Bull. Kitakyushu Mus. Nat. Hist., 15: 43-118. March 28, 1996

Gondwanan Evolution of the Troidine Swallowtails (Lepidop- tera: Papilionidae): Cladistic Reappraisals Using Mainly Immature Stage Characters, with Focus on the Boisduval

Michael J. Parsons

Entomology Section, Natural History Museum of Los Angeles County 900 Exposition Blvd., LosAngeles, California 90007, U.S.A.*' (Received December 13, 1995)

Abstract In order to reappraise the interrelationships of genera in the tribe , and to test the resultant theory of troidine evolution against biogeographical data a cladistic analysis of troidine genera was performed. Data were obtained mainly from immature stages, providing characters that appeared to be more reliable than many "traditional" adult characters. A single cladogram hypothesising phylogenetic relation ships of the troidine genera was generated. This differs markedly from cladograms obtained in previous studies that used only adult characters. However, the cladogram appears to fit well biogeographical data for the Troidini in terms of vicariance biogcography, especially as this relates to the general hypotheses of Gondwanaland fragmentation and continental drift events advanced by recent geological studies. The Ornithoptera is shown to be distinct from . Based on input data drawn equally from immature stages and adult characters, a single cladogram hypothesising the likely phylogeny of Ornithoptera was generated. With minor weighting of a single important adult character (male forcwing uppersidc sex-brand), a further two clado grams were generated, one of which is similar to hypotheses proposed by previous workers. Based on these findings, and on ecological data, notably larval foodplant relations with Aristolochiaceac, as well as present-day biogeographical data, a new theory of the origin and evolution of Ornithoptera is presented which fits well Gondwanan vicariance events ascertained by geological studies: essentially that Ornithoptera evolved on northward drifting , allopatrically from Troides on the Indian plate, and therefore that Ornithoptera did not reach the Australian subrcgion via TroiV/w-likc ancestors in Southeast Asia as has been previously postulated.

Key Words androconia, , Aristolochiaccae, Atrophaneura, Australia, , , biogeography, Chilasa, classification, continental drift, Cressida, , foodplant, genitalia, Gondwanaland, Indonesia, Irian Jaya, larvae, , , , Ornithoptera, ova, , Papilioninac, Papilionini, Pararistolochia, , , Parnassiini, , phylogenetic, pupae, sphragis, swallowtail, Thottea, , Troides, Zcrynthiini.

*' Present address: Department of Entomology and Ncmatology, University of Florida, Bldg. 970, Hull Road, Gainesville FL 32611-0620, U. S. A. 44 Michael J. Parsons

Contents Introduction 44 "Total Evidence" and Improving Analyses 46 The Tribe Troidini Ford, 1944 48 Method 49 Recognition ofTroidini Terminal Taxa 49 Methodology, Rationale and Selection ofOutgroups forTroidini Analysis 50 Selection of Characters for Troidini Analysis 53 Enumeration and Discussion ofTroidini Characters 54 The genus Ornithoptera Boisduval, 1832 68 Generic Distinctness of Ornithoptera and Troides 71 Outgroup Selection for Analysis of Ornithoptera 75 Enumeration of Ornithoptera Characters 77 Results 79 Enumeration ofTroidini Cladcs 79 Enumeration of Ornithoptera Cladcs 91 Discussion 94 Troidine Evolution and Gondwanan Palacobiogcography 94 Ornithoptera Origin and Biogcography: A New Hypothesis 97 Evidence from Foodplant Relations 101 Evolution: Also Part of the Gondwanan Picture 103 Older, More Ancient Origins 105 Concluding Remarks 107

INTRODUCTION

Like that of Munroe (1961), the early work of Igarashi (e.g. 1963), culminating in his more recent publications (e.g. Igarashi 1979 and 1984), can be regarded as pioneering in classifying the family Papilionidae. Igarashi's research was particu larly important because he emphasised the use of immature (or early) stages (eggs, larvae and pupae) to study the classification of the family. His work is, therefore, as much a foundation to the understanding of papilionid systematics as those of Munroe and Ehrlich (1960) and Munroe (1961) who concentrated on adult characters. Reliable phylogenetic reconstructions are of great importance in biogeographical studies (e.g. Hennig, 1966; De Jong, 1979). Only characters that are truly effective and/or useful in such systematic analyses will provide 'reasonable' (i.e. generally accepted) results. These must be selected and defined, which is a highly subjective process, limited by constraints on a researcher's knowledge of the particular group under scrutiny. As it is impossible to know which characters provide the 'best' (most unequivocal or truly reliable) results, this selection process represents a fundamental problem in any method of systematic study or classification (cladistic or otherwise), and renders all resultant hypotheses open to legitimate speculation. Nevertheless, it follows that the broader a researcher's knowledge is (encompassing Reappraisal of Troidini 45

other related groups, i.e. outgroups, etc.), the more likely it is that they will be able to make effective choices in character selection and thereby, in theory, improve the likelihood of obtaining results that better ascertain the actual course of events in the evolution of any study group. Only those facies that most assuredly reflect unidirectional evolutionary changes (rather than reversals, or those that have evolved independently several times) will provide the strongest corroboration of the 'correct' structures of evolutionary trees. Preferably, therefore, such characters should be relatively stable, not subject to rapid change, and so will provide greater reliability in their use in ascertaining sequences of evolution. Problems of character selection are perhaps more evident than ever in modern phylogenetic studies. For example, recent analyses of the as an entire group (including two cladistic studies) have resulted in at least three markedly different scenarios for their evolution at the family level: Kristensen (1976); Scoble (1986); Shields (1989). Even for the most popular and well-studied family, Papilionidae, Hauser (1993), while critically re-reviewing their classification, found that, contrary to previous findings, the subfamily Parnassiinae does not comprise a monophyletic group because the characters used were insufficiently well- known and/or rather poorly analysed. In their cladistic study of American swallow tails, Tyler etal. (1994) concluded that "with a judicious choice of these [characters] and the right taxa, you can make the papilionid phytogeny look almost any way you please." Likewise, even in studies of bird and human evolution, recent new fossil finds (providing still further characters for analysis) have only served to continue to challenge conventional theories of the historical lineages of these groups. It need hardly be emphasised that the latter are groups with at least a good base of adequate fossil evidence, whereas in the (notably butterflies) this is extremely scant at best. The -feeding papilionids in New Guinea were found to be particularly good candidates for cladistic study because reasonable data on their immature stages and ecology is available, and outgroups for these groups can be selected with some confidence. In addition, in carrying out a study of the foodplant relations of New Guinea Troidini, it was found necessary to address some apparent taxonomic problems in the classification of the 'birdwing' genera, most notably the validity of Ornithoptera as a genus distinct from Troides (Parsons, 1991). For example, in recent cladistic analyses, Hancock (1983) and Miller (1986, 1987b) merely saw fit to lump the two taxa, thereby obscuring the great importance of their relationships with other troidine genera. As Hauser (1993) pointed out: where characters are insufficiently well known and/or are poorly analysed then the results can likewise be expected to be poor or incorrect. For example, he found that the forewing cubital crossvein does not represent an autapomorphy of the , and that the Parnassiinae are paraphylctic as the subfamily presently stands because of poor interpretation ofadult characters, especially confusion overvalid synapomor- 46 Michael J. Parsons phic characters to truly define Parnassiinae.

"Total Evidence" and Improving Analyses Authors such as Kluge (1989) and Tyler etal. (1994) have convincingly argued that the only really effective way to reconstruct the phylogenies of organisms is to include all possible characters, using the so-called "principle of total evidence" (Kluge, 1989). Adult characters have been the prime (often only) focus of attention for Lepidopterists in most systematic and phylogenetic studies. Characters of male butterfly genitalia, venation and wing pattern have traditionally been selected (including most of the recent cladistic analyses of butterflies) primarily because they are the easiest to study. However, Homma (1954), for example, suggested that the alimentary canal in butterflies is of taxonomic use, and some analyses have also included adult butterfly sensory and ambulatory appendages. Nevertheless, as my own studies of many different New Guinea butterfly groups have repeatedly demon strated (e.g. Parsons, 1986a, b, 1989a, b, 1991), genital and wing pattern facies often change rapidly in response to external selection pressures, and can respond markedly differently, even between closely related groups. This is hardly surprising consider ing the high mobility of the as an adult, leading to its interaction with a complex array of environmental factors, all of which potentially influence its external facies. This frequently results in adult characters which, although they appear 'good,' are found to be taxonomically far less reliable when studied in the context of a large number of groups. Similar views were expressed by Tyler et al. (1994). Courtship and mimicry are factors that obviously play an important role in modifying the external facies of adult butterflies. For example, because of mimicry, the venation and wing patterns of some New Guinea butterflies have become highly modified, and often convergent, demonstrating how rapidly changeable and 'plastic' such adult characters are, even intraspecifically. The modification of overall wing shape can affect the positions, lengths and connections between the veins of the adult butterfly wing. As Vane-Wright (1976) noted, Mynes anemone Vane-Wright, 1976, in New Guinea is a particularly good example of this, the adults appearing very atypical of their genus through their mimicry of members of the ithomiine nymphalid genus Tellervo Kirby, 1894. In fact, Van Eecke (1915) erroneously established a new New Guinea genus, Bigaena, which he considered was closely related to the morphine nymphalid genus Hyantis Hewitson, 1862, but which was actually the mimetic female of the satyrine nymphalid Mycalesis drusillodes (Oberthur, 1894). During the course of studying New Guinea butterflies I have also found that sex- limited mimicry has been responsible for the erroneous description of the males and females of a single species as two separate species in several cases (Parsons, unpublished). Extreme sexual dimorphism is obvious in various New Guinean genera, most notably Ornithoptera (Troidini: Papilionidae) where field evidence has shown (Straatman, 1979, and pers. obs.) that the females can apparently be Reappraisal of Troidini 47

Miillerian mimetic mimics of one another. Some adult characters can, therefore, present great problems in the assessment and interpretation of the extent of changes in them. As a result, many facies of adults are more likely to introduce homoplasy ('noise') into cladistic analyses. Where higher taxa (tribes, subfamilies, etc.) are concerned, this can be particularly detrimental, although genital and wing pattern facies are obviously useful in ascertaining the relationships of taxa at the intrageneric level. In effect, different characters can each be of maximum value at an appropriate level in the heirarchy, so by employing them at the wrong level in analyses only erroneous results will be achieved. When attempting to construct phylogenetic trees it is arguably more effective to use a smaller number of reliable characters, than a larger number of characters of questionable status (e.g. Shields, 1989). In this context, butterfly immature stages (eggs, larval, pupal), and related life history data, appear to provide character sets of great systematic value. This was well stated by Brown and Freitas (1994): "In principle, juvenile stages should be useful in the understanding of relationships among taxa, especially at the genus-level and above, because juveniles, like embryonic or ontogenetic stages, conserve characters of older sister lineages, diverging less than the more specialized adults." Tyler et al. (1994) also stated the case well: "These stages — especially first instar larvae and pupae — are likely to show relationships at these levels [genera, tribes] far better than the adults, simply because many of their characters are 'conservative' and evolve slowly, being less subject to change under fluctuating environmental pressure." The same views were presented in Brown et al. (1995), and papilionid pupae (notably troidine and leptocircine) were mentioned as being particularly useful in providing reliable characters for taxonomic study. Waterhouse (1937) had earlier expressed similar sentiments, particularly with regard to the classification of Australian subregion : "Since larvae and pupae do not seem to have altered as much as the perfect during the process ofevolution, these will be found to provideexcellent characters, which then may be correlated with structural differences in the adults." He noted also that "great weight should be given to the life-histories" of the Indo- Australian Papilionidae in future revisionary studies. Thus papilionid pupae appear particularly useful for classification of the family. Although subject to selection pressures resulting in their crypsis (etc.), butterfly pupae may be viewed as relatively simple sessile 'containers,' inside which larval to adult development takes place. Consequently, overall pupal morphology (but also including particular facies) can highlight well many relationships between butterfly groups (e.g. Chapman, 1895; Bell, 1912; Mosher, 1969). Waterhouse (1937) stated he had found that hesperiid larvae and pupae provide very good characters for making generic distinctions, and Bell (1921: 792) used characters of the pupa (and eggs) of various Indian genera as a means of uniting them as several natural groups. 48 Michael J. Parsons

As is being increasingly realised, therefore, the immature stages of butterflies are often crucial in providing the necessary corroborative and/or conclusive data for better resolving cladograms based solely on adult characters. It is likely that pupal morphology, for example, could eventually provide a good basis for the long needed tribal classifications of the Hesperiidae and Pieridae. Very good recent examples of this include better resolution of the systematic and interfamilial relationships of the Nymphalidae by Harvey (1991), especially using larval sctal characters. The recently described larval and pupal characters of neotropical Anetia Hubner, 1823, have shed much new light on the determination of the systematic position of this nymphalid genus within the subfamily Danainae, and have provided additional data which challenges previous hypotheses arrived at from earlier cladistic work (Brovver et al., 1992). Also, based primarily on cladistically analysed larval characters, the systematic positions of the neotropical genera Antirrhea Hubner, 1822, and Caerois Hubner, 1819, were found by De Vries et al. (1985) not to belong in the nymphalid subfamily Satyrinae as traditionally thought, but instead had to be transferred to the . Apart from the importance of immature stage facies, ecological and behavioural 'characters' may also be of value. Although difficult to quantify and utilise in systematic analyses, such data has the potential to serve as corroborative information key to the understanding of the evolution of some butterfly groups. They are more likely to be unknown to any taxonomist unable or unwilling to venture into the field to observe them. For example, the close relationship in New Guinea and Australia between Ornithoptera and their aristolochiaceous foodplants implies that the speciation of these papilionids has been highly influenced by that of their larval foodplant vines, and the limited ranges of these foodplants have apparently had a direct bearing on the biogeography of the Ornithoptera. This must obviously, therefore, be considered when postulating their evolutionary relationships.

The Tribe Troidini Ford, 1944 Prior to the study of Tyler et al. (1994), Miller's (1986, 1987b) work was the most recent detailed cladistically-based attempt at attaining a better understanding of the classification and relationships of the Papilionidae. Unlike the study of Tyler et al., Miller's did not incorporate any detailed immature stage characters, and relied primarily on traditionally used adult characters (male and female genitalia, etc.). As noted above, these are frequently subject to change, rendering them difficult to interpret and rather unreliable ('unpredictable') as characters for use in analyses of the higher taxonomic categories. Thus Miller did not incorporate important generic level details of immature stages into his data. Moreover, in his analysis of the tribe Troidini, Papilionini was used as the outgroup, all character states for Papilionini being scored as plesiomorphic, so that the rooting of the resultant cladogram for Troidini was not in the Zerynthiini. As noted by Hauser (1993) (and Reappraisal of Troidini 49

I concur), the Parnassiinae are paraphyletic, the Aristolochia-keding zerynthiines standing out as a well defined group, presently regarded as a tribe within subfamily Parnassiinae. However, Miller (1987b: 416) found that he could not reconcile the homology of characters such as the fleshy larval tubercles of Troidini with 'Parnas siinae' as the Parnassiini (sensu stricto) entirely lack these. Nevertheless, despite the contrary belief of Miller, the Troidini larval and pupal tubercles are clearly homologous with those of Zerynthiini. The setae of the early instar larval tubercles of both groups (as well as those of Papilionini) are also homologous, despite Miller's doubts.

METHOD

Recognition of Troidini Terminal Taxa Based on studies of troidine immature stages and adults, and in consideration of the work of others, I treat the tribe Troidini as comprising 8 genera: Euryades C. & R. Felder, 1864, Cressida Swainson, 1832, Pharmacophagus Haase, 1891, Ornithoptera Boisduval, 1832, Battus Scopoli, 1777, Parides Hubner, 1819, Troides Hubner, 1819, and Atrophaneura Reakirt, 1865. All of these taxa possess a sufficient number of distinctive characters to establish them as monophyletic units. The larval and pupal morphologies of the taxon antenor (Drury, 1773) from Madagascar (the sole troidine in the Afrotropical Region) are particularly distinct and clearly demonstrate that the species warrants full generic status (pers. obs.: preserved eggs, larvae and pupae in my own archives). These stages were unavailable to Miller (1986) and Hancock (1980 and 1988). Therefore, Pharmacophagus, to which antenor belongs, is certainly a valid genus as maintained by Miller, and not a subgenus ofAtrophaneura as Hancock argued. Based on the total evidence from the immature stages of Trogonoptera Rip- pon, 1898, I consider that the taxon is, at best, a subgenus of Troides (see below). For similar reasons, Pachliopta Reakirt, 1865, is considered here a subgenus of Atrophaneura. Unlike Munroe (1961), Miller (1986, 1987b) or Hancock (1988) I find only characters supporting the unity of Atrophaneura and Pachliopta. Initially, Hancock (1980) correctly treated Pachliopta as a synonym of Atrophaneura and pointed out that Munroe's (1961) separation of Pachliopta (as a group of 13 species) was largely based on adult characters (e.g. genitalia) that should be considered specialised only at the group (not genus) level. Based on his cladistic analysis of the Papilionidae, Miller (1986, 1987b) arrived at an entirely different interpretation of Atrophaneura. He treated Pachliopta as a distinct genus and lumped two species groups of Old World Atrophaneura under the New World neotropical genus Parides. However, such an arrangement is untenable based on clear and important homologies (representing autapomorphies) of larval and pupal characters in Atrophaneura and Pachliopta, and the absence of these in other troidine taxa (e.g. Figs. 1 and 13). In his reappraisal of 50 Michael J. Parsons

Atrophaneura, Hancock (1988) treated Atrophaneura, Pachliopta and Pharmacophagus as three subgenera within Atrophaneura. He concluded that his subgenus Atrophaneura contains two species-groups: lalreillei with 15species from South-east Asia and Japan, nox with 12 South-east Asian species. Hancock stated that his subgenus Pachliopta contains two species-groups: coon with 4 South-east Asian species, polydorus with 12 Indo-Australian species. His subgenus Pharmacophagus contains two species-groups, each represented by the single species antenor and hector. Hancock's placement of the Indian region species hector (Linnaeus, 1758) together with antenor in Pharmacophagus was clearly influenced by convergence of its adult characters to resemble those of antenor. His systematic arrangement is untenable on the basis of the immature stages of hector which clearly place it within the genus Atrophaneura (sensu stricto), whereas those of Pharmacophagus are so distinct as to obviously place the genus elsewhere. In particular, as stressed by Davidson and Aitken (1890), the pupal morphology and maculation of hector is almost identical to that of its close relative A. (Pachliopta) aristolochiae. Various other taxa have, at one time or another, been named and described as valid genera or subgenera of the Troidini: Aetheoptera Rippon, 1890, Pompeoptera Rippon, 1890, and Schoenbergia Pagenstecher, 1893 (= Ornithoptera), Ripponia Haugum and Low, 1975 ( = Troides), Panosmia Wood-Mason & de Niceville, 1887, and Losaria Moore, 1902 (=Atrophaneura). All recent authors of detailed systematic studies of the Papilionidae have treated these taxa, established on adult autapomor- phies, as at most subgenera of the genera under which they have been placed. I regard them as no more than species groups, their group names having no useful taxonomic value. For example, I agree with Miller's (1987b: 425) comments on the invalidity of Ripponia. Therefore, this and the other taxa mentioned above are not treated as distinct in my data matrix (Table 1).

