Pattern in and 291 Geological signal and dispersal noise in two contrasting groups in the Indo-Australian tropics: R-mode analysis of pattern in Lepidoptera and cicadas

Jeremy D Holloway Department of Entomology, The Natural History Museum, Cromwell Road, London, SW7 5BD, UK

Key words: phenetic biogeography, cladistic biogeography, panbiogeography, cluster analysis, R- mode pattern analysis, Lepidoptera, cicadas, Sundaland, , Melanesian archipelagos

Abstract the Indo-Australian tropics would probably be aware of the interest shown by biogeographers Biogeographic methodology and philosophy are reviewed in the results of their work and the possibility for within the context of obtaining an objective means of iden- some mutual benefit to be gained by the sharing tifying geological signal in a complex archipelagic system where dispersal has played a major role in the development of ideas and data They might also be puzzled at of biogeographic pattern A method of R-mode analysis is the frequent lack of agreement amongst explored that associates phylogenetically related groups of biogeographers on the best way to approach (clades) which share common features in terms of data on plant and distributions, particu- geographical representation of endemics and more wide- larly where the objective was to derive from spread species Such associations of clades, perhaps com- patible with the panbiogeographic concept of generalised them information on Earth history tracks, can provide the basis for an independent analysis of Indeed there has been a danger in the not too area relationships, facilitating the identification of contrast- distant past that biogeographers would polarise ing patterns in the associations which may reflect spatially into two or three extreme groups One rather or temporally different episodes in the past geography of the archipelago The method is applied to data for Lepidop- traditionalist group was accused by the other, tera and cicadas on the basis of areas of endemism recog- more iconoclastic groups of embracing biologi- nised for the former and illustrates both the differences in cal dispersal to explain all biogeographic pat- pattern between the two, due to much more restricted pow- tern Each dispersal event was unique and there- ers of dispersal of cicadas, and common features that prob- fore did not generate scientifically testable hy- ably have a geological basis A second analysis for data, based on their much smaller areas of endemism, re- potheses Such scenarios were condemned as veals a lack of clear groupings similar to that of the Lepidop- story-telling, and dispersal was dismissed as ran- tera at a grosser geographical scale, indicating that similar dom and therefore uninformative stochastic pattern generation through dispersal over the In attempts to secure biogeography on a geological template may occur, but in a more localised fash- ion The results are assessed in relation to geological events, sound methodological basis of hypothesis test- in particular, the independent evolution of groups in the ing and falsification, there developed a counter- regions of Sundaland, the Sulawesi area and inner and outer tendency to reject any notion of biological dis- Melanesian archipelagos Evidence is presented for associa- persal and veer to another extreme where mod- tion of such groups on broader northern ( and ern biogeographic pattern was considered to outer Melanesian archipelagos) and southern (Sundanian, Banda arc, inner Melanesian archipelago and ) have developed by a process of fragmentation, axes Sulawesi (or its components) appears to have inter- or vicariance, of both Earth and life together, acted with both axes in a complex fashion with the potential to discover, using the new methodology, one unique set of area relation- ships from biological data that would encapsu- Introduction late earth history Conflicting biogeographic pat- terns in even very small archipelagos led some Geologists investigating the tectonic history of biogeographers, who might be termed geologi-

