New species ofAristolochiaandPararistolochia(Aristolochiaceae) from Australia and New Guinea

Botanical Journal of the Linnean Society (1996), 120: 199–238. With 47 figures

New species of Aristolochia and Pararistolochia (Aristolochiaceae) from Australia and New Guinea MICHAEL J. PARSONS Entomology Section, Natural History Museum of Los Angeles County, 900 Exposition Blvd., Los Angeles, CA 90007, U.S.A.

Received January 1995, accepted for publication November 1995

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 to Aristolochia, whereas the other is clearly accommodated in Pararistolochia, the latter previously regarded as being solely a tropical African genus. The distinction between Pararistolochia and Aristolochia is discussed. It is concluded that the Australia and New Guinea taxa were found to substantiate circumscription of Pararistolochia. Two new species of Aristolochia (A. nauseifolia and A. goliathiana) are described, as well as 12 species of Pararistolochia (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 of P. meridionaliana (subsp. popondettensis and subsp. milnensis) are described. Eleven other taxa from Australia and New Guinea are newly combined under Pararistolochia: P. deltantha F. Muell., 1868; P. dictyophlebia Merr. & Perry, 1948; P. dielsiana O.C. Schmidt, 1923; P. engleriana O.C. Schmidt, 1927; P. gracilifolia O.C. Schmidt, 1923; P. laheyana Bail., 1912; P. momandul K. Sch., 1889; P. praevenosa F. Muell., 1861; P. pithecurus Ridl., 1914; P. schlechteri Laut., 1905; as well as P. decandra Ding Hou, 1983, from Borneo. ©1996 The Linnean Society of London

ADDITIONAL KEY WORDS: — Afrotropical – Indo-Australian – Indonesia – Irian Jaya – Ornithoptera – Papilionidae – Papua New Guinea – Sepik – taxonomy – Troides.. CONTENTS Introduction . . . . . . . . . Genus Pararistolochia . . . . . . . New combinations of taxa . . . . Descriptions of new species . . . . Rationale for descriptions . . . Floral morphology and terminology Leaf terminology . . . . . . Aristolochia . . . . . . . . Pararistolochia . . . . . . . Acknowledgements . . . . . . . References . . . . . . . . . .

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M. J. PARSONS INTRODUCTION

The Aristolochiaceae of tropical Australia and New Guinea (i.e. the Australian subregion of the Indo-Australian region as approximately bounded by Wallace’s Line) have attracted a certain level of interest among entomologists in recent years because of their importance as larval foodplants of the spectacular birdwing butterflies Troides and Ornithoptera of the swallowtail family Papilionidae (e.g. Igarashi, 1977; Haugum, 1981; Straatman & Inoue, 1984; Weintraub, 1995). This prompted my own studies of these vines two decades ago. Because of the particular closeness of the relationship between the birdwings and their aristolochiaceous larval foodplants it is particularly desirable to have a clear understanding of the specieslevel taxonomy of these plants. It is frustrating when even the most basic taxonomy of these plants is confused. Most species are poorly described, while others are unnamed. During the course of this on-going study, and whilst employed by the Papua New Guinea Government, I undertook a detailed conservation study of the Ornithoptera. This included an extensive survey of their larval foodplants (Parsons, 1983). I have continued herbarium and fieldwork studies of Australian subregion Aristolochia to the present (e.g. most recently in Indonesian New Guinea-Irian Jaya: Parsons, 1992, 1995), and have been able to compare and contrast these species with similar fieldwork elsewhere in the Indo-Australian region (e.g. southernmost China, Malaysia, Brunei, Australia), as well as in the New World tropics, and North America. Lauterbach (1914) and Schmidt (1923) provided important early studies of New Guinean Aristolochiaceae, and Schmidt (1932) and Jones & Gray (1988) laid foundations for the identification of Australian Aristolochiaceae. Some important type localities of Aristolochia species collected by Ledermann in the Sepik River catchment area of what was then German New Guinea (now East and West Sepik Provinces, Papua New Guinea), and described by Schmidt (1923), were identified and discussed by Frodin & Gressitt (1982) and Veldkamp, Vink & Frodin (1988). Hou (1983a, 3b, 1984) further improved knowledge of Malesian (i.e. including New Guinean, but not Australian) Aristolochiaceae. However, his revisionary work for Flora Malesiana fell short of being a definitive means of identifying many of the New Guinea taxa as it was necessarily based on only rather limited pressed herbarium material. Parsons (1983) and Jebb (1991) provided illustrated synopses of the New Guinean region Aristolochiaceae. However, both these unpublished documents were essentially working papers reflecting incomplete states of knowledge at the time of their restricted release. Consequently, they contain incorrect determinations of species. Their main strengths are that they illustrated certain hitherto poorly known taxa, most notably their flowers, which provide the key characters by which species can most easily and reliably be recognized. Having travelled and collected extensively in New Guinea, and in particular Papua New Guinea, I have been able to ascertain hitherto unrealized problems of taxonomy in the Aristolochiaceae. As these result from factors such as hybridization, Hou was understandably entirely unable to appreciate them from rather scant herbarium material. As many Australian subregion Aristolochia are difficult, if not impossible, to identify without flowers, Hou’s problems were compounded by the paucity of fertile material for study and the usually rather inadequate herbarium sheet data. In addition, a thorough revision of the New Guinean Aristolochiaceae

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requires an accurate knowledge of Australian taxa and would be incomplete without including these species. Where fertile material has been available to field collectors, the basic specimen sheet descriptions of the flowers of these vines are invariably insufficient to identify accurately poorly known taxa to species. Part of the reason for this is that field observers have obviously found it difficult to describe succinctly, yet accurately, the usually rather complex and unusual Aristolochia flowers. Where material has been fertile, the results from pressing (rather than alcohol preserving) flowers have been rather disappointing as these lose much of their structure and colour, becoming very difficult to comprehend structurally. An almost complete lack of alcohol preserved specimens, or good colour photographs from the field has, therefore, been a serious shortcoming until now, and Hou (1984: 64) stressed this in a note to future collectors of Malesian specimens, whilst Jebb (1992) noted the importance of making voucher specimens in general for New Guinean Aristolochia. As a result of these problems, I experienced many early difficulties in identifying New Guinean Aristolochiaceae to species when comparing the newly studied live material and the newly gained ecological data, with the results finally presented in Hou’s series of publications. In fact, the published descriptions of most of the New Guinean taxa became an admixture of characters derived from several different species because these facies were summarized from mixtures of herbarium material. Another major set-back to the study of New Guinea Aristolochia has resulted from the apparent loss of a large number of O.C. Schmidt’s type specimens. Schmidt (1923) described six Aristolochia species from the Sepik River region of the then German occupied northern portion of the eastern half of Papua New Guinea. These taxa (A. lauterbachiana, A. novoguineensis, A. ledermannii, A. dielsiana, A. gracilifolia and A. crassinervia) are important because, as Hou (1984: 108) pointed out, they were almost certainly lost in Berlin during World War II, and duplicates either do not exist or have not yet been relocated. In the absence of type material, Schmidt’s rather basic original written descriptions provide little assistance in determining which taxa they represent, especially since the material on which he based his work was crucially (flowers) incomplete in a number of cases. These descriptions are critically analysed below in the light of my field knowledge of the Sepik River region Aristolochiaceae, and species from other areas throughout Papua New Guinea. Hou (1984) found that he could not resolve to species Schmidt’s taxa A. lauterbachiana and A. novoguineensis, or ascertain whether they are truly distinct. From their descriptions, their single-lobed perianth clearly places them in the first of two distinctive groups of Australian subregion Aristolochiaceae. The first group is characterized by the 1-lobed (unilabiate) perianth of the flower, the second by a distinctly 3-lobed (trilabiate) perianth.

GENUS PARARISTOLOCHIA

Pararistolochia Hutch. & Dalz. Fl. W. trop. Africa. 1: 75 (1927). (Type: P. triactina Hook.f.). Pararistolochia was originally erected by Hutchinson & Dalziel (1927) for a group of Afrotropical region (Ethiopian) Aristolochia. Hutchinson (1969: 498) stated that he was struck by a few African species included in Aristolochia, and that “For these I established a different genus … distinguished by having perfectly actinomorphic

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3-lobed calyx (perianth) and very elongated, ribbed, hard, indehiscent fruit in contrast to that of Aristolochia (sensu stricto) with rounded dehiscent fruits.” In the second edition of Flora of West Tropical Africa, Hutchinson & Dalziel (1954) provided a check list of 12 Pararistolochia species. In a dedicated study of Pararistolochia, Poncy (1978) noted that the genus was not always recognized by botanists because it had been poorly defined. Nevertheless, based on a detailed morphological study of all characters (mainstem, leaves and petioles, cauliflory, inflorescences, flowers, pollen), as well as the general ecologies and biogeography of the nine Afrotropical region species listed, Poncy was confident of the distinctness of Pararistolochia from Aristolochia. As he correctly pointed out, all Pararistolochia species (i.e. also including those recognized here as occurring within the Australian subregion) are confined to the humid tropics, whereas Aristolochia has greatly broadened its range into the subtropics, and even (weakly) into certain temperate regions of the world, as well as containing species that have become adapted to markedly arid world regions (e.g. Arizona, USA; Mexico; Australia, and likely elsewhere). Huber (1960) pointed out that, in the Aristolochiaceae, cleistocarpy was previously only credited to Pararistolochia, but that it also occurs in New Guinea A. momandul, and that the species, like Pararistolochia, has a 6 [actually 3]-lobed perianth (Fig. 18). Hou (1983a) noted that his newly described Bornean endemic, Aristolochia decandra, is “closely related to those [species] of Pararistolochia of tropical Africa by: 1) a flattened stem, appearing as ‘8’ on cross section, 2) by being cauliflorous, 3) having a 3-lobed perianth, and 4) in the higher number [10] of stamens.” In fact, the ‘figure 8’ condition of the A. decandra stem (the medullary rays radiating in two directions from a central line when viewed in cross section) referred to by Hou (1983a) and figured photographically for two Malesian species by him (Hou, 1983b: 245, fig. 7a, and 1984: 102, fig. 16) is characteristic of all the Australian subregion species with a 3-lobed perianth, as opposed to radiating uniformly in a normal rounded stem as in typical Aristolochia. Hou’s (1983b) illustration of the cross-section of the stem of A. singalangensis from W. Sumatra, confirms that this species (a representative of the characteristic ‘dutchman’s pipe’ flowered group of Aristolochia), like Pararistolochia, also possesses the bi-directional medullary ray character [e.g. as figured for African P. zenkeri (Engl.). Hutch. & Dalz. by Hutchinson & Dalziel (1954)]. Nevertheless, as Hou pointed out, the fruit of A. singalangensis is dehiscent, being the only Malesian species to split from the fruit apex towards its base. In fact, this particular mode of dehiscence also defines the ‘dutchman’s pipe’ flowered group of Aristolochia centred in mainland South-east Asia and the Oriental region. Therefore, the bi-directional medullary ray character cannot be used as a definitive character of Pararistolochia as it is shared with the ‘dutchman’s pipe’ group of Aristolochia (which probably represents a valid species group of Aristolochia based on the medullary ray character, plus the distinctive general facies of its flowers). In Hou (1984: 64) the editor of Flora Malesiana (C.G.G.J. van Steenis) was motivated to add a note into the text body “Whether the West African genus Pararistolochia can be maintained is liable to doubt. KEAY (Fl. W. Trop. Afr. ed. 2, 1, 1, 1954, 77) distinguished it from Aristolochia by: ‘fruit indehiscent, elongated, strongly ribbed, cucumber-like’, but these characters seem also to occur in the New Guinean A. dielsii [sic! i.e. dielsiana] SCHMIDT … Editor.” Actually, in effect this highlighted the incorrect placement of the taxon dielsiana in Aristolochia, rather than pointing out the supposed taxonomic weakness of Pararistolochia. Following Hou’s

