolochia
linearifolia Griseb., Cat. Pl. Cub. 115 (1866)).
Distribution: Australia (Queensland, Cape York Peninsula).
Etymology: named after the Cape York Birdwing butteries,
that often exclusively feed on Aristolochia species.
Aristolochia paradisiana (Mich.J.Parsons) Buchwalder &
Wanke, comb. nov.
Pararistolochia paradisiana Mich.J.Parsons, Bot. J. Linn. Soc.
120(3): 223 (1996).
Type: Papua New Guinea: East Sepik Province, Lonem
Village, near Maprik, growing in mature, well spaced
secondary forest, ~200 m, 28 Mar. 1988, M.Parsons 4 (holo:
LAE (69996); iso: A, K, CANB, UPNG).
Distribution: Papua New Guinea (East Sepik Province).
Aristolochia peninsulensis (Mich.J.Parsons) Buchwalder &
Wanke, comb. nov.
Pararistolochia peninsulensis Mich.J.Parsons, Bot. J. Linn. Soc.
120(3): 234 (1996).
Type: Australia: Queensland, Iron Range, rainforest, 40 m,
20 Nov. 1985, B. Gray 4239 (holo: QRS (080850)).
Distribution: Australia (Queensland, Cape York Peninsula).
Aristolochia pithecurus Ridl., J. Bot. 52: 296 (1914)
Pararistolochia pithecurus (Ridl.) Mich.J.Parsons, Bot. J. Linn. Soc.
120(3): 207 (1996).
Type: Papua New Guinea: Central Province, Sogeri region,
Mount Koroko, 2500 ft, Forbes 621 (holo: BM; iso: L (0360038
!), WRSL).
Distribution: Papua New Guinea (Central Province).
Aristolochia praevenosa F.Muell., Fragm. 2: 166 (1861)
Pararistolochia praevenosa (F.Muell.) Mich.J.Parsons, Bot. J. Linn.
Soc. 120(3): 207 (1996).
Type: Australia: Queensland, Clarence River, Beckler s.n.
(lecto (designated by Parsons (1996a)): MEL (MEL1553302)).
Distribution: Australia (Queensland, Wet Tropics, Border
Range to New South Wales).
Aristolochia preussii Engl., Bot. Jahrb. Syst. 24: 492 (1898)
Pararistolochia preussii (Engl.) Hutch.& Dalziel, Bull. Misc. Inform.
Kew 1928: 24 (1928).
Type: Cameroon: between Barombi and Kumba, 1046 ft,
Preuss 108 (holo: B, destroyed?). Neotype (designated here):
Brenan 9484 (P (P00482859) Cameroun: Divison Kumba,
Banga, Bakundu Forest Reserve, 18 Mar. 1948.
Distribution: Cameroon, Equatorial Guinea, Gabon.
Note: we neotypify with the specimen Brenan 9484
(P00482859), which was already veried by Keay as a
representative specimen of the species.
Aristolochia promissa Mast., Gard. Chron. 11: 494 (1879)
Pararistolochia promissa (Mast.) Keay, Kew Bull. 1952: 160, t.
1 (1952).
Type: Cameroon: Victoria, Kalbreyer 7 (holo: K).
Aristolochia agellata Stapf, Bull. Misc. Inform. Kew 1906:
80 (1906).
Type: Gold Coast: Aburi Gardens, W.H.Johnson 487, 1060
(syntypes: K).
Aristolochia talbotii S.Moore, Cat. Pl. Oban 17: 93 (1913);
Pararistolochia talbotii (S.Moore) Keay, Kew Bull. 1952: 161 (1952);
Aristolochia talbotii S.Moore var. longissima S.Moore, Cat Pl. Oban
17 : 94 (1913).
Type: Nigeria: Oban, Talbot 128, Talbot 2310, Talbot 1642
(syntypes: BM (BM 000528393 !), K (K000350286 !)).
Aristolochia tenuicauda S.Moore, Cat. Pl. Oban 17: 94 (1913);
Pararistolochia tenuicauda (S.Moore) Keay, Kew Bull. 1952: 160
(1952).