Biogeography and Geological Evolution of SE , pp 291-314 Edited by Robert Hall and Jeremy D Holloway © 1998 Backhuys Publishers, Leiden, The Netherlands 292 J( D( Holloway cal dispersalists, to postulate complex scenarios formative about the juxtaposition of land at the of island or terrane integration that would, if time when that dispersal occurred But subse- taken to their limits, rival the story-telling of the quent dispersal events equally will obscure such biological dispersalists Page and Lydeard (1994) pattern, possible overlaying it with pattern re- have reviewed the debate in relation to bioge- flecting a more recent juxtaposition of lands ography in the Caribbean archipelago Any methodology of pattern analysis must there- The philosophies of three different biogeo- fore optimise the possibility of recognising se- graphic schools: cladistic/vicariant, dispersalist/ quential, overlaid patterns Also, terrane accre- migrationist and panbiogeographic, have been tion in areas such as the Sunda shelf and New contrasted by Wilson (1991) in relation to a Guinea may lead to biotic assemblages that are number of criteria, such as methods of analysis disharmonic and of diverse affinity (Polhemus, used, assumptions about dispersal, evolution 1996) and other process factors, attitudes to fossil evi- Dispersal, though random, stochastic, need dence, non-structural physical factors such as not be uninformative (see also Jong, 1998 this climate, and the explanatory and predictive volume) Its frequency will relate to factors such powers of the models From these philosophies as distance and area among the islands existing has emerged a general consensus on the need to at any point in geological time This was mod- separate as far as possible the analysis of bio- elled for island systems by MacArthur and geographic pattern from considerations of the Wilson (1967), shown to hold in the establish- processes that have led to development of such ment of the biota of Norfolk Island where the pattern But disagreement has remained on the source areas are unambiguous (Holloway, 1977, means whereby this may best be done, eliminat- 1996), and thence applied to help understand ing the likelihood of circularity of argument, yet the biological enrichment of Sulawesi not introducing such constraints in the cause of (Holloway, 1991) and to highlight anomalies in methodological rigour that the results foreclose the biogeographical affinities of elements of the on some process option in subsequent interpre- biota of Lord Howe Island (Holloway, 1977, tation Crisci et al (1991) stated, “Although it is 1979) valid to investigate the existence of a unique This stochastic basis for interaction between pattern of interrelationships among areas of en- island biotas, with likelihood of dispersal from demism, it cannot be accepted as an a priori as- one to another based on their distance and re- sumption of the analysis” spective areas, offers one means whereby bio- Arguments with much of the middle ground geographic pattern in dispersive groups of or- excluded are never ultimately very productive in ganisms can provide pointers to geological his- a real and messy world, and there are signs of a tory in complex archipelagic systems But this is welcome return to a more pragmatic approach realised through the methods of phenetic, rather to analysis of biogeographic pattern in relation than cladistic biogeography There is the addi- to earth history (e(g(, Page, 1989; Page and tional prospect, discussed by Roger Butlin at the Lydeard, 1994; Wagner and Funk, 1995) There meeting that led to this book, of modelling dis- are still many potential pitfalls in the interpreting persal and speciation processes over different of biogeographic pattern: for example, the types geological scenarios for the region, contrasting of distribution patterns in the New Zealand biota the resulting distribution patterns with actual that have been related to tectonic arc structures ones Again, phenetic methods of analysis may in the islands (e(g(, Craw, 1989) are also manifest be as appropriate for this as cladistic ones Such in the distributions of plants introduced by Euro- modelling has yet to be undertaken in a biogeo- peans, and suggested for these to have a cli- graphic context, but has been deployed to help matic basis by Wilson et al (1992) understand Polynesian colonisation of the Pa- There are a number of problems in attempting cific and options for coconut dispersal across it to investigate biogeographic pattern in relation (Levison et al , 1969; Ward and Brakefield, 1992) to geological history in the Indo-Australian trop- Phenetic