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 EA 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
Australian Systematic Botany 27(1) 48-55 https://doi.org/10.1071/SB13042
Submitted: 27 September 2013 Accepted: 12 May 2014 Published: 30 June 2014
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.
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) first 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 plant–butterfly interaction. Caterpillars of a dozen Papilionidae genera (Allancastria, Archon, Battus, Cressida, Euryades, Losaria, Luehdorfia, 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 world’s largest butterflies, Ornithoptera alexandrae and O. goliath (Straatman and Inoue 1984; Jebb 1993; Parsons 1996a). Each of the 10–13 Ornithoptera taxa is popular among collectors for their iridescent bright colours and their size (Parsons 1992). Because of excessive collecting and habitat loss, birdwing butterflies are listed in CITES (mostly appendix II) and on the IUCN Red List (most listed as Vulnerable or Endangered). As a consequence, the Commonwealth Scientific and Industrial Research Organisation (CSIRO) and many Australian NGOs have taken initiatives to involve the public in conservation efforts, for example, by planting native butterfly 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 butterflies a natural resource and established the Insect Farming & Trading Agency to develop this resource sustainably (Hutton 1985).
Nevertheless, all protection measures require correct identification and accurate species names. This is especially true for the host plants that have received considerably less attention than the butterflies. Indeed, Jebb (1993) highlighted several incorrect identifications 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 butterflies feed, hampering conservation measures of these prominent flagship taxa. As a consequence of co-evolution, the distribution of the butterflies 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.
All species belonging to the subg. Pararistolochia are treated here, providing new combinations and new names. The taxonomic affiliation according to different authors has been critically revised and morphological characters used by these authors for their respective classifications 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 first 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 five or six were characteristic for other species. Masters (1879) suggested that his newly described A. promissa Mast. had close affinities 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, flattened stem, cauliflorous, 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.
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Huber (1960) was the first 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 affinities 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, flower, fruit and seed. When González and Stevenson (2002) published the first cladistic analysis of the Aristolochioideae on the basis of morphology, they followed the classification of Schmidt (1935), with a subgeneric rank for the monophyletic Pararistolochia. Later, the monophyly of Pararistolochia was confirmed 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.
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 field 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. Pflanzenfam. 2nd edn, 16b: 241 1935
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 flattened stem, an actinomorphic, three-lobed perianth, and a longitudinally more or less ribbed berry with a fleshy endocarp (Hutchinson and Dalziel 1927, 1928; Schmidt 1935; Huber 1985). The majority of the species are cauliflorous, except the Australasian species A. deltantha, A. ornithopterae, A. laheyana, A. sparusifolia, A. praevenosa, A. momandul and the African A. goldieana, which also flower on young stems. In comparison to species of subg. Siphisia and subg. Aristolochia that have six anthers, except for Aristolochia subsect. Pentandrae Duch. displaying five 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 confirmed 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).
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Aristolochia alexandriana (Mich.J.Parsons) Buchwalder & Wanke, comb. nov.
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.
Type: Australia: Queensland, State Forest Reserve 310, Rainforest, 780 m, 8 Nov. 1977, 17°13′S 145°42′E, B. Gray 784 (holo: QRS (017289).
Distribution: Australia (Queensland, Wet Tropics).
Aristolochia biakensis (Mich.J.Parsons) Buchwalder & Wanke, comb. nov.
Type: Indonesia: New Guinea, Jappen-Biak Island, Saroerai bij Seroei, 27 Jul. 1939, Aet & Injan 173 (Exp. Ir L.J. van Dijk) (holo: BO (BO1330–2332)).
Distribution: Indonesia (Yapen and Biak Island).
Aristolochia ceropegioides S.Moore, J. Bot. 58: 269 (1920)
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)
Type: Indonesia: western Borneo, Kalimantan, H. Winkler 1256 (holo: L (L0360036 !)).
Distribution: Indonesia (Borneo, Kalimantan).