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revisionary studies of Indo-Australian region Aristolochiaceae, Huber (1985) published his findings on the seed anatomy of representatives of all Aristolochiaceae, except Saruma, and contrasted these with the classification of the family based on other characters. From his results, Huber, like Poncy (1978), accepted Pararistolochia as a distinct genus and, furthermore, proposed that Aristolochia (sensu lato) could be subdivided into still more genera. As hinted at in his earlier 1960 work cited above, Huber (1985), noting data in Hou (1983b and 1984), suggested that African Pararistolochia also contains at least five Malesian species (he included by name A. decandra, A. dielsiana, A. momandul and A. engleriana), and that the same geographical disjunction is also known in other closely related plant genera. As its fruit was unknown, and Jongkind (1990) was not sure of whether it belonged in Pararistolochia, the author preferred to describe a new aristolochiaceous liana from Gabon, tropical West Africa, under Aristolochia. Apart from the extent of the syrinx, bract size and bract presence or absence, and the cauliflorous habit, Jongkind maintained that “the more or less equally 3-lobed perianth limb, the large number of anthers and stigma lobes or the very long cucumber-shaped fruits are not universally manifest in Pararistolochia”. Nevertheless, only one species was mentioned as occupying a supposedly intermediate position between Aristolochia and Pararistolochia, and Jongkind (like Huber, 1985) rightly pointed out that the “Asian” (actually Australian subregion) species could possibly help to solve this “taxonomical dilemma”. In light of the above comments by previous authors, it is interesting to note that the numerous stable characters of the Australian subregion group of species with trilabiate perianths only taxonomically strengthens, rather than invalidates, the generic distinctness of Pararistolochia. Furthermore, this provides another extremely important example of the biogeographical link of the Australian subregion with Africa in terms of plate tectonics and Gondwana (to be discussed in detail by Parsons in prep.: Revision of the Australian and New Guinean Aristolochiaceae). From my own studies of herbarium material of the African taxa I concur with those authors (e.g. Hutchinson & Dalziel, Huber, Poncy, Verdcourt) who consider Pararistolochia to be clearly distinct from Aristolochia. Moreover, my cladistic analyses of the troidine (tribe Troidini) papilionid butterflies whose larvae feed on these vines also clearly point to Pararistolochia being distinct on the basis of this evidence (Parsons, in press). In particular, Ornithoptera (a very distinct genus of troidine papilionids in New Guinea and Australia) are primarily Pararistolochia feeders, whereas members of the related, mostly Oriental subregion genus Troides are primarily Aristolochia feeders. Weintraub’s (1995) independent research on troidines and their aristolochiaceous foodplant relations in the Oriental subregion has provided similar findings and, notably, he also recognized Pararistolochia as being distinct in a cladistic analysis of the Aristolochiaceae. Certain species seemingly diffuse the distinction between Aristolochia and Pararistolochia. Nevertheless, if all characters of such taxa are assessed as complete sets then it is invariably quite clear as to which of the two genera they belong. The elongate, succulent, indehiscent fruit of Pararistolochia, as opposed to the rounded, dry basket-like dehiscent fruit of Aristolochia, is the best, most effective, character by which the two genera may be told apart. However, there are a few species where this distinction is slightly obscured. For example, in New Guinea, Aristolochia crassinervia and A. goliathiana both possess unilabiate flowers which clearly place them in Aristolochia, whereas their fruits show a definite secondary specialization towards

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succulence and indehiscence as in Pararistolochia. However, A. crassinervia and A. goliathiana are noteworthy since their fruits still clearly retain the six cleavage lines by which they do eventually dehisce once almost dry (photographically well illustrated by Hou, 1983b: 235, figs 4a and 4b, and Hou, 1984: 104, fig. 17). These fruits are, however, obviously adapted to be consumed (whilst still succulent and newly turning orange) by vertebrates (e.g. fruit bats). The presence of edible pulp and very tough wingless seeds within them also confirms this. It is relevant and important to mention here that Hou’s (1984: 97) statement that the fruits of New Guinea P. momandul ultimately dehisce is incorrect. The observation arose from the misinterpretation by Hou (1983b: 241) of an ink drawing, by myself, of P. pithecurus (assumed to be A. momandul) which I sent Hou (along with other drawings). Unfortunately, he erroneously assumed that the strong shadowing of the prominently longitudinal 6-ribbed fruit represented the fruit dehiscing, whereas this was not the case. Like all Pararistolochia fruits, those of P. momandul and P. pithecurus are succulent and indehiscent when ripe, the hard deltoid seeds being dispersed in the faeces of birds and mammals who consume the fruit. In fact, those of P. pithecurus resemble the well known star fruit, Averrhoa carambola (Oxalidaceae), commercially grown throughout the Indo-Australian region, except that the star fruit is only 5-ribbed. Despite previous speculation as to whether Pararistolochia is a truly taxonomically valid grouping, Verdcourt (1986, and pers. comm., 1985) recognized the genus in his contribution to the study of the flora of tropical East Africa. He included an improved description of Pararistolochia, stating that it is “unquestionably abundantly distinct and worthy of generic rank.” In his key, Verdcourt straightforwardly and unambiguously separated Pararistolochia from Aristolochia as follows: Fruit indehiscent, elongate, strongly ribbed; perianth regular 3-lobed or irregular with 1 or 3 long tails from the mouth of the limb; stigmatic lobes 6–12; anthers 6–24; flowers mostly on the old wood ........................................................1. Pararistolochia Fruit dehiscent, short; perianth irregular with a single lobe [long lip], or 2-lipped, with a broad standard and long keel; stigmatic lobes up to 6; anthers up to 6; flowers borne on leafy shoots ............................................................................ 2. Aristolochia This key can be used, entirely unmodified, to place all Australian subregion species to their relevant genera. All those whose flowers bear trilabiate perianths key out to Pararistolochia, whereas those with unilabiate perianths key to Aristolochia. In fact, apart from its distinctive characters of mostly cauligerous flowering habit, the 3-lobed perianth and indehiscent succulent fruit, this group of species in the Australian subregion is also characterized by the possession by all species of sclerophyllous leaves that are extremely tough, strong and which (when mature) are frequently brittle in texture. This is also true of Afrotropical region Pararistolochia. In marked contrast, the mature leaves of Australian subregion Aristolochia (sensu stricto) are soft and glabrous in texture. Based on evidence from related genera (Thottea, Holostylis and Euglypha e.g. see Pfluge-Gregory, 1956) it appears that six anthers and style lobes represents the plesiomorphic (i.e. ancestral) condition in both Aristolochia and Pararistolochia. Therefore, those species of Pararistolochia (mainly in the Afrotropical region) which possess higher anther and style lobe numbers can be considered to be derived in this respect, just as certain Aristolochia species possess reduced (5 or exceptionally 3) anther and style lobe numbers. All of the known Australian subregion Pararistolochia have the

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more unspecialized number of six stigmatic lobes and six anthers. Pararistolochia decandra Ding Hou from Borneo is unusual in its ten stigmatic lobes and ten anthers. A similar trend of increase in stigmatic lobes and anthers is present in the African species, many of which often possess nine stigmatic lobes and nine anthers. Other similarities between Afrotropical region and Australian subregion Pararistolochia are noteworthy. For example, the bud, flower, fruit and seed of Papua New Guinea P. meridionaliana are very similar to those of P. triactina, the tropical African type species for Pararistolochia. In addition, the utricle of Australian species P. laheyana Bail. bears a pair of ventral swellings that also very closely resemble those of P. triactina. Pararistolochia, therefore, contains the following alphabetically arranged 23 Australian subregion taxa (full synonymies will be presented by Parsons in prep.): Previously described taxa formally combined below: P. P. P. P. P. P. P. P. P. P. P.

decandra (Ding Hou, 1983) M.J. Parsons deltantha (F. Muell., 1868) M.J. Parsons dictyophlebia (Merr. & Perry, 1948) M.J. Parsons dielsiana (O.C. Schmidt, 1923) M.J. Parsons engleriana (O.C. Schmidt, 1927) M.J. Parsons gracilifolia (O.C. Schmidt, 1923) M.J. Parsons laheyana (Bail., 1912) M.J. Parsons momandul (K. Sch., 1889) M.J. Parsons praevenosa (F. Muell., 1861) M.J. Parsons pithecurus (Ridl., 1914) M.J. Parsons schlechteri (Laut., 1905) M.J. Parsons

Taxa newly described below: P. P. P. P. P. P.

P. P. P. P. P. P.

alexandriana M.J. Parsons, sp. nov. australopithecurus M.J. Parsons, sp. nov. biakensis M.J. Parsons, sp. nov. kepara M.J. Parsons, sp. nov. manokwariensis M.J. Parsons, sp. nov. meridionaliana M.J. Parsons, sp. nov. : subsp. meridionaliana M.J. Parsons, subsp. nov. : subsp. popondettensis M.J. Parsons, subsp. nov. : subsp. milnensis M.J. Parsons, subsp. nov. paradisiana M.J. Parsons, sp. nov. peninsulensis M.J. Parsons, sp. nov. tithonusiana M.J. Parsons, sp. nov. sepikensis M.J. Parsons, sp. nov. sparusifolia M.J. Parsons, sp. nov. linearifolia M.J. Parsons, sp. nov.

Acronyms used for herbaria containing material discussed or described below are — Australia: QRS = Division of Forest Research, Commonwealth Scientific and Industrial Research Organisation (CSIRO), Atherton; BRI = Queensland Herbarium, Indooroopilly; CANB = Australian National Herbarium CSIRO, Canberra; DNA = Northern Territory Herbarium, Darwin; MEL = National Herbarium of Victoria, South Yarra, Victoria. Papua New Guinea: BFC = The Forestry College of Papua New Guinea herbarium, Bulolo; IFTA = Insect Farming and Trading Agency, Bulolo; LAE = Papua New Guinea National Herbarium, Forestry

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Research Institute, Lae; UPNG = University of Papua New Guinea Herbarium, Waigani. Indonesia: BO = Herbarium Bogoriense, Bogor. Poland: WRSL = Museum of Natural History, Wroclaw University. Germany: B = Botanischer Garten und Botanisches Museum, Berlin-Dahlem. The Netherlands: L = Rijksherbarium, Leiden. United Kingdom: BMNH = British Museum of Natural History, London; K = Herbarium of the Royal Botanical Gardens, Kew, Richmond. United States of America: A = Herbarium of the Arnold Arboretum, Harvard University, Cambridge, Massachusetts. When referring to habitat in the descriptions, I follow Beehler’s (1978) categories of the various forest zones in mainland New Guinea (altitudinal ranges approximate): Lowland 0–500 m), Hill (500–1000 m), Lower montane (1000–1600 m), Mid montane (1600–2100 m), Upper montane (2100–2500 m).

NEW COMBINATIONS OF TAXA

Pararistolochia decandra (Ding Hou) M.J. Parsons comb. nov. Aristolochia decandra Ding Hou, Blumea 28 (1983) 343.-Holotype: Hans Winkler 1256 (L) Kalimantan (Western Borneo). Pararistolochia deltantha (F. Muell.) M.J. Parsons, comb. nov. Aristolochia deltantha F. Muell., Fragm. (1868) 179.-Lectotype (here designated): Dallachy s. n. (MEL 1553306) Rockingham’s Bay, Queensland, Australia. Aristolochia deltantha var. deltantha: Bail., Queensland Agric. J. 28 (1912) 199. Pararistolochia dictyophlebia (Merr. & Perry) M.J. Parsons, comb. nov. Aristolochia dictyophlebia Merr. & Perry, J. Arnold Arbor. 29 (1948) 152.-Holotype: Clemens 4901 (A; iso. BRI), Ogeramnang, Huon Peninsula, Morobe Province, Papua New Guinea. Pararistolochia dielsiana (O.C. Schmidt) M.J. Parsons, comb. nov. Aristolochia dielsiana O.C. Schmidt, Bot. Jahrb. 58 (1923) 490.-Holotype: Ledermann 9169 (B, n. v.), Etappenberg (Base Mountain), East Sepik Province, Papua New Guinea. Pararistolochia engleriana (O.C. Schmidt) M.J. Parsons, comb. nov. Aristolochia engleriana O.C. Schmidt, Fedde Rep. 23 (1927) 288, nom. nov. Aristolochia ledermannii O.C. Schmidt, Bot. Jahrb. 58 (1923) 489, non Engler 1911.-Holotype: Ledermann 12055 (B, n. v.), Schr¨aderberg (Schrader Mountain), East Sepik Province, Papua New Guinea. Pararistolochia gracilifolia (O.C. Schmidt) M.J. Parsons, comb. nov. Aristolochia gracifolia O.C. Schmidt, Bot. Jahrb. 58 (1923) 490.-Holotype: Ledermann 12458 (B, n. v.), Felsspitze (Rocky Peak), West Sepik Province, Papua New Guinea. Pararistolochia laheyana (Bail.) M.J. Parsons, stat. nov., comb. nov. Aristolochia deltantha F. Muell. var. laheyana Bail., Queensland Agric. J. 28 (1912)

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199.-Holotype: C. T. White s.n. (BRI 333077) MacPherson Range, Queensland, Australia. Pararistolochia momandul (K. Sch.) M.J. Parsons, comb. nov. Aristolochia momandul K. Sch., in K. Sch. & Hollr., Fl. Kais. Wilh. Land. (1889) 105.-Holotype: Hollrung 520 (B, n. v.; iso. BO, WRSL) Konstantinhafen, Astrolabe Bay, Madang Province, Papua New Guinea Pararistolochia praevenosa (F. Muell.) M.J. Parsons, comb. nov. Aristolochia praevenosa F. Muell., Fragm. 2 (1868) 166.-Lectotype (here designated): Beckler s.n. (MEL 1553302) Clarence River, Queensland, Australia. Pararistolochia pithecurus (Ridl.) M.J. Parsons, comb. nov. Aristolochia pithecurus Ridl., J. Bot. 52 (1914) 296.-Holotype: Forbes 621 (BM; iso. L, WRSL), Sogeri region, Mt. Koroko, 2500 ft., Central Province, Papua New Guinea. Pararistolochia schlechteri (Laut.) M.J. Parsons, comb. nov. Aristolochia schlechteri Laut., in K. Sch. & Laut., Nachtr. Fl. Deut. Schutzgeb. S¨udsee. (1905) 260.-Holotype: Schlechter 14156 (part of the type and carbon rubbing print of leaves, WRSL), Ramu River Valley, near Madang, Madang Province, Papua New Guinea.