methods have been applied to data ics If the geological hypotheses current today sets for mobile animal groups in the Indo-Aus- for the history of the archipelago are taken at tralian tropics Holloway and Jardine (1968) de- face value, then biological dispersal is likely to rived distance measures between major islands be the primary means whereby ancestral ranges in terms of their faunal (species-level) similari- of higher taxa (natural groupings of species) are ties using data for , birds and bats established Thereafter, fragmentation and They used non-metric multidimensional scaling vicariance can lead to biogeographic pattern in- to find two-dimensional plots of points repre- Pattern in Lepidoptera and cicadas 293 senting the islands that best summarised this ar- five points in geological time from the mid- ray of ‘faunal distances’ The plots for butterflies Cenozoic to the present and birds showed high correlation with current Most recent references to the analysis of geography, support for the predominance of a Holloway and Jardine (1968) have been in rela- stochastic, dispersal mode of pattern generation tion to its use of phenetic methodology (cluster within the group analysis of the matrix of faunal dissimilarity co- The authors went on to suggest that, where efficients that formed the basis for the non-met- the plots departed from such correlation, this ric multidimensional scaling just described) to might reflect some aspects of past geography, generate just such a tree structure, and this has particularly where plots for different groups of been compared with results deriving tree struc- organisms showed the same trends Thus, close tures using the methods of cladistics (e(g(, association of peninsular , , Holloway (1991), using Lepidoptera data for the and was interpreted in terms of same area) A point of interest in this compari- faunal intermingling facilitated by union of these son relevant to the chapter by Jong (1998 this lands on the Sunda shelf during Pleistocene low volume) is that the phenetic analysis of Lepidop- sea-levels Sulawesi was placed well separate tera distributions, reflecting overall faunal simi- from Borneo, with the Philippines in an interme- larities, paired the northern and southern diate position, perhaps indicative of greater iso- Moluccas, whereas the cladistic analyses, reflect- lation for the former in the past, interaction be- ing phylogenetic relationships, did not, group- ing primarily through the latter, perhaps through ing them independently with greater exposure of geological structures be- (Holloway, 1991): this is entirely consistent with tween the Minahasa peninsula of Sulawesi, the conclusions of Jong Sangihe and Mindanao Relationships between But the primary purpose of Holloway and the northern and southern Moluccas, New Jardine (1968) was to illustrate the complemen- Guinea, the Bismarck Islands and the Solomons tarity of two approaches to analysis of distribu- also departed from current geography, with the tional data, classifying areas in terms of relation- scaling method indicating high stress values be- ships of their biotas (Q-mode), and taxa in terms tween the two-dimensional summary and the of their distribution amongst areas (R-mode) raw data, perhaps a measure of the complex This distinction perhaps also encapsulates the geological history of what is now known to be a divergence between the cladistic and composite zone of several different structures, panbiogeographic schools of biogeography, a even within the island of New Guinea itself, as distinction rarely referred to in the literature, ex- will be seen later ceptions being Simberloff and Connor (1980) Indeed, the history of the archipelago is as and Page (1989) much one of fusion and convergence of terranes It is the purpose of this chapter to return to and other structures such as island arcs as of this complementarity and to suggest it offers an their fragmentation and divergence (Polhemus, escape from the search for a unique dichoto- 1996) Hence the focus of the cladistic approach mous tree hypothesis of area relationships that is to biogeography, attempting to portray area re- often the focus of cladistic biogeographic meth- lationships in the form of a dichotomous tree ods It does not foreclose on the possibility dis- structure from analysis of biogeographic pattern