Aristolochia deltantha F.Muell., Fragm. 6 (46): 179 (1868)
Type: Australia: Queensland, Rockingham’s 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)
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)
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)
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.
Type: Mondah forest, 10 km N of Libreville, Gabon, 2009, D. Nguema et al. 1050 (holo: MO; iso: LBV, WAG) (non Aristolochia fimbriata 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)
Type: Nigeria, Thompson s.n. (holo: K).
Distribution: Cameroon, Guinea, Nigeria, Sierra Leone.
Aristolochia gracilifolia O.C.Schmidt, Bot. Jahrb. Syst. 58: 490 (1923)
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)
Type: Gabon: Chaillu Mts., Songou Mt, between Dibandi and Mouyanama, ~20 km E of Mimongo, 1°37′S, 11°46′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)
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 final conclusion.
Aristolochia kepara (Mich.J.Parsons) Buchwalder & Wanke, comb. nov.
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.
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)
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
Type: in Africa occidentalis tropicae regione dicta Gabon, 1854, Aubry-Lecomte s.n. (holo: P (P00487096 !); iso: MPU (MPU018708))
Type: Gaboon river, 27 Nov. 1881, H. Soyaux 317 (iso: P (P00487098 !)).
Type: Cameroon: Lolodorf, Mar. 1896, G. Staudt 186 (holo: B destroyed?; iso: P (P00487095 !)).
Type: Côte d’Ivoire: 4 Apr. 1909, A. Chevalier 21145 (holo: P? n.v.).
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).
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)
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.
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.
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.
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.
Type: Papua New Guinea: Northern Province, Hegata Village, immediately out of Popondetta, 100-m elev., 16 Mar. 1988, Parsons 2 (holo: LAE (69992)).
Distribution: Papua New Guinea (Northern Province, Oro Province).
Aristolochia momandul K.Schum, in K.Schumann & M.Hollrung, Fl. Kais. Wilh. Land.: 105 (1889)
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.
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 butterflies, that often exclusively feed on Aristolochia species.
Aristolochia paradisiana (Mich.J.Parsons) Buchwalder & Wanke, comb. nov.
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.
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)
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)
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)
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 verified by Keay as a representative specimen of the species.
Aristolochia promissa Mast., Gard. Chron. 11: 494 (1879)
Type: Cameroon: Victoria, Kalbreyer 7 (holo: K).
Type: Gold Coast: Aburi Gardens, W.H.Johnson 487, 1060 (syntypes: K).
Type: Nigeria: Oban, Talbot 128, Talbot 2310, Talbot 1642 (syntypes: BM (BM 000528393 !), K (K000350286 !)).
Type: Nigeria: Oban, Talbot 2318 (BM (BM000528395)).
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)
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.
Type: Papua New Guinea: West Sepik, Bewani Subprovince, Mount Yungat, northern slopes Bewani Mountains, strangler in 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.
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.
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
Type: Gabon: 1°N, 1862, Mann 1851 (holo: K (K000350283 !); iso: P (P00482857 !)).
Type: Uganda: Emin Pascha expedition, 1890–1891, Stuhlmann s.n. (holo: B destroyed?).
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)
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).
References
Bentham G, Hooker JD (1880) Aristolochiaceae. Genera Plantarum 3, 121–125.Bliss BJ, Wanke S, Barakat A, Ayyampalayam S, Wickett N, Wall PK, Jiao Y, Landherr L, Ralph PE, Hu Y, Neinhuis C, Leebens-Mack J, Arumuganathan K, Clifton SW, Maximova SN, Ma H, dePamphilis CW (2013) Characterization of the basal angiosperm Aristolochia fimbriata: a potential experimental system for genetic studies. BMC Plant Biology 13, 13
| Characterization of the basal angiosperm Aristolochia fimbriata: a potential experimental system for genetic studies.Crossref | GoogleScholarGoogle Scholar | 23347749PubMed |
Condamine FL, Sperling FAH, Wahlberg N, Rasplus JY, Kergoat GJ (2012) What causes latitudinal gradients in species diversity? Evolutionary processes and ecological constraints on swallowtail biodiversity. Ecology Letters 15, 267–277.