DESCRIPTIONS OF NEW SPECIES

Rationale for descriptions This work is a precursor to a more complete revision of Australian subregion species which will contain complete keys to all species, as well as ecological details of all species described below (Parsons in prep.: Revision of the Australian and New Guinean Aristolochiaceae). A family description may be found in Hou (1984). For the new taxa numbered 4 through 7 brevity in description is achieved by comparing them with the first newly described Pararistolochia species (meridionaliana sp. nov.). P. meridionaliana is described in greater detail to provide a baseline description with which other taxa are then compared. Where applicable, the main differences for each new species are also emphasized by their comparison with their nearest relatives (particularly for new taxa numbered 8 through 14). Besides working with incomplete or infertile material, previous authors of new taxa notably neglected to make comparative descriptions, or to state accurately how their new taxa are related to, or differ from, their nearest relatives. As the leaves of Australian subregional Pararistolochia are frequently very variable in size and shape, whilst all are coriaceous (sclerophyllous), the level of confidence of making accurate species determinations is usually extremely low when this must be done with only foliage available. For this reason, the formal descriptions below particularly emphasize the characteristic or unique features of each species.

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Floral morphology and terminology The flowers of Pararistolochia and Aristolochia are three-dimensionally complex, serving as bulbous-ended, funnel-shaped, tubular traps for insects to enable pollination. Therefore, floral terminology is important and is provided in Figure 1 (based on that of Pfeifer, 1966). A noteworthy floral feature of some Australian subregional Pararistolochia is the ‘filament’. This is a connate filiform process formed from the greatly extended apices of the three true lobes of the perianth which remain fused or twisted (Figs 1 and 17–22), thereby holding the apices of the lobes together and forming three apertures to the flower between each lobe (rather than a single aperture where the lobes are able to fully open out). The filament is a modification of the extremely elongate perianth lobes characteristic of P. schlechteri and P. meridionaliana, defined here as ‘tendrils’ (Figs 8–13). The distinction between the filament and tendrils is important as, although each represents the same part of the flower, the filament forms an upright, non-fleshy, very narrow and linear, single, thread-like process, whereas the tendrils are horizontal, fleshy, distinctly tapered, free extensions of each perianth lobe. The filament has clearly been secondarily lost by some species. Examples exist in both filament-bearing and freely-lobed flower types, of flowers in which the perianth lobe bases have become so prominently developed laterally that they form ‘pseudo-lobes’. Good examples of these flower types are, respectively, P. biakensis (Fig. 22) and P. schlechteri (Fig. 13). For this reason, the perianth tube of P. schlechteri was originally described by Lauterbach (as A. schlechteri) as being uniquely 6-lobed. The gynostemia of all Australian subregional Pararistolochia are very similar, resembling that of P. meridionaliana(Fig. 7), so these are not repetitively described for each species. The gynostemium comprises closely united styles, stigmas and anthers. The style is apically divided into six distinct stigmatic lobes, below which is a ring of six pairs of adnate anthers above the short stipe-like stalk.

Leaf terminology Leaves of Pararistolochia and Aristolochia are typically heart-shaped (e.g. Figs 27–46). The term sinus (e.g. Hou, 1984) is used to describe the U- or V-shaped condition of the portion of the leaf where the petiole joins the lamina, the leaf auricles (lobes) standing either side. Obviously, where the sinus is deep the auricles are long and prominent, sometimes even overlapping one another above the petiole.

Aristolochia Aristolochia nauseifolia Parsons, sp. nov. (Figs 4 and 27) Folia ubi obtrita odorem nauseosum emittentia, caudex proprie amplus dauciformis; A. puberae Duchartre valde affinis sed foliis majoribus perlongioribus angustioribusque, plantis non omnino pubescentibus; A. thozetii F. Muell. arcte affinis sed foliis typice latioribus, lobo perianthii unilabiato breviore.

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Figure 1. Details of floral morphology of Pararistolochia pithecurus (Ridl.) M.J. Parsons comb. nov.

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HERBACEOUS CLIMBER to 1 m long with woody, carrot-like rootstock. STEMS slender, c. 1.5 mm in diameter. LEAVES with petiole 11–25 mm, sparsely pubescent or almost glabrous; lamina linear-lanceolate, with prominently laterally dilated, rounded auricles, soft, glabrous, basally cordate, to 11 cm long by 1.5 cm across widest part of lamina, 2.5 mm across auricles; sinus broadly U-shaped, to 1 cm deep; some basal leaves shorter and deltoid, to 5 cm long by 2.3 cm; veins reddish tinged, basal veins 2 pairs, the inner pair ascending to c. 1/2 of the blade, the outer one much shorter, with branches extending upward or to the auricles, elevated and prominent beneath, distinct above; lateral veins hardly distinguishable from the veinlets; veinlets closely reticulate, elevated and prominent beneath, faint or invisible above. INFLORESCENCES sessile, singly in leaf axils. FLOWERS purple, to c. 2.5 cm long overall, utricle 5 mm long by 4 mm wide. PERIANTH tubular, shortly unilabiate, tube 5 mm long, limb 12 mm long. Pedicel 4 mm long, ovary 4 mm, glabrous. FRUIT ovoid, 15 mm long by 10 mm, pendulous, dehiscent when dry brown and ripe, opening to an inverted parachute.

Holotype. Australia, Queensland, 3 km from Kubin Village, Moa Island [middle Torres Strait], 30 m alt., 14.xi.1989, B. Gray 5007 (QRS 091512). Other material examined. AUSTRALIA, Queensland: Thursday Island, Jaheri 108917 (BO 1330-2294); Northern Territory: Elcho Island, J.R. Maconochie 2222 (DNA 9797, K). Distribution and habitat. Islands of the Northern Territory and the Torres Strait of northern Queensland, Australia; 0–50 m. According to Jones & Gray (1988), and G. Sankowsky (pers. comm., 1991), the species also occurs in central coastal mainland Queensland, in the vicinity of Marlborough, but I am not aware of any voucher material from these areas. T. Lambkin (pers. comm., 1988) stated that, during a visit to Murray Island in the Torres Strait, he located an Aristolochia, the leaves and fruit of which he sketched, and which closely resemble those of the type of A. nauseifolia. Aristolochia nauseifolia grows rapidly during the summer wet season, becoming dormant in the dry season, the stems dying back to the rootstock. In central coastal Queensland the species grows in open forest, the stems twining around grass and other low herbage (Jones & Gray, 1988). The specific epithet, nauseifolia, was chosen to describe the aromatics of the leaves which, when crushed, emit a nauseous odour (described by Jones & Gray for central coastal Queensland specimens, and as a ‘strong smell’ in the collecting data of the holotype). From my experience of other species, this is assumed to be similar to the acrid odour emitted by leaves of certain south American introduced ornamental vines if crushed, as well as by northern Queensland populations of Australian Figures 2–10 Aristolochia flowers. When figures drawn from photographs, these are the author’s unless otherwise specified. 2. A. pubera after Jones & Gray (1988: photo pl. on p. 151), Australia. 3. A. thozetii from A. Hiller photo, Macham’s Beach, Cairns, Australia. 4. A. nauseifolia sp. nov. holotype, B. Gray QRS 091512, 3 km from Kubin Village, Moa Island [middle Torres Strait] Queensland, Australia. 5. A. goliathiana sp. nov. from M. Parsons photos, Manki Ridge, Bulolo, Morobe Province, Papua New Guinea. 6. A. crassinervia from photos of living material of M. Parsons UPNG 15903, Lonem Village, near Maprik, East Sepik Province, Papua New Guinea. (Figures 7–10. Pararistolochia flowers. 7. P. meridionaliana sp. nov. subsp. meridionaliana gynoecium from photo of living material of M. Parsons UPNG 15906, Pongani to Afore Road just before Ondoro Village, Northern Province, Papua New Guinea. 8. meridionaliana subsp. P. meridionaliana, as above. 9. P. meridionaliana subsp. popondettensis subssp. nov. (perianth lobes not in fully open position) from photo of living holotype of M. Parsons LAE 69992, Hegata Village, immediately out of Popondetta, Northern Province, Papua New Guinea. 10. P. meridionaliana subsp. meridionaliana as above.

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indigenous Aristolochia pubera Duchartre. Aristolochia nauseifolia is closely related to A. pubera (Fig. 2) and A. thozetti F. Muell. (Fig. 3), both of which are semi-arid (eucalypt savannah habitat) adapted endemic Australian herbaceous, ground sprawling twiners. However, A. nauseifolia has broader leaves than typical A. thozetti, and the perianth limb of its flower (Fig. 4) is shorter, resembling that of A. pubera. Its leaves (Fig. 27) are, however, larger, far more elongate and narrower than those of A. pubera, and A. nauseifolia is not wholly pubescent as is typical A. pubera. Aristolochia goliathiana Parsons, sp. nov. (Figs 5, 28 and 29) Aristolochia crassinervia sensu Ding Hou, 1984. Fl. males. ser. 1, 10: 104, pro parte, excl. typus, non Schmidt (1923). In forma floris pro maxima parte A. tagalae Cham. simulans; A. crassinerviae O.C. Schmidt valde affinis sed foliis perminoribus sinubus albis auriculis plerumque imbricatis, flore pro parte maxima atroroseomarronino, lobo perianthii unilabiato longo typice irregulariter torsivo, fructu majore. or LIANA to 5–10 m tall, occasionally to 20 m, the leaves and stems smelling of orange-peel when crushed. STEMS round, 0.5–2 cm in diameter; bark straw yellow, thick, corky, longitudinally irregularly deeply dissected; branches forming mainly in canopy and at base of mainstem, initially smooth, green, soon turning pustular and corky. LEAVES with petiole 6–8 cm, lamina sparsely pubescent or almost glabrous; when mature dark green above, pale green below, white above at base of sinus and petiole, subcoriaceous, to 14–17 cm long by 8–9 cm wide; base cordate, sinus narrow, to 2.5–3 cm deep; auricles usually overlapping, rounded; apex acuminate or shortacuminate; glabrous above, minutely hairy beneath; basal veins 2 or 3 pairs, the inner pair similar to the midvein, ascending to near the apex of the blade, the outer one much shorter, with branches extending upward or to the auricles, elevated and prominent beneath, distinct above; lateral veins hardly distinguishable from the veinlets; veinlets closely reticulate, elevated and prominent beneath, faint or invisible above; young leaves pale yellow, soft, fleshy. INFLORESCENCES cauligerous, rarely in axils of leaves, racemiform, rhachis up to 3 cm long, internodes very short or condensed, sometimes the bracteate flowering rhachis very short, flowers appearing fascicled; bracts lanceolate, 5–10 mm long, sparsely minutely pubescent on both surfaces, glabrescent. FLOWERS to c. 6.2 cm long overall; almost wholly dark pinkishmaroon, but perianth limb with white areas, utricle almost wholly white. PERIANTH without basal stipe and with a contraction between perianth and ovary, glabrous outside, venation rather faint; utricle subglobose, c. 7 mm in diameter, hairy inside, with two glandular round bodies (c. 0.5 mm in diameter); tube c. 12 mm long by 2.5 mm wide, glandular-hairy inside; limb 1-lipped, apically or almost wholly spirally twisted, up to 45 mm long by 4 mm wide with glandular hairs on the inner surface. Pedicel c. 20 mm long. STAMENS 6; anthers oblong, c. 0.7 mm long. GYNOSTEMIUM c. 1 mm long; ovary c. 20 mm long, glabrous; style 6-lobed; lobes triangular, c. 0.5 mm long, the basal part of each lobe slightly extended outward (c. 0.3 mm long and wide). FRUIT pendulous, fleshy pale green when mature, pale orange when ripening, hard, dark brown when ripe, tardily dehiscent, cylindric or oblong, not ridged or angular, 5.5 cm long by 3 cm wide (overall 9.5 cm long, including stalk) minutely granular. SEEDS triangular or deltoid, 7 mm long by 6–7 mm wide, not winged, rather smooth or obscurely muriculate on both surfaces; funicle broadened and covers the upper surface. TWINER