Methodology, Rationale and Selection of Outgroups for Troidini Analysis I use the cladistic method in this analysis primarily as a strict and repeatable means of investigating the phylogenetic interrelationships of the Troidini. As noted above, most taxa mentioned have, at one time or another, been regarded as 'good' genera. I have followed the general methodology and format used by other entomol ogists, notably Miller (1987b), Pape (1992) and Brown (1993). This uses com puter analysis running algorithms that construct phylogenetic trees based on the presence of shared derived characters. The data matrix editing facility of the cladogram editing/analysing program, MacClade 3.04, was used to create and edit the numerically defined character matrix (data set), and to generate NEXUS files also readable by PAUP. The data comprised 44 selected characters, 12 of which are multistate (9 three state; 3 four state: Table 1). The MacClade matrix editor was also used to export the data set to an input file that could be read by the MS-DOS based computer program Hennig86 Reappraisal of Troidini 51

(Farris, 1989) for analysis. The data set was run using the Hennig86 option (ie*) which includes the "Branch and Bound" procedure by which the program searches for the shortest tree length and guarantees, therefore, to find the most parsimonious cladograms (trees) (where all characters were ordered and each was equally weighted = default options). For further confirmation, the data was also run using the successive character weighting procedure ('xsteps w'; in combination with 'mh*;bb;*' and 'cc;' until weights no longer change). The procedure weights characters according to their best fit to the cladograms generated by multiplying consistency and retention indices and scaling these in the range 1-10, thereby giving higher priority to clades based on more reliable characters. For comparative purposes, further analyses of the data matrix were made using the Macintosh-based computer programs PAUP 3.1.1 (Swofford and Begle, 1993), which also permits branch and bound searching, and various consensus (most parsimonious of all parsimonious trees: e.g. Nelsen [ = strict]), searches. My polarisation and scoring of characters as to their plesiomorphic (0=less specialised/primitive) or apomorphic (1, 2, etc. = more specialised/derived) states, are based on the method of out-group comparisons (Hennig, 1966; Watrous and Wheeler, 1981), in which in-group character states (i.e. for Troidini) were determined by comparison with the character state in the outgroup. This outgroup was used to root the cladogram. Considering the problems of papilionid and systematics highlighted by Hauser (1993), care must be taken in the selection of any outgroups for the cladistic analysis of Troidini. For example, I am mindful that even within the Troidini as the tribe presently stands, important characters such as the elaborate alate external sphragis of Cressida and Euryades imply that such taxa may be better separated from Troidini into a tribe of their own: Cressidini. This was suggested by Ford (1944), and later accepted by Ehrlich (1958), but was subsequently refuted by Munroe (1961) who returned Cressida and Euryades to Troidini. Reservations about the value of the form of the sphragis can be raised because an apparently homologous external form of sphragis occurs in Latreille, 1804 (Parnassiini), whereas in all other troidines and the Zerynthiini, sphragides are more internal, being primarily simple and plug-like (e.g. Orr, 1988). This and other similar concerns about characters are discussed further below. Both Hancock (1983) and Miller (1986, 1987b) used Papilionini (undoubtedly the sister-group to Troidini) as the outgroup to polarise characters for Troidini, as they supposed it to be the more plesiomorphic taxon of the two tribes. Thus, Hancock (p. 4) and Miller (p. 416) both rejected use of any taxa of the subfamily Parnassiinae for this purpose. Hancock's reasoning was based on his assertion that the "red-tuberculate" condition of the papilionid larva is "polyphyletic" (i.e. homoplasious for the Zerynthiini and Troidini) and not, therefore, a reliable plesiomorphy. Similarly, Miller rejected the homology of the larval and pupal tubercles in Zerynthiini and Papilionini. I contend, however, that they are indeed 52 Michael J. Parsons homologous. Moreover, I accept the views of Munroe and Ehrlich (1960) and Munroe (1961) that the ancestral papilionid was an Aristolochiaceae-feeder. My reasons for this are similar to theirs. I note in summary here, however, that on the basis of their latitudinal gradients in species numbers (e.g. Slansky, 1972) and feeding specialisations (e.g. Scriber, 1973), the ancestral swallowtail was most likely a tropical Aristolochia-keding species. Important corroborative evidence for this is the fact that a significant majority (5 out of 7) of swallowtail genera in the Parnassiinae (undoubtedly plesiomorphic compared to Troidini) are all Aristolochia- feeders as larvae: Hubner, 1822, Luehdorfia Cruger, 1878; Westwood, 1851; Bhutanitis Atkinson, 1873; Ochsenheimer, 1816. Bhutanitis is selected to (mainly) represent Zerynthiini as the outgroup for comparison in this study because this tribe (presently classified in Parnassiinae) is traditionally regarded as being subordinate in rank to Papilioninae, to which Troidini clearly belong. Moreover, Bhutanitis is a very coherent taxonomic unit whose member species are few (4) and, as larvae, feed exclusively on Aristolochiaceae (Aristolochia): an important character shared by the Troidini. Bhutanitis is restricted to the Oriental subregion, and its ecology and immature stages have been well documented by Lee (1986a, b, 1986c) and Igarashi (1989), while its adults were taxonomically well examined by Saigusa and Lee (1982). The genus is a good representative of its tribe, Zerynthiini, in general, but I score overall pupal form in the data matrix (Table 1) based on Luehdorfia. This is reasonable since the Luehdorfia pupa apparently possesses a greater number of plesiomorphic characters, whereas the Bhutanitis pupa exhibits a number of apomorphic characters, clearly having become cryptically specialised as a dead-twig/ stick mimic. Accepting Hauser's (1993) warning that Parnassiinae are paraphyletic, at least Bhutanitis out of the 7 parnassiine genera appears worthy of direct comparison with Troidini due to the larger number of characters shared by Zerynthiini and Troidini, as opposed to Parnassiini and Troidini. For sake of completeness, and in order to see where a further outgroup with apomorphic characters would terminate, I have included a representative of Papi lionini. I selected Papilio Linnaeus, 1758 (sensu striclo: i.e. as defined by Linnaeus' type species, P. machaon) because the genus, apart from being well-known, is traditionally classified in the same subfamily, Papilioninae, as the Troidini, and in the tribe Papilionini, presently regarded as the sister tribe of Troidini. Papilio, representative of the Papilionini, was traditionally regarded as being more advanced than Troidini in various apomorphic characters, although, like Bhutanitis, it shares various synapomorphies with Troidini. I stress here that I do not include in Papilio species such as the Oriental clytia Linnaeus, 1758, and the New Guinea region laglaizei Depuiset, 1877 (and sibling relatives moerneri Aurivillius, 1919, and toboroi Ribbe, 1907), which arguably belong in a distinct genus, Chilasa Moore, 1881, based on their distinctive immature stage characters. Species such as laglaizei arguably Reappraisal ofTroidini 53 exhibit plesiomorphic, rather troidine-like, larval and pupal characters (pers. obs., and Eliot in Corbet and Pendlebury, 1978, apparently following Igarashi, 1976). Therefore, it would be interesting to specifically revise Chilasa and cladistically compare these data against Troidini and other papilionid groups.

Selection of Characters for Troidini Analysis Characters to effectively group genera of the tribe Troidini require careful con sideration and selection. As emphasised above, some characters contradict others, and some are difficult to polarise. I concur with Shields (1989) that it is preferable to base phylogenies on a suite of fewer characters that are (or at least appear to be) reasonably easy to comprehend. I am also particularly sympathetic to views of Tyler et al. (1994), especially in light of the cautions raised by Hauser (1993), that papilionid characters should be selected and scored with care for them to be of any real value. Like Tyler et al., I do not believe it necessary for characters to be analysed as discrete sets for adults, immature stages, behaviour (etc.), and I too am confident that certain ecological, behavioural and immature stage data can be united with adult morphological characters into a single data matrix for combined analysis. As noted above, authors such as Kluge (1989) and Tyler et al. (1994) maintain that this is the only effective way to analyse data ("principle of total evidence"). In constructing the data matrix for cladistic analysis of the Troidini plus outgroups I have attempted to conservatively use characters which I perceive (i.e. my bias as to their use) as being less prone to ambiguity, and therefore of assessed greater value, and which can thus be polarised with confidence. Hopefully, therefore, such characters are less likely to introduce homoplasy into the analysis. This rule has been especially strictly applied in the choice of adult characters. For example, based on knowledge gained from extensive studies of the adult male genitalia and the adult wing patterns of numerous groups of New Guinea butterflies, I have, because of their 'plasticity,' severely limited the inclusion of such characters in the data matrix for analysis of Troidini. This has resulted in my rejection of many of the adult characters (especially male genitalia) used by previous authors, and a more 'cautious' polarisation of those that I have adopted. For example, I have been far more conservative in my use of characters of the adult female bursa copulatrix than Tyler et al. (1994). I only observe apparently reliable broad trends in the shape and condition of the bursa, whereas the signum is apparently easily transformed or lost. As another example: complete red prothoracic ("pronotal") collars are present in most (but not all) Troides, including T. (Trogonoptera) brookiana (Wallace, 1855) and T. (T.) trogon (Vollenhoven, 1860), as well as in most Atrophaneura, and in Pharmacophagus, as well as a number of Parides, but are absent in Ornithoptera, Battus and Euryades. In Cressida the collar is red, but is indistinct and divided by black scaling dorsally as in some Parides. In Euryades there is only a trace of red in the region of the prothorax. As the value of this character is, therefore, 54 Michael J. Parsons difficult to interpret it is omitted from the data matrix used in the present analysis. Apart from selecting characters which usefully group taxa into broad categories, I have also included certain important facies by which particular terminal clades (e.g. genera and subgenera) can clearly be recognised. These are, therefore, single autapomorphies which are flagged by PAUP as "uninformative" as they obviously do not assist in the resolution of relationships between taxa. There are eight such characters (Nos. 5, 10, 13, 14, 18, 26, 30 and 34) in the matrix (Table. 1). They arc included to facilitate the definition and discussion of each clade. Other, less well defined autapomorphies were omitted from the matrix, but these are discussed below under their respective clades. Characters of Troidini that appear particularly useful for defining groups within the tribe, and which can apparently be used with most confidence, are those of the first instar larva and pupa. Theoretically these stages are subject to minimal environmental influences and this is reflected in the general phenotypic uniformity and stability of these stages. Comparisons of Troidini pupae illustrate particularly well the close relationships of some of the genera and, in conjunction with distributional data, suggest the relevant distances between others. For example, the pupal morphologies of Pachliopta and Trogonoptera show that these taxa merely represent species groups (subgenera) of Atrophaneura and Troides, respectively. Ornithoptera is clearly distinct from Troides and does not deserve to be subsumed in synonymy (Parsons, 1991: 83). Characters of the genus are, therefore, scored alongside those of taxa of equal taxonomic rank in the matrix in Table 1. Those of Pachliopta and Trogonoptera are included in this analysis for completeness. The various characters scored for neotropical Battus and Parides were based on photo graphic and preserved immature stage material in my own collection (e.g. gift from the Late Hamilton Tyler of copies of good colour slides and photographic prints by Paul Spade for Mexican species), plus the plates in Tyler et al. (1994). The latter publication also enabled neotropical Euryades characters to be scored. Monotypic Madagascan Pharmacophagus antenor was scored from preserved adult and im mature stage material in my collection. Cressida, Ornithoptera, Troides, Trogonoptera, Atrophaneura, Pachliopta were all scored from personally collected field data, plus some obtained from the literature: notably Igarashi (1963, 1979, 1984) for his illustrations of many papilionid immature stages, and Straatman (1968), Straatman and Nieuwenhuis (1961), Jumalon (1966, 1970), Koiwaya (1989), Goh (1994) and Weintraub (1995) for some important Oriental subregion papilionid immature stages (e.g. Atrophaneura and Trogonoptera).

Enumeration and Discussion of Troidini Characters Traditionally used characters of adult genitalia, wing pattern and wing venation are deliberately omitted unless they appear to be reasonably easy to interpret and so can be polarised with relative confidence. Immature stage characters are favoured, Reappraisal of Troidini 55 but are still conservatively selected. All multistate characters are ordered to reflect assumed increasing apomorphy, all "0" states being considered plesiomorphic.

FOODPLANTS 1. Primary foodplant genus: Q=Aristolochia, 1=Pararistolochia, 2=Thottea, 3 = None of these (Rutaceae, etc.). This an important character which clearly distinguishes genera such as Bhutanitis, Parides, Battus, Euryades, Cressida and Pharmacophagus, all of which are primarily Arislolochia feeders. Troides (but not including the solely Aristolochia- feeding Trogonoptera) have been recorded on certain Thottea species in the Oriental Region, but the genus specialises mainly in Arislolochia. Ornithoptera are primarily Pararistolochia-feeders (Parsons, in press), except that 0. priamus

Atrophaneura Ornithoptera Papilio Papilio (Pachliopta) (Chilasa) (Sugura) Fig. I. Comparison of tribe Troidini and other papilionid (as indicated) first instar larval tubercles and configurations of tubercle setae (after, Igarashi, 1984: not to scale). Parentheses denote taxa considered in this study to be of subgeneric (species group) status. A) Bhutanitis (Zerynthiini: Parnassiinae) and Battus. B) Euryades. C) Cressida. D) Parides, Troides and Trogonoptera. E) Atrophaneura and Pachliopta. F) Ornithoptera. G) Papilio subgenus Chilasa (Papilionini: Papilioninae). H) Papilio euchenor (Papilionini: Papilioninae) (subgenus? Sugura Okano, 1983). 56 Michael J. Parsons

(Linnaeus, 1758) also utilises Arislolochia throughout its range, and 0. goliath Oberthur, 1888, and 0. victoriae (Gray, 1856) have become wholly Arislolochia adapted. Atrophaneura (plus Pachliopta) are primarily Thottea feeders. Of rele vance, for example, was the observation by Straatman and Nieuwenhuis (1961) that A. aristolochiae (Fabricius, 1775) in Sumatra would oviposit on the natural Arislolochia foodplant of Troides (Trogonoptera) brookiana, but that this proved fatal to the resultant larvae. Some Atrophaneura species have, however, extended their ranges northwards and eastwards by adapting to feed on Arislolochia, and even Pararistolochia. It is not certain whether the polarity of Pararistolochia, and Thottea could be reversed above, but Pararistolochia seems to be a more likely ancestral foodplant, and could perhaps even be considered as the plesiomorphic character state for Papilionidae as a whole.

IMMATURE STAGES

OVUM 2. Ovum glue thickness: 0= Thin, l=Thick. This character defines a synapomorphy which unites all genera of the Troidini, except Troides, in which the thin condition of the glue is apparently a reversal. In Troides (minus Trogonoptera) the red glue covering the egg is much thinner than in other Troidini. The ovae of Papilio and Bhutanitis are very thinly covered by a layer of pale yellow glue. 3. Ovum glue ribbedd: 0= No, l=Yes (Figs. 2 and 3). This character defines a synapomorphy which unites all the troidine genera except Ornithoptera and Troides (in which the unribbed condition is apparently a reversal). The thick, waxy-looking, reddish or orange glue covering the troidine ovum is deeply furrowed, or sculpted into fairly regular vertical ribs, comprising granulose sections, in the other troidine genera (Figs. 2 and 3). Within Troides, subgenus Trogonoptera is notable (from colourphotographic figure in Goh, 1994) for its weakly-ribbed ovum which possesses granulosestriations as in Pharmacophagus. The ova of Papilio and Bhutanitis are evenly covered with glue. 4. Ovae grouped: 0= Yes, l=No. This character is obviously basedon the female's oviposition behaviour. It defines a synapomorphy which unites Troidini+ Papilio (sensu striclo). However, females of the troidine genus Battus, like Bhutanitis and other zerynthiines, lay ovae in closely grouped batches, often numbering 20-40. This is, therefore, an apparent reversal in Battus. The resulting larvae are gregarious during their early instars. Tyler et al. (1994) also regarded grouped ova as the plesiomor phic condition. Although ova are laid singly in Baroniinae, grouped ova are prevalent in Aristolochiaceae-feeding Zerynthiini, so it appears reasonable to Reappraisalof Troidini 57

assume that ancestral Papilionidae laid ova in batches, and that solitary egg placement is a specialisation. If it is found that the New Guinea species laglaizei, moerneri, and toboroi do belong in Papilio (rather than in a separate genus: presently Chilasa), then along with certain neotropical species (e.g. anchisiades Esper, 1788), these species with grouped ova would have to be considered here. However, I consider that they warrant separation from Papilio on this and other characters, so Papilio, like most Troidini, is scored as laying solitary eggs. 5. Ovum diameter: 0 = Less than 2.2 mm, 1= Between 2.8 to 4.8 mm. This character distinguishes an autapomorphy which defines Ornithoptera, in which the apomorphic condition of the egg is extremely large for a butterfly (and indeed any insect). A silkmoth species is wrongly credited in the Guinness Book of Records as laying the world's largest insect egg, the title correctly belonging to whose egg may reach a maximum diameter of 4.8 mm (Parsons, 1991). Even the eggs of the smallest Ornithoptera species, 0. richmondia (Gray, 1852) and 0. meridionalis (Rothschild, 1897), are markedly larger than the eggs of all other troidines: notably Troides (the only other genus of Troidini which comprises large species).

LARVA 6. First instar larva head chaetotaxy. Number of setae per cephalic lobe: 0 = Less than 14, 1= 14 or more. As noted by Igarashi (1984) and adopted by Tyler et al. (1994), the number of head setae of first instar larvae is apparently a useful character. Igarashi recognised three specialised conditions of increasing numbers of setae. The third category (not included here) was considered by Igarashi to be the most advanced, and is a state where numerous setae are present on the head, obscuring any ground plan pattern present in Igarashi's first or second categories. The division between states "0" and "1" of 14 setae (as selected by Tyler et al.) is based on the fact that most parnassiines, zerynthiines and troidines possess less than this number (although , Baroniinae, is specialised, bearing numerous Y-shaped setae), whereas most papilionines possess greater numbers of setae. Within the Troidini, Atrophaneura (but perhaps not all species?) bears 23 setae per lobe, whereas Pachliopta (but perhaps not all species?) has 13, as do Troides and Ornithoptera. 7. First instar larva body subdorsal tubercle posteriorly-directed basal seta: 0=Absent, 1= Present. As Igarashi (1984) demonstrated, the subdorsal tubercle morphology of the first instar papilionid larva is distinctive and, therefore, provides very important taxonomic characters (Fig. 1) for systematic and phylogenetic analy ses (including characters 8-11 below). Igarashi noted progressive specialisa- 58 Michael J. Parsons

tion (i.e. an apparent sequential transition) of the condition of these body tubercles and their setal pattern when various papilionid larvae are compared with one another. The ancestral papilionid larva was apparently devoid of prominently raised tubercles, these being developed in the zerynthiine papilionids, reaching their zenith in troidines, and undergoing subsequent reduction in groups of 'higher' Papilioninae such as Papilio and Graphium. Nearly all Troidini are well characterised by their possession of a distinctly separate, posteriorly-directed basal seta to the subdorsal tubercle. The basal seta is, however, lacking in Battus which notably also has the plesiomorphic condition of a single long apical seta as in Bhutanitis. The character represents, therefore, a synapomorphy of Troidini, except for an apparent reversal in Battus. The condition in Papilio is that the entire tubercle is semichitinous and the basal seta appears to have been secondarily lost. In the later instars of many Papilio species the tubercles become fully rigidly chitinised. 8. First instar larva body subdorsal tubercle sclerotised apex: 0 = Absent, 1= Present. This character (Fig. 1), illustrated by Igarashi (1984), defines a synapo morphy which clearly unites Troidini + Papilio. The troidine genus Battus (Fig. 1A) possesses, however, the plesiomorphic condition, lacking the sclerotised apex to the subdorsal tubercle. 9. First instar larva body subdorsal tubercle sclerotised apex size: 0= Small, 1= Large. This character (Fig. 1), illustrated by Igarashi (1984), distinguishes an apomorphy which groups the 'higher Troidini' (i.e. minus Euryades and Cressida, but also, by default, Battus which lacks a sclerotised apex) + Papilio, whose subdorsal tubercles bear large sclerotised apices. 10. First instar larva body subdorsal tubercle sclerotised apex elongate: 0 = No, 1 =Yes. This character (Fig. 1), illustrated by Igarashi (1984), distinguishes an autapomorphy that clearly defines Ornithoptera. The sclerotised apex of the tubercle is longest in Ornithoptera. In Papilio a similar apomorphic condition exists in the New Guinea species P. euchenor Guerin-Meneville, 1829 (Fig. 1H), but the tubercle is not heavily chitinised as in Ornithoptera. Together with its specialised pupal characters, this has prompted the placement of euchenor in its own monotypic genus Sugura by Okano (1983), but further study is required to better endorse this taxon. 11. First instar larva body subdorsal tubercle spinose setae: 0 = Singular, 1= Multiple (more than 3). As in character 7 above^ this character (Fig. 1), illustrated by Igarashi (1984), is a synapomorphy of Troidini + Papilio, with an apparent reversal in Battus, which possesses the plesiomorphic condition and is notably similar to Reappraisal of Troidini 59