Type: Nigeria: Oban, Talbot 2318 (BM (BM000528395)).
Aristolochia congolana Hauman, nom. inval., Fl. Congo Belge 1: 384
(1948), nom. inval .
Distribution: Cameroon, Central African Republic, Republic
of the Congo, Democratic Republic of the Congo, Gabon, Ivory
Coast, Nigeria.
Aristolochia schlechteri Lauterb., Nachtr. Fl. Schutzgeb.
Südsee 260 (1905)
Pararistolochia schlechteri (Lauterb.) Mich.J.Parsons, Bot. J. Linn.
Soc. 120(3): 207 (1996).
Type: Papua New Guinea: Madang Province, near Madang,
Ramu River Valley, Schlechter 14156 (holo: WRSL).
Distribution: Papua New Guinea (Madang Province).
Aristolochia sepikensis (Mich.J.Parsons) Buchwalder &
Wanke, comb. nov.
Pararistolochia sepikensis Mich.J.Parsons, Bot. J. Linn. Soc. 120(3):
231 (1996).
Type: Papua New Guinea: West Sepik, Bewani Subprovince,
Mount Yungat, northern slopes Bewani Mountains, strangler in
Aristolochia subgenus Pararistolochia Australian Systematic Botany 53
lower montane forest, 700 m, 20 Sep. 1982, K. Karenga (holo:
A (56530); iso: K, LAE ?).
Distribution: Papua New Guinea (Sandaum Province, West
Sepik Province).
Aristolochia sparusifolia (Mich.J.Parsons) Buchwalder &
Wanke, comb. nov.
Pararistolochia sparusifolia Mich.J.Parsons, Bot. J. Linn. Soc.
120(3): 228 (1996).
Type: Australia: Queensland, State Forest Reserve 143, Bushy
Logging Area, Mount Lewis, 800 m, 2 Dec. 1982, B. Gray 2876
(holo: QRS (072007); iso: BRI (AQ0462911)).
Distribution: Australia (Queensland, Wet Tropics).
Aristolochia tithonusiana (Mich.J.Parsons) Buchwalder &
Wanke, comb. nov.
Pararistolochia tithonusiana Mich.J.Parsons, Bot. J. Linn. Soc.
120(3): 222 (1996).
Type: Indonesia: Irian Jaya, Wamaré to Mokwam foot track,
open lower montane primary forest, ~1500 m, 22 Nov. 1992,
M. Parsons 005 (holo: BO).
Distribution: Indonesia (Irian Jaya, Manokwari).
Aristolochia triactina Hook. f., Trans. Linn. Soc. London 25:
186. 1865
Pararistolochia triactina (Hook. f.) Hutch. & Dalziel, Fl. W. Trop.
Afr. 1: 77, f. 21 (1927).
Type: Gabon: 1
N, 1862, Mann 1851 (holo: K (K000350283
!); iso: P (P00482857 !)).
Aristolochia stuhlmannii Engl. Panzenw. Ost-Afrikas 2: 169 (1895).
Type: Uganda: Emin Pascha expedition, 18901891,
Stuhlmann s.n. (holo: B destroyed?).
Aristolochia schweinfurthii Engl., Bot. Jahrb. Syst. 24: 492
(1898); Pararistolochia schweinfurthii (Engl.) Hutch. & Dalziel, Kew
Bull. 1928: 24 (1928).
Type: eastern Sudan: Monbuttu-Land, Apr. 1870, Schweifurth
3507 (holo: B destroyed?; iso: K (K000350263 !)).
Distribution: Angola, Cameroon, Central African Republic,
Democratic Republic of the Congo, Equatorial Guinea, Gabon,
Ruanda, Sudan, Uganda.
Aristolochia zenkeri Engl., Bot. Jahrb. Syst. 24: 490 (1898)
Pararistolochia zenkeri (Engl.) Hutch. & Dalziel, Bull. Misc. Inform.
Kew 1928: 24 (1928).