cussed earlier that data to hand may contain sev- in plant and animal groups with taxa endemic to eral different sets of area relationships that may those areas needs to be examined carefully It derive from different periods in geological time, may thus not be possible to represent the hy- offering a series of biological shapshots of past potheses of geological evolution of the Indo- geography: the dispersive vicariance of Duffels Australian archipelago in dichotomous tree (1983) form, though attempts have been made (Turner, In addition, by comparing data from the Lepi- 1995; Boer, 1995b) Yet this is seen as an impor- doptera (the speciality of the author), offering a tant prerequisite for application of cladistic wide range of dispersal abilities across the taxa, biogeographic methods (Morrone and Carpen- with those from the cicadas, with a much higher ter, 1994) Page and Lydeard (1994: their Fig 1) degree of local endemism and very few ex- present what are stated to be geological area tremely widespread taxa and therefore generally cladograms for the Caribbean from an earlier probably much less dispersive, it is possible to paper by Rosen These lack a time dimension, compare pattern in a group where stochastic each perhaps representing a snapshot of geo- pattern generation predominates with one graphical juxtaposition of areas of endemism at where vicariant pattern generation is far more 294 J( D( Holloway prevalent, as suggested by Boer (1995b) by a multiplicity of contrasting patterns, possibly Though groups of the second type may prof- of different ages and certainly blurred by disper- fer pattern that is much more informative in sal events In addition, there is no concordance terms of geology, their limited powers of disper- of pattern between montane-restricted and low- sal mean that their diversity in archipelagos will land restricted groups (Holloway, 1970, 1986a), be very much lower than in mainland areas and though this would be predicted under a purely lower than in other, more mobile groups among vicariant model of biological response to geo- the islands, and hence the sample of area logical change cladograms available for analysis of congruence An R-mode approach sets out to identify dis- will be relatively small An extreme example of tinct groupings of patterns and will therefore of- this is shown by major freshwater fish groups, fer a better prospect for at least a preliminary virtually restricted to continental shelf areas in biogeographic analysis in the Indo-Australian the Indo-Australian tropics (Darlington, 1957) tropics Identification of such groupings could Thus, the cicadas are represented in the archi- provide a prelude to a set of Q-mode area analy- pelagos east of the Sunda shelf and north of ses incorporating taxa from each general pattern Australia mostly by two major lineages (Boer, so identified: the complementarity of methodol- 1995b) The Lepidoptera are represented by ogy advocated by Holloway and Jardine (1968) hundreds, though a number of families, such as Trees for these taxa could be assessed for, and the , Bombycidae, Eupterotidae ‘cleaned’ of paralogy using the approach of Nel- and Limacodidae, also exhibit significantly re- son and Ladiges (1996), prior to an analysis of duced diversity in the more isolated archipela- congruence in area-relationships However, this gos Page and Lydeard (1994) advocated select- chapter will focus only on the preliminary R- ing groups with maximal endemism amongst mode analysis, as the sample of cladistic analy- their taxa but, as will be seen in the analyses, ses of groups is still relatively small When more groups showing endemicity in the remoter ar- analyses are available, it may prove that some of chipelagos tend to have taxa that are uninforma- the R-mode derived groups are distinguished tively widespread in those adjacent to continen- from each other merely by differential paralogy tal areas, and those with endemism in the latter rather than by inherently different structure of rarely penetrate the remoter archipelagos area relationships, but this should not be an a priori assumption The development of paralogy itself may be of interest for investigation of as- An R-mode method of pattern analysis pects of speciation and enrichment of biotas in biodiversity studies (e(g(, the widely different The phylogenetic data