Bhutanitis (Fig. 1A). As Igarashi demonstrated, the general trend in first larval instar tubercle spinose setae is to become more abundant, with single setae (unisetose), or groups of 2 or 3, in the least specialised papilionids (Baroniinae and Parnassiinae), these not being raised on tubercles as in Papilioninae (or at least not as prominently so in Zerynthiini of Parnassiinae). 12. First to fifth instar larvae spinose setal retention of tubercles: 0=Full, increasing in number, 1=Partial, decreasing in size and number up to fourth instar, 2 = Entirely lost by second instar. In the outgroup, Bhutanitis, as in all zerynthiines, the larval tubercles fully retain their spinose setae, and these increase in number. In all other troidines, except Euryades and Parides, these setae are lost by the second instar. Unique in the Troidini, Parides has the plesiomorphic condition persisting up to the fourth instar. Similarly, Euryades is unique in the Troidini for its complete retention of spinose setae throughout all instars, being entirely zerynthiine-like in this character. In Papilio, as in many other Papilioninae, the gradual reduction in setal size and number accompanies the gradual loss of the entire tubercle (character 13 below). 13. First to fifth instar larval tubercle development: 0= Remain prominent, 1= Degenerate. This character, tabulated by Igarashi (1984) distinguishes an autapomor- phy which clearly defines Papilio. Gradual loss of tubercles is a specialisation related to overall reduction in the larval profile and, therefore, increase in crypsis. Zerynthiini and Troidini advertise their presence as larvae, whereas those of Papilionini mainly do not. 14. Fourth and fifth instar larvae length of lateral prothoracic tubercles compared to other body tubercles: 0= Approximately same length, 1=3-4 times longer, resembling 'pseudo-antennae.' This character distinguishes an autapomorphy which defines Battus. The lateral tubercles of the prothoracic segment of larval instars 4 and 5 are 3-4 times longer than the remainder of the abdominal tubercles, resembling antennae, their function apparently being sensory. A similar elongation of these tubercles is present in the genus Sericinus (Parnassiinae: Zerynthiini), as well as in the troidine Ornithoptera vicloriae (in which this is obviously a specialisation within its genus). 15. Mature larva tubercle size: 0=Prominent, l=Larger, 2= Vestigial or absent. This character, like 12 above, shows that Euryades possesses the plesiomor phic condition very similar to that of Bhutanitis and other zerynthiines. Otherwise tribe Troidini might be defined by the state " 1" apomorphic tubercle condition of character 15. The typical apomorphic condition in Papilio (i.e. Papilionini) is for the tubercles to be absent. 16. Mature larva tubercle spinose setae: 0= Prominent, 1= Reduced, 2= Entirely 60 MichaelJ. Parsons

fleshy and non-setose. Like 15 above, this character defines an autapomorphy which clearly unites the troidine genera, with the exception of Euryades which possesses the ple siomorphic condilion. Otherwise tribe Troidini might be defined by the state "2" apomorphic tubercle condition of character 16. 17. Mature larva presence oi distinctly shorter subdorsal abdominal tubercles: 0 = None obviously shorter than the rest, 1= Shorter on segments 3 + 4, 2sShorter on segments 2 + 3 and 5 + 6, 3 = Most reduced or absent. This character distinguishes autapomorphic states which define Pharma cophagus ("1") and Trogonoptera ("2"). Parides exhibits some variation in Reappraisal of Troidini 61

subdorsal abdominal tubercle length in some species, but the majority, like the remaining troidine genera, clearly exhibit the plesiomorphic condition. There is frequently a reduction of tubercles in Battus by the final instar, but this is a much more uniform reduction of all tubercles, as opposed to specific tubercles in Pharmacophagus and Trogonoptera. In some Battus species this is extreme so that all the subdorsal tubercles become small and insignificant, while the lateral tubercles become vestigial, or are entirely lost. 18. Mature larva angle of subdorsal abdominal tubercles in relation to body: 0= Remain erect (60-90°), 1= Fully recumbent (ca. 180°) and posteriorally- directed. This character distinguishes an autapomorphy which defines Trogonoptera. The subdorsal abdominal tubercles of the mature larva uniquely point back wards and lie appressed to the body segments (e.g. photograph in Goh, 1994). 19. Mature larva ground colour: 0 = Dark striated, 1= Uniform. This character defines an apomorphy which apparently arose convergently in Euryades and Ornithoptera+Pharmacophagus. Papilio (Papilionini) also possesses the apomorphic condition. The dark striated ground colour condition is not always obvious in the troidine genera for which I score it, but it does occur in most species of each genus. Battus provides a very well defined example of the plesiomorphic condition of this character as do some Troides (eg. Fig. 25). 20. Larva with most instars bearing red spots or patches: 0= Present, 1=Absent. Excepting Baroniinae, the plesiomorphic condition of the papilionid larva is considered to be dark, spotted with red or orange. The red markings warn of its close chemical association with Aristolochiaceae. Of all the taxa in this analysis, only Papilio autapomorphically lacks these red spots as it is not an Aristolochiaceae-feeder and its larvae are primarily cryptically coloured (green and brown). 21. Larval abdominal saddle form, when present: 0= Absent, 1= Segments 3 & 4 irregularly (e.g. Fig. 5), 2= Segments 3 & 4 as a distinct diagonal lateral line (Figs. 25 and 26), 3= Segment 4 only (Figs. 23 and 24). As Igarashi (1984) pointed out, the configuration of the pale mid- abdominal larval markings (forming a distinctive 'saddle,' e.g. Figs. 23-26) is a useful diagnostic character in Papilionidae, being present in many troidine and Fig. 2. Pharmacophagus antenor ovum: lateral view. Fig. 3. Ditto: dorsal. Fig. 4. P. antenor fourth instar larva: lateral. Fig. 5. P. antenor fourth instar larva detail of saddle markings: lateral. Fig. 6. P. antenor fifth instar larva: lateral. Fig. 7. P. antenor pupa detail of prothorax and head: dorsal. Fig. 8. P. antenor pupa detail of thorax and head: lateral. Fig. 9. P. antenor detail of pupal dorsolateral abdominal tubercle (segment 5): dorsal. Fig. 10. Ditto: lateral. 62 Michael J. Parsons

papilionine genera, but absent throughout all instars in Euryades, Battus and Bhutanitis (Zerynthiini), for example. State "3" of this character distinguishes an autapomorphy which defines Ornithoptera. I assume that the normal extent of the saddle (across segments 3 and 4: Figs. 25 and 26) has been second arily reduced to 4 only in Ornithoptera (Figs. 23 and 24). In addition, some Ornithoptera species — meridionalis, paradisea Staudinger, 1893, tithonus de Haan, 1840, and rothschildi Kenrick, 1911 — lack a saddle, but only in their later larval instars. In Parides, Troides (plus Trogonoptera) and Atrophaneura (plus Pachliopta) the saddle characteristically forms a broad or narrow line laterally and diago nally across segments 3 and 4 (Figs. 25 and 26).

PUPA 22. Pupa overall shape in dorsal profile: 0= Elongate-ovoid, narrow dorsally across wings (Figs. 27 and 29), 1= Squat, broad dorsally across wings, distinctly diamond-shaped (Fig. 32). The overall shape of the pupa is an important character and shows clear general trends. The most plesiomorphic forms (not included here) are repre sented by Baroniinae and Parnassiinae (Parnassiini). These are squat, ovoid, weakly C-shaped in lateral profile, the thoracic region not as distinctly demar cated as in the next, more advanced, pupal shape. This next shape is well defined in Luehdorfia (Parnassiinae: Zerynthiini), Cressida (Fig. 31), Euryades, Ornithoptera (Fig. 29) and Pharmacophagus (Fig. 27), the sequence exhibiting a definite progressive increase in the typical S-shaped lateral profile of the pupa (character 23 below). Many papilionines (including Papilio, but notably the laglaizei group in New Guinea) retain this general pupal shape, while others broaden laterally as in the next troidine pupal group. This group are also S- shaped in lateral profile, but more weakly so, and are characterised by a much greater lateral expansion of the wings at about abdominal segment 3: Battus, Parides, Troides (plus Trogonoptera) and Atrophaneura (plus Pachliopta) (Fig. 34). Certain zerynthiines (Bhutanitis, Sericinus, Parnalius), like some papilionines (notably Oriental subregion clytia group) have evolved cryptic, markedly cylindrical, stick-like pupae. This is a clear specialisation in relation to camouflage permitting pupation on exposed twigs (etc.). In light of a parallel specialisation in Bhutanitis I consider it necessary to use its close relative Luehdorfia to score Bhutanitis (i.e. Zerynthiini) as plesiomorphic "0". 23. Pupa overall shape in lateral profile: 0= Short, weakly S-curved, abdomen and 'ventral margin' (legs plus wing profile) mildly curved (Fig. 31 and 33). 1= Elongate, strongly S-shaped, abdomen and 'ventral margin,' prominently outwardly curved (Figs. 28 and 30). This character defines a synapomorphy which groups Pharmacophagus + Ornithoptera separately from the remainder of the troidine genera. It is impor- Reappraisal of Troidini 63

tant to state here that the synapomorphy for this pair of genera is also representative of several other pupal characters (apart from 22 above) which could be used to further endorse the monophyly of this clade. In particular, the overall shape and combination of subtle (i.e. difficult to effectively define and quantify) characters of the pupae are more similar to each other than to any of the other troidine pupae (Figs. 27-30). The cuticular condition and coloura tion of the Pharmacophagus and Ornithoptera pupae also exhibit important similar ities. In all other troidine genera, except Cressida (Fig. 31) and Atrophaneura (plus Pachliopta) (Fig.34), the pupal cuticle bears a distinctive reticulate pattern and mild rugosity. For example, as Igarashi (1984) noted, the brown form of the Euryades pupa resembles similar dark forms of Battus pupae in colour and in the "minute reticular pattern" of the pupal cuticle. The possibly homologous, most plesiomorphicexpression of this character may well be the strongly rugose, thick pupal cuticle of Baronia (Baroniinae) and Luehdorfia (Zerynthiini). The 'milder' condition in the Troidini is especially notable across the wings in reticulately patterned genera because, by comparison, it is clearly absent in the cuticle of the wings in Cressida (Fig. 31), Pharmacophagus (Figs. 27 and 28), Ornithoptera (Figs. 29 and 30) and Atrophaneura (plus Pachliopta) (Fig. 34). The pupae of the latter four genera are also characterised by an obvious waxy gloss that is only weakly apparent in other troidine genera. However, in considera tion of other characters, the loss of a distinctly reticulate and rugose cuticle, and apparently reciprocal gain in cuticle glossiness, were likely independently derived characters for Atrophaneura. It is also reasonable to assume this in light of the (other) extremely specialised modifications of the colour and form of the Atrophaneura pupa (Figs. 13 and 34). As the above mentioned characters are difficult to interpret effectively, I have omitted them from the data matrix. 24. Pupa primary dorsolateral abdominal tubercles present on segment numbers: 0=4-7, 1=5+ 6. State "2" of this character distinguishes an apomorphy which defines Troides. The restriction of primary dorsolateral abdominal tubercles to only segments 3 and 4 in Troides (coupled with their increase in size) is a specialisa tion from the predominant plesiomorphic condition of segments 4-7 in Papilionidae. 25. Pupa dorsolateral abdominal tubercle overall shape: 0= Small and conical (Figs. 29-31), 1=Large, broad, and truncate or spatulate. (Figs. 27, 28, and 32-34) This character defines an apomorphy which groups the troidine genera, except Euryades, Cressida(Fig. 31) and Ornithoptera (Figs. 29 and 30). Like the outgroups Bhutanitis and Papilio, the latter possess the plesiomorphic condition. Pharmacophagus can also be scored for this despite the fact that its tubercles are specialised, being rounded and irregularly nodular as highlighted in character 26 below. Munroe (1961) considered that the larva and pupa of Cressida larva 64 Michael J. Parsons

and pupa (Fig. 31) are "primitive, having tubercles low and unspecialized." 26. Pupa dorsolateral abdominal tubercle apices: 0= Rounded or tapered to a point (Figs. 30-34), 1= Bluntly truncate, deltoid and irregularly nodular (Figs. 9 and 10). This character distinguishes an autapomorphywhich defines Pharmacophagus. In dorsal profile, these are uniquely deltoid and nodular (Figs. 9, 10, 27 and 28). In all other Troidini the tubercles are tapered, even when they are slightly truncate or spatulate (Battus, Parides, Atrophaneura). 27. Pupa abdominal segment 4 distinctly laterally flanged: 0= Absent (Figs. 30 and 33), 1= Minimally flanged (Fig. 28), 2= Large and semispatulate or spatulate (Fig. 34). This character distinguishes an apomorphy which defines Pharmacophagus through its possession of the autapomorphic state "1" (Fig. 28). The weak lateral flange in the Pharmacophagus pupa is one character suggesting a relation ship of the genus with the prominently flanged Atrophaneura (plus Pachliopta) pupae (Fig. 34). 28. Pupa frontal profile: 0= Square and simple with small vertically-directed lateral tubercles (e.g. Figs. 7 and 27), 1= Ornate with medium to large laterally- directed lateral tubercles (Figs. 29 ad 32). This character defines an apomorphy which groups the troidine genera, except Cressida (Fig. 31) and Pharmacophagus (Fig. 27) which possess the ple siomorphic condition. As in character 22 above, I use the zerynthiine Luehdorfia, rather than specialised Bhutanitis, to polarise the character. 29. Pupa saddle marking: 0= Present (e.g. Figs. 27, 29 and 32), 1= Highly modified or secondarily lost. According to my polarisation of this character, based on Bhutanitis, the saddle ground plan consists of a simple paler pigmented patch dorsally across the metathoracic segment and abdominal segments 1-4. Most troidines (notably Battus, Parides, Troides plus Trogonoptera, Pharmacophagus and Ornithoptera) apomorphically possess a characteristic pair ofnarrow, slightly distally diverg ing, subdorsal dark lines across the saddle, from the metathoracic segment to the commencement of abdominal segment 4 (e.g. Figs. 28 and 30). The character is somewhatobscure in some Battus and Parides, but is otherwiseeasilyscored for these genera. In Cressida (Fig. 31) and Atrophaneura (plus Pachliopta) (Figs. 13 and 34) pupae the saddle has become highly modified, but the divergent lines are recognisable, albeit rather obscured by the additional disruptive pattern of horizontal reddish-brown lines. In Euryades, as in many Papilionini (e.g. Papilio machaon), the saddle is so obscured as to be effectively unrecognisable. 30. Pupa overall colour and main markings: 0= Yellow, pale brown, pinkish-brown or green marked with dark brown, 1= Pure white marked with maroon red, brownish-mauve and bright yellow (Fig. 31). Reappraisal of Troidini 65

This character distinguishes an autapomorphy which defines the genus Cressida. 31. Pupa presence of mauve or purple pigment at apices of some protuberances, and/or laterally on raised margin of wings, and onto abdomen: 0= Absent, 1= Often present. Although not present in all species (apparently because of specific varia tion), or all pupal forms of some species (e.g. in some species of those genera with dimorphic pupae), this character is typical of Euryades, Battus, Parides and Troides (plus Trogonoptera). Conversely, mauve or purple pigment is always absent in Bhutanitis (and other zerynthiines), Cressida, Ornithoptera, Atrophaneura (plus Pachliopta), and Papilio. 32. Pupa colour morphism: 0= Monomorphic, l=Distinctly dimorphic. This character is difficult to score effectively. For example, Ornithoptera pupae are primarily monomorphic. That of 0. richmondia is monomorphic, and is surprisingly distinctively pale green. However, pupae of 0. priamus poseidon Doubleday, 1847, in mainland New Guinea include an additional form which tends slightly towards a greenish hue. This form is likewise present in Troides, but more frequently and distinctly so, and Troides also has a distinctive orange pupal form, thereby being effectively trimorphic. In the neotropics, Battus and Parides have distinctly dimorphic (green/brown) pupae to the same degree as in Papilio (e.g. Clarke and Sheppard, 1972; Hazel and West, 1979; Sims and Shapiro, 1983; Tyler el al., 1984). Euryades, Cressida, Pharmacophagus, Atrophaneura and Pachliopta are all monomorphic. Monomorphism is scored as the plesiomorphic condition as the exposed pupae of Zerynthiini (i.e. including Bhutanitis) are all monomorphic. Terrestrial Parnassiini pupae cannot be con sidered for effective comparison as they are hidden (unexposed). Nevertheless, they are also monomorphic. 33. Pupation distance from foodplant: 0 = Near, l=Far. This character, a result of behaviour of the final instar larva during the prepupal phase, distinguishes an autapomorphy of Troidini that has apparently reversed in Troides (plus Trogonoptera) and Atrophaneura (plus Pachliopta). For example, Troides do not wander far away, if at all, from their foodplants, whereas Ornithoptera do. West and Hazel (1979) and Sims and Shapiro (1983) provided details of pupation sites in Battus. This character is also useful in conjunction with the following character 34. 34. Main pupation site: 0 = Stems and twigs, l=Leaves. This character (as for 33 above, resulting from behaviour of the final instar larva during the prepupal phase) distinguishes an autapomorphy which defines Ornithoptera. Ornithoptera invariably pupate underneath leaf midveins of other than their foodplant vines, whereas genera such as Atrophaneura, Troides and Parides, Battus (etc.), usually select stems and twigs, often of the foodplant, on 66 MichaelJ. Parsons

which to pupate. The latter is true of the outgroup Bhutanitis (Zerynthiini) (e.g. Igarashi, 1989), and of the zerynthiine genus Luehdorfia (e.g. Ishii and Hidaka, 1982).

ADULT 35. Adult female presence and form of sphragis: 0= Small and plug-like, 1= Large and alate, 2=Absent. The presence of a small, generally unstructured and plug-like sphragis appears to be universal in the Papilionidae (e.g. Orr, 1988), except where it is apparently secondarily lost. Hence sphragis presence is treated here as the plesiomorphic condition. This character could be further divided into several other subcategories using size and shape. Euryades and Cressida are clearly specialised in the shape of their prominently alate external sphragides. Never theless, their exact homology is not clear (as they are not exactly morphological ly similar) and the occurrence of large external sphragides in the Papilionidae is sporadic. For example, large scoop-shaped external sphragides are otherwise present only in Parnassius (Parnassiinae). Furthermore, as Orr (1988) and Hauser (1993) both implied, it is difficult to argue any real trend in sphragis evolution in relation to its size and shape. However, it does appear to be true that a simple plug-like sphragis was evolved early on in the butterflies and, therefore, that its loss (e.g. in Papilio) is a specialisation. The Bhutanitis sphragis (well figured by Saigusa and Lee, 1982) is irregular and somewhat intermediate between plug and alate form sphragides. However, it is scored as "0" here because it is distinctly less well developed than in Euryades and Cressida. 36. Adult female bursa copulatrix: 0= Small or vestigial, 1= Large and thinly membranous, 2= Larger, thickly corrugated, membranous and distinctly double chambered. Miller (1987b: 416) rightly cautioned that this character, in general, and its polarity, are difficult to interpret. Nevertheless, based on the outgroup Bhutanitis and other zerynthiines, the plesiomorphic condition appears to be a small bursa and sclerotised ductus (notably in Bhutanitis as illustrated by Saigusa and Lee, 1982, and in Luehdorfia as illustrated by Miller, 1987b). The apomorphic conditions show a progressive enlargement of the bursa and loss of a chitinised ductus (state "1"), ultimately resulting in a very large, thickly corrugated, membranous and distinctly double chambered bursa (state "2") as exhibited by Ornithoptera and Troides (plus Trogonoptera). Interestingly, Battus shows a definite tendency towards this latter state (e.g. good figures in Miller, 1987b). This is an important character, the apomorphic condition of which groups the 'higher Troidini' (i.e. minus Euryades and Cressida) + Papilio. Euryades and Cressida share the plesiomorphic condition of the bursa copulatrix. Both genera entirely lack signa in the bursa, and the bursa is so small as to be truly Reappraisal of Troidini 67

vestigial by comparison with all the other taxa under analysis. Baronia (Baroniinae) possesses a typical pair of spinose signa in the bursa, so the absence of signa in these genera, and in zerynthiines such as Bhutanitis and Luehdorfia, is apparently a specialisation. 37. Adult female foretibia and foretarsus presence of spines and sensilla: 0 = Only on first and second tarsomeres, the sensilla concentrated on the first tarsomere, and occasionally the second, 1= Spines in a tightly spaced row, running length of tibia and tarsus, sensilla evenly spaced between these. This character was established by Miller (1986, 1987b) and defines an important apomorphy which strongly supports the monophyly of the Troidini. In fact, it is the only such character which unequivocally does so. I include it here as it appears unambiguous and easy to interpret and I agree that it is one of the best indicators of troidine monophyly. 38. Adult male pseuduncal suture: 0= Absent, l=Complete, 2= Incomplete. Hauser stressed caution in interpreting this character as considerable differences exist in the degree of fusion within the Troidini and Papilionini. Like the form of sphragis (character 35 above) the male pseuduncal suture provides a synapomorphy (state "1") which suggests the grouping of Euryades + Cressida. The remainder of the troidine genera possess the state "2" synapo- morphic condition. In most troidine genera, and Papilio, the suture is ap parently secondarily partly fused. Its complete absence is seemingly the plesiomorphic condition as, within the Papilionidae, a pseuduncus (or super- uncus) is restricted to the Papilioninae (e.g. Miller, 1987b; Hauser, 1993). 39. Adult male discrete hindwing androconial brush at vein 1A+2A: 0= Absent, l=Fine and weakly developed, 2= Coarse and well developed. This character supports the monophyly of the clade Troidini + Papilio. It is usually associated with spatulate androconial scales on the dorsal surface of the abdomen. Its absence is considered to be the plesiomorphic condition. Its development, along with pouched inner marginal hindwing androconia (charac ter 40 below), is a specialisation of Papilioninae. It is present in all Troidini and is also scored as being present in Papilio as various species possess the androconial brush (e.g. Miller, 1987b). Its development in Troidini reaches its extreme in Ornithoptera, notably 0. paradisea and 0. tithonus. 40. Adult male hindwing (pouch) androconial scales: 0= Absent, l=Coarse, spatu late and fixed in position, 2= Fine, hair-like and deciduous. The presence of pouched hindwing androconial scales in the adult male defines the monophyly of Troidini. This character is relatively easily observed as Troides (including Trogonoptera), Atrophaneura (including Pachliopta) and Parides males possess fine and deciduous androconial scales in the pouch at the hindwing inner margin, whereas the opposite is true in the remaining troidine genera. Although various species of the subgenus Pachliopta of Atrophaneura have 68 Michael J. Parsons

these scales secondarily reduced and often vestigial (resembling state "1"), I have scored Pachliopta as possessing the state "2" condition as this is clearly a further derived condition (i.e. modification) of state "2". Moreover, the male of A. (Pachliopta) neptunus (Guerin-Meneville, 1840) possesses a normal well- developed hindwing pouch and androconia, whereas the hindwing pouches and androconial scales of some members of the subgenus Atrophaneura are also variously modified: e.g. in A. (Atrophaneura) kageni (Rogenhorfer, 1889) and its close allies, priapus (Boisduval, 1836), luchti (Roepke, 1935), and sycorax (Grose- Smith, 1885) (which are all notable Troides mimics). In Ornithoptera, for example, the pouch (or wing flap) apparently contains androconial scales, but these are spatulate and differ little from ordinary scales (e.g. Miller, 1987b: 415). 41. Adult antenna presence of ventral sensory setae (sensilla): 0=ln patches, l=In pits. Like character 40 above, this adult character also supports the monophyly of the Troidini. It has apparently been secondarily lost in Battus which lacks the pits. 42. Adult labial palpus number of segments: 0=3, 1=2. This character was established by Miller (1986, 1987b) and supports the monophyly of the Troidini. It has either been plesiomorphically retained, or has otherwise been secondarily reversed, in Pharmacophagus and Battus which, like Bhutanitis and Papilio possess a 3-segmented labial palpus. As Miller sug gested, it seems clear that the majority of troidines have lost the basal segment of the labial palpus as a specialisation. 43. Adult red body scaling on abdomen: 0= Absent, 1= Present and extensive on abdomen. This character defines an apomorphy which appears in Cressida, Pharmaco phagus and Atrophaneura (plus Pachliopta). When compared with many other characters, it seems distinctly homoplasious, suggesting that the apomorphic condition probably arose independently (and convergently) at least twice in Troidini. 44. Adult wing red scaling: 0= Present, 1=Absent. The apomorphic loss of red adult wing scaling groups Ornithoptera + Troides (plus Trogonoptera). Nevertheless, based on other characters, it appears likely that this was derived separately by Ornithoptera and Troides. The remainder of the troidine genera all bear clearly defined red spots or patches on their wings, representing the plesiomorphic condition.