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Holotype. Papua New Guinea, Morobe Province, Wau Sub-District, Kulolo Creek, 149°18' E, 9°38' S, on edge of creek in secondary forest, 4800 ft alt., H. Streimann & A. Kairo NGF 45343 (LAE); iso.: A, BFC, BRI, CANB 218091, K, L. Other material examined. PAPUA NEW GUINEA, Central Province: Sogeri Plateau, Straatman (UPNG 00189). Morobe Province: Boana, 3000 ft, Clemens 41750 (A); Palenkwa, Golden Pines, Wau subdistrict, 3000 ft, Streimann & Kairo NGF 39387 (L, CANB 206546); Garaina, 2500 ft, Hutton (LAE). Milne Bay Province: Raba Raba subdistrict, Biniguni Mayu I track, 90 m, Streimann & Leach 28577 (BRI, CANB 303823, K, L); Obulaku Village and vicinity [Kiriwina, Trobriand Islands], Frodin 09190 (UPNG 4099). SOLOMON ISLANDS: Fiu River, Malaita, B. Stone 2400 (K, L); Tarahao, N.W. Santa Ysabel, W. Beer’s collectors BSIP 7825 (K, LAE). Distribution and habitat. Mainland New Guinea, and some satellite islands, including Kiriwina in the Trobriands, Papua New Guinea; also certain islands of the Solomons, including Malaita and Santa Ysabel. A. goliathiana occurs throughout much of the central cordillera of mainland New Guinea in marginal and advanced secondary forest, as well as primary lower montane to mid montane forest, at altitudes recorded between 90–1800 m. In mainland Papua New Guinea the species most frequently occurs in hill forest and lower montane forest from 500–1500 m. The late R. Straatman (in litt., 1979) stated that he had found A. goliathiana (as sp. indet.: no voucher) growing it about 2500 m near Nabire in Irian Jaya, Indonesia. On Kiriwina Island the species can be found growing in areas of open secondary vegetation in coral-based soils, and sprawling over the raised coral outcrops that comprise most of the island, from almost sea level to an elevation of about 5 m. The specific epithet, goliathiana, was chosen to emphasize the importance of this species as a primary larval foodplant of the Goliath Birdwing Butterfly, Ornithoptera goliath Oberth¨ur, 1888. Aristolochia goliathiana has been confused in the past with A. crassinervia. Nevertheless, from its original description, and the geographical position of its type locality, A. crassinervia is a distinctly different species as was confirmed by study of living material in the East Sepik Province, Papua New Guinea (Fig. 6) for which eight specimen sheets from a plant originally from Drekikir, East Sepik Province, Papua New Guinea (M. Parsons 5: A, CANB, K, LAE, UPNG 15903) were made. These are, to date, the only authentic specimens of A. crassinervia presently in collections. Although the type for A. crassinervia is apparently lost, and the original specimen was infertile, Schmidt’s description contains enough information to be reasonably certain of the true identity of the species based on the Drekikir specimen. Several, very obvious characters clearly separate A. goliathiana from A. crassinervia. For example, in his original description Schmidt (1923) recorded that the leaves of the type of A. crassinervia were 5–19 cm long by 4–12 cm wide. Although similar, the leaves of A. goliathiana, however, only ever attain a size of about 14 by 8 cm. They exhibit a degree of clinal variation and, although heart-shaped, become distinctly more rounded moving eastwards along the New Guinea mainland Central Cordillera, so that on Kiriwina Island in the Trobriands and at Alotau (Milne Bay Province, Papua New Guinea) (e.g. UPNG 4099), and in the Solomons (e.g. BSIP 7825, Santa Ysabel Island) they are very rounded (Fig. 28). Whereas the leaves of A. crassinervia are distinctively wine-red above at the base of the sinus where it joins the petiole, those of A. goliathiana are characteristically white. In A. crassinervia the leaf

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auricles never overlap as they frequently do in A. goliathiana. The contrast in the foliage of the two species is also obvious in its overall appearance as the leaves of A. crassinervia tend to hang down from, or sprawl irregularly throughout, the canopy, whereas those of A. goliathiana form more regular alternating rows at the tops of its mainstems and in the canopy. Jebb (1991: 7) pointed out that A. goliathiana (as A. crassinervia) can be identified by “its very prominent 2° venation on the lower surface contrasting with the almost flat upper surface, although other species may also have prominent venation, the upper surface is usually bullate, and the leaf less leathery.” When mature, the leaves of A. goliathiana hardly vary in shape, but the younger leaves may be slightly more proportionately elongate and, like the youngest shoots are, distinctively, pale yellow, soft and fleshy. When crushed, the leaves and stems of A. goliathiana have perhaps the most pungent and pleasant odour of all the New Guinean Aristolochiaceae, being much stronger than that of A. crassinervia. The leaves of A. goliathiana emit a strong, lasting orange peel-like odour. This is also true of any section of the stem, which smells even more strongly. The stems, even when dead and dry for over three years, give off this scent when broken open. The flowers of A. goliathiana and A. crassinervia differ markedly in size, shape and colouration. Those of A. goliathiana (Fig. 5) more closely resemble A. tagala Cham. in their general morphology and their notably long perianth limb. The flower of A. crassinervia (Fig. 6) is, however, distinctly more erect with a very short and broadly flared perianth and short erect limb, and is almost wholly pale pink. In contrast, the obviously more elongate A. goliathiana flower is mostly dark pinkish-maroon, its long perianth limb characteristically irregularly spiralled. The A. goliathiana fruit is very similar to that of A. crassinervia, but is much larger. That of A. crassinervia is only about 3 cm long by 1.3 cm wide, and the fruit has an overall length of about 5.5 cm, including its stalk. Pararistolochia Pararistolochia meridionaliana Parsons, sp. nov. (Figs 7, 8, 10) Aristolochia dielsiana sensu Ding Hou, 1984. Fl. males. ser. 1, 10: 105, pro parte, excl. typ. non Schmidt (1923). Pararistolochiae schlechteri Laut. arcte affinis sed foliis permajoribus auriculis non prominentibus, 3 cirrhis perianthii longioribus latioribusque ventraliter involutis, basibus loborum perianthii concavis. Slender climbing vine or liana. STEMS to 15–40 m high, rounded, up to 4–6 cm in diameter; bark of oldest stems straw yellow, thick, corky, longitudinally irregularly deeply dissected; branches forming mainly in canopy and at base of main stem. LEAVES with petiole 2.5–8 cm long, often densely pubescent; lamina when young basally cordate, somewhat lanceolate, densely pubescent, pale yellow, supple; when mature dark green above, paler below, strongly coriaceous, broad-ovate, spadeshaped, 19–37 (–45) cm long by 4.5–23.5 (–40) cm wide; base shallowly or extremely mildly subcordate; auricles broad and distinct, sinus 0.5–3 cm deep; apex acuminate, rarely apiculate; pubescent on both surfaces, or only ventrally pubescent in very old leaves (upper pubescence lost), especially on midvein, veins and veinlets; veins elevated and prominent beneath, slightly elevated above; basal veins 1 pair,

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ascending upward to 1/2–3/4 of the blade, each vein with a weaker, short basal branch; lateral veins 3–5 pairs; veinlets crossbar-like or reticulate, slightly elevated, prominent beneath, main veins distinct above. INFLORESCENCES mainly cauligerous, rarely in axils of terminal leaves, solitary or paired on a brachyblast arising from the (old) stem, rachides up to 45 cm long, with spaced internodes usually every c. 15 mm; bracts very small, c. 1 mm, densely pubescent. Pedicel c. 12 mm long. FLOWER BUDS densely pubescent, elongate-pyramidally pointed, with 3 short, angular basal prominences. FLOWER to 7 cm overall. PERIANTH tube weakly S-curved, c. 25 mm long, enlarging throughout to c. 20 mm in diameter at well-rounded throat, 3-lobed, lobes narrowly triangular, each c. 50 mm overall length, margins ventrally and inwardly rolled, medially constricted to 3 filiform irregularly curving tendrils c. 25 by 2 mm; utricle obovoid or ellipsoid c. 12 mm long by 8 mm wide, densely woolly hairy inside over an irregularly dark purple-black basal half, upper half near syrinx smooth, white or pale greenish-white, no glandular bodies visible. Utricle and perianth externally greenish-white, main veins weakly purple highlighted, throat bright yellowish-green with zone of prominent, white, downwardly-directed setae, aperture equilaterally Y-shape constricted by prominent intrusion of bright sulphuryellow medial portion of each lobe base, perianth lobes dark lime-green above, tendrils pale orange, lobe midveins and a ring around throat dark purple-maroon. GYNOSTEMIUM 3.5 3 5.5 mm long, lower part c. 1 mm, stipe-like; ovary c. 12 mm long, densely pubescent; style 6-lobed, lobes linear 2.5–3 mm long, no annular ring at base. FRUIT a pendulous prominently 6-ribbed capsule, to 35 cm long by 6 cm wide, hard and green when mature, black and soft when ripe. Holotype. Papua New Guinea, Central Province, along Hiritano Highway, in disturbed forest by the roadside, H. Hopkins & M. Parsons 913 (LAE 69991); iso.: K, UPNG (13444, duplicates for chemical analysis — see Rali and Griffin, 1991). Other material examined. PAPUA NEW GUINEA, Central Province: Brown River Forestry, near Teak Forest, 30 m, R. Carver 73 (LAE 67855); Mamai Plantation, Port Glasgow, Straatman M6 (LAE); Brown River, c. 100 ft, Hutton 73 (LAE). Northern Province: Pongani to Afore Road just before Ondoro Village, 460 m, M. Parsons 3 (K, UPNG 15906). Distribution and habitat. Endemic to south-eastern mainland Papua New Guinea. Three subspecies are recognized here (see below). P. meridionaliana subsp. meridionaliana occurs either side of the Owen Stanley Range in the Central Province in the region of Brown River, north-west of the capital, Port Moresby, and in the Northern Province around the hill station of Afore on the Managalase Plateau. It occurs in advanced secondary forest, often in marginal situations, and primary hill forest. Present records show that subsp. meridionaliana ranges from several metres above sea level (e.g. Brown River) to 800 m (e.g. Managalase Plateau). The specific epithet, meridionaliana, was chosen to emphasize the importance of this species as a primary larval foodplant of the delicately tailed Meridion Birdwing Butterfly, Ornithoptera meridionalis (Rothschild, 1897). Pararistolochia meridionaliana has been confused in the past with Aristolochia (i.e. Pararistolochia) dielsiana. Nevertheless, from its original description, and the geographical position of its type locality, P. dielsiana is a distinctly different species as was confirmed by study of herbarium material from the East Sepik Province, Papua New Guinea. Several, very obvious characters clearly separate P. meridionaliana from P. dielsiana. For example, in the

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original description, Schmidt (1923) recorded that the leaves of the type of P. dielsiana were only 9–11.5 cm long by 2.3–4 cm wide. Although similar, the leaves of P. meridionaliana (Fig. 34) attain much larger sizes. For example, in an extreme case, the late R. Straatman (in litt. 1979) recorded one leaf from an Afore specimen as being 1 m long by 0.7 m wide (although I have never found them to be quite as large in the same area). Pararistolochia meridionaliana is geographically restricted to the southeasternmost ‘arm’ of mainland Papua New Guinea, whereas typical P. dielsiana was collected in westernmost Papua New Guinea on Etappenberg (Base Mountain), Westkette (West Range) in the extreme west of the East Sepik Province in the general region of the May River headwaters. The flowers of P. dielsiana are impossible to comprehend accurately from the original description of the species, and in the absence of the actual type specimen. However, an apparently authentic specimen of P. dielsiana has since been collected from the Hunstein River region, near Ambunti, East Sepik Province by Hoogland & Craven (CANB 10753). The flowers of this specimen are notably different in the position of their perianth lobes and the morphology of the throat in life. The elongate, tendril-like lobes are apically separate, apparently basally connate, and have moved to a position superior to the throat, one to each of three ovate apertures comprising the flared portion of the throat. Pararistolochia meridionaliana is also related to P. schlechteri from which it differs from the type material (WRSL) studied, including Schlechter’s original drawings; also by its much larger leaves, and in the markedly different flower. Whereas the flower of P. schlechteri Laut. (Fig. 13) has perianth lobes in which the tendrils are much narrower and dorsally inwardly rolled, that of P. meridionaliana (Figs 8 and 10) is broader, being ventrally inwardly rolled. The perianth lobe bases of P. meridionaliana are concave, whereas in P. schlechteri they are medially extremely extended, forming three additional, elongate, triangular, non-tendrilled ‘pseudo-lobes’ between each true lobe (making the perianth tube effectively sixlobed). Pararistolochia meridionaliana Parsons subsp. popondettensis Parsons subsp. nov. (Figs 9, 34) Pararistolochiae meridionalianae subsp. meridionalianae arcte affinis sed foliis magis elongatis, ad 33 cm longis et 20 cm latis, interdum ad 50 cm longis et 25 cm latis, sinubus ad 3.5 cm profundis, floris faucis ore subdeltoideo, pallide flavo, maculis atromarroninorubris dense maculato, lobulis perianthii interdum sparse pubescentibus praecipue secus costas, capsulis ad 25 cm longis. Very similar in all characters to P. meridionaliana subsp. meridionaliana, but differing in the following features: LEAVES usually more elongate, to c. 33 cm long by 20 cm wide (occasionally up to 50 by 25 cm), sinus to 3.5 cm deep. FLOWER with mouth of throat subdeltoid, pale yellow, heavily blotched and mottled with dark maroon-red; perianth lobes dorsally maroon-red, sometimes sparsely pubescent, especially along midveins. FRUIT to only c. 25 cm long. Holotype. Papua New Guinea, Northern Province, Hegata Village, immediately out of Popondetta, 100 m, 16. iii. 1988, M. Parsons 2 (LAE 69992). Figures 11–13. Pararistolochia flowers. 11. P. alexandriana sp. nov. holotype, from photo of living material of M. Parsons LAE 69994, Ahora Village, north of Popondetta town, Northern Province, Papua New Guinea. 12. P. kepara holotype from photo of living material of M. Parsons LAE 69995, near Kamondo Village, near Kokoda, Northern Province, Papua New Guinea. 13. P. schlechteri holotype after drawing by R. Schlechter (WRSL).