Type: Cameroun: Bipinde, Dec. 1896, Zenker 1226 (holo: B
destroyed?; iso: BM (BM000528397 !), HBG (HBG502751)).
Distribution: Angola, Cameroon, Democratic Republic of the
Congo, Ivory Coast, Nigeria, Republic of the Congo.
Acknowledgements
We thank the curators of the Herbaria A, B, BO, BRI, BRLU, CNS, DR,
G, K, KEP, L, MO, NHM, NY, PRE, S, SING, WAG and WRSL for
providing specimens for this study. Special thanks go to the curator of B,
Robert Vogt, for checking whether types were still present in this herbarium.
Field research was conducted under permit ATH 12/009, granted by the
Queensland Government (Department of National Parks, Recreation, Sport
and Racing). We are grateful to Darren Crayn (Australian Tropical
Herbarium, James Cook University, Cairns) and Taylor Feild (James Cook
University, Townsville) for their support. We thank Frank Müller (TU
Dresden) for taxonomic advices. We very much appreciate comments
and corrections by Favio González (National University of Colombia,
Bogotá) and two anonymous reviewers, as well as those by the editorial
board. This study has been co-supported by a grant from the Deutsche
Forschungsgemeinschaft (NE 681/11-1).
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www.publish.csiro.au/journals/asb
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The troidine papilionid genus Ornithoptera is shown to be distinct from Troides In order to reappraise the interrelationships of the member species of Ornithoptera , and to test the resultant theory of their evolution against biogeographical data, the genus is revised using Hennigian cladistic methodology through computer analysis. The input data for this was drawn equally from immature stages and adult characters. A single cladogram hypothesising the likely phylogeny of the Ornithoptera species was generated. With minor weighting of a single important adult character, a further two cladograms were generated, one of which is similar to hypotheses proposed by previous workers. Based on these findings, and on ecological data, notably larval foodplant relations with Aristolochiaceae, as well as present-day biogeographical data, a new theory of the origin and evolution of Ornithoptera is presented which fits well Gondwanan vicariance events ascertained by geological studies: essentially that Ornithoptera evolved on northward drifting Australia, allopatrically from Troides on the Indian plate, and therefore that Ornithoptera did not reach the Australian Subregion via Troides -like ancestors in Southeast Asia as has been previously postulated. As ‘living fossils' representative of an ancient, Gondwanaland-derived, austral faunal element, the Ornithoptera should be regarded even more highly in terms of their ‘conservation value'.
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In preparation for a forthcoming revision of the family Aristolochiaceae in Australia and New Guinea, a total of 14 new species and two new subspecies are described. Due to the fact that Aristolochiaceae within the region fall into two distinctive, well-defined groups of species, it was considered necessary to examine the affinities of these within the family. One group belongs toAristolochia, whereas the other is clearly accommodated inPararistolochia, the latter previously regarded as being solely a tropical African genus. The distinction betweenPararistolochiaandAristolochiais discussed. It is concluded that the Australia and New Guinea taxa were found to substantiate circumscription ofPararistolochia. Two new species ofAristolochia(A. nauseifoliaandA. goliathiana) are described, as well as 12 species ofPararistolochia (P. alexandriana, P. australopithecurus, P. biakensis, P. kepara, P. manokwariensis, P. meridionaliana, P. paradisiana, P. peninsulensis, P. tithonusiana, P. sepikensis, P. sparusifolia, P. linearifolia). Two subspecies ofP. meridionaliana(subsp.popondettensisand subsp.milnensis) are described. Eleven other taxa from Australia and New Guinea are newly combined underPararistolochia: P. deltanthaF. Muell., 1868;P. dictyophlebiaMerr. & Perry, 1948;P. dielsianaO.C. Schmidt, 1923;P. englerianaO.C. Schmidt, 1927;P. gracilifoliaO.C. Schmidt, 1923;P. laheyanaBail., 1912;P. momandulK. Sch., 1889;P. praevenosaF. Muell., 1861;P. pithecurusRidl., 1914;P. schlechteriLaut., 1905; as well asP. decandraDing Hou, 1983, from Borneo.