The genus Ornithoptera Boisduval, 1832 Ornithoptera comprises large to extremely large butterflies, particularly the fe males which are markedly phenotypically different from their males. The genus is Reappraisal of Troidini 69

Table 1. Character distribution matrix for cladistic analysis of the papilionid tribe Troidini (Papilioninae) plus outgroups Bhutanitis (Bhut) and Papilio (Papil). For details of characterstates and polarity sec also text. 0 scores represent the assumed plesiomor phic condition, 1-3 increasingly more apomorphic states. Characters for Pharmacopha gus first instar larva were unknown (indicated as ?) but were scored anyway based on knowledge of other taxa.

CO is 33 •C § CHARACTERS OQ 1I a I 1 1 ^ 5 FP _ Primary foodplant genus: 0=Aristolochia, etc. 1 0 3 0 0 0 1 0 0 0 0 2 2 Ovum glue thickness: 0=Thin, l=Thick. 2 0 0 1 1 1 1 1 1 0 1 1 1 S 3 Ovum glue ribbed: 0=No, l=Ycs. 3 0 0 1 1 1 0 1 1 0 1 1 1 > Ovae grouped: 0=Yes, l=No. 4 o 0 1 1 1 1 1 0 1 1 1 1 1 _ Ovum sire: 0=<2.2mm, 1=2.8-4.8mm. 5 0 0 0 0 0 1 0 0 0 0 0 0 1st instar larva number ofsetae per cephalic lobe. 6 0 1 0 0 0? 0 0 0 0 0 1 0 1st instar larva subdorsal tubercule basal seta. 7 0 0 1 1 1? 1 0 1 1 1 1 1 1st instar larva subdorsal tubercule sclerotised apex. 8 0 1 1 1 1? 1 0 1 1 1 1 1 1st instar larva subdorsal tubercule sclerot. apex size. 9 0 1 0 0 1? 1 0 1 1 1 1 1 1st inst. larva subdors. tuberc. sclerot. apex elongate. 10 0 0 0 0 0? 1 0 0 0 0 0 0 1st instar larva body subdorsal tubercle spinose setae. 11 0 1 1 1 1? 1 0 1 1 1 1 1 1-5 instar larvae spinose setal retention of tubercles. 12 0 1 0 2 2 2 2 1 2 2 2 2 1-5 instar larval tubercle development. 13 0 1 0 0 0 0 0 0 0 0 0 0 4&5 instar length of lateral prothoracic tubercles. 5 14 0 0 0 0 0 0 1 0 0 0 0 0 Mature larva tubercle size. 0=Prom., l=Larger, etc. 15 0 2 0 1 1 1 1 1 1 1 1 1 Mature larva tubercles setae. 0=Prom., l=Reduc, etc. 16 0 1 0 2 2 2 2 2 2 2 2 2 Mature larva shorter subdorsal abdominal tubercles. 17 0 3 0 0 1 0 0 0 0 2 0 0 Mature larva subdorsal abdominal tubercles angle. 18 0 0 0 0 0 0 0 0 0 1 0 0 Mature larva ground colour: 0=Striated, l=Uniform. 19 0 1 1 0 1 1 0 0 0 0 0 0 Larva at most instars with red spots or patches. 20 0 1 0 0 0 0 0 0 0 0 0 0 _ Larvalabdominalsaddle form, when present. 21 0 1 0 1 1 3 0 2 2 2 2 2 ~ Pupa dorsal profile: 0=Elongate, l=Squat. 22 0 0 0 0 0 0 1 1 1 1 1 1 Pupa lateral profile: 0=Weak S-sh., l=Prom. S-sh. 23 0 0 0 0 1 1 0 0 0 0 0 0 Pupa primary dorsolateral abdominal tubercles. 24 0 0 0 0 0 0 0 0 1 1 0 0 Pupa dorsolateral abdominal tubercle overall shape. 25 0 0 0 0 0 0 1 1 1 1 1 1 Pupa dorsolateral abdominal tubercle apices. 26 0 0 0 0 1 0 0 0 0 0 0 0 Pupa abdominal seg. 4 distinctly laterally flanged. < 27 0 0 0 0 1 0 0 0 0 0 2 2 a. 3 Pupa frontal profile: 0=Square & simple, l=Ornate. 28 0 1 1 0 0 1 1 1 1 1 1 1 a Pupa saddle marking: 0=Present, l=Modif. or lost. 29 0 1 1 1 0 0 0 0 0 0 1 1 Pupa overall colour and main markings. 30 0 0 0 1 0 0 0 0 0 0 0 0 Pupa presence ofmauve or purple pigment. 31 0 0 1 0 0 0 1 1 1 1 0 0 Pupa colour morphism: 0=Monomorphic, l=Dimorph. 32 0 1 1 0 0 0 1 1 1 1 0 0 Pupation distance from foodplant: 0=Near, l=Far. 33 0 1 1 1 1 1 1 1 0 0 0 0 _ Main pupation site: 0=Leaves, l=Stems and twigs. 34 0 0 0 0 0 1 0 0 0 0 0 0 ~ Female sphragis: 0=Plug, l=Large/alate, 2=Absent. 35 0 2 1 1 0 0 0 0 0 0 0 0 Female bursa copulatrix: 0=SmVvest., l=Large, etc. 36 0 1 0 0 1 2 1 1 2 2 2 1 Female foretibia and tarsus spines and sensillae. 37 0 0 1 1 1 1 1 1 1 1 1 1

h- Male pseuduncal suture: 0=Absent, l=Compl., etc. 38 0 2 1 1 2 2 2 2 2 2 2 2 1j 3 Male discrete HW androconial brush at vein 1A+2A. 39 0 1 1 1 1 2 1 1 1 1 1 1 a Male hindwing (pouch) androconial scales. 40 0 0 1 1 1 1 1 2 2 2 2 2 < Antenna presence of ventral sensillae. 0=Patches, etc 41 0 0 1 1 1 1 0 1 1 1 1 1 Labial palpus number ofsegments: 0=3,1=2. 42 0 0 1 1 0 1 0 1 1 1 1 1 Red body scaling extensive and onto abdomen. 43 0 0 0 1 1 0 0 0 0 0 1 1 Wing red scaling: 0=Present, 1=Absent. 44 0 0 0 0 0 1 0 0 1 1 0 0 70 Michael J. Parsons renowned for this pronounced sexual dimorphism, as well as the beauty and sizeof its members (Fig. 20). Consequently, Ornithoptera are highly economically important, being in great demand by collectors and forming a significant part of the world insect trade (Parsons, 1992a and 1995a). The genus contains 10 species: alexandrae (Rothschild, 1907); chimaera (Rothschild, 1904); goliath Oberthur, 1888; meridiona- lis (Rothschild, 1897); paradisea Staudinger, 1893; priamus (Linnaeus, 1758); richmondia (Gray, 1852); rotfischildi Kenrick, 1911; tithonus de Haan, 1840; victoriae (Gray, 1856). All, except richmondia, priamus, goliath and victoriae, are endemic to the island of New Guinea (Figs. 16and 18), the western half of which includes Irian Jaya in Indonesian territory, the eastern half of which belongs to the Independent State of New Guinea (PNG). O. richmondia is endemic to Australia, being limited to a small contiguous area comprising subcoastal southern Queensland and subcoastal northernmost New South Wales (NSW), approximately between latitudes 25-30°S (Maryborough to Clarence River). 0. priamus is widespread throughout the Austra lian subregion, ranging from Sulawesi through New Guinea to the Solomons and north-eastern Australia (much of coastal and subcoastal Queensland). The range of 0. goliath is centred on mainland New Guinea, but to the west the species also occurs on Seram and Waigeo Islands in Indonesia, and to the east, in PNG, on Goodenough Island in the D'Entrccasteaux Group. 0. victoriae occurs in (within PNG territory) and throughout the remainder of the Solomons Group. '0. allotlei,' an hybrid between 0. priamus and 0. victoriae, has been recorded from Bougainville Island. 0. tithonus and 0. rotfischildi are apparently absent in PNG and are only known from western Irian Jaya. The smallest Ornithoptera is meridionalis, the female of which has a wingspan of about 13 cm. The largest, 0. alexandrae, has females which can attain wingspans of about 25 cm, being the world's largest butterfly species. Ornithoptera species have usually been divided among three subgenera (e.g. Haugum and Low, 1979). Part of the history of the usage of subgenera was discussed by Szent-Ivany (1970). In fact, the subgeneric names applied to the Ornithoptera are clearly unnecessary taxonomic divisions to describe what are, at best, species groups within the genus (Fig. 16). The systematic position of 0. alexandrae has been the subject of debate by many authors (e.g. Jordan, 1909- 10; Zeuner, 1943; McAlpine, 1970). However, this was discussed well by Blandin (1973) who suggested that the similarities (mainly their elongate wing shape) between 0. alexandrae and 0. victoriae were the result of convergence, rather than derivation. The two species have, therefore, traditionally been treated as representatives of the subgenus Aelheoptera Rippon, 1890. Schmid (1970) endorsed this in his phylogenetic appraisal of the Ornithoptera but, like Zeuner, he considered that the hybrid allotlei was a 'linking' species, so most of his conclusions are invalid. Blandin suggested that the normally broad wings of the Ornithoptera are modified in different ways to achieve the elongate wings of alexandrae and victoriae. Jordan had previously drawn Reappraisal of Troidini 71 attention to the greater similarity between the forewing venation of alexandrae and priamus, than between alexandrae and victoriae. In fact, the evolution of the specialised wing shape and pattern in alexandrae somewhat parallels the similar specialised development of these facies in the Malaysian Troides (Trogonoptera) brookiana (Wallace, 1855) and T. (T.) trogon (Vollenhoven, 1860). Unlike other members of Troides, their forewings are particularly elongate and are marked with iridescent green (rather than white) scaling. Therefore, although it is parsimonious to regard wing shape as homologous in alexandrae and victoriae, they nevertheless appear to have evolved this similar character state independently of one another. All previous conclusions regarding the interspecific relationships of Ornithoptera have, until now, been based primarily on adult characters. For example, Darby (1982 and 1985) carried out studies of Ornithoptera female genitalia. These are robust and heavily sclerotised, being fairly distinct for all species, and they will likely provide further taxonomically useful characters for analysis. However, no studies have been carried out that include morphological characters provided by Ornithoptera immature stages to test the correctness of, or better corroborate, previous findings, so these are employed here. Those of all but 0. rothschildi (which are only known in outline) have been described. The immature stages of meridionalis were recorded by Szent-Ivany and Carver (1967), Straatman (1967) and Szent-Ivany (1970); priamus and victoriae by Straatman (1970); alexandrae by Straatman (1971); priamus by Borch and Schmid (1973); paradisea by Borch and Schmid (1975); goliath and chimaera by Straatman and Schmid (1975); tithonus by Parsons (1995) (the latter four species being illustrated with colour photographic plates). Brief, but incorrect, statements made by Meek (1913) and Straatman (1971), respectively, that the larva and pupa of alexandrae are very similar to those of victoriae were, no doubt, biased by the earlier taxonomic arrangements drawn from adult characters. Later, however, Straatman and Schmid (1975) did state that the pupa of 0. goliath closely resembles that of 0. alexandrae. In fact, both the larval and pupal stages of 0. alexandrae most closely resemble those of 0. priamus, and it is the immature stages of 0. victoriae that distinctly differ from all other Ornithoptera species.

Generic Distinctness of Ornithoptera and Troides Earlier authors (e.g. Rothschild, 1895) often treated Ornithoptera as subordinate in rank to Troides, and this arrangement was recently readopted by Hancock (1983) and Miller (1986 and 1987). However, Parsons (1991: 83), maintained that the synonymy of Ornithoptera with Troides is untenable based on the differences in their general biology, and in many important morphological features of the adults and immature stages of both genera. For example, the two genera are easily separated by the general features oftheir adult venation, secondary sexual characters (Figs. 11, 20 and 21) and genitalia, as well as by behavioural, ecological and immature stage differences. Although Straatman (1976), and Sands and Sawyer (1977), have 72 Michael J. Parsons shown that intcrgeneric hybridization between 0. priamus and T. oblongomaculatus (GoEZE, 1779) does take place in the wild in New Guinea, only males result, confirming that the parent species are genetically distinct from one another. Troides has been variously treated taxonomically. ROTHSCHILD (1895) placed all of the large 'birdwing' butterflies of the Indo-Australian Region under Troides, while Jordan (1909-10 in Seitz) considered all to belong to Papilio Linnaeus, 1758. Rippon (1898-1906) believed Troides to represent a tribe containing the following genera: Aetheoptera, Pompeoptera Rippon, 1890, Schoenbergia Pagenstecher, 1893, Ornithoptera and Trogonoptera. In his revision of the birdwings, Zeuner (1943) treated the group as four genera (Ornithoptera, Schoenbergia. Trogonoptera, and Troides), while Munroe (1961) recognised only Troides and Ornithoptera as genera, and treated Trogonoptera as a subgenus of Troides. D'Abrera (1975) followed Munroe. but reinstated Trogonoptera at the generic level, while Haugum and Low (1975) erected a new monotypic genus, Ripponia, for the species hypolitus (Cramer, 1775). Xicui.escu (1980) argued that Trogonoptera is a valid genus. Nevertheless, reverting to Roth schild's (1895) lumping of taxa. Hancock (1983) and Miller (1986 and 1987), once again, placed all birdwings (i.e. including Oriiilhoplera) under Troides. The distinctness of Ornithoptera species from those of Troides is well illustrated by

Vein CuA2

Vein CuA2 =

Abdominal adroconial Troides Ornithoptera

A B D E Fig. 11. Diagrammatic representations of transverse sections of Troides (A-B) and Ornithop tera (C-E) male hindwing anclroconia showing the basic differences of these structures in each genus. The brush stands on vein IA+2A. A) Position of androconia in relation to abdomen. B) Configuration of androconia in Troides with hindwing flattened out. C) Ditto in Ornithoptera. D) Ornithoptera brush in retracted position. E) Ditto when fully erect and displayed. Reappraisal of Troidini 73 their different adult morphologies. Troides (plus Trogonoptera) males can be easily distinguished from those of Ornithoptera by the differences in structure of their hindwing inner marginal androconia (Fig. 11). This was noted by Miller (1987), but was not regarded by him as significant. Munroe (1961: 33) correctly inter preted these differences and used them to key out Ornithoptera and Troides as separate genera. Hauser (1993) pointed out that the occurrence of "bristle-like scent scales" on the ventral surface of the male hindwing anal region appears to be a reliable autapomorphous character supporting the monophyly of the subfamily Papilioninae in general. In Troides the margin forms a well-defined, flap-like fold or pouch densely packed with extremely fine, 'woolly,' white, hair-like, androconial scales arising from the wing upperside in the region of vein 1A+ 2A. These scales are deciduous and easily fragmented, even adhering to a smooth needle point when it contacts them (e.g. Zeuner, 1943; Parsons, 1983). It is notable, and particularly relevant that, as observed by Miller (1987), the only other troidine genera possessing similar male androconia are certain Atrophaneura Reakirt, 1865, and the neotropical genus Parides Hubner, 1819. No Troides species bear male forewing sex-brands, as in one species group of Ornithoptera and, although Ornithoptera males do possess flap-like inner marginal hindwing pouches, none of these contain androconia like those of Troides (the androconial scales being spatulate or, if hair-like, then extremely sparse). Instead, in Ornithoptera, the primary androconia comprise an erectile, retractable, brush-like inner marginal row of long, stiff, robust hair-scales arising from the wing underside along vein 1A+ 2A (Fig. 11C-D). These show a definite transition, increasing in size and robustness from priamus, alexandrae and victoriae (shorter, finer and sparser), through rothschildi and goliath (medium), reaching their greatest development in meridionalis, paradisea and tithonus (Figs. 20 and 21). In all Ornithoptera species, the brushes clearly serve to interact with other spatulate androconia which often form discrete and differently coloured sex-brands on the uppersides of the male abdomen. This is notable in 0. paradisea, for example, in which species the central portion of the abdomen often collapses forming a 'trough' as it dries in dead adult males as the cuticle of the tergites is thinner in this region (possibly in relation to the assumed pheromone secreting function of the overlying scales). The hindwing brushes obviously collect pheromones from these abdominal sex-brands and disperse these scents when erect and fanned out. Such brushes are also present in Troides, but the scales are much sparser, weaker and are not fully erectile. Therefore, as is clear in Figs. 11 and 20, the main difference in emphasis has been for Troides to develop the pouch androconia on the upper surface of the wing, whereas Ornithoptera has developed the vein 1A+2A brush androconia on the opposite (underside) side of the hindwing. As pointed out by D'Abrera (1975), Troides adults are not as sexually dimorphic as those of Ornithoptera and, unlike Ornithoptera, Troides tends to be somewhat 74 MichaelJ. Parsons polymorphic in both sexes. Most members of the genus are generally rather similar in appearance, being basically dark brown or black, the forewings often striped with white or yellow along the veins, the hindwings with translucent yellow to golden- yellow patches of various extent. The Troides abdomen is usually dark brown dorsally, yellowish ventrally, and a collar of bright red is often present on the pro- thorax (the "pronotal collar" of some authors). Ornithoptera, however, are characte rised by green or blue scaling in the males and much more distinctly differently coloured and patterned females. As Darby (1982) pointed out, in the female genitalia, the Ornithoptera lamella antevaginalis is usually much larger than that of Troides, with a bulbous fold (the operculum) that completely closes the ostium bursae, whereas the ostium is only partially closed in Troides. In addition, the Ornithoptera bursa is distinctly double chambered, with an additional apical appendix bursa, whereas that of Troides (sensu stricto) is only very obscurely double chambered, the appendix bursa only very mildly defined (so that Darby quite reasonably regarded the Troides bursa as being single-chambered). The distinctness of Ornithoptera from Troides is especially well illustrated by notable differences in their immature stages. For example, the smallest Ornithoptera eggs (ca. 3.0 mm diameter: 0. richmondia and 0. meridionalis) are almost one-third as big again as the largest Troides egg (ca. 2.0 mm diameter: T. oblongomaculatus) (pers. obs.). This obviously affects the female's capacity for oogenesis, and even the physical capacity of its abdomen to contain a certain number of mature eggs. For example, my dissections of females have shown that the 'carrying capacity' of Ornithoptera for mature ovae is one-third (e.g. 0. goliath) to two-thirds (e.g. 0. priamus) less than in Troides females whose abdomens can contain up to about 30 mature eggs at any one time during the ongoing cycle of oogenesis. This translates into a fecundity in Troides that is approximately double that of 0. priamus, and probably about three times the fecundity of the remaining Ornithoptera (Parsons, unpublished). The morphology of Ornithoptera and Troides larvae and pupa are markedly different (compare Figs. 23/24 with Figs. 25/26, and Figs. 29/30 with 32/33). As Igarashi (1979 and 1984) pointed out, the chitinous apices of the tubercles in the first larval instar of Ornithoptera are distinctly longer than in Troides (and all other troidine genera, including Cressida Swainson, 1832, and Atrophaneura). As noted by Munroe (1961), Ornithoptera later instar fleshy larval tubercles are longer, slenderer and more pointed than in Troides. In the mature larvae of the two genera the Ornithoptera larval ground colour never bears the body striations present in Troides, and the saddle mark in Troides characteristically forms an oblique lateral band on abdominal segments 3 and 4, but only a band on 4 in Ornithoptera. In Ornithoptera the apices of the tubercles are so acutely tapered as to be thread-like in all species (especiallyobvious in instars 1-4), whereas the tubercles of Troides, although tapered, are always more apically rounded. Considering also other troidine genera: Parides species possess very pointed subdorsal tubercles, but are never as acutely tapered as Reappraisal of Troidini 75 in Ornithoptera. Other Parides exhibit a mixture of apically rounded and longer, more tapered tubercles very similar to Troides (photos in Brown et al., 1981). Troides pupae (e.g. Fig. 32) are very broad (laterally, across the wings), being squat with broad, blunt dorsolateral abdominal tubercles. The Ornithoptera pupa (e.g. Fig. 29) is slenderer and more elongate than in Troides, with smaller, more sharply pointed dorsolateral abdominal tubercles. Like that of Parides, the Troides pupal cuticle bears a reticulate pattern and accompanying rugosity. The pattern is especially obvious across the wings in the two genera, for example. Conversely, however, the Ornithoptera pupal cuticle lacks this reticulate pattern and accompanying rugosity, being instead smoother with a reciprocal increase in glossiness that is only mildly observable in the Troides pupa. The Ornithoptera pupa saddle marking is similar to that of Troides in its overall appearance, but the divergent lines are broader and bolder. The Ornithoptera pupa is also unique for its tribe (an autapomorphy not included in the data matrix in Table 1) in that the dark markings laterally on abdominal segment three resemble a simple eye-spot (although the pair of abdominal 'eye-spots' so formed are not prominent and probably do not serve any protective function). Davidson and Aitken (1890) and Bell (1911) noted that the pupa of (Cramer, 1779), when stimulated by touch, can make "a husky squeaking noise produced apparently by friction of the abdominal rings" (Davidson and Aitken), or "a loud hissing sound... produced by rubbing the abdominal segments 8, 9, 10, 11 together at the margins by a contracting motion repeated at short intervals." (Bell). Ornithoptera pupae never stridulate. It is noteworthy that the clear distinction between Ornithoptera and Troides is also upheld by the immature stage behaviour of the two genera. Troides species (like Atrophaneura), almost invariably pupate on twigs and stems, very near, or belonging to, their foodplants. However, Ornithoptera larvae almost invariably wander a number of metres away from their foodplants, and always select the undersides of leaves on bushes or trees other than the foodplant on which to pupate, doing so along the leaf mid-vein.