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Other material examined. PAPUA NEW GUINEA, Northern Province: Popondetta, Straatman 231 (LAE 90452); Hegeta Village, Popondetta, Benjamin sheet No. 238317 (LAE 67855); Idua-Haijo logging area, 3 km north-east of Hohota Village, J. Waikabu & A. Kairo sheet No 228552 (LAE 70277); Jonita area, Nr. Popondetta, M. Parsons 015 (A, K, LAE). Distribution and habitat. Endemic to the hinterlands of the Popondetta region of the Northern Province, mainland Papua New Guinea. Subspecies popondettensis occurs in primary and secondary forest on the lowland coastal plain, often in marginal situations, rarely to within 1 km of the coast and not far into the surrounding foothills; 10–500 m. The subspecific epithet, popondettensis, is derived from the town of Popondetta, the hinterlands of which contain this subspecies of P. meridionaliana. Subsp. popondettensis is a primary larval foodplant of Queen Alexandra’s Birdwing Butterfly, Ornithoptera alexandrae (Rothschild, 1907). Pararistolochia meridionaliana Parsons subsp. milnensis Parsons subsp. nov. Pararistolochiae meridionalianae subsp. meridionalianae valde affinis sed floris fauce rotundato albo crassinervato maculis maculis amorphis coccineis maculato, zonis pilorum alborum deorsum curvatorum indistincta debilibi, apertura parva rotundata profunda perexpansa lobulorum basibus non intrusis, basibus lobulorum medialiter pallide flavis, lobulis et lobulorum cirrhis dorsaliter omnino sparse pubescentibus totius coccineis, longioribus et parum tenuioribus, ad 6.5 cm adjecte longis, fructu ad 20 cm tantum longo. Very similar in most characters to P. meridionaliana subsp. meridionaliana, but differing in the following features: FLOWER throat rounded, white, heavily veined and blotched with crimson-red, zone of downwardly-directed white setae more diffuse and not as prominent, aperture small, round and deep, broadly flared and not constricted by prominent intrusion of each lobe base, lobe bases medially pale yellow; perianth lobes and tendrils dorsally entirely sparsely pubescent, wholly bright crimson-red, longer and slightly more slender, each to c. 6.5 cm overall length. FRUIT up to only c. 20 cm long. Holotype. Papua New Guinea, Milne Bay Province, Pini Range, near Alotau, Milne Bay, c. 150 m, 1. vi. 1991, M. Parsons 023 (LAE 69993). Distribution and habitat. Endemic to south-easternmost mainland Papua New Guinea and presently known only from the region of the type locality in primary and secondary rainforest, often in marginal situations; 20–150 m. The subspecific epithet, milnensis, is derived from Milne Bay, the hinterlands of which contain this subspecies of P. meridionaliana. As in the case of P. meridionaliana subsp. meridionaliana in the Central Province, subsp. milnensis is a primary larval foodplant of Ornithoptera meridionalis. Pararistolochia alexandriana Parsons, sp. nov. (Figs 11, 35) Pararistolochiae meridionalianae Parsons subsp. popondettensi Parsons valde affinis sed foliis minoribus constanter magis elongatis, flore minore dense pubescenti atroroseorubro, lobulis perianthii brevioribus basaliter ad faucem angustioribus, cirrhis perianthii

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minus evolutis, marginibus perianthii loborum ventraliter perinvolutis (partibus maximis lobulorum ventraliter superpositis). Very similar in characters to P. meridionaliana subsp. popondettensis, but differing in the following features: STEMS to only 4 cm in diameter. LEAVES always elongate, but usually only to c. 25 cm long by 16 cm wide, sinus to 1.5 cm deep. INFLORESCENCES much more numerous, frequently longer, rachides up to c. 50 cm, with spaced internodes more numerous and usually every c. 8 mm; bracts larger (c. 2–5 mm long). FLOWERS smaller, usually more abundant, mouth of throat equilaterally Y-constricted, white, densely pubescent, heavily veined with dark pinkish-red; perianth lobes shorter, each to c. 4 cm overall length, more evenly tapering, margins more strongly inwardly rolled, overlapping most of the lobe length ventrally, lobes basally narrower at throat and with less distinct tendrils, dorsally entirely densely pubescent, dark pinkish-red. FRUIT only c. 15 cm long by 4 cm wide. Holotype. Papua New Guinea, Northern Province, Ahora Village, north of Popondetta town, supporting trees forming secondary forest up to 10 m tall, c. 100 m, 14. iii. 1988, M. Parsons 1 (LAE 69994). Distribution and habitat. Sympatric with lowland populations of P. meridionaliana subsp. popondettensis in the hinterlands of Popondetta on the surrounding lowland coastal plain in primary and secondary rainforest, often in marginal situations. rarely to within 1 km of the coast; 20–150 m. The specific epithet, alexandriana, was chosen to emphasize the importance of this species as a larval foodplant of Ornithoptera alexandrae. Until now, P. alexandriana has been overlooked because of its similarity to P. meridionaliana subsp. popondettensis, which like P. alexandriana, but unlike the other two subspecies of P. meridionaliana, is also a primary foodplant of O. alexandrae. Further confusion has arisen from the fact that typical P. meridionaliana normally possesses lime-green flowers (e.g. north and south of the Owen Stanley Range), whereas on the Popondetta Plain its flowers (Fig. 9) are maroon-red and thus more closely resemble the flowers of P. alexandriana (Fig. 11). Also, the two species often hybridize where they are sympatric, producing flowers which exhibit a range of intermediate characters (presumably depending also on the extent of backcrossing and recrossing). Besides the morphological differences outlined above, the flowers of P. alexandriana emit a very pleasant and fairly obvious scent, lacking in all subspecies of P. meridionaliana. Pararistolochia kepara Parsons, sp. nov. (Figs 12, 43) Pararistolochia meridionalianae Parsons subsp. popondettensi Parsons arcte affinis sed foliis minoribus constanter magis elongatis, inflorescentiis perpaucioribus saepe brevioribus, flore typice flavovirenti sed disperse pallide turcosomaculato dorsaliter omnino dense pubescenti setis atroroseorubris. Very similar in characters to P. meridionaliana subsp. popondettensis, but differing in the following features: CLIMBER to 20 m tall. STEMS to only 2 cm in diameter. LEAVES always elongate, but usually only to c. 22 cm long by 9 cm wide, basally subcordate, sinus barely 1 mm deep, usually absent in squarely truncate young terminal leaves. INFLORESCENCES much less numerous, frequently shorter, rachides up to c. 20 cm, with irregularly spaced internodes every c. 8–15 mm; bracts larger (c. 2–5 mm long). FLOWERS utricle 13 mm long, perianth tube c. 26 mm long, lobes and tendrils to c. 6.5 cm, more evenly tapering, margins more strongly inwardly rolled, overlapping

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most of lobe length ventrally, lobes basally narrower, throat rounded and perianth midveins only slightly raised and intrusive; perianth lobes and tendrils dorsally entirely heavily pubescent with dark pinkish-red setae, lime-green with pale turquoise highlights, except apical 12 mm of tendril suffusing to dark pink, main lateral veins and marginal side veins of lobes minutely dotted with dark pinkish-red; interior of apex of utricle below syrinx heavily dark red speckled. FRUIT to only c. 20 cm long, more prominently ribbed. Holotype. Papua New Guinea, Northern Province, near Kamondo Village, near Kokoda, 1 km U.T.M. grid square EL 8714, on supporting tree in primary forest, c. 400 m, 18. v. 1991, M. Parsons 017 (LAE 69995); iso. K. Distribution and habitat. Endemic to mainland Papua New Guinea in the south-eastern approach into the Kokoda Valley, Northern Province, and presently only known from this type locality area in the vicinity of the villages of Kepara, Sengi and Kamondo, where it grows in advanced secondary forest and primary forest; 300–500 m. The specific epithet, kepara, was chosen from the village name of Kepara at the approximate centre of the type locality area for the species. Besides the morphological differences between the flowers of P. kepara (Fig. 12) and P. meridionaliana (Fig. 10) outlined above, those of P. kepara, as in P. alexandriana, emit a very pleasant and fairly obvious perfume. This character, and the densely pubescent strongly rolled perianth lobes, together with the leaf shape of P. kepara, show that this is a sister species to P. alexandriana. It is not known whether P. kepara is utilized by any Ornithoptera species, but it is likely a primary larval foodplant of the common Priam Birdwing, Ornithoptera priamus (Linnaeus, 1758), as the butterfly and plant are sympatric. If O. alexandrae truly occurs in the Kokoda area — as, based on present day distributional data, the butterfly’s type locality was apparently erroneously stated by its collector, Meek (1913) to be Biagi near Kokoda (which is not far from the type locality area of P. kepara) — then P. kepara may eventually also prove to be an important larval foodplant for O. alexandrae. Pararistolochia manokwariensis Parsons, sp. nov. (Figs 15, 36, 37) Pararistolochiae meridionalianae Parsons ubique simulans sed foliis saepe minoribus apicibus insigniter acuminatis auriculis bene evolutis ventraliter involutis supra petiolis imbricatis; floribus perpaucioribus omnino pubescentibus pilis tenuibus lobulis perianthii typice latis subdeltoideis marginibus extimis crassissimis ventraliter involutis. Generally similar in characters to P. meridionaliana subsp. meridionaliana, but differing in the following features. CLIMBER to c. 10 m tall. STEMS to only 2 cm in diameter. Figures 14–21. Pararistolochia flowers. 14. P. tithonusiana sp. nov. holotype, including bud (bottom) from photo of living material of M. Parsons (BO), Wamar´e to Mokwam foot track, c. 1500 m, Irian Jaya, Indonesia. 15. P. manokwariensis sp. nov. holotype, bud only, from photo of living material of M. Parsons (BO), Wamar´e to Mokwam foot track, c. 500 m Irian Jaya, Indonesia. 16. P. paradisiana sp. nov., including bud (bottom) from D’Abrera (1975: photo pl. p.99), 17. P. pithecurus comb. nov. from photo of living material, Bulolo area, Morobe Province, Papua New Guinea. 18. P. momandul comb. nov. photo of living material of M. Parsons (K), near Sakita (Perombaturu) Village, Kumusi Timber Area, Northern Province, Papua New Guinea. 19. P. sparusifolia sp. nov. after Jones & Gray (1988: photo pl. on p. 151), Mt. Lewis, Queensland, Australia. 20. P. australopithecurus sp. nov. after Jones & Gray (1988: photo pl. on p. 151) Mt. Bartle Frere, Queensland, Australia. 21. P. sepikensis sp. nov., including bud (right), based on holotype, K. Karenga NGF 56530 A, Mt. Yungat, northern slopes Bewani Mts., Bewani Subprovince, West Sepik Province, Papua New Guinea.

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LEAVES with petiole to 7 cm long; lamina narrowly ovate, apex acuminate, to c. 45 cm long by 17–20 cm wide, base cordate, sinus to 4 cm deep, auricles often apically inwardly curved, overlapping above petiole. INFLORESCENCES cauligerous, 1 or 2 on very short, bracteate rachides (flowers appearing almost sessile), rhachis to 12 mm long, bracts minute and internodes condensed, pubescent. Pedicel c. 7 mm long. FLOWERS less abundant, wholly finely pubescent (adhering to fabric), arising at irregular intervals all along main stem, from just above ground level into canopy. PERIANTH tube S-curved, c. 15 mm long by 4 mm wide, slightly constricted below throat, then broadly flaring to throat, perianth lobes subdeltoid in dorsal profile, downwardly angled and ventrally inwardly rolled in lateral profile, ventral margin c. 3 mm long outer margin c. 20 mm broad, mouth of throat equilaterally broadly Y-constricted at each lobe base (i.e. not constricted at each lobe midvein); utricle c. 10 mm long by c. 6 mm wide. Utricle and perianth externally white, main veins weakly highlighted by pink, especially along basal region of tube; throat white, forming a sharply defined, almost completely circular base to each perianth lobe, remainder outer portion of lobe maroon-red to pinkish-mauve. Ovary c. 9 mm long. FRUIT only immature specimen studied, but probably very similar to that of the following species when mature.