for Indo-Australian insect species-richness of Australian versus New groups, particularly Lepidoptera, that are accu- Guinea genera in the cicada tribe Chlorocystini mulating do not lend themselves easily to Q- referred to later) mode cladistic biogeographic analysis Deriva- The initial R-mode method of Holloway and tion of a single pattern of area relationships is Jardine (applied to further Lepidoptera data by virtually impossible using Component Analysis Holloway, 1973, 1974, 1979) classified taxa into (too much sympatry, too many widespread spe- faunal elements merely on similarity of presence cies), and Parsimony Analysis methods yield re- or absence in the areas incorporated in the study sults that appear to offer little advance over (effectively areas of endemism) The results of phenetic ones (Holloway, 1991) Morrone and such analyses are not dissimilar to the sort of Carpenter (1994) compared these and a further track analyses that have been made by method in relation to a range of data sets and panbiogeographers (e(g(, Craw, 1989) An exten- concluded that there still remained considerable sion of this (Holloway, 1969) grouped higher problems in their application, and that these de- taxa, mostly genera, in terms not just of pres- rived primarily from the effects of dispersal ence or absence in each area but also of richness There are also major computational problems in in species – a faunal centre analysis This ap- the application of such methods Nelson and proach grouped genera together that shared Ladiges (1996) have suggested that their proto- centres of richness, or massing centres, as well col for eliminating geographical paralogy (rep- as overall distribution etition) in cladograms circumvents these prob- A modified method is presented here that in- lems Turner (1995: p 103 et seq ) encountered corporates aspects of both these approaches It similar problems in his analyses of Australasian classifies higher taxa as in the second method, plant data The data suggest there may indeed but their component species are first assigned to Pattern in Lepidoptera and cicadas 295 distributional categories (faunal elements or (1996) All these groups are restricted to the generalised tracks) identified by the first Repre- Indo-Australian tropics and subtropics The data sentation of species in each higher taxon across are drawn from the following: these categories is tabulated and converted to a Arctiidae: Byrsia, Neoscaptia (Holloway, 1984) percentage Pairwise comparisons of the higher : Macrauzata (Inoue, 1993a); taxa give a measure of similarity of representa- (Watson, 1957) tion across the categories for each pair These Geometridae: Tanaorhinus rafflesii group similarity coefficients are then submitted to clus- (Holloway, 1982); Crasilogia, Nadagarodes, ter analysis to identify significant groupings of Polyacme (Holloway, 1984); Astygisa vexillaria the higher taxa, where similarity of representa- group, Bracca (3 clades), Probithia (Holloway, tion of their component species amongst the 1991); Ectropidia, Zeheba (Holloway, 1991, distributional categories is high 1993); Omiza, Peratophyga, Petelia medardaria The single-link cluster analysis method is group (Holloway, 1993); Dindica (Inoue, 1990); used A linkage diagram is also constructed to Dasyboarmia (Sato, 1987; Holloway, 1993) assess further the structure and cohesion of the Hesperiidae: Matapa (Jong, 1983) single-link clusters This approach, although Lasiocampidae: Arguda, Lajonquierea, somewhat cumbersome, gives better retrieval of Radhica, Syrastrena (Holloway, 1987a) the information content of the array of similarity Limacodidae: Susica (Holloway, 1982, 1986b); coefficients than do averaging and centroid clus- Darna (Holloway, 1986b; Holloway et al(, tering methods, and is an ad hoc and illustrative 1987); Setora, Thosea (Holloway et al , 1987); version of the non-hierarchic clustering method Narosa concinna group (Holloway, 1991) of Jardine and Sibson (1968) A more detailed : Drupadia (Cowan, 1974); Curetis discussion of the merits of this approach was (Eliot, 1990); Allotinus, Logania, Miletus (Eliot, presented by Holloway (1977, pp 163-169) 1986); Caleta, Catochrysops, Catopyrops, Danis, At this stage no information on phylogenetic Jamides (bochus and celeno groups), structure within the higher taxonomic groups is