Outgroup Selection for Analysis of Ornithoptera The same methodology as that outlined above for the cladistic analysis of the Troidini was used to study the Ornithoptera. From the data presented above, Ornithoptera is obviously clearly distinct from Troides and does not deserve to be subsumed in synonymy with that genus. Furthermore, in a complete analysis of the Troidini, using data drawn primarily from troidine immature stage characters, the results (Figs. 12, 13 and 15) suggest that Ornithoptera is phylogenetically more plesiomorphic than Troides, and indeed shares greater affinities with Madagascan endemic Pharmacophagus Haase, 1891, the relationship of Pharmacophagus and Ornitho ptera being closer to Australian subregion restricted Cressida (rather than any of the 'higher troidines' such as Troides). Therefore, I have used Cressida as the outgroup to 76 Michael J. Parsons

Table 2. Character distribution matrix for cladistic analysis of Ornithoptera. 0 scores represent the assumed plesiomorphic condition, 1-3 increasingly more apomorphic states. FP= Foodplant, EC= Ecology. Foodplant and early stage characters for rothschildi are based on scant data, so certain characters (indicated as ?) arc scored based on knowledge of other taxa.

.s 3s CO c 0) Q> c o 1 0) ,J0 .9 CO CO & CO 8 1 2 •2 o CHARACTERS t co s §> 2 •5 35 S FP _ Foodplant main genus. 0=Pararistolochia,etc. 1 0 0 0 0 1 1 0? 0 0 0 0 1st inst. larv. tuberc. scler. apic. 0=Short, etc. 2 0 2 2 2 2 1 1? 1 1 1 0 1st instar larva saddle form. 0=Divided, etc. 3 0 0 0 1 2 1 0? 0 0 0 0 1st instar larva ground colour. 0=Black, I=Red. 4 0 0 1 1 1 0 0? 0 0 0 1 2 Mature larva colour morph. 0=Di, l=Mono. 5 0 0 0 1 1 1 1? 1 1 1 1 DC Mature larva ground colour: 0=Black, l=Red. 6 0 0 0 0 1 0 0 0 0 0 0 < -J Mature larva saddle presence: 0=Yes, l=No. 7 0 1 0 0 1 0 1 0 0 1 1 Mature larva saddle form: 0=Sm, l=Broad. 8 0 0 0 1 0 1 0 0 0 0 0 Mature larva tubercle apices red: 0=Yes, etc. 9 0 2 0 0 2 2 2 2 1 0 0 Larva lateral prothoracic tubercles long. 10 0 0 0 0 1 0 0 0 0 0 0 Pupa overall colour. 0=Yellow, l=Green. 11 0 1 0 0 0 0 0? 0 0 0 0 2 Pupa melanism: 0=None, l=Some, 2=Much. 12 0 0 0 0 0 1 2? 2 2 1 1 3 Q. Pupa frontal profile: 0=Simple, l=Raised. 13 0 0 0 1 1 1 1? 1 1 1 1 Pupadorsolateralabdominal tubercle form. 14 0 0 0 0 1 0 0? 0 0 0 0 ~~ Male hindwing shape: 0=Short, l=Long, etc. 15 0 0 0 1 1 0 0 0 1 2 2 Male hindwing und. black spots form. 16 0 0 0 2 2 1 1 1 1 3 3 Male hindwing upp. ground colour. 17 0 0 0 1 1 2 2 2 2 2 2 Male hindwing upp. transluscent yellow. 18 0 0 0 0 0 1 1 1 1 1 1 Male hindwing und. androconial brush form. 19 0 0 0 0 0 1 0 1 1 1 1 Male forewing upp. sex-brand form. 20 0 1 1 1 2 0 0 0 0 0 0 Male forwing termen shape: 0=Acute, etc. 21 0 0 0 1 1 0 0 0 0 0 0 Male forewing yellow scaling. 0=None, etc. 22 0 0 0 1 2 1 1 1 2 1 1 Male forewing upperside irrid. markings form. 23 0 1 1 0 0 0 0 0 0 0 0 £j Male forewing underside black spots. 24 0 0 0 0 0 0 1 1 1 2 2 3 Male abdomen und. black stripe. 0=No, etc. 25 0 0 0 0 1 0 0 0 0 0 0 < Male valva distal point. 0=0,1=1,2=2. 26 0 1 1 1 1 2 1 2 2 2 2 Male genitalia form of harpe.0=Short,etc. 27 0 0 1 2 2 2 2 2 2 2 2 Male valva basal spines. 0=2 vestigial, etc. 28 0 0 0 2 1 1 2 2 2 2 2 Female hindwing upperside pale markings. 29 0 0 0 1 1 0 0 0 0 0 0 Female hindwing spots shape and position 30 0 0 0 0 0 1 1 1 1 1 1 Female wing base colour. 0=Brown, etc. 31 0 0 0 0 1 0 0 0 0 0 0 Female forewing vein R3 origin. 32 0 0 0 0 1 0 1 1 0 1 1 Female abdomen hirsute. 0=No, l=Yes. 33 0 0 0 0 0 0 1 1 0 0 0 Female lamella antevaginalis. 0=Keeled, etc. 34 0 0 0 0 1 0 0 0 0 0 0 Female thorax red scaling. 0=Yes, l=No. 35 0 0 0 0 1 0 0 0 0 0 0

EC - Altitudinal Range. 0=0-1200m, 1=>1200 m. 36 0 0 0 0 0 0 1 1 0 0 0 Reappraisal of Troidini 77 define and polarise characters for the Ornithoptera in the data matrix (Table 2). Hancock (1991) used Troides and Trogonoptera to polarise his characters as his earlier studies (e.g. Hancock, 1983), like those of Miller (1986 and 1987), had established Troides as the plesiomorphic congener of Ornithoptera. As I have shown above (also Parsons, 1991: 83), this is untenable based on an entire suite of characters from egg to adult.

Enumeration of Ornithoptera Characters Ornithoptera characters are fully tabulated in the data matrix in Table 2. Apart from selecting those which usefully group taxa into broad categories, I have also included certain important facies by which particular terminal clades (i.e. species) can be clearly recognised. These are, therefore, single autapomorphies which are flagged by PAUP as "uninformative" as they obviously do not assist in the resolution of relationships between taxa. There are eight such characters (Nos. 6, 10, 11, 14, 25, 31, 34 and 35) in the matrix, all of which, except No. 11, apply to 0. victoriae and which, therefore, demonstrate well the distinctness of the species (e.g. Fig. 16). Character 11 (pupa pale apple-green) is a distinct autapomorphy for 0. richmondia alone. 1. Foodplant main genus. 0 = Pararistolochia, 1=Arislolochia. I polarise this character this way because, from the frequency of usage of Pararistolochia as foodplants (e.g. Parsons in press), it appears that this was the foodplant of the Ornithoptera common ancestor (Fig. 15). 2. First instar larva length of sclerotised apices of tubercles. 0=Short, 1= Medium, 2= Long (Fig. IF). Based on the other troidine genera, a shortly sclerotised tubercle apex is the plesiomorphic condition (particularly notable in 0. meridionalis), with a progres sive elongation in other taxa. 3. First instar larva form of saddle. 0= Broadly divided middorsally, 1=Joined middorsally, 2= Absent. Presence of a saddle is plesiomorphic for the subtribe Troiditi to which Ornithoptera belongs, so its complete loss by 0. victoriae is apparently apomorphic. This loss is complete, continuing through to the fifth instar (character 8 below). 4. First instar larva ground colour. 0= Black, l=Red. 5. Mature larva colour morphism. 0=Dimorphic, 1= Monomorphic. 6. Mature larva ground colour. 0=Black, l=Red. 7. Mature larva saddle present. 0= Yes (Figs. 23 and 24), 1=No. 8. Mature larva saddle form. 0= Small or absent, 1= Large and broad. 9. Mature larva tubercle apices orange or red. 0= Yes (Figs. 23 and 24), 1=Some, 2 = Absent. 10. Mature larva lateral prothoracic tubercles long. 0= No (Fig. 24), l=Yes. 11. Pupa overall colour. 0=Yellow and brown, l = Pale apple-green. 78 Michael J. Parsons

12. Pupa extent of melanistic markings. 0= None, l = Minor (Figs. 29 and 30), 2= Extensive. 13. Pupa frontal profile. 0= Simple, l=Raised and 'ornate' (e.g. Fig. 29). 14. Pupa dorsolateral abdominal tubercle form. 0= Acutely pointed (Figs. 29 and 3), 1=Apically rounded. 15. Male hindwing shape. 0= Short, 1= Elongate (Fig. 20), 2=Tailed. 16. Male hindwing underside black spots form. 0= Large and full row (Fig. 20), 1= Reduced, rounded, tornally absent, 2= Lost due to pattern modification, 3= Lost due to termen modification. 17. Male hindwing upperside extent of iridescent green or blue scaling. 0= Predomi nantly uniform green, 1= Modified or reduced by black, 2= Modified or reduced by yellow scaling. 18. Male hindwing upperside extent of translucent yellow scaling. 0= Small and macular, 1= Extensive panels. 19. Male hindwing underside androconial brush. 0=Scales coarse, long, orange or brownish, 1= Scales fine, long, orange or brownish, 2= Scales finer, longer, pure white (Figs. 20 and 21). 20. Male forewing upperside sex-brand presence and position. 0= Absent, 1= Present subterminal, 2 = Present terminal. Based on all other Troidini, which lack a discrete sex-brand on the male forewing, its presence in four species of Ornithoptera must be the apomorphic condition. 21. Male forewing termen shape. 0=Acute, l=Obtuse. 22. Male forewing upperside presence of yellow scaling. 0= Absent, 1= Traces, 2= Obvious subapically, even macular. 23. Male forewing upperside iridescent scaling. 0=Broad bands or panels, especial ly in median region, 1= Reduced to distinctly narrower bands paralleling costa and inner and terminal margins, or even lost at inner and terminal margins. 24. Male forewing underside black spots. 0= Distinct subterminal band, 1= Distinct postmedian band, 2= Lost. 25. Male abdomen presence of black midventral stripe. 0= Absent, 1= Present. 26. Male valva distal margin number of tooth-like processes. 0= None, 1= 1, 2= 2. 27. Male genitalia form of harpe of valva. 0= Short and apically broad, 1= Short and distinctly apically waisted, 2= Long and apically waisted or pointed. I score the progressive elongation of the harpe of the male valva in this way because this appears to be the general trend. 28. Male genitalia form of basal spines of valva. 0= Paired, short and vestigial hooks, 1= Single basal, more elongated, 2= Single or paired, prominent. 29. Female hindwing upperside pale markings divided by broad dark brown regions along veins. 0= No, l=Yes. 30. Female hindwing upperside spot shape and position. 0= Subdeltoid, often Reappraisal of Troidini 79

distally concave and subterminal, 1=Ovate or rounded and postmedian. 31. Female wing basal colouration. 0= Dark brown, l=White or yellow patches. 32. Female forewing vein R3 origin. 0= Inside cell apex, l=Outside cell at junction of base of connate portion of R4 and R5 with discocellular, or arises on it. 33. Female abdomen presence of extensive black hair-like scales. 0= No, l=Yes. 34. Female genitalia: keel of antevaginalis. 0= Present, 1=Absent, instead notched (Darby, 1982). 35. Female thorax presence of red hair scales. 0= Yes, l=No. 36. Altitudinal range. 0= Lowland, mainly 0-1200 m, 1= Montane, mainly above 1200 m.

RESULTS

Enumeration of Troidini Clades Analysis of the 44 character data matrix (Table 1) using Hennig86 yielded a single most parsimonious cladogram (Fig. 12). The same cladogram was obtained using PAUP to analyse the data set, defining the characters as ordered in a 'Branch and Bound' search. Consequently, no further analysis was considered necessary. The various clades for the cladogram (Figs. 12, 13 and 15), and the characters which define their monophyly, are discussed below.

CLADE 1— OUTGROUP Bhutanitis (Zerynthiini) Bhutanitis is plesiomorphic for all characters used in this analysis, being the outgroup used to polarise characters and root the computer-generated trees.

CLADE 2—Troidini + outgroup Papilio (representing tribe Papilionini) Synapomorphies: Character 38, states 1 and 2.—Adult male pseuduncal suture present. Character 39, states 1 and 2.—Adult male discrete hindwing androconial brush at vein 1A+ 2A present. This clade traditionally represents the subfamily Papilioninae (with other papilionine genera included), the integrity or validity ofwhich is not under scrutiny here.

CLADE 3 — OUTGROUP Papilio Synapomorphies: Character 1, state 3.— Primary foodplant genus Rutaceae (etc.). Character 13, state 1.—First to fifth instar larval tubercles gradually degenerate. Character 15, state 2. —Mature larva size ofmost tubercles vestigial or absent. Character 17, state 3.—Mature larva presence of distinctly shorter subdorsal abdominal tubercles. Most reduced or absent. 80 Michael J. Parsons

i 1: 0*1 3: l->0- 5: 0->1 0*1 ,10: 0*1 0*1 ,21: 1*3 0*1 ,34: 0*1 \*V ,36: 0->1 .30: 1*2 ,44: 0->1

HENNIG86 ANALYSIS: mhennig length 106ci 55 ri76 trees I bb file 0 from mhennig I tree bb length 106ci 55 ri 53 trees 1 tplot file 0 from bb 1 tree

PAUPANALYSIS: Tree length = 106 Consistency index (CI) =0.557 Homoplasy index (HI) =0.443 CI excludinguninformativecharacters =0.515 HIexcluding uninformative characters =0.485 Retention index (RI) = 0.530 Rescaledconsistency index (RC) =0.295

Fig, 12. Cladogram hypothesising phylogenetic relationships of the tribe Troidini (Papi lioninae), plus outgroups Bhutanitis (Bhut) and Papilio (Papil). Character changes ("all possible," analysed using MacClade 3.04), and their directions, are shown for each cladc at each branch. Cladc numbers in circles; synapomorphic characters for internal branches, and autapomorphic characters for terminal clades, in bold italics; asterisks alongside state changes denote character reversals. Reappraisal of Troidini 81

Atrophaneura Subgenus Atrophaneura (L) Subgenus Pachliopta (R)

Troides Subgenus Troides (L) Subgenus Trogonoptera(R)

Parides

Battus \\

Ornithoptera

Deciduous hair-like HW androconia in adult male Pharmacophagus

Fig. 13. Pupal relationships of tribe Troidini (Papilioninae), plus plesiomorphic outgroup Zerynthiini (Parnassiinae) based on cladistic analysis in the present study. 82 Michael J. Parsons

(3 -o

9 CO <« (3 o O .(0 • E CO Z •HS0 a := o w co < co 0) CO O (0 I* E < (0 co a 3 co 3« rE . t lil (0 < S

Fig. 14. Area cladogram for the tribe Troidini (Papilionidae) based on results of the present cladistic analysis.

Character 20, state 1. — Larva at most instars without red spots or patches. Character 35, state 2. — Female sphragis absent. As other studies have shown, Papilio (sensustricto) (Papilionini) is the apomorphic sister group to the Troidini. Its position in the cladogram generated from my data matrix also suggests this (e.g. Figs. 12, 13 and 15). Note that Fig. 15 shows Papilio branching to the extreme right as PAUP draws the tree in its "standard" (default) mode in the "Tree-Order" Options. With the addition and correct scoring of other papilionine genera (notably Chilasa) to the data matrix their fundamental relationship with Troidini may yet change.

CLADE 4—Tribe Troidini Synapomorphies: Character 37, state 1.—Female foretibia and foretarsus spines in a tightly spaced row, running length of tibia and tarsus, sensilla evenly spaced between these. Character 40, states 1 or 2. — Adult male hindwing (pouch) androconial scales present. Although Miller (1987b) stated that "Understanding relationships within the Troidini has proved to be an especially difficult problem", he maintained that the monophyly of the tribe is quite certain based on several known autapomorphic adult structures. However, three of these he rightly regarded as homoplasious, implying that they are difficult to interpret. Of the other four characters Miller selected as synapomorphies defining Troidini, apparently the best was his adult character 100 ("Female with a row of closely spaced spines running the length of the [fore] tibia."). Reappraisal of Troidini 83

Distribution of Troidini on Gondwanan plates

Madagascan plate

Indian (and/or S.E. Asian plate

Australian plate

S. American plate

Subgenera

Genera

PERIOD OF INIT IAL GONDWANAN FRAGMENTATION

Subtribes

Tribes (Subfamilies)

Aristolochiaceae-feeding common ancestor

Fig. 15. Summary of concepts of the phylogenetic and systematic relationships, foodplant relations, and Gondwanan origins, of the tribe Troidini (Papilionidae) based on results of the present cladistic analysis of the tribe.