Holotype. Indonesia, Irian Jaya, Wamar´e to Mokwam foot track, open primary hill forest, c. 500 m, 22. xi. 1992, M. Parsons 002 (BO); iso.: K. Other material examined. INDONESIA, Irian Jaya: Manokwari, Afanshari, c. 20 m, W. Vink 1969 (BO 956244); Warsamson Valley, east of Sorong, Jayapura, 50 m, W. Vink 17561 (LAE, L). Distribution and habitat. Endemic to the subcoastal region and foothills of the Arfak Mountains of mainland Irian Jaya, Indonesia; c. 20–1100 m. P. manokwariensis occurs in advanced lowland secondary forest (e.g. in the region of the Manokwari University area at about 20 m altitude: pers. obs., 1992), and primary hill forest. At Bini Village in the Arfaks (900–1100 m) Sius Mandosir collected fruits of manokwariensis (seen., 1992). The specific epithet, manokwariensis, was chosen from the town of Manokwari, in the hinterlands of which this species occurs. It shares a sibling relationship with the following species: their common ancestry assumed from the very close morphological similarities of their flowers (see below). Pararistolochia tithonusiana Parsons, sp. nov. (Figs 14, 38) Pararistolochiae manokwariensi Parsons valde affini sed foliis insigniter minioribus atroviridibus saepe apiculatis coriaceoribus nervis infra prominentibus minus flavis pubescentia densiore, flore circa duplo majore lobulis perianthii dorsaliter rubrobrunneis fauce laete sulphureo. Generally similar in characters to P. manokwariensis, but differing in the following facies: CLIMBER to c. 15 m tall. LEAVES with petiole to 6 cm long, densely pubescent; lamina smaller, to 32 cm long by 15 cm wide, more strongly coriaceous, veins and veinlets more prominent beneath, more densely pubescent, basal veins 1 very prominent pair, ascending upward to 3/4 of the blade, other lateral veins barely more prominent than crossbar-like veinlets running at right angles between midvein and main pair of basal veins, veinlets reticulate; sinus to c. 4.5 cm deep, auricles prominent, inwardly directed, c. 3 cm wide; apex often apiculate. INFLORESCENCES

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larger. Pedicel 28 mm long. FLOWER perianth tube c. 35 mm long, ventral margin of lobe 12 mm long; utricle 15 mm long by 10 mm wide. Utricle and perianth externally white, tube main veins slightly more strongly and evenly highlighted by pale maroon, throat forming a sharply defined, almost completely circular base to each perianth lobe, bright sulphur-yellow, weakly speckled by brownish-maroon setae, changing to white in lower portion of throat, remainder of each lobe outwardly dark brown with reticulate network of blackish-brown veins. Ovary 18 mm long. FRUIT ovoid or slightly pyriform, prominently 6-ribbed, to c. 12 cm long by 6.5 cm wide, green and hard when mature, bright orange and soft when ripe. Holotype. Indonesia, Irian Jaya, Wamar´e to Mokwam foot track, open lower montane primary forest, c. 1500 m, 22. xi. 1992. M. Parsons 005 (BO). Distribution and habitat. Endemic to the Arfak Mountains of mainland Irian Jaya, Indonesia. P. manokwariensis is presently known to occur in open lower montane primary forest at about 1500 m, but the altitudinal range of the species will likely be found to be much broader once further collections are made. The specific epithet, tithonusiana, was chosen to emphasize the importance of this species as a larval foodplant of the Tithonus Birdwing Butterfly, Ornithoptera tithonus de Haan, 1840. Pararistolochia tithonusiana shares a sibling relationship with P. manokwariensis and it is possible that the two are sympatric in some localities as their type localities are very close to one another. However, P. manokwariensis appears to be a predominantly lowland species, whilst P. tithonusiana is a mid to upper montane species. Pararistolochia manokwariensis has large tough pale green leaves and small flowers, about half the size of P. tithonusiana, whilst the similar but larger-flowered P. tithonusiana has smaller, notably much thicker and even tougher darker green leaves, the leaf veins being even more prominent, densely pubescent, and less yellow. The main leaf differences were well illustrated in a composite colour photographic plate, which also included a dorsal view of the flower of P. tithonusiana (Parsons, 1995: 264). Even in smaller buds of P. tithonusiana the throat of the perianth tube is creamy-yellow internally and the perianth lobes are more broadly and definitely brown, not distinctly reddish as in those of P. manokwariensis. The flowers of both species are covered with setae that permit them to adhere easily to fabric. Pararistolochia paradisiana Parsons, sp. nov. (Figs. 16, 30, 31) Pararistolochia momandul K. Sch. (Fig. 18) et P. pithecuro (Fig. 17) arcte affinis sed foliis in catervis 6–10-foliatis dactyliformibus aggregatis, catervis foliorum in caulibus terminalibus effuso-dependentibus alternantibus, auriculis foliorum interdum nullis saepe minutis raro valde ventraliter involutis supra petiolis imbricatis, flore lobulis perianthii libere aperientibus filis terminalibus nullis. Slender climbing vine. STEMS to 5–10 m high; c. 1 cm in diameter. LEAVES aggregated in hand-shaped groups of 6–10 leaves, the groups arranged in alternate rows on terminal stems; petiole 1–2 cm long; lamina coriaceous, ovate-lanceolate, to 22–26 cm long by 6.5–9 cm; base shallowly subcordate; auricles often minute, sometimes absent, occasionally overlapping tightly above petiole at join with blade, sinus 1–3 mm deep; apex apiculate; smooth on both surfaces; veins elevated and prominent beneath, slightly elevated above; basal veins 1 pair, ascending upward to 1/3–1/2 of the blade, a pair of short weak veins below these at auricles; lateral veins 4 of 5 pairs; veinlets crossbar-like or reticulate, slightly elevated and prominent

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beneath, main veins distinct above. INFLORESCENCES in upper leaf axils, sometimes cauligerous, solitary or paired, rachis up to 8 cm long, with spaced internodes every c. 5 mm; bracts minute and pubescent. FLOWER c. 37 mm overall. PERIANTH tube weakly S-curved, c. 15 mm long by 7 mm wide, enlarging throughout, broadly flaring to c. 12 mm in diameter at deep circular throat; perianth 3-lobed, subdeltoid, apically rounded and medially indented at apex of midvein, 12 mm long; utricle obovoid, 10 mm long by 7 mm wide. Utricle and perianth externally white, strongly brownishpurple veined, throat white, heavily finely dark brown speckled, with zone of downwardly-directed prominent white setae immediately above small circular aperture, perianth lobes above pale yellow, obscurely brownish veined, sinus between each lobe dark purple-black margined, tapering away almost at apex of lobe. FRUIT small and densely brownish pubescent when first forming; when mature a hard, green, pendulous, elongate-ovoid, prominently 6-ribbed capsule, to 11.5 cm long by 4.5 cm wide, bright orange and soft when fully ripened. Holotype. Papua New Guinea, East Sepik Province, Lonem Village, near Maprik, growing in mature, well-spaced secondary forest, c. 200 m, 28. iii. 1988, M. Parsons 4 (LAE 69996); iso. A, K, CANB, UPNG. Other material examined. PAPUA NEW GUINEA, East Sepik Province: Amuk River, Amahop, Maprik Subdistrict, M. Benjamin sheet No. 231911 (LAE 67796); Imbiap Village, Maprik, P. Clark NGF 49885, sheet No. 232392 (LAE); Maprik, 100 m, Hutton NGF 49852 (LAE); Maprik, 100 m, Hutton NGF 49851 (LAE). West Sepik Province: Creek 6 km N. of Bewani, c. 40 km SSW of Vanimo, Bewani Sub Province, 130 m, J. Wiakabu et al. NGF 50018 (A, LAE, QRS 091892). Distribution and habitat. Endemic to mainland New Guinea, throughout the Torricelli and Bewani Ranges of the East and West Sepik Provinces, Papua New Guinea, and apparently westwards throughout much of northern Irian Jaya as the species occurs in the Arfak Mountains (pers. obs., 1992). Besides the Maprik area, I have also located P. paradisiana at Ambunti on the middle Sepik River in the Waskuk Hills behind the town. The species occurs in advanced secondary forest and primary forest in lowland and hill situations; 100–500 m. The specific epithet, paradisiana, was chosen to emphasize the importance of this species as a larval foodplant of the swordtailed Paradise Birdwing Butterfly, Ornithoptera paradisea Staudinger, 1893. The leaves and leaf habit of P. paradisiana are distinct from all other known New Guinea species of Pararistolochia. When living, they are very glossy on both surfaces, but more so on the upper side. The leaf auricles may be entirely absent, but are usually present and are characteristically minute. They exhibit varying degrees of development, and in the most extreme form are sharply inwardly directed so that they overlap just above the petiole, giving the leaf the appearance of being merely basally tapered (Fig. 30). The leaves vary only slightly in shape depending on their overall width, but they are always elongate-ovate. They Figures 22–26. Pararistolochia flowers. 22. P. biakensis sp. nov. based on holotype, AET & INJAN BO 1330-2332, Saroerai bij Seroei, Jappen-Biak [Island], Irian Jaya, Indonesia. 23. P. peninsulensis sp. nov. after Jones & Gray (1988: photo pl. on p. 152) Iron Range rainforest, Queensland, Australia. 24. P. linearifolia sp. nov., including bud from pressed specimen (right), after Jones & Gray (1988: photo pl. on p. 151) Iron Range open forest, Queensland, Australia. 25. P. praevenosa comb. nov., from photo of living material, collection A. Hiller, Mt. Glorious, Queensland, Australia. 26. P. paradisiana, sp. nov. characteristic ‘hand’ of leaves, from photo of living material, near Ambunti, East Sepik Province, Papua New Guinea.

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have the characteristic habit of forming ‘hands’ of 6–10 leaves in alternate rows on terminal stems that sprawl throughout the supporting canopy (Fig. 26). The herbarium specimens (e.g. LAE) of Bewani P. paradisiana are of fertile material, but as this is pressed it shows only limited characters. However, a living flower was illustrated with two good colour photographic plates by D’Abrera (1975), and D. Neville has photographed a flower of this species from the Arfak Mountains (Irian Jaya) (seen 1992). The flower is rather similar to those of P. momandul (Fig. 18) and P. pithecurus (Fig. 17) except that the perianth lobes are not apically bonded by a filament as in those species. In the Arfaks, the perianth lobes are slightly more elongate than topotypical specimens. Pararistolochia australopithecurus Parsons, sp. nov. (Figs 20, 45, 46) Pararistolochiae pithecuro Ridl. (Fig. 17) valde affinis sed foliis saepe magis auriculiformibus, flore majore lobulis perianthii insigniter latioribus magis deltoideis filis terminalibus longioribus magis spiralibus. Twining vine with small root tubers. STEMS to 8–10 (–30) m tall; c. 1 cm in diameter. LEAVES with petiole to 4.5 cm long, pubescent; lamina coriaceous, heart-shaped, ovate to oblong-lanceolate, to 40 cm long by 14 cm wide; base cordate, auricles prominent and well rounded, sinus to 3.5 cm deep; apex acuminate; glabrous above, pubescent beneath; veins elevated and prominent beneath, less so above; basal veins 1 pair, ascending upwards to c. 1/2–2/3 of the blade, basally branched; lateral veins 4 or 5 pairs; veinlets reticulate, prominent beneath, distinct above. INFLORESCENCES cauligerous, sometimes almost at ground level, rarely in axils of terminal leaves, solitary or usually 2 or 3, paniculiform, rhachis short (to 2 cm long), bracts minute and pubescent. Pedicel 18 mm, ovary 11 mm. FLOWER 5–6 cm overall; perianth 3-lobed, tube S-curved, c. 30 mm long by 9 mm wide, enlarging throughout, but slightly constricted below throat, broadly flaring to up to 1.8 cm in diameter; perianth lobes subdeltoid, apically bluntly pointed, 16 mm long, lobes forming 3 apertures and apically tied by maroon filament, to 30–40 mm long by 1.5 mm wide, expanding to c. 3 mm wide over spirally twisted apical half; utricle obovoid, 17 mm long by 11 mm wide. Utricle and perianth externally white, strongly brownish-purple veined; throat bright sulphur-yellow, pubescent with white setae; margin of each lobe, outwardly rolled, c. 1.5 mm broad, dark purple-black, pubescent with purple setae to 1.5 mm long. FRUIT a pendulous capsule to 8 cm long by 3.5 cm wide, elongate-ovoid, longitudinally prominently 6-ribbed, hard and green when mature, bright orange and soft when fully ripened. Holotype. Australia, Queensland, State Forest Reserve 310 (17° 13'S, 145° 42'E), rainforest, 780 m, 8. xi. 1977, B. Gray 784 (QRS 017289). Other material examined. AUSTRALIA, Queensland: Beatrice River, 700 m, R. Collins (QRS 024799); State Forest Reserve 755, Boonjee L.A., 720 m, B. Gray 331 (QRS Figures 27–31. Aristolochia leaves (numbers inside figures are leaf length). 27. A. nauseifolia sp.nov. holotype, B. Gray QRS 091512, 3 km from Kubin Village, Moa Island [middle Torres Strait] Queensland, Australia. Smaller leaf is basal form (left). 28. A. goliathiana sp. nov., eastern rounded-leaved form, D.G. Frodin UPNG 4099, Obulaku Village, Trobriand Is., Milne Bay Province, Papua New Guinea. 29. A. goliathiana sp. nov. isotype (A), western more elongate-leaved form, Streimann & Kairo NGF 45343, Kulolo Creek, Wau Subdistrict, Morobe Province, Papua New Guinea. Figs 30, 31. Pararistolochia leaves (numbers inside figures are leaf length). 30. P. paradisiana sp. nov. (note minute overlapping auricles), near Ambunti, East Sepik Province, Papua New Guinea. 31. P. paradisiana sp. nov. isotype (A), J. Wiakabu et al. NGF 50018 LAE, Creek 6 km N. of Bewani, c. 40 km SSW of Vanimo, Bewani Sub Prov., West Sepik Province, Papua New Guinea.