| What causes latitudinal gradients in species diversity? Evolutionary processes and ecological constraints on swallowtail biodiversity.Crossref | GoogleScholarGoogle Scholar | 22251895PubMed |
Ehrlich PR, Raven PH (1964) Butterflies and plants: a study in coevolution. Evolution 18, 586–608.
| Butterflies and plants: a study in coevolution.Crossref | GoogleScholarGoogle Scholar |
González FA, Stevenson DW (2000) Perianth development and systematics of Aristolochia. Flora 195, 370–391.
González FA, Stevenson DW (2002) A phylogenetic analysis of the subfamily Aristolochioideae (Aristolochiaceae). Revista de la Academia Colombiana de Ciencias Exactas. Físicas y Naturales 26, 25–60.
Hooker JD (1865) Description of some new and remarkable species of Aristolochia from western tropical Africa. Transactions of the Linnean Society of London 25, 185–187.
| Description of some new and remarkable species of Aristolochia from western tropical Africa.Crossref | GoogleScholarGoogle Scholar |
Huber H (1960) Zur Abgrenzung der Gattung Aristolochia L. Mitteilungen Botanische Staatssammlung München 3, 531–553.
Huber H (1985) Samenmerkmale und Gliederung der Aristolochiaceen. Botanische Jahrbücher für Systematik 107, 277–320.
Huber H (1993) Aristolochiaceae. In ‘The Families and Genera of Vascular Plants. Vol. 2’. (Eds K Kubitzki, JG Rohwer, V Bittrich) pp. 129–137. (Springer: Berlin)
Hutchinson J, Dalziel JM (1927) Aristolochiaceae. In ‘Flora of West Tropical Africa. Vol. 1’. pp. 75–78. (Royal Botanic Gardens, Kew: London)
Hutchinson J, Dalziel JM (1928) Tropical African Plants: II. Bulletin of Miscellaneous Information 1928, 22–32.
| Tropical African Plants: II.Crossref | GoogleScholarGoogle Scholar |
Hutton A (1985) Butterfly farming in Papua New Guinea. Oryx 19, 158–162.
| Butterfly farming in Papua New Guinea.Crossref | GoogleScholarGoogle Scholar |
Jebb M (1993) ‘Aristolochia in New Guinea.’ (Christensen Research Institute: Madang, Papua New Guinea)
Keay RWJ (1952) Revision of the ‘Flora of West Tropical Africa’ – I. Kew Bulletin 7, 149–165.
| Revision of the ‘Flora of West Tropical Africa’ – I.Crossref | GoogleScholarGoogle Scholar |
Leal ME, Nguema D (2011) Aristolochiaceae. In ‘Flore du Gabon. Vol. 42’. (Eds MSM Sosef, J Florence, H Borobou, L Ngok Banak) pp. 5–10. (Markgraf: Weikersheim, Germany)
Ma JS (1992) A taxonomic revision on genus Aristolochia subgenus Pararistolochia. Acta Phytotaxonomica Sinica 30, 508–514.
Masters MT (1879) New garden plants. The Gardeners Chronicle 11, 494
Neinhuis C, Wanke S, Hilu KW, Müller K, Borsch T (2005) Phylogeny of Aristolochiaceae based on parsimony, likelihood, and Bayesian analyses of trnL–trnF sequences. Plant Systematics and Evolution 250, 7–26.
| Phylogeny of Aristolochiaceae based on parsimony, likelihood, and Bayesian analyses of trnL–trnF sequences.Crossref | GoogleScholarGoogle Scholar |
Ohi-Toma T, Sugawara T, Murata H, Wanke S, Neinhuis C, Murata J (2006) Molecular phylogeny of Aristolochia sensu lato (Aristolochiaceae) based on sequences of rbcL, matK, and phyA genes, with special reference to differentiation of chromosome numbers. Systematic Botany 31, 481–492.