According to Miller, these are relatively few and are located on only the first and second larsomercs in [all?] other papilionids, with sensilla concentrated on the first tarsomere, and occasionally the second. Miller noted that, in the Troidini. tin- spines are more numerous and are arranged in a light row running the length of the tarsus, with sensilla evenly spaced between them. Even the immature stage characters apparently do not provide synapomorphies that unequivocally characterise the monophyly of Troidini. For example, characters 84 Michael J. Parsons of the first instar larval tubercles (notably the posteriorally-directed basal seta and discretely sclerotised apex) would provide excellent support of Troidini as a monophyletri group if Battus did not possess the plesiomorphic zerynthiine-like simple unisetose tubercle form in its first larval instar. Likewise, the prominent and naked (fleshy) tubercles of the fifth instar troidine larva would be an excellent synapomor phy defining the monophyly of the tribe if it were not for Euryades which possesses the plesiomorphic zerynthiine-like character of prominently spinose tubercles right through to its final larval instar (the spinose setae not being lost as in all other troidines). Nevertheless, such homoplasy is not great and, when considered in combination, these immature stage characters (like those of the adult mentioned above) do endorse the monophyly of Troidini. Prominent naked tubercles was a character numbered 103 by Miller, but was poorly defined by him and was additionally coupled with the separate character state of the dorsolateral pupal tubercles on abdominal segments 4-7. However, both larval and pupal tubercles are homologous with similar tubercles in Zerynthiini and Papilionini and are also categorisable as exhibiting a number of states within the Troidini. In most troidines, the later instar larvae bear a distinctive median saddle mark, although this can be entirely lost in certain species. Pupae provide the some of the best characters indicating the overall intratribal relationships of the troidines, and the level of distinctness of each genus (Fig. 13). The pupae of Battus are generally similar to those of Parides. Parides pupae resemble those of Troides in their general form, colouration and pattern, particularly their saddle markings. Atrophaneura pupae (e.g. Fig. 34) fall into the general form of Battus/Parides/Troides grouping (e.g. Figs. 32 and 33), but their colouration and often highly ornate flange-like extensions are specialised modifications which obscure this relationship. The pupae of Euryades, Cressida (Fig. 31), Pharmacophagus (Figs. 27 and 28) and Ornithoptera (Figs. 29 and 30) resemble one another in overall form and, apart from their increasingly larger size, are morphologically similar to the Luehdorfia (Zerynthiini) pupa: notably Euryades and Cressida. The pupa of Pharmacophagus resembles that of Ornithoptera as is clear from a comparison of Figs. 27/28 with Figs. 29/30. The early divergence of Euryades and Cressida from the remaining troidine genera has been determined by most previous studies using adult characters. My character set apparently corroborates this as the cladogram shows first Euryades, then Cressida, diverging (Figs. 12, 13 and 15). However, apart from characters 35 (sphragis large and alate) and 38 (male pseuduncal suture complete) few synapomorphies define their relationship as monophyletic. Using only adult characters in analyses, other authors such as Hancock (1983) and Miller (1986, 1987b), have assumed a closer relationship between Euryades and Cressida than my cladogram suggests. However, even Miller (1987b) found that some of the adult characters suggesting a "close" relationship between the two genera were weak, or had been misinterpreted by Reappraisal of Troidini 85 previous authors. Their pupal characters show only superficial similarities. In fact, Euryades and Cressida are each distinctive genera on the basis of their larval and pupal characters. Their poorly definable relationship is perhaps to be expected, considering the length of time over which they have been separate (assuming, as most authors now do, that their common ancestor was distributed in the palaeocontinent of Gondwanaland and that their separation commenced during its fragmentation between 60- 80 million years ago). Based on the two above mentioned adult characters, and using traditional evolutionary taxonomic methodology, there is reason to regard Euryades and Cressida as belonging in their own tribe: Cressidini. This is possible because the large and alate apomorphic state of the sphragis (character 35) appears to be homologous in these taxa. Alate, spatulately-paired sphragides are unique to Euryades and Cressida and do not occur in any other butterflies. A phylogenetic hypothesis that treats Euryades+ Cressida as a monophyle tic clade is easily supported from my data matrix in re-analysis using Hennig86 and PAUP 3.1.1 by weighting character 35. However, on present evidence, Troidini appears to represent a single tribe comprising three subtribes: Euryaditi, Cressiditi and Troiditi (Fig. 15).

CLADE 4a — Genus Euryades Euryades is poorly characterisable, the genus lacking any really distinctive autapomorphic characters, although in overall shape and colouring its pupa is reasonably distinct (Fig. 13, and photographic plate 20B in Tyler el al., 1994). For example, one notable apomorphy (not incorporated into the data matrix used for analysis), when present (in green morph pupae) is the broad irregular mauve middorsal band that runs across the methathoracic segment and abdominal segments 1-3 (general position of the saddle in other troidine pupae). The simple angular front (i.e. frontal profile) is similar to those present in the pupae of some Battus and Parides species. Some characters, such as the zerynthiine-like retention of spinose setae on the body tubercles throughout all larval instars, implies that Euryades is relatively more plesiomorphic than Cressida, as reflected in the cladogram hypothesis ing their phylogenetic relationships (Figs. 12, 13 and 15). In addition, although not scored in the data matrix, Euryades is the only troidine possessing a distinctly parnassiinc/zerynthiine-like hindwing, in which the ground colour is predominantly yellow, the red spots being distinctly macular and arranged in two well separated (subterminal and median) rows. This is apparently a plesiomorphic trait that is also retained by many members of the sister tribe of Troidini, the Papilionini. Miller (1987b: 492) pointed out, and Tyler et al. (1994: 73) photographically illustrated, that the sensory maxillary palpi of Euryades and Cressida first instar larvae bear multiple conical sensilla in much greater numbers than in the other Troidini for which the details of this character are known. For example, Battus and Parides possess between 10-20 sensilla, the plesiomorphic condition, according to Tyler et 86 Michael J. Parsons al., being 8 sensilla (e.g. in Zerynthiini and Papilionini). However, whereas Euryades possesses over 40 sensilla, Cressida bears over 250 sensilla, and the broad and flat condition of the palpus (certainly in Cressida, but not in Euryades) is hardly comparable. Therefore, I find it difficult to accept this as indicating any relation ship, and the maxillary palpi of Cressida certainly appear to be extremely apomorphic relative to the plesiomorphic condition of this character in Papilionidae in general. Although I have not incorporated it into my data matrix, the configuration of over 40 sensilla in the maxillary palpi of the Euryades first instar larva certainly represents a further distinctive autapomorphy for the genus.

CLADE 4b — Stem to Troidini minus Euryades Synapomorphy: Character 15, state 1. — Mature larva tubercle very prominent. Prominent larval tubercles define the monophyly of this clade. Except for Parides (and Euryades before this branch), character 12 (first to fifth instar larvae spinosity of tubercles entirely lost by second instar) is common to all troidine genera, otherwise it would also be a definitive synapomorphy at this branch in the cladogram.

CLADE 4c—Genus Cressida (Monotypic) Autapomorphy: Character 30, state 1.— Pupa overall colour and main markings pure white marked with maroon red, brownish-mauve and bright yellow. The Cressida pupa (Fig. 31) is particularly distinctive in shape, colouration and markings. The specific form ofthe Cressida first larval instar body subdorsal tubercle apices with only four setae, subtended by a notably small rounded chidnous tubercle apex: Fig. IB) is also an autapomorphy that defines the genus, but this was not scored as such in the data matrix. No other Troidini have this specific configuration of chitinised tubercle apices. Although I have not incorporated it into my data matrix, as mentioned under Euryades above, the configuration of over 250 sensilla in the maxillary palpi of the Cressida first instar larva represents yet another distinctive autapomorphy for the genus.

CLADE 4d — Subtribe Troiditi (6 genera) No unequivocal synapomorphies within my data matrix define the monophyly of this clade. The simple plug-like plesiomorphic state of the sphragis is the only character which is common to all troidine genera at this branch (the sphragides of Euryades and Cressida being large and alate). Reappraisal of Troidini 87

CLADE 4e — Pharmacophagus + Ornithoptera Synapomorphy: Character 23, state 1. — Pupa overall shape in lateral profile elongate, strongly S- shaped, abdomen and 'ventral margin' prominently outwardly curved. As discussed above under the description of character 23, the strongly S-shaped pupal lateral profile is a synapomorphy of clade 4e, and is also representative of a number of other apparently important synapomorphies, all of which endorse the monophyly of Pharmacophagus plus Ornithoptera. In this sense, character 23 can be interpreted as encompassing a suite of several subtle characters that are apparently autapomorphic at this branch of the clade. As these are difficult to clearly define they have, however, been omitted as separate characters in the data matrix.

CLADE 4f— Genus Pharmacophagus (Monotypic) Autapomorphies: Character 17, state 1.—Mature larva presence of distinctly shorter subdorsal abdo minal tubercles on segments 3 + 4. Character 26, state 1.—Pupa dorsolateral abdominal tubercle apices bluntly trun cate, deltoid and irregularly nodular (Figs. 9 and 10). Character 27, state 1.— Pupa abdominal segment 4 minimally laterally flanged. Pharmacophagus is well defined by the autapomorphies listed above. The overall pupal morphology (Figs. 27 and 28) clearly resembles that of Ornithoptera (Figs. 29 and 30). The strongly S-shaped lateral profile of the Pharmacophagus pupa (Fig. 28) most closely resembles that of 0. paradisea (Fig. 30). In addition, another apomor phy for the genus (not incorporated into the data matrix) is the shape of all subdorsal tubercles in the mature larva. These are distinctly club-like, being medially distinctly narrower than at the apex (Figs. 6 and 22). Although Cuban Battus devilliers (Godart, 1823) also possesses prominently clubbed subdorsal tubercles (colour photographic plate in Tyler et al., 1994: pi. 31a), this is a specialisation at the species (rather than genus) level, as all other known Battus larvae possess tubercles with the plesiomorphic tapered condition.

CLADE 4g—Genus Ornithoptera (10 species) Autapomorphies: Character 1, state 1.— Primary foodplant genus Pararistolochia. Character 5, state 1.— Ovum size extremely large 2.8-4.8 mm diameter. Character 10, state 1.—First instar larva body subdorsal tubercule sclerotised apex elongate. Character 21, state 3. — Larval abdominal saddle on segment 4 only. Character 34, state 1.— Main pupation site leaves. Character 39, state 2.—Adult male discrete hindwing androconial brush at vein 1A+ 2A coarse and well developed. 88 Michael J. Parsons

Ornithoptera is well defined by the distinct autapomorphies listed above and as discussed in detail in the section above comparing the generic distinctness of Ornithoptera with Troides. For example, two other apomorphies for the genus (not incorporated into the data matrix) are the shape of all subdorsal tubercles in the mature larva which are so finely tapered as to be thread-like (especially obvious in instars 1-4), and pupal abdominal segment three which bears simple lateral 'eye- spots.'

CLADE 4i —Genus Battus (12 species) Autapomorphy: Character 14, state 1.—Fourth and fifth instar larvae length of lateral prothoracic tubercles compared to other body tubercles 3-4 times longer, resembling 'pseudo- antennae.' As mentioned above, the several plesiomorphic characters of Battus, such as eggs laid in batches, the simple, unisetose form of the first instar larval subdorsal tubercles, entirely saddle-less condition of all larval instars, as well as the adult characters of ventral antennal sensilla in simple patches (rather than in paired pits), and three-segmented labial palpi, all suggest that, on traditional taxonomic grounds, Battus warrants distinction higher than that of a genus. Indeed, by using only adult characters in analyses, other authors such as Hancock (1983) and Miller (1986, 1987b), have found that Battus diverges early enough from the remaining troidine genera to warrant subtribal status: Battiti. Nevertheless, the results of my analysis suggest that, despite its character reversals, Battus clearly belongs within the subtribe Troiditi (Fig. 15). This is especially clear, for example, from the pupal morphology (Fig. 13). More importantly, if my polarisation of characters and resultant phy logenetic hypothesis (cladogram) is correct, then my data has highlighted an ap parent case of atavism in the Troidini. This was unexpected, but is very interesting since the evidence for this is strong in the number of characters involved, their distinctness, and the fact that they occur throughout a range of stages in Battus. This raises the possibility that the apparent character reversals in Battus, although not as impressive as the presence of a moth-like frenulum and retinaculum wing- coupling device present in the Australian endemic pyrgine hesperiid Euschemon raffiesia (W.S. Macleay, 1826), may be equally important in terms of comprehension of the genetic evolution of Battus.

CLADE 4j — 'Higher troidines' Synapomorphies: Character 21, state 2.—Larval saddle form on abdominal segments 3 & 4 as a distinct diagonal lateral line. Character 40, state 2. —Adult male hindwing (pouch) androconial scales fine, hair like and deciduous. Reappraisalof Troidini 89

The two autapomorphic characters defining the monophyly of this clade are distinct and apparently reliable. Whereas character 40 (Miller's, 1987b: 424, character 143 "Male hind wing margin with deciduous scales") was found to be homoplasious by Miller, it becomes a shared homologue (and, therefore, a true synapomorphy) at node 4j in the cladogram representing my hypothesis of Troidini phylogeny. Similarly, Miller (1987b: 422) found that his character 131 ("male hind tibia swollen, covered with a mat of short spines") was homoplasious in his cladogram: as he stated "the least congruent with the phylogenetic hypothesis proposed for the Troidini and deserves further investigation." Miller noted that his character 131, although somewhat variable and sporadic in occurrence, is nevertheless present in many Pachliopta, some Atrophaneura, some Troides, and a few Parides (sensu stricto). Again, according to the results of my analysis, this character becomes a shared homologue and thus another valid synapomorphy for my clade 4j, adding still further support to the hypothesis that Parides+ Troides (plus Trogonoptera) -{-Atrophaneura (plus Pachliopta) is a true monophylum (to the exclusion of the other troidine genera). It is apparent from my analysis (Figs. 12 and 15) that Parides and Troides share a reasonably close and important relationship. For example, as Miller (1987b) noted, the adult male hindwing androconial apparatus (my character 40, his character 143, as outlined above) is exactly similar in both genera. Miller, apparently unaware of my published observations on Troides courtship, and the function of the deciduous male hindwing androconia (Parsons, 1983), provided an exactly similar account of courtship in Parides based on observations by P. Feeny of one species. This included the fact that, as in Troides, the Parides male androconial scales remain physically attached to the female's antennae after the looping courtship flight of the male. One group of predominantly black Parides species with yellow hindwing panels and dense red prothoracic scaling (the "pronotal collar" of some authors, as noted above) are effectively small neotropical 'versions' of Troides (e.g. plate 56 of Tyler etal., 1994). The relationship is also particularly obvious from the immature stages of Parides and Troides, most notably the pupal morphology of the two genera (Fig. 13).

CLADE 4k — Genus Parides (34 species) No unequivocal autapomorphies within my data matrix define the monophyly of Parides as a terminal clade. Miller (1987b: 428) found only two adult characters to define his Parides subgenus Parides (i.e. Parides sensu stricto) noting that the unique adult feature is the dorsoventral orientation of the signum. As determined by character 12, state 1, the subdorsal tubercles of the larval instars 2 and 3 retain their apical setae (i.e. a character reversal). Even the chitinous tubercle apices can be retained up to the second instar on all, or certain, tubercles in some Parides species. This latter character is not scored in my data matrix, but is an autapomorphy for the 90 Michael J. Parsons genus.

CLADE 41 — Troides plus Atrophaneura No unequivocal synapomorphies withinmy data matrix define the monophyly of this clade. Nevertheless, the Troides and Atrophaneura pupae (Fig. 15) are the 'squattest' and broadest of all the troidine genera, so a pupal length to width ratio (not incorporated into the data matrix used for analysis) could be used as a synapomorphy at this branch in my cladogram. The character might, for example, be scored as an apomorphy, state 2, for character 22 in the present data matrix.

CLADE 4m — Genus Troides (32 species) Synapomorphy: Character 24, state 1.—Pupa primary dorsolateral abdominal tubercles present on segments 5 + 6. The Troides pupa is distinctive in overall shape and exhibits a number of subtle synapomorphic characters that have not been incorporated into the present data matrix. The large size and broad triangular shape of the primary dorsolateral abdominal tubercles on segments 5+ 6 provide the most obvious synapomorphic pupal character defining the monophyly of Troides.

CLADE 4n — Subgenus Troides (30 species) No unequivocal autapomorphies within my character set define the monophyly of this terminal clade. Nevertheless, adult characters such as absence of red wing scaling (character 44) and overall similarity and uniformity of wing colour and markings might be used to provide further autapomorphies to better define the subgenus Troides.

CLADE 4o — Subgenus Trogonoptera (2 species) Autapomorphies: Character 17, state 2.—Mature larva presence of distinctly shorter subdorsal abdo minal tubercles on segments 2 + 3 and 5 + 6. Character 18, state 1.—Mature larva angle of subdorsal abdominal tubercles in relation to body fully recumbent (ca. 180°) and posteriorally-directed (especially those on segments 2-7 which lie almost appressed to the body). This terminal clade is well defined by the two autapomorphies listed above. In addition, the Trogonoptera first instar larval subdorsal tubercles are somewhat intermediate to those of Troides and Ornithoptera in general form, having a rather Ornithoptera-Mke arrangement of setae but differing from all other Troidini in that they possess no clear distinction between the chitinised apex and the fully membranous base. Reappraisal of Troidini 91

CLADE 4p — Genus Atrophaneura (43 species) Synapomorphies: Character 1, state 2. — Primary foodplant genus Thottea. Character 27, state 2. —Pupa abdominal segment 4 distinctly laterally flanged by semispatulate or spatulate extensions. This clade is well defined by the two synapomorphies listed above. Others defining this clade could easily be established from the pupal shape, colour and markings which are unique among the Troidini (Figs. 13 and 31), and indeed for the entire family Papilionidae. For example, the horizontally 'zebra-striped' modifica tion of the saddle markings (not incorporated into the data matrix used for analysis) could be used as a synapomorphy at this branch of the present cladogram.

CLADE 4q — Subgenus Atrophaneura (17 species) No unequivocal autapomorphies within my character set define the monophyly of this clade. The most consistent and distinctive character (not incorporated into the data matrix used for analysis) that apparently defines the subgenus Pachliopta (in comparison with subgenus Atrophaneura) is the more ornate, laterally flanged front of the pupa, that of Atrophaneura being much simpler (Figs. 13 and 31).

CLADE 4r— Subgenus Pachliopta (17 species) Apart from the above mentioned pupal front autapomorphy, which defines the taxa Pachliopta and Atrophaneura as two separate terminal clades with value as subgenera, no other reliable autapomorphies within my character set define the monophyly of this clade. Nevertheless, as mentioned under character 40 above, subgenus Pachliopta is characterised in the majority of its members by secondary modification of its hindwing androconial scales.

Enumeration of Ornithoptera Clades In an initial analysis of the data, the matrix was run unordered and the cladograms rooted using a hypothetical ancestor, based on character states of outgroups, its character states all plesiomorphic "0" (Table 2). This yielded a single cladogram of tree length 69 (Fig. 16). Not represented in the cladogram in Fig. 16 is the possible importance of the male forewing sex-brand (character 20), a clearly defined sexual feature of richmondia, priamus, alexandrae and victoriae, which is not only absent in all other Ornithoptera, but indeed all other members (i.e. therefore genera) of the entire tribe Troidini (Fig. 12). Although, based on the results of the initial analysis of the unweighted data set, an hypothesis of richmondia + priamus + alexandrae + victoriae as representing a natural group is not parsimonious, and despite the fact that several characters (5, 13, 17, 22, 28: Table 2) suggest that such a clade might not be valid, it is nevertheless another plausible evolutionary scenario for the group. Thus, in order to acertain other potential interspecific relationships of the 'lower clades' 92 Michael J. Parsons

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PAUP ANALYSIS Tree length = 69 Consistency index (CT)= 0.725 richmondia-like Homoplasy index (HI) = 0.27S CI excluding uninformative characters= 0.689 common ancestor HI excluding uninformative characters = 0.311 Retention index (RI) = 0.740 Rescaled consistency index (RC) = 0.S36

Fig. 16. Initially generated single cladogram hypothesising phylogeny of species of the genus Ornithopterabased on cladistic analysis, using Hennig86 and PAUP, of 36 larval, pupal and adult characters. Character changes ("all possible," analysed using Mac Clade 3.04), and their directions, are shown for each clade at each branch. Cladc numbers in circles; synapomorphic characters for internal branches, and autapomorphic characters for terminal cladcs, in bold italics; asterisks alongside state changes denote character reversals; bullets denote species particularly restricted in geographical range. Reappraisal of Troidini 93

CD .JO

PAUP ANALYSIS Tree length = 83 Consistency index (CI) =0.723 Komoplasy index (HI) = 0.277 CI excluding uninformative characters= 0.693 HI excluding uninformative characters=0.307 Retention index (RI) = 0.723 Rescaled consistency index (RC) = 0.523 B

Fig. 17. Cladograms hypothesising phylogeny of species of the genus Ornithopterabased on cladistic analysis, using Hennig86 and PAUP, of 36 larval, pupal and adult characters when weighting character 20: male sex-brand (otherwise all characters unordered and unweighted). A) The first of these cladograms accepted as being another plausible evolutionary scenario to that presented in the initially generated single cladogram. B). The second of these cladograms rejected here as not adequately defining the 'lower cladcs' (richmondia,priamus, alexandraeand victoriae).