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002409); E/P 29 State Forest Reserve 650, Mt. Fisher, 1200 m, K. Sanderson 882 (QRS 024804). Distribution and habitat. Endemic to the region between the Mulgrave and South Johnstone Rivers, Queensland, Australia (Jones & Gray, 1988); 0–1200 m. Jones & Gray further stated that P. australopithecurus is an understorey vine in rainforest from sea level to over 1000 m altitude. P. australopithecurus occurs on Mt. Bartle Frere and in its hinterlands, near Babinda, South of Cairns, north Queensland, as well as on the Evelyn Tableland, about 30 km to the south-west. The holotype was collected in the State Forest Reserve 310 from the region of Little Mulgrave on the CairnsAtherton, Gilles Highway (the northern region of the Bellenden Ker Range). G. Sankowsky (pers. comm., 1991) noted that P. australopithecurus occurs all the way up Mt. Bartle Frere (c. 300–1200 m) in moist montane primary forest, and that the species also occurs at localities on the nearby Russell River and Woopen Creek. The specific epithet, australopithecurus, was chosen to emphasize the importance of this species as the nearest relative of P. pithecurus (Fig. 17) which is endemic to the Central Cordillera of New Guinea. Elliot & Jones (1982) and Jones & Gray (1988) both provided a basic description of P. australopithecurus (as Aristolochia spec. indet. from Mt. Bartle Frere). Both works included a good colour plate of the living flowers and foliage from the same original photograph. Pararistolochia australopithecurus differs from New Guinea P. pithecurus in only a few key features, otherwise the two species share a close (likely sibling) relationship. The leaves of P. australopithecurus are frequently more heart-shaped than those of P. pithecurus, and its flowers (Fig. 20) are larger with notably broader, more deltoid perianth lobes, and with a longer filament which is more strongly apically spirally twisted than in P. pithecurus. Pararistolochia sparusifolia Parsons sp. nov (Figs 19, 44) Pararistolochia momandul K. Sch. (Fig. 18) arcte affinis sed foliis typice sagittatis sinibus latis auriculis latis prominentibus, flore lobulis perianthii longioribus filis terminalibus brevioribus colore omnino magis viridi. Similar in characters to P. australopithecurus, but differing in the following features: Slender glabrous, wiry climber, or sometimes ground sprawling creeper, seldom to more than 8–10 m tall. Root system with only a few small, thickened, yam-like tubers. LEAVES with petiole to 4.0 cm long, pubescent; lamina glossy, lanceolate, narrowly spearhead-shaped, to 18 cm long by 4 cm wide. INFLORESCENCES in axils of terminal leaves, often on new growth, solitary or paired, sessile or on bracteate rhachis 7 cm long, bracts 2 mm long, pubescent. Pedicel short, c. 8 mm long. FLOWERS smaller, usually more abundant, 35–43 mm overall. PERIANTH tube c. 22 mm long by 6 mm, enlarging throughout, flaring to 12 mm in diameter at throat; lobes almost hemispherical, 10 mm long; filament to c. 15 mm long, apical portion (c. 5 mm long) pink, often with the 3 sections splayed in different directions; utricle 11 mm long by 6 mm wide. Utricle and perianth externally white, tinged with pale green, main veins of tube weakly highlighted by maroon; throat bright sulphuryellow; margin of each lobe barely thickened. Ovary c. 8 mm long. FRUIT to only 5 cm long by 2.5 cm wide, elongate-ovoid, yellow and soft when ripe. Figures 32–35. Pararistolochia leaves. 32. P. biakensis sp. nov. holotype (broader-leaved form), AET & INJAN BO 1330-2332, Saroerai bij Seroei, Jappen-Biak [Island], Irian Jaya, Indonesia. 33. P. biakensis sp. nov. (more lanceolate-leaved form), as above. 34. P. meridionaliana subsp. popondettensis M. Parsons subsp. nov. 015 (K), Jonita area, near Popondetta, Northern Province, Papua New Guinea. 35. P. alexandriana holotype, M. Parsons LAE 69994, Ahora Village, north of Popondetta town, Northern Province, Papua New Guinea.

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Holotype. Australia, Queensland, State Forest Reserve 143, Bushy Logging Area [Mt. Lewis], rainforest, 800 m, 2. xii. 1982, B. Gray 2876 (QRS 072007). Other material examined. AUSTRALIA, Queensland: Mt. Lewis, G. Sankowsky (QRS 071824); Mt. Lewis, 1000 m, B. Gray 131 (QRS 024800); Exp. E/P 18, North Mary Logging Area R 143, Mt. Lewis, 1000 m, K. Sanderson 406 (QRS 024801); State Forest Reserve 143, North Mary Logging Area, 1100 m, B. Gray 1069 (QRS 048959); Mt. Lewis, North Mary Logging Area, Parish of Riflemead, 1200 m, R. Collins 20162 (QRS 087165); Summit of Mt. Lewis, 4020 ft. (QRS 024797); Mt. Lewis, 900 m, G. Sankowsky (QRS 068509). Distribution and habitat. Endemic to Mt. Lewis in the Daintree National Park near Mossman, Queensland, Australia; 300–1320 m (summit). Pararistolochia sparusifolia occurs in the understorey of moist primary rainforest. The specific epithet, sparusifolia, was chosen to emphasize the normally spearheadshaped leaves of this species. The nearest relative of P. sparusifolia is P. momandul K. Sch. (Fig. 18) which is endemic to the lowland hill forests of Papua New Guinea. Elliot & Jones (1982) and Jones & Gray (1988) both provided a basic description of P. sparusifolia (as Aristolochia spec. indet. from Mt. Lewis). Both works included a good colour plate of the living flowers and foliage from the same original photograph. Pararistolochia sparusifolia differs from P. momandul in its leaves which are characteristically spearhead-shaped, with broad sinus and broad prominent auricles, whereas the leaves of P. momandul are broader and more ovate. The flower of P. sparusifolia differs from P. momandul mainly in its shorter apical filament, longer perianth lobes, and its more overall greenish colour, as opposed to pinkish-mauve in P. momandul. Pararistolochia sepikensis Parsons sp. nov. (Figs 21, 42) Pararistolochiae momandul K. Sch. (Fig. 18) valde affinis sed foliis magis apiculatis, flore lobulis perianthii ad basem typice magis lateraliter evolutis. Similar in characters to P. australopithecurus, but differing in the following features: Slender glabrous, wiry climber. LEAVES with petiole 1.5–2.5 cm long; lamina to 22 cm long by 6.5 cm wide, apiculate, broadly spearhead-shaped, more narrowly lanceolate when young (17 cm long by 4.5–5.0 cm wide); auricles short, fairly distinctively subtriangular, sinus to 6 mm deep. FLOWER greenish-purple; utricle c. 11 mm long; perianth tube c. 15 mm long by c. 5 mm wide, broadly flaring at throat; lobes elongate, c. 8 mm along ventral margin; filament fine and connate over its entire length of c. 16 mm. FRUIT unknown. Holotype. Papua New Guinea, West Sepik Province, Mt. Yungat, northern slopes Bewani Mts., Bewani Subprovince, strangler in lower montane forest, 700 m, 20.ix.1982, K. Karenga sheet No. 56530 (A); iso.: LAE (apparently misplaced or missing: M. Kuduk, pers. comm., 1995) Distribution and habitat. Apparently endemic to the West Sepik Province of mainland Papua New Guinea, (but probably occurs elsewhere in north-western Papua New Guinea and across the border into north-eastern Irian Jaya). According to the Figures 36–38. Pararistolochia leaves. 36. P. manokwariensis sp. nov. holotype (auricles barely meet), M. Parsons (BO), Wamar´e to Mokwam foot track, c. 500 m Irian Jaya, Indonesia. 37. P. manokwariensis sp. nov. holotype (auricles overlapping), as above. 38. P. tithonusiana sp. nov. holotype, M. Parsons (BO), Wamar´e to Mokwam foot track, c. 1500 m, Irian Jaya, Indonesia.

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holotype label data, P. sepikensis occurs in hill forest (probably advanced secondary forest and primary forest) at about 700 m, but the altitudinal range of the species will likely be found to be much broader once further collections are made. The specific epithet, sepikensis, was chosen from the type locality province(s) in Papua New Guinea (of which there are two halves: the East and West Sepik, named for the large Sepik River which flows through them). The description of this species is based on herbarium specimens only (A, LAE). Its flower is very distinct, being similar to that of P. momandul (Fig. 18), but is notable for the much greater lateral development of its perianth lobe bases (although not as prominently as in the following species, P. biakensis Fig. 22). The flower of P. sepikensis is described on the specimen as “perianth tube greenish purple outer surface, yellow at the base of opening [each of the three apertures] and purple towards the [lobe] tips around inner surface.” Based on knowledge of other species, Figure 21 approximately depicts the flower as it would appear in life. Pararistolochia biakensis Parsons, sp. nov. (Figs 22, 32, 33) Pararistolochiae schlechteri Laut. (Fig. 13) arcte affinis sed flore lobulis perianthii ad basem lateraliter perevolutis, filis terminaliter connatis. Similar in characters to P. australopithecurus, but differing in the following facies: Slender glabrous, wiry climber. LEAVES with petiole 1.5–2 cm long; lamina to 15 cm long by 5 cm wide, spearhead-shaped, blade slightly wasted above auricles; auricles prominent, slender, well-rounded, sinus to 13 mm deep. FLOWER green; utricle c. 11 mm long; perianth tube c. 15 mm long by c. 6 mm wide, very broadly flaring at throat; lobes elongate, c. 17 mm along ventral margin; filament c. 32 mm long, sections at extreme apex slightly splayed. FRUIT unknown. Holotype. Indonesia, Nieuw Guinea, Jappen-Biak [Island], Saroerai bij Seroei, AET & INJAN 173 (Exp. Ir L.J. VAN DIJK), 27.vii.1939 (BO 1330–2332) (flower and foliage). Cotype BO 1392-43 (foliage). Distribution and habitat. Endemic to Biak Island, north-western Irian Jaya, Indonesia. The habitat and elevation for the holotype are not stated on the label data. The specific epithet, biakensis, was chosen from Biak, the name of the type locality island. The description of this species is based on the only known specimens comprising the holotype and cotype sheets in the Herbarium Bogoriense. It is very distinct in its flower which is unique in the extreme lateral development of its elongate perianth lobe bases, yet still retaining the well developed apical filament. In general, the flower resembles that of P. schlechteri (Fig. 13), but the lobe bases are even more laterally extended and, whereas the true tendrilled perianth lobes are separate Figures 39–47. Pararistolochia leaves. 39. P. peninsulensis sp. nov. Webb & Tracey CANB 288295, Mt. Tozer, Iron Range Area, Cape York Peninsula, Queensland, Australia. 40. P. peninsulensis sp. nov. B. Gray 4239, Iron Range, rainforest, 40 m, Queensland, Australia, QRS 080850. 41. P. peninsulensis sp. nov. (CANB 288295) (as 39 above). 42. P. sepikensis sp. nov. K. Karenga NGF 56530 A, Mt. Yungat, northern slopes Bewani Mts., Bewani Subprovince, West Sepik Province, Papua New Guinea. 43. P. kepara sp. nov. holotype, M. Parsons LAE 69995, near Kamondo Village, near Kokoda, Northern Province, Papua New Guinea. 44. P. sparusifolia sp. nov. holotype, B. Gray QRS 072007, State Forest Reserve 143, Bushy Logging Area [Mt. Lewis], rainforest, Queensland, Australia. 45. P. australopithecurus sp. nov. (auricles reduced) R. Collins QRS 024799, Beatrice River, Queensland, Australia. 46. P. australopithecurus sp. nov. (auricles prominent) K. Sanderson QRS 024804, Mt. Fisher, Queensland, Australia. 47. P. linearifolia sp. nov. B. Hyland QRS 067813, National Park Reserve 8, Parish of Weymouth, Queensland, Australia. Smaller leaf (right) is a basal form from holotype B. Gray QRS 091513, Old Wolfram diggings, 3 km from Kubin Village, Moa Island, Queensland, Australia.