| Molecular phylogeny of Aristolochia sensu lato (Aristolochiaceae) based on sequences of rbcL, matK, and phyA genes, with special reference to differentiation of chromosome numbers.Crossref | GoogleScholarGoogle Scholar |
Parsons MJ (1992) The World’s largest butterfly endangered: the ecology, status and conversation of Ornithoptera alexandrae (Lepidoptera: Papilionidae). Tropical Lepidoptera 3, 33–60.
Parsons MJ (1996a) New species of Aristolochia and Pararistolochia (Aristolochiaceae) from Australia and New Guinea. Botanical Journal of the Linnean Society 120, 199–238.
Parsons MJ (1996b) A phylogenetic reappraisal of the birdwing genus Ornithoptera (Lepidoptera: Papilionidae: Troidini) and a new theory of its evolution in relation to Gondwanan vicariance biogeography. Journal of Natural History 30, 1707–1736.
| A phylogenetic reappraisal of the birdwing genus Ornithoptera (Lepidoptera: Papilionidae: Troidini) and a new theory of its evolution in relation to Gondwanan vicariance biogeography.Crossref | GoogleScholarGoogle Scholar |
Poncy O (1978) Le genre Pararistolochia, Aristolochiaceae d’Afrique tropicale. Adansonia, Sér. 2 17, 465–494.
Ross EM, Halford DA (2007) Aristolochiaceae. Pararistolochia linearifolia. In ‘Flora of Australia. Vol. 2 Winteraceae to Platanaceae’. (Ed. AJG Wilson) p. 250. (Australian Biological Resources Study and CSIRO Publishing: Melbourne)
Schmidt OC (1935) Aristolochiaceae. In ‘Die natürlichen Pflanzenfamilien. Vol 16b’, 2nd edn. (Eds A Engler, K Prant) pp. 204–242. (Engelmann: Leipzig, Germany)
Simonsen TJ, Zakharov EV, Djernaes M, Cotton AM, Vane-Wright RI, Sperling FAH (2010) Phylogenetics and divergence times of Papilioninae (Lepidoptera) with special reference to the enigmatic genera Teinopalpus and Meandrusa. Cladistics 26, 1–25.
Straatman R, Inoue S (1984) Identified hostplants of Ornithoptera species. Transactions of the Lepidopterological Society of Japan 34, 124–126.
von Mueller FJH (1860–1861) Aristolochiaceae. Aristolochia praevenosa. In ‘Fragmenta Phytographiae Australiae, Vol. 2’. p. 166. (Auctoritate Gubern. Coloniæ Victoriæ, Ex Officina Joannis Ferres: Melbourne)
Wanke S, González F, Neinhuis C (2006) Systematic of Pipevines: combining morphological and fast-evolving molecular characters to investigate the relationship within subfamily Aristolochioideae (Aristolochiaceae). International Journal of Plant Sciences 167, 1215–1227.
| Systematic of Pipevines: combining morphological and fast-evolving molecular characters to investigate the relationship within subfamily Aristolochioideae (Aristolochiaceae).Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2sXhtFylurk%3D&md5=eba7e00a8f60ad8bb69f1a507bd88b94CAS |
Wanke S, Jaramillo MA, Borsch T, Samain MS, Quandt D, Neinhuis C (2007) Evolution of the Piperales: matK gene and trnK intron sequence data reveal lineage specific resolution contrast. Molecular Phylogenetics and Evolution 42, 477–497.
| Evolution of the Piperales: matK gene and trnK intron sequence data reveal lineage specific resolution contrast.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD28Xht1CgsbnO&md5=fa8365a9b2e63a0666744ffee597e070CAS | 16978885PubMed |