(richmondia+ priamus+ alexandrae+ victoriae),the data were re-run, forcing these four species to be a natural group. This was achieved by simply weighting character 20 (presence of male sex-brand) in PAUP. In re-running the data set the final weight used was 6, the minimum number of cladograms (two) remaining constant (from three) at this weight for character 20, as well as after the weight was increased. The two cladograms generated were of longer tree length: 83 as opposed to 69 in the first cladogram (Fig. 16). The first (Fig. 17A) of the subsequent two cladograms obtained by weighting the sex-brand character apparently best fits the available evidence of the relationships of richmondia, priamus,alexandraeand victoriae,because 0. victoriaepossesses numerous apomorphic characters, whereas the immature stages and adults of alexandraeshare many more affinities with those of priamus. Consequently, together with the initially generated cladogram in Fig. 16, I accept the cladogram in Fig. 17A (over that in Fig. 17B), as another hypothesis of the phylogeny of the Ornithopteraworthy of serious considera tion. Additional searching would likely provide other characters (possibly of the female genitalia) to further support the Fig. 17A hypothesis. It differs little from Hancock's (1991) cladogram hypothesising Ornithopteraphylogeny. However, Hancock's methodology (which he omitted to describe) apparently did not involve use of computer analysis. Furthermore, he did not present his data for analysis as a complete matrix, so it is not clear how he scored characters across all taxa (the values 94 Michael J. Parsons

of which affect the overall positions of clades, depending on how they are scored). One of the main differences between Hancock's (1991) concept of the Ornitho ptera, and my own, was his treatment of the taxon priamus as comprising four species, whereas I do not consider that the data presently available shows that the three additional taxa (two of which are insular in origin), are more than just well defined subspecies of priamus. Various lepidopterists have long debated whether priamus represents one or more species. Like numerous other butterflies of the Indo- Australian Region, the taxon is perhaps best regarded as a 'superspecies' on the 'verge' (in the geological timescale) of speciating into several distinct species (notably the taxa croesus, aesacus and urvillianus) in several of its outermost, more isolated island locations. However, the monomorphic green pupa of 0. richmondia (unique for the genus), the unmarked (saddleless) dark brown larva, and the sterility of Fi hybrids from artificial flight-cage hybridizations between priamus and richmondia (Common and Waterhouse, 1981), provide good reasons for considering richmondia to be distinct from priamus, as has been suggested by certain other authors, notably Common and Waterhouse (1981). Furthermore, as Zeuner (1943: 138) pointed out, the male genitalia of richmondia, especially in the shape of the harpe (plesiomorphically rounded and unspecialised), are notably different from all other priamus-\\kc taxa. Zeuner considered, therefore, that these genitalic differences indicated an early separation of richmondia from the priamus stock. Otherwise the adults of richmondia differ very little phenotypically from those of priamus (Table 2). In the case of priamus, its large distribution, and oligophagy within the Aristolochiaceae (in marked contrast to its more foodplant specialised and more geographically restricted relatives), are indica tive of the long period of time (compared to its relatives) that priamus must have required to become oligophagous and widely distributed. The fact that richmondia and priamus are, from the results of my analysis, the most plesiomorphic of the Ornithoptera (e.g. Fig. 16) is important as this is congruent with my concepts of the Gondwanan vicariance biogeography (e.g. Nelson and Platnick, 1980 and 1981) of the genus.

DISCUSSION

Troidine Evolution and Gondwanan Palaeobiogeography A diagrammatic summation of my interpretation of the phylogenetic relation ships of the Troidini, including their foodplant relations and vicariance biogeography (e.g. Nelson and Platnick, 1980, 1981), based on the analysis of my data set as it presently stands, is provided in Fig. 15. I assume from evidence of the present-day distributions of the Troidini (Figs. 14 and 18), in relation to the known events of plate tectonics, that Troides (i.e. plus Trogonoptera) and Atrophaneura (i.e. plus Pachliopta) were isolated together on the Indian and / or South-east Asian plate (the latter postulated by Ridd, 1971) during, or at least relatively soon after, the main period of Reappraisal of Troidini 95 fragmentation and commencement of drift of the plates comprising Gondwanaland. If a South-east Asian plate did once form part of Gondwanaland as Ridd convincing ly argued, then this may explain the present-day geographical position of the subgenus Trogonoptera (of Troides) in Sumatra, and Palawan. Similarly, it appears that Ornithoptera and Cressida were isolated and drifted on the Australian plate. The results of my cladistic analysis of the important papilionid tribe Troidini differ markedly in key areas from those of Miller (1987b), as Miller's did from those of Hancock (1983). For example, Miller concluded that genus Parides comprises subgenera Parides, Atrophaneura and Panosmia, and that Pachliopta represents a distinct genus. Moreover, Miller's hypothesis of troidine phylogenetic relation ships (and as depicted in his figure 186 area cladogram for the Troidini) does not suggest a picture of troidine vicariance biogeography that is congruent with that suggested by evidence of their present distributions. Other important differences between my results and thoseof Miller and Hancock are that Euryades and Cressida are not as closely related as adult characters alone would suggest, and that Battus, although possessing a number of apparently plesiomorphic characters (possibly, as suggested above, through atavistic recurrence of these), is otherwise typically troidine, and belongs in the subtribe Troiditi. From my present analysis, Battus becomes the sister group to Parides+ Troides+Atrophaneura, as opposed to belonging in its own separate subtribe, Battiti, as Miller and Hancock both concluded. In spiteof the use ofstrict cladistic principles in the phylogenetic analysis of the Troidini, it is clear that paucity ofdata, coupled with the misinterpretation ofknown characters (see Hauser, 1993), caused Ornithoptera to be erroneously lumped with Troides. Miller (1987b) subsumed Ornithoptera intosynonymy with Troides, not even according the taxon subgeneric recognition. I maintain that such arrangements are untenable as is particularly obvious from pupal characters, but also from a suite of others, including those of adults. My findings confirm that Ornithoptera is a valid taxon at the generic level, and moreover suggest that the true relationship of Ornithoptera lies closer to Pharmacophagus than Troides (sensu stricto). Furthermore, the oriental taxon Atrophaneura, minus neotropical Parides, is re-established as a monophyletic unit, whereas it became paraphyletic in Miller's (1986, 1987b) study. The close relationship of Parides and Troides noted by Miller (1987b: 424) is, however, endorsed by my results and further highlights the biogeographical import ance of the Parides/Troides and Pharmacophagus/Ornithoptera relationships in the context of vicariance biogeography. Miller (1987a) (and subsequently Brown et al., 1995, and Weintraub, 1995) argued that foodplant shifts at lower taxonomic levels in the Troidini fail to support the hypothesis of parallel cladogenesis (i.e. true stepwise coevolution) of the butterflies and their foodplants. This, despite suggestions in the early literature (particularly by Ehrlich and Raven, 1965) that this tribe of swallowtail butterflies 96 Michael J. Parsons provides a notable and, therefore, classic example of the general concept of the coevolution of insects and their foodplants. Berenbaum (1995) pointed out that Miller's work (notably 1987a) is "widely cited as the death knell for the notion of any form of coevolution between swallowtails and their hosts; yet that particular study dismisses parallel cladogenesis of swallowtails and hosts based on a phylogene tic study of only a small section of the family." I would further argue that this "falling out of favor" (as Berenbaum aptly put it) of the strict coevolutionary theory is hardly surprising for two main reasons: 1) that the taxonomic (and, therefore, by default, the phylogenetic) resolution of 'real' (i.e. valid and evolutionarily informa tive) taxa in both the Troidini and Aristolochiaceae has been extremely poor (and still requires much research and further refinement), and 2) that, given the great length of time that the relationship has been in progress, it can be expected that the overall 'picture' of relationships (which was presumably once more clearly defined) has become 'variously blurred.' By further clarifying knowledge of the taxonomy and phylogenetics of the Troidini and Aristolochiaceae, existing views of this picture can doubtless be greatly enhanced. Most authors do agree, however, that there are key compatibilities observable between the phylogenies of the Troidini and Aristolochiaceae as they are presently perceived. As Feeny (1995) stated "At the very least and regardless of the polarity of host shifts, however, it is possible that a majority of swallowtails has not strayed far from a single major phyletic lineage of plants." Exact congruences of cladograms (e.g. Page, 1994), may never be completely attained, yet the evidence (especially including that of the intimate chemical relations between these butterflies and their foodplants) does, I contend, indeed point to their 'coevolution.' Whether or not it has been one of a true 'chemical weapons-based arms race' (in the sense that the butterflies adapt, at each step, to the 's attempts to develop chemical defenses against them) is highly uncertain, but studies such as those of Feeny (1975, 1995) may eventually shed better light on this. In addition, as Weintraub (1995) rightly pointed out, many of the early literature records have 'muddied' the pool of data on troidine foodplants, many incorrect foodplants being recorded because they were based on erroneous determinations. My findings, like those of Weintraub, have shown that troidine foodplant relations are indeed far more restricted (to Aristolochiaceae) than previous data have suggested. Based on the characters of Papilio (Table 1), the Papilionini have retained a number of obviously plesiomorphic characters, whereas the Troidini have equally specialised in many respects, notably in adult colour and pattern. This is a result of the latter tribe having remained in close association with Aristolochiaceae, and so having more fully and universally developed mimetic assemblages (etc.). Where only the adult characters have been studied, it is mainly this 'homogenisation' and general convergence in adult phenotypes through mimicry that has led to problems in the systematics and classification of the Troidini, their relationships being obscured Rea| )oraisai oi Vnidini 97 in this wav. Although not due to mimicry, the large size of the adults of both Ornithoptera and Troides. and their superficially similar adult facies. has been the primary stimulus for taxonomists to regard these taxa as being congeneric. Otherwise there are only relatively few characters to suggest that the relationship between the two genera is as elose as has been supposed, whereas many more characters suggest otherwise. A similar, but converse, argument was used by Miller (1987b) to point out that superficial differences in adult morphology "including dramatic differences in size and pigmentation patterns, all of which are autapomorphic", has been an important reason why previous authors had overlooked the close (likely sister-generic) relationship of Troides and Parides (sensu Miller, 1987b).

Fig. 18. World distribution ofgenera of the tribe Troidini (Papilioninae).

Ornithoptera Origin and Biogeography: A New Hypothesis

"It is perhaps the most surprising feature in the distribution of Troides and its allies that Troides proper (except 7". oblongomaculalus) has not been able to advance father east than Ilalmaheira, Obi and Ceram, whilst the eastern group of Ornithoptera, though evidently established on Ilalmaheira and Obi since the late Tertiary, has only reached Ceram, although the present configuration of the island groups would afford means of dispersal for strong-Hying insects such as the birdwing butterflies. This fact appears in an even stranger light if one recalls that the second group of phylogcnetically old forms of Ornithoptera is confined to the Solomon Isles and North-east New Guinea, whilst New Guinea itself is the native country of the Schoenbergia-groups." (F. E. Zeuner, 1943: 167).

Ornithoptera and Troides were the subject of a classic study by Zeuner (1943) (reviewed by Corbet, 1944, which he aptly termed "paleontology without fossils"). 98 Michael J. Parsons

I include above the quote from Zeuner's work as it highlights, and exactly pertains to, the discussion that follows. In addressing the questions raised by his particularly perceptive statement, any responses should, to reflect any credibility, fit the available biogeographical evidence as closely as possible. Zeuner hypothesised that the evolution and speciation of all the birdwing groups took place in successive waves from an origin in Sundaland. Racheli (1980) concurred with this concept, but Holloway (1973) considered that their origin was centred on the Asian mainland. Toxopeus (1950) postulated an Aru-Merauke ridge for dispersal of 0. priamus into Australia. Zeuner concluded that the Ornithoptera probably evolved during the Pleistocene (2 million years before present — mybp), and that 0.5-1 million years were required for the specific characters to become stabilised for some species. He placed particular emphasis on past fluctuations in sea level during each glacial period, which successively connected Australia to New Guinea (and the Aru Islands), via what is now the Torres Strait. The theory as he described it:

"each time the sea levelwas low, the low country was populated with recently developed forms from the islands. This process has been continuing at least since the mid Tertiary, and the extraordinary variety of closely-allied species, species-groups, and genera is the result. The regional character of the fauna was intensified by the barriers afforded by the sea to the west, south-cast, and by high mountains to the north and north-west."

As priamus is the most widespread and oligophagous Ornithoptera species, it is reasonable to assume that, together with closely related endemic Australian sister species, richmondia, these represent the relatively most primitive lineage of their genus. My reasons for assuming that richmondia is the extant representative closest to ancestral Ornithoptera are, in part, stated above. The suite of apparently plesiomor phic features possessed by richmondia, together with its small adult size (assuming the ancestral condition was as such), as well as the immediate geographical proximity of closely related priamus (and no other species), strongly suggest that the Ornithoptera originated in Australia. Data for Ornithoptera larval foodplant specialisations not only corroborates this, but also suggests that the role of continental drift must be carefully considered in the evolution of the Australian subregion Troidini. Tropical rainforest lianas of the genus Pararistolochia are the primary larval foodplants of Ornithoptera. The aristolochiaceous genus was considered to be restricted to the Afrotropical Region, but was recognised by Parsons (in press) to also occur in the Australian subregion, being represented by at least 20 species throughout New Guinea and Australia. Moreover, because the Australian subre gion members of Pararistolochia had been taxonomically 'buried' in the literature within Aristolochia, the biogeographical significance of this went entirely unrecognised (a case ironically similar, albeit coincidental, to that of Ornithoptera). The existence of Pararistolochia in two remotely separate continental faunal regions (Fig. 19) is clearly biogeographically important in its own right, as well as from the aspect of Reappraisal ol 1 99

Fig. 19. World distribution of genera of Aristolochiaceae, larval foodplants of the troidine swallowtails. Note the distribution of Pararistolochia, primary foodplants of Omithopterc in the Australian subregion. their relationship with the Ornithoptera as their foodplants. Now, therefore, a far more interesting picture of the Aristolochiaceae has emerged which exhibits notable parallels with some of the distinctive Troidini distributions (Fig. 18). For example, the occurrences of related troidines Euryades and Cressida are also widely separated into two of the world's major faunal regions: respectively, the Neotropical Region (South America) and Australian subregion (New Guinea and Australia). Moreover, very similar data are known for other insect groups. For example, a winged but bulky, stag beetle was only recently discovered on Mt. Lewis in eastern central Queensland. Australia. Remarkably, it was shown by Moore (1978) to be con generic with Sphaenognathus Bcqiet, 1838, a genus otherwise only known from South America. The specialisation of Ornithoptera on Pararistolochia, and the apparently great age of the insects and plants which these genera represent (e.g. Forbes, 1932: Thorne, 1963, 1974). as well as the very specific chemical 'dependence' of all Troidini on their foodplants (e.g. FEENY, 1995), suggests that, as in other Troidini (e.g. Brown et al., 1995). the relationship is very ancient. If this is so, then the present-day distribu tions of the Aristolochiaceae and their herbivores may be considered to be indicative of past events of continental movements, if any. In the case of Ornithoptera the evidence is extremely impressive that fragmentation and drift ol" the resulting Gondwanan plates has played a fundamental vicariance role similar to that proposed by FOODEN (1972) affecting the aboriginal land mammal faunas of the region. It can be theorised that, after break-up of the Gondwanan continents. Ornithoptera arrived at 100 Michael J. Parsons its easterly destination via drift of the Australian plate, while Troides separately rafted to its westerly position, either on the Indian plate, or on the South-east Asian plate postulated by Ridd (1971) (Fig. 15). Only after subsequent dispersal into the Indonesian Archipelago by both genera independently of each other did their ranges begin to become sympatric, with presently minor sympatry only in New Guinea. It appears, therefore, that Ornithoptera did not reach the Australian subregion via Southeast Asia from Troides-Y\ke ancestors as has been assumed in previous studies. Furthermore, it seems likely that the allopatric evolution of already well-differentiated Ornithoptera and Troides would have been a direct result of the break-up of Gondwana land, as the separation of the two stocks from ancestral populations would have subjected them to evolution in isolation. Moreover, similar to Miller (1987), the results of my cladistic analyais suggests that the sister group of Indo-Australian Troides (sensu stricto, i.e. minus Ornithoptera) + Atrophaneura is the neotropical genus Parides (Fig. 15). This is important as it further endorses the hypothesis presented above. Importantly, an exactly similarscenario for the arrival and evolution of palms in the Indo-Australian Region was proposed by Dransfield (1981). He suggested that they reached Malesia "twice" in this way, arriving on the Indian plate and spreading from the north-west into the archipelago via the Asian mainland, and also on the Australian plate, radiating from the the south-east via New Guinea. A similar course of evolutionary events has been suggested for mammals in the region. For example, Menzies and Dennis (1979) pointed out that the New Guinea marsupial fauna is very similar to that of Australia, obviously being derived from it, whereas the circa 50 species of closely related rodents (all of which belong to endemic genera, except cosmopolitan Rattus), and the bats, of the New Guinea fauna today are of South-east Asian origin. The splitting of Pararistolochia (and likely also Arislolochia) into geographical isolates during the fragmentation events affecting Gondwanaland also explains their present-day distributions. In addition, it can be postulated from their present day distributions and foodplant relations that, along with Troides, Atrophaneura and Thottea (Aristolochiaceae) evolved in isolation on the Indian and/or South-east Asian plate, while Cressida apparently 'accompanied' Ornithoptera in isolation on the Australian plate. Similarly, Coleman and Monteith (1981) noted that a strikingly close association exists between diurnal uraniid moths and their predominant Omphalea (Euphorbiaceae) foodplants. They pointed out that this plant genus contains only about 20 widely disjunct species, yet it hosts uraniids in such widely separated countries as Jamaica, Cuba, Trinidad, Madagascar and Australia. They also speculated, therefore, that the uraniid moth/ Omphalea relationship dates from before the breakup and drifting apart of the Gondwanan continents. Reappraisal of Troidini 101