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in P. schlechteri, they remain as a connate filament in P. biakensis as is characteristic of the P. momandul group proper. Based on knowledge of the floral morphology of other species, Figure 22 approximately depicts the P. biakensis flower as it would appear in life. On the type sheet it is simply described as “groen” (green), which may also be a unique character depending on the extent and hue of this. Based on knowledge of all other known Pararistolochia species of the region, it is likely that the external venation is at least slightly highlighted with purple-maroon. Hou (1983b: 241) included P. biakensis in P. momandul (as “Aristolochia momandul”) which represented a mixture of various distinct taxa. In assessing the unusual flower of biakensis, he pointed out that Ridley (1914) described his taxon Aristolochia pithecurus (i.e. P. pithecurus) (Fig. 17) as having ‘four’ lobes, one of which was the filament typical of P. momandul, P. pithecurus and related species. Ridley did not, however, suggest that the filiform process arose from between any of the other lobes, or that it was one of them, merely (because of the vagueness of his description) leaving its exact position and origin open to speculation. Apparently because of this, Hou (1984: 97) erroneously described A. momandul ( = P. momandul) (Fig. 18) as occasionally possessing a fourth “filiform lobe” (in addition to its usual filament). The three above mentioned species are all part of a distinct species group (Parsons, in prep.), being characterized by their possession of a filament (Fig. 1). Pararistolochia peninsulensis Parsons, sp. nov. (Figs 23, 39–41) Pararistolochiae praevenosae F. Muell. (Fig. 25) affinis sed foliis majoribus magis variabilibus auriculis prominentibus, flore fauce atromarroninorubro lobulis brevibus, lobulo primo libero, lobulo secundo et lobulo tertio connato (lobulo latiore formantibus). Similar in characters to P. australopithecurus, but differing in the following: Slender twisting vine to 10–15 m tall arising from yam-like rootstock. STEMS 1–1.5 cm in diameter, young stems wiry, pubescent. LEAVES with petiole 1–2.5 cm long, pubescent; lamina glabrous, broadly ovate to heart-shaped, to 20 cm long by 6–8 cm wide; auricles broad, usually fairly prominent, sinus to 15 mm deep. FLOWER 35–43 mm long overall; wholly pubescent. PERIANTH tube strongly S-curved, c. 15 mm long, broadly flaring to c. 14 mm in diameter, c. 26 mm from apex to apex across lobes; lobes short, c. 6 mm long, indistinct, asymmetrical, one broader than remaining pair which together resemble an opposing single larger lobe directed towards utricle as the margin between each bears no sinus, apices of each lobe slightly hooked. Utricle and perianth tube externally white, heavily maroon blotched, longitudinal and reticulate veins prominently dark maroon highlighted; throat deep maroon, heavily pubescent with dark maroon setae. FRUIT slightly longer and narrower, to 10 cm long by 3 cm wide, orange and succulent when ripe. SEEDS horseshoe-shaped. Holotype. Australia, Queensland, Iron Range, rainforest, 40 m, 20.xi.1985, B. Gray 4239 (QRS 080850). Other material examined. AUSTRALIA, Queensland: Mt. Tozer, Iron Range Area, Cape York Peninsula, 400 m, Webb & Tracey 8708 (CANB 288295); Northern lower slopes of Mt. Tozer, 130 m, B. Hyland 6185 (QRS 024798). Distribution and habitat. Endemic to the Iron Range on the east coast, very near the apex of the Cape York Peninsula, Queensland, Australia; 0–400 m. Recorded in all

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types of lowland rainforest, both coastal and inland (Jones & Gray, 1988). Of the three herbarium sheets of P. peninsulensis, the Mt. Tozer specimen (CANB 288295) was collected at about 400 m elevation in “Rainforest with affinity to simple evergreen types. Gully site on shallow soils derived from granite”, whilst a specimen from the lower slopes of Mt. Tozer at 130 m elevation (QRS 024798) was collected in “Poor low heathy forest”. The specific epithet, peninsulensis, was chosen to emphasize that the species is endemic to northernmost Cape York Peninsula. The closest relative of P. peninsulensis is P. praevenosa (Fig. 25), which is endemic to the hill forests of south-easternmost Queensland and north-easternmost New South Wales, Australia. Jones & Gray (1988) gave a basic description of P. peninsulensis (as Aristolochia spec. indet. from Iron Range rainforest), and provided a good colour plate of the live flowers and foliage. Leaf variation (Figs 39–41) is notable in P. peninsulensis as well as in individual specimens. For example, CANB 288295 which has both broadly (Fig. 39) and fairly narrowly (Fig. 41) hastate leaf forms on the same stem. Pararistolochia peninsulensis differs from P. praevenosa in its larger, more variable leaves, which have prominent auricles, whereas those of P. praevenosa are smaller and ovate, lacking distinct auricles. The flowers of the two species are similar, but in P. peninsulensis (Fig. 23) the throat is deep maroon red, and the perianth lobes are short, with only one distinct, the other two merging into a broader lobe. In P. praevenosa (Fig. 25) the throat is yellow and waxy looking, the three perianth lobes well defined, with thick, fleshy, dark glossypurple margins. Pararistolochia linearifolia Parsons, sp. nov. (Figs 24, 47) Pararistolochiae praevenosae F. Muell. (Fig. 25) valde affinis sed foliis lanceolatis, perianthiis florum pertubularibus lobulis perianthii vestigialibus. Herbaceous twiner with bulbous rootstock, the 1–3 ovoid tubers to 25 mm long by 11 mm wide. STEMS slender to 3 m long; c. 1–2 mm in diameter. LEAVES with petiole to c. 8 mm long, smooth or sparsely pubescent; lamina linear-lanceolate, to 17 cm long by 1.5 cm wide, basally cordate, auricles indistinct, with shallow sinus to 1–5 mm deep, glabrous above, pubescent beneath (mildly adhering to fabric) (some leaves, especially basally, may be hastate or deltoid c. 5 cm long by 2 cm wide); veins elevated and prominent beneath, less so above; basal veins 1 pair, ascending upwards to c. 1/8 of the blade, basally branched; lateral veinlets numerous, to c. 18, in fairly regular alternating pairs at right angles to very linear midvein, connected by arched lateral veinlets at blade margin; veins and reticulate veinlets prominent beneath, distinct above. INFLORESCENCES solitary, sessile, in leaf axils all along main stems. Pedicel to c. 8 mm long. FLOWER 3–4 cm long overall; perianth very tubular, trumpetlike, barely apically 3-lobed; tube prominently S-curved and parallel-sided below throat, c. 15 mm long by 5 mm wide; perianth lobes subdeltoid, vestigial c. 2.5 mm long; utricle ovoid, c. 9 mm long by 5.5 mm wide. Utricle and perianth externally creamy-white, pinkish tinged, longitudinal veins prominently highlighted with maroon, reticulate veins maroon blotched; throat bright sulphur-yellow, lobes glossy, broadly margined with dark maroon. Ovary c. 3 mm long. FRUIT a pendulous capsule, barely longitudinally 6-ribbed, to 2.5 cm long by 1 cm wide, orange and succulent when ripe. Holotype. Australia, Queensland, Old Wolfram diggings, 3 km from Kubin Village, Moa Island, open forest, 30 m, 14. ii. 1989, B. Gray 5006 (QRS 091513).

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Other material examined. AUSTRALIA, Queensland: National Park Reserve 8, Parish of Weymouth, open forest, B. Hyland 11567 (QRS 067813); Vrylia Point, grassy woodland, A. Morton (QRS 066474). Distribution and habitat. Endemic to northernmost Cape York Peninsula, and Moa Island in the Torres Strait (type), Queensland, Australia; 0–80 m. Pararistolochia linearifolia grows among grasses and other herbage in open forests at Iron Range, the plants withstanding destruction by bush fires almost annually (Jones & Gray, 1988). Jones & Gray (1988) gave a basic description of P. linearifolia (as Aristolochia spec. indet. from Iron Range open forest) and provided a good colour plate of the flowers and foliage. G. Sankowsky (pers. comm., 1991) pointed out that he collected living material of P. linearifolia at the Iron Range about 60 km north of Moreton (Telegraph Station) (i.e. on the Cape York Road at almost exactly 12°S). He stated that the species occurs at the mouth of the Pascoe River (Weymouth Bay, about 20 km north of Mt. Tozer), and is very common in the Glennie Tableland a further 20 km north of the Pascoe River. ‘Vrylia Point’ (namely Vrilya Point) (QRS 066474) is on the west coast of the Peninsula, opposite Jardine River National Park. With its uniquely narrowly parallel-sided, elongate mature leaves (for which the specific epithet, linearifolia, was chosen), and very tubular, trumpet-like flowers, this species has clearly specialized to be dry-adapted. From its floral morphology (Fig. 24), P. linearifolia is more closely related to P. praevenosa (Fig. 25) than to P. peninsulensis (Fig. 23). This is interesting, considering that P. linearifolia and P. praevenosa have such widely disjunct distributions at either end of Queensland, being separated by a distance of about 1600 km. Nevertheless, there is one tentative record of P. praevenosa from the Atherton Tableland near Cairns (K 1930), so P. praevenosa may be more widely distributed than the present rather sparse records show. Besides the obvious differences in their leaves, the P. linearifolia flower has vestigial perianth lobes, whereas in P. praevenosa the three perianth lobes are well-defined.

ACKNOWLEDGEMENTS

Many people have graciously given me their time and assistance during my years of researching Aristolochiaceae. Thanks to their kindness and encouragement, the difficult task of preparing this manuscript was made somewhat easier. Friends and colleagues during my four years of early fieldwork in Papua New Guinea include Ted Henty, Alister Hay, Neville Howcroft, Paul Katik, Robert Johns, David Frodin, Heiner Streimann and Aubiter Kairo, all of whom helped in one way or another. Follow-up work was greatly facilitated by the correspondence and mail loans kindly provided by Max Kuduk and Wayne Takeuchi at the LAE Herbarium. In Australia, Gary Sankowsky, Anthony and Kate Hiller, Donald Sands, Mark Daish, John Connors and David Jones all assisted me to obtain specimens and photographs of living or herbarium material of Australian species. In the Netherlands, Ding Hou, Rijksherbarium, Leiden, patiently discussed with me, and corresponded about, his revisionary research during the 1980s. This and his assistance with photographic material and certain reprints, including his own publications, were invaluable aids to my own work. In the USA, Robert Gustafson, Botany Section, Los Angeles County Museum of Natural History (LACM), kindly assisted with practical advice, including

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improvements to the manuscript, as did Gary Wallace, Rancho Santa Anna Botanic Garden, Claremont, California. At the LACM Entomology Section Brian Brown and Brian Harris provided invaluable support which enabled the completion of this research. Marshall Johnston, Botany Department, University of Texas, very kindly provided the Latin translations, for which I am extremely grateful. Emily Wood of the Arnold Arboretum Herbarium, Harvard University, Cambridge, Massachusetts, was most helpful in efficiently facilitating loans of material from that institution. In Poland, Wanda Stojanowska of the Museum of Natural History, Wroclaw University kindly assisted with type material in the WRSL collection. I also thank the anonymous reviewers who rightly encouraged me to improve the manuscript and provided useful suggestions for doing so, as well as Robert Mill (Royal Botanic Garden, Edinburgh, UK) for his most helpful editing of the final draft. Lastly, but by no means least, I wish to extend my very special thanks to my wife Maria who has given me her support and constant encouragement during all of my personally funded research, trying though this has often been.

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Pfeifer HW. 1966. Revision of the North and Central American hexandrous species of Aristolochia (Aristolochiaceae). Annals of the Missouri Botanic Gardens 53: 115–196, 59 figs. Pfeifer HW. 1970. A taxonomic revision of the pentandrous species of Aristolochia. University of Connecticut. Poncy O. 1978. Le genre Pararistolochia, Aristolochiaceae D’Afrique tropicale. Adansonia (Ser. 2). 17: 465–494. Rali T, Griffin F. 1991. A phytochemical investigation of Aristolochia sp. the food plant of larval Ornithoptera priamus (Lepidoptera: Papillionidae [sic!]). Science in New Guinea 17: 143–146. Schmidt OC. 1923. Neue Aristolochiaceen Papuasiens. Engler Botanische Jahrb¨ucher vol. 58: 488–491. Schmidt OC. 1932. Zur Kenntnis der Gattung Aristolochia in Australien. Feddes Repertorium 30: 72–75. Straatman R, Inoue S. 1984. Identified hostplants of Ornithoptera species. Tyˆo to Ga 34: 124–126. Veldkamp JF, Vink W, Frodin DG. 1988. XI. Ledermann’s and some other German localities in Papua New Guinea. Flora Malesiana Bulletin 10: 32–38. Verdcourt B. 1986. Aristolochiaceae. In: Flora of Tropical East Africa. Aristolochiaceae. Rotterdam and Boston: Balkema AA Weintraub JD. 1995. Host plant association patterns and phylogeny in the tribe Troidini (Lepidoptera: Papilionidae). In: Scriber JM, Tsubaki Y, Lederhouse RC, eds. Swallowtail Butterflies: Their Ecology and Evolutionary Biology. Gainesville FL: Scientific Publishers.307–316