Evidence from Foodplant Relations Based on the scenario of the origins of Ornithoptera and Troides suggested above, several of the present-day larval foodplant relations of the Australian subregion Troidini seemingly do not fit a postulated ancestral 'plan' of Pararistolochia only for Ornithoptera, Arislolochia only for Troides, and Thottea only for Atrophaneura. Some of the seemingly 'anomalous' relations are, perhaps, more easily explained than others. Firstly, the fact that feeds as a larva only on Arislolochia tagala and one other Arislolochia species (presently unnamed), as opposed to Pararistolochia, is .'uncharacteristic' for its genus. This is based on the assumption that ancestral Ornithoptera exclusively utilised Pararistolochia (Fig. 15) and did not come into contact with Arislolochia species of Oriental subregion origin until northward drifting Australia contacted postulated existing Melanesian arcs (e.g. Holloway, 1984; Nix and Kalma, 1972). The distinctiveness of 0. victoriae immature and adult stages (Table 2), and its geographical restriction to the archipelago (Fig. 16), imply that the species has been long isolated from the remainder of Ornithoptera. Conversely, the greater similarity of 0. alexandrae and 0. priamus immature stages to one another (and to the remaining Ornithoptera) suggest a much closer relationship between these two species than to 0. victoriae (best illustrated by the Fig. 17A cladogram). Therefore, it is reasonable to assume that the utilisation of Arislolochia by ancestral victoriae could have been through switching from Pararistolochia to Aristolochia foodplants. This would have permitted the species to colonise an archipelagic region where Pararistolochia was unavailable, but through which Aristo lochia was probably already radiating from the west. The fact that 0. priamus does not utilise two Arislolochia species in the A. crassinervia group is significant, but may be due (in part) to the plants being chemically toxic to priamus (Straatman, 1970). This is surprising considering the oligophagy for aristolochiaceous foodplants otherwise exhibited by priamus, but competitive exclusion by 0. goliath and 0. victoriae for A. crassinervia and its sibling may have played an important role in precluding priamus from also utilising these as foodplants. Spade et al. (1988) found that there is no unequivocal evidence from their data to suggest that a group of neotropical (Mexican) troidines are "partition ing" foodplants to avoid interspecific competition, but that this was implied by regional shifts in foodplant choice. They even suggested in their study that partitioning by Battus and Parides between two distinct taxonomic groups of neotro pical Aristolochia (hexandrous and pentamerous species) is possible. My data for Australian subregion Troidini shows some very interesting similarities and parallels to Spade's et al. findings, suggesting that such partitioning is also in operation in the New Guinea troidine foodplant relations. The wide range of an undescribed foodplant (Parsons, in press) that is sibling to A. crassinervia throughout New Guinea and many of the satellite islands, and well into the Solomons, implies that the vine may have been the first of the two crassinervia 102 Michael J. Parsons group species to have dispersed into emerging Miocene 'proto' New Guinea (e.g. Nix and Kalma, 1972) from the Oriental subregion. Based on the foodplant specialisa tions of 0. victoriae and O. goliath, this dispersal apparently predated that of A. tagala. The fact that larval goliath specialises in feeding on both of the crassinervia subgroup Aristolochia species also adds weight to these assumptions as, within its species group, most of the immature stage and adult facies of goliath are more plesiomorphic (Table 2) than those of the other member species: rolhschildi, chimaera, tithonus, paradisea and meridionalis (Figs. 16 and 17A). Based on its present-day ecology (e.g. success in disturbed marginal secondary forest areas and production of abundant, year-round, wind-borne seed), Arislolochia tagala is apparently the most recent member of its genus to enter New Guinea. The immigration and spread of this foodplant from Oriental subregion, where it also occurs extensively, would have prepared the way for the penetration of Troides into the region. That Troides has, on the geological timescale, not been long in New Guinea is indicated by the fact that it is represented there by only a single species (oblongomaculalus). Moreover, the distribution of oblongomaculalus (like that of Troides as a whole: India, South-east Asia, South-eastern China, Philippines, Indonesia and mainland New Guinea) attenuates from west to east and, in New Guinea, is restricted to the mainland. Unlike 0. priamus, therefore, T. oblongomaculalus has not yet crossed the many sea water barriers to any of the main islands of the easternmost archipelagos where A. tagala is, nevertheless, widespread and abundant. Other apparent contradictions in New Guinean and Australian troidine larval foodplant relations are suggested by those of Atrophaneura polydorus (Linnaeus, 1763), which is widespread in the Australian subregion, and by rclictual monotypic , which is restricted to south-eastern New Guinea and north-eastern Australia. In the case of A. polydorus, it is likely that, just as 0. priamus has apparently successfully expanded its range of larval foodplants from Pararistolochia to also include oriental A. tagala, so A. polydorus conversely increased its foodplant spectrum from A. tagala to also include certain Pararistolochia species. This would have occurred in recent geological time when the Australian plate and emerging Miocene 'proto' New Guinea came into contact with the Melanesian arcs (e.g. Nix and Kalma, 1972). A. polydorus utilises several Pararistolochia species in New Guinea and Australia, but its primary foodplant is A. tagala. Based on the distribution and foodplant relations of Atrophaneura as a whole, the genus can be presumed to have originated in the Oriental subregion and ancestrally utilised Thottea and/or Aristolochia as larval foodplants. Cressida is at least as phylogenetically old as Ornithoptera, and likely older (Fig. 12). Based on its present-day distribution, Cressida was apparently also restricted to the Australian fragment of Gondwanaland. Possibly, therefore, ancestral Cressida, like Ornithoptera, may have been restricted to Pararistolochia larval foodplants during the early part of its isolation on northward drifting Australia. However, today Cressida only utilises a notably distinct group of endemic Australian Aristolochia Reappraisal of Troidini 103 species. Its specialisation on these small, herbaceous, arid-adapted Aristolochia species (pubera, thozetii and holtzei) restricted to north-eastern tropical / subtropical Australia strongly suggests that these foodplants are also of Australian/Gondwanan origin. Therefore, it is possible that Aristolochia species endemic to Australia were utilised early on in the evolution of Cressida, even perhaps as a means of avoiding competition with ancestral Ornithoptera. If it was 'always' an Aristolochia-feeder, ancestral Cressida may have utilised typical large vine-like Aristolochia foodplants which necessarily evolved into smaller herbaceous perennial species as the Australian climate gradually desertified. Some evidence suggesting that at least the herbaceous Australian Aristolochia are chemically quite distinct from species of obvious Oriental origin, such as A. tagala, was obtained by simple experiment. When provided with A. tagala, Cressida larvae from Port Moresby, PNG, fed only very reluctantly and eventually died (pers. obs.). It is also interesting to note that, besides a typical vine like species ( in Madagascar), the Afrotropical Region is home to a small (possibly relictual) group of three arid-adapted perennial herbaceous Aristolochia species which may, or may not, eventually be shown to share taxonomic links with the Australian species of very similar habit. If there are such links, then this would suggest that ancestral Cressida began to utilise perennial herbaceous Aristolochia at the very beginning of its evolution. Having elucidated apparent anomalies in the foodplant relations of some Australian subregion Troidini it is relevant to point out that other key larval foodplant switches in the more 'primitive' Papilionidae are well known. For example, the large genus Pamassius Latreille, 1804, and monotypic Hypermnestra Menetries, 1846, unlike all other Aristolochiaceae-feeding members of their subfamily Parnassiinae, have clearly adapted (evolved) to now utilise succulent- leaved plants of several unrelated families. This obviously permitted these parnas- siines to radiate into the temperate Holarctic regions of the world (e.g. Hancock, 1983).

Birdwing Evolution: Also Part of the Gondwanan Picture If a South-east Asian plate did once form part of Gondwanaland as Ridd (1971) convincingly argued, then this may explain the present-day geographical position of the subgenus Trogonoptera (of Troides) in Sumatra, Borneo and Palawan. Reviewing geological evidence, Audley-Charles (1987) further suggested that a series of Gondwanan continental fragments (comprising land blocks that now represent much of Tibet+ Burma+ Malay Peninsula, and most of the major Indonesian islands) rifted from the northern margin of the Australian plate to drift northwards, arriving between South-east Asia and New Guinea/Australia in the late Cretaceous to early Tertiary (circa 90-80 mybp). Ridd illustrated Borneo and Sumatra as belonging to the South-east Asian plate and, according to Zeuner (1943) and Haugum and Low (1978-85: Vol. 2: 88), several characters suggest that Trogonoptera is ancestral to 104 Michael J. Parsons

Troides. Although this cannot be proven, it is likely that Trogonoptera represents a sister group to Troides in which certain plesiomorphic character-states have been retained. For example, in Trogonoptera (from colour photographic figure in Goh, 1994) the ovum appears to be weakly ribbed, with granulose striations as in Pharmacophagus (Figs. 2 and 3) Parides, Atrophaneura and most other troidines, this condition apparently being the plesiomorphic state in Troidini. The morphology of the Trogonoptera first larval instar subdorsal tubercles, and the presence of iridescent green wing scaling in the adult males (entirely absent in Troides), are characters more similar to those of Ornithoptera, which is ancestral to Troides (Fig. 12). From colour photographic figures in Igarashi (1979) and Goh (1994), Trogonoptera first larval instar subdorsal tubercle apices are clearly only weakly chitinous, and the setae are more widely spaced as in Ornithoptera (contrary to Igarashi, 1984: 49, who stated that Trogonoptera first larval instar subdorsal tubercles resemble those of Troides). It is possible, therefore, that certain of the vicariance events of Gondwanan fragmentation may also explain the distinctiveness and geographical position of Trogonoptera. Hancock (1980) also concluded that the phylogeny of the Indo-Australian Troidini (he discussed Cressida, Atrophaneura and Troides) supports the suggestion by Ridd (1971) that South-east Asia was also part of Gondwanaland, but for different reasons to those outlined above. He maintained that they are derivable from the more primitive South American Euryades and Parides, and represent a dual invasion before the break-up of Gondwanaland, their present-day distributions suggesting that the two invading ancestors, Cressida and Atrophaneura / Troides, followed different dispersal routes. Hancock presumably considered that ancestral Cressida drifted with Australia, although he did not state this. He maintained that the Atrophaneura/ Troides ancestor appears to have dispersed via India (as Atrophaneura) to South-east Asia. He argued that the presence of Pharmacophagus antenor, the only troidine in the Afrotropical Region, supports the suggestion that dispersal was via India. With the post-Gondwanaland unification of India and Asia, Atrophaneura was able to radiate throughout the Indo-Australian Region, the most easterly representatives belonging to the specialised polydorus group. Hancock (1988) further refined his theories, suggesting that the Troides lineage radiated from Sundaland, and Atrophaneura from further north, its spread to Sundaland appearing to have resulted in the differentia tion of Pachliopta (the numerous specialisations of the latter group suggesting that it is a coloniser adapting to new environments). Evidence from the present-day distributions of Cressida and Euryades, as well as from the sole Australian subregion representative genus, Tellervo Kirby, 1894, of the otherwise neotropical nymphalid subfamily Ithomiinae, can be cited as strong additional support for the evolutionary origins of the birdwing genera newly suggested above. The highly disjunct geographical position of the wholly Australian subregion restricted Cressida, in relation to that of its nearest relative, South American Euryades, is indicative of the effect on these genera of the fragmentation and drift of Reappraisal ofTroidini 105

Gondwanan continental plates. Smart (1978) recognised this when he pointed out from this evidence that the affinities of Cressida and Euryades with Parnassiinae "strongly suggest that all the subfamilies and tribes of Papilionidae arose relatively early from theircommon ancestral stock ... It has possibly not been pointed out before that Euryades and Cressida occupy the same general areas inhabited by present day marsupials." Likewise, the geographical distributions of the New Guinea region restricted Tellervo (Ambon, Seram, Kai and Aru, eastwards throughout mainland NG, to the Bismarcks, Solomons and the of Queensland in northern Australia), and the remaining Ithomiinae restricted to tropical South America, further strengthen the argument for a truly Gondwanan component in the New Guinea butterfly fauna. Emmel et al. (1974) noted that chromosomal evidence suggests that Tellervo is a long isolated genus, and that "the ancestors of the genus Tellervo reached Australia at a period near the beginning of differentiation within the Ithomiinae" (although Kitching, 1981, accepted the concepts of previous authors and postulated that Tellervo entered Australia from New Guinea). Of relevance is the absence, apart from Pharmacophagus antenor in Madagascar, of any troidine papilionids in the entire Afrotropical Region. This is remarkable considering that the African mainland is home to about 12 endemic species of Pararistolochia and 3 Aristolochia species. Such 'negative' evidence, although difficult to assess, may also be eventually found to fit the 'Gondwanan picture' of troidine evolution as it is further refined. As analysis of butterfly data is becoming more refined and detailed, evidence for other possible Gondwanan groups is slowly emerging. For example, in a biogeog raphical study of Sulawesi Hesperiidae De Jong (1990) noted that the genera Taractrocera Butler, 1870, and Cephrenes Waterhouse & Lyell, 1914, indicate a stronger than previously suspected Australian element in the Sulawesi fauna. He suggested that, for Taractrocera + Ocybadistes Heron, 1894, -\-Suniana Evans, 1934, the group originally dispersed from Australia westwards only once, and that if it were Oriental, at least 2-6 dispersal events to the east would be needed to explain their present-day distributions. De Jong came to a similar conclusion about Cephrenes and made the important point that the Australian origin of the Taractrocera species in and west of Wallacea is only apparent because phylogenetic relationships were taken into account, whereas counting species and calculating dissimilarity of some sort would not have revealed this, nor his conclusion of a single dispersal event eastwards across Wallace's Line for Taractrocera + Ocybadistes + Suniana.

Older, More Ancient Origins Of obvious relevance to the hypothesis of the origins of Indo-Australian Region troidines proposed above are the ages of these papilionid genera, as their ancestral taxa must at least be as old as the main period of tectonic events affecting Gondwanaland. For example, Troides and Ornithoptera would presumably have had 106 Michael J. Parsons

an origin a minimum of about 80-60 million years ago (Late Cretaceous), if they evolved from a Gondwanan common ancestor at the time when fragmentation and drifting apart of the palaeocontinent was taking place. Based on their separate studies of papilionids Miller (1986, 1987) and Tyler etal. (1994) came to much the same conclusion. Similarly, in light of the "isolated, probably heliconiine genus Cethosia" in the Indo-Australian Region, Benson et al. (1976), in their study of the coevolution of passifloraceous-feeding nymphalids with their foodplants, were forced to consider that the relationship "Perhaps dates the tribe [Heliconiini] from before the South American-Antarctica-Australia fission over 60 million years ago." Like troidines, heliconiines are also butterflies which clearly share intimate chemical relationships with their larval foodplants (reflected in the aposematic colouration, behaviour, etc., of their adults and immature stages). Previous authors, such as Zeuner (1943), have assumed that the birdwing genera are far less ancient than proposed above, but evidence is accumulating that Lepidoptera (and notably the butterflies) may be somewhat older order than was initially believed. For example, many of the oldest butterfly fossils (Early Oligocene: ca. 38 mybp), look remarkably 'modern' in their facies (e.g. Scudder, 1875, 1889; Zeuner, 1942, 1961; Brown, 1976; Shields, 1976; Smart, 1978), implying a much earlier period during which their major differentiations took place. Even the oldest known fossil butterflies — two papilionid species dating back to 48 mybp — were readily comparable to Baronia brevicornis Salvin, 1893, by Durden and Rose (1978), and probably belong in the Baroniinae, according to Scott (1985). Baroniinae is considered to represent the most plesiomorphic papilionid subfamily and contains only B. brevicornis which occupies a relict position in a small area of southern subcoastal Mexico. This species is thought to be very close in its appearance to the first papilionid and, therefore, to the first true butterfly. Tindale (1980) suggested that the butterflies are a very archaic group, noting that Durden and Rose's finding strengthen the view of a Mesozoic origin of the lepidopteran stem. Observing that most entomologists consider Oligocene insect fossils fit comfortably into modern genera, Vane-Wright (1991) considered it likely that "most (if not all) modern butterfly genera would have been established by the time the east Sulawesi fusion event took place, and that many present day species lineages would also have been established." Whalley (1985) suggested that the earliest known fossil of a scale covered wing, with a microstructure of the scales very similar to that of modern Lepidoptera, and dating from the Lower Jurassic (ca. 180 mybp), could be that of a moth, but might also be of a trichopteran. Whalley (1986) further noted that modern lepidopteran families, including the major butterfly lineages, were differentiated by the Paleocene- Eocene (ca. 65-60 mybp). Razowski (1974) considered the origin of the Lepidop tera at the beginning of the Jurassic. From his study of fossil Lepidoptera, Shields (1976) concluded that butterfly radiation probably occurred even earlier, during the Reappraisal ofTroidini 107

Triassic (ca. 200 mybp), with the major radiation of the Lepidopteran families already completed by the Upper Jurassic-Early Cretaceous (ca. 120 mybp). These findings corroborate the time period postulated above for the origin of the Indo- Australian troidine genera relative to the geological time-scale. Forbes (1932) pointed out that Aristolochiaceae is considered to be among the most primitive ofall the flowering plants and gave his reasons for suggesting a Jurassic (ca. 190-136 mybp) origin for the Papilionidae. From the botanical point of view, Hawkes and Smith (1965) argued that representatives of some modern genera of flowering plants must have been in existence on Gondwanaland at least by early Cretaceous times (120-100 mybp), if not earlier. Like Hawkes and Smith, Truswell et al. (1987) pointed out that palynological evidence now available suggests that some angiosperms, including some of the allegedly primitive ones [i.e. Aristolochiaceae], may have had at least pan- tropical distributions in the late Cretaceous and early Tertiary. Audley-Charles (1987) also noted that the oldest fossils identified as angiosperms are about 120 million years old, and exhibit such well developed features that they imply that Angiospermae must have originated at least as far back as the lateJurassic. In fact, Melville (1975), in his study of Australian relict plants and their bearing on angiosperm evolution, concluded that the characteristic Australian flora must have evolved insitu from ancestors of Permian age (280-225 mybp). Such facts, therefore, only add weight to the concept that evolution of the Ornithoptera and other proposed Gondwanan butterfly groups may be very much earlier than has been traditionally assumed.

CONCLUDING REMARKS From immature stage characters and ecological data my findings re-establish that Ornithoptera should be regarded as a valid taxon, and that the generic level is acceptable rank. In spite of the previous use of strict cladistic principles in the classification of Troidini, it is clear that paucity of data, coupled with the misinter pretation and /or neglect of known characters, caused Ornithoptera to be erroneously lumped with Troides. Hauser (1993) came to a similar conclusion about other papilionid taxa in his preliminary studies of the swallowtails. Apparently, the large size of both Ornithoptera and Troides, and their superficially similar adult facies, has been the primary stimulus for taxonomists to regard the taxa as being congeneric. A similar, but converse, argument was used by Miller (1987) to point out that superficial differences in adult morphology "including dramatic differences in size and pigmentation patterns, all of which are autapomorphic", has been an important reason why previous authors had overlooked the close (likely sister-generic) relation ship of Troides and Parides (sensu Miller, 1987). Otherwise only relatively few characters indicate that the relationship between Ornithoptera and Troides is as close as 108 Michael J. Parsons has been supposed, whereas many more suggest otherwise. Zeuner (1943) postulated a recent and relatively rapid evolution in the birdwing genera Ornithoptera and Troides, which he considered were closely related. I suggest that this hypothesis does not fit the available evidence, particularly the ecological data from the field (unavailable to Zeuner), as well as the somewhat better, more detailed, present-day knowledge of the geology of the Australian subregion. Rather, conversely to Zeuner, and other more recent authors, I find that the data indicate Ornithoptera and Troides are each as ancient as general phylogenetic studies of the Papilionidae have suggested. This suggests that, even in the Jurassic, as Gondwana land began to fragment, plant families such as Aristolochiaceae and already well- diversified groups of ancestral troidines were widely extant in the palaeocontinent, and their populations would have been subjected to division as the continental plates began to separate. As a result, it appears that Ornithoptera underwent its evolution on the Australian plate allopatrically from Troides on the Indian plate. Only much later in geological time did the events of continental drift bring the two genera together from opposite directions. Following subsequent extensions in their ranges, most notably Troides eastwards throughout the Indonesian archipelagos, the two genera became weakly sympatric in New Guinea. As stated above, the Ornithoptera are well known to collectors and are of great economic importance within the global insect trade. Consequently, they have received a notable level of attention by regional and international conservation organisations (e.g. Parsons, 1992a, 1995a and 1995b). In a world where habitats and species diversity are rapidly diminishing, and in the face of a continuing decline in conservation funding, it is inevitable that difficult choices must be made regarding which fauna and flora to research and conserve, or to simply protect. Such selections inevitably involve prioritising species. In light of evidence of their ancient Gondwanan origins, I suggest that the Ornithoptera, as well as their Pararistolochia foodplants, should be regarded even more highly in terms of their 'conservation value.' In a sense, these insects and plants are 'living fossils' representative of an ancient and obscure, or otherwise now naturally scarce, austral faunal and floral element.

ACKNOWLEDGEMENTS

Many people have kindly given me their time and assistance during my research. To my wife, Maria, my thanks for her constant advice, support and encouragement. Friends and colleagues at the Natural History Museum of Los Angeles County (LACM) Entomology Section who helped in numerous ways include Roy Snelling, Brian Brown, Brian Harris and Arthur Evans. Their help greatly hastened the completion of this work. Brian Brown kindly read through the manuscript, and provided helpful suggestions for its improvement. My thanks also go to Barbara Reappraisalof Troidini 109

Allen of the Exhibits Section for kindly allowing me use of her computer graphics equipment to prepare some of the figures; also to Jim Angus and Don Reynolds of the LACM Molecular Laboratory for generously allowing me use of their computers. In Azuza, Rosser Garrison was of great help in innumerable ways, including permission to use his library, as well as in assisting me to obtain certain literature. I am also most grateful for the hospitality that he and his wife, Jo, provided on many occasions.

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Numbers in square brackets refer to approximate larval or pupal lengths in millimeters. Fig. 20. Newly eclosed male of Ornithoptera tithonus in lateral view drying wings and hindwing inner marginal androconial brushes (Mokwam Village, Arfak Mountains, Irian Jaya, Indonesia). Note large size of the androconial brushes. Fig. 21. Ditto: dorsal. Fig. 22. Pharmacophagus antenor comparison of three larval instars (to same scale): A) second: dorsal [13]; B) third: dorsal [25]; C) fifth: lateral [58]. Fig. 23. Ornithoptera priamus third instar larva dorsal (Brown River area, Central Province, PNG) [28]. Fig. 24. Ditto: but fifth instar lateral [90]. Fig. 25. mature fifth instar larva lateral (Xishuangbanna area, Yunnan Province, S. China) [50]. Fig. 26. Troides oblongomaculalus fifth instar larva dorsolateral (Popondetta area, Northern Province, PNG) [50]. Fig. 27. Pharmacophagus antenor pupa dorsal [40]. Fig. 28. Ditto: lateral [40]. Fig. 29. Ornithoptera paradisea pupa dorsal (Maprik area, East Scpik Province, PNG) [60]. Fig. 30. Ditto: lateral [60]. Fig. 31. Cressida cressida pupa lateral (Mt. Glorious, Nr. Brisbane, Queensland, Australia) [28]. Fig. 32. Troides oblongomaculalus pupa dorsal (Bulolo area, Morobc Province, PNG) [48]. Fig. 33. Ditto: lateral [48]. Fig. 34. Atrophaneura polydorus pupa lateral (Popondetta area, Northern Province, PNG) [28]. Reappraisal of Troidini 117 118 Michael J. Parsons

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