Nacaduba, , Psychonotis, Udara (sub- incorporated in the analysis However, the ap- genera Udara, Perivaga, Selmanix) (Hirowatari, plication of this R-mode method is facilitated by 1992); Callictita (Parsons, 1986) advances in the systematics of the Lepidoptera : Lacera (Holloway, 1979); since the earlier analyses referred to above, and serva group, Avitta, Ophyx (Holloway, 1984); also improved data on their distribution The ar- Aegilia, Paectes cristatrix group (Holloway, rival of cladistic methodology and the sharp- 1985); Chasmina (Holloway, 1989); Anomis ened concept of monophyly associated with it subgenus Rusicada (Holloway and Nielsen, 1998) has meant that it is possible to identify and se- : Besida, Cerura kandyia group, lect more reliably groups that are likely to be Phalera subgenus Erconholda, Phalera grotei monophyletic The same applies to the cicada group, Teleclita strigata group (Holloway, 1987b) data : Euploea (2 clades), Ideopsis, Parantica (4 clades) (Ackery and Vane-Wright, 1984); Tellervo (Ackery, 1987); Ptychandra The data (Banks et al , 1976); Chersonesia, Cyrestis (Holloway, 1973); Idea (Kitching et al , 1986); Lepidoptera Polyura (2 clades) (Smiles, 1982) Papilionidae: (3 clades) (Saigusa et Genera, subgenera or clades (monophyletic al , 1977) groups of species within genera) with five or : (2 clades) (Yata, 1990) more species (up to about 30) were selected Pyralidae: Vitessa (Munroe and Shaffer, 1980) from recent taxonomic publications Data for 85 Saturniidae: Attacus + Coscinocera (Peigler, 1989) monophyletic groups were located Phyloge- Thyrididae: Herdonia (Inoue, 1993b); Misalina netic hypotheses (cladograms) were available (Whalley, 1976) for 38 of these: the rest are from recent revisions The distributional categories recognised in- where the morphological definition of the clude endemism to individual islands or tight is- groups is unambiguous However, a number of land groups, together with wider categories these are rather large, and the method might (e(g(, Sundaland, Melanesian, Coral Sea, Indo- most appropriately be applied to groups of a Australian) identified by the faunal element more uniform size range (e(g(, 5-15 species), analyses for Lepidoptera by Holloway (1973, equivalent to the subtrees of Nelson and Ladiges 1974, 1979) 296 J( D( Holloway

WIDESPREAD Wide Indo-Australian Wide mainland Asia Wide Oriental SE Asian mainland Himalaya + SE Asian mainland / Sundaland Lesser Sunda / Borneo Sumatra Java Philippines Sulawesi WEBER’S LINE Wide Melanesian New Guinea and Moluccas Moluccas New Guinea Bismarcks Solomons Fiji New Caledonia Australia n Udara: Udara 423122 1111 n Prosotas 441 21 11111 n Jamides: celeno gp 636125135 4141 n Avitta 1321111 111 n Nacaduba 73   4121  11 322211 Zeheba 1111 1111 n Euploea: clade 2112 3 1 4     1   1115 1 211 1 ¡ Erconholda 1131  ¡ Besida 1111 1 Ptychandra 16  PHILIPPINES Udara: Selmanix 114  l Graphium: eurypylus gp 1121211  ¡ Omiza 11111  ¡ Setora 211 221  ¡ Phalera: grotei gp   1    1      1 1  1    1    ¡ Tanaorhinus: rafflesii gp   1    1      1 1  1    ¡ Byrsia 11 111 ¡ Idea 113111 11 l Curetis 221214211 1 ¡ Eurema: sari + candida gps   1    1     1 1 1 1   1  ¡ Logania 21112 11 l Tridrepana 24146321 126 l Herdonia 122451111 2 ¡ Caleta 12112 1 ¡ Bracca: emolliens gp 2 123 22111 l Polyura: delphis gp 271121  Matapa 4111  ORIENTAL l Peratophyga 11622 1 l Allotinus 221124243  l Miletus 11149151  Susica 1311  l Drupadia 11115 5  l Chersonesia 221  Syrastrena 1112   Darna 1115173121  Thosea 241414232 21 Ideopsis 121 11 Parantica: aglea gp 121 111 l Macrauzata 12211  Dasyboarmia 1123111  SUNDANIAN MONTANE Lajonquierea 1511  Dindica 122911121  n Probithia 1111 11 Radhica 131  Arguda 12132  n Anomis: Rusicada 342111 221 Cyrestis 12111 3 Pithecops 1111 1

50 60 70 80 90 100 PERCENTAGE SIMILARITY

Fig1 Single-link dendrogram and raw data for most of the Oriental cluster from the R-mode analysis of Lepidoptera taxa In the table of raw data the numbers of species of each taxon falling into each distributional category (listed at top) are indicated Symbols are used to facilitate cross-reference with the clusters recognised on the linkage diagram of Figs3 and 4 and discussed in the text The rest of the dendrogram and data are illustrated in Fig2 Pattern in Lepidoptera and cicadas 297

ORIENTAL/WIDE CLUSTER (FIG1) Wide Indo-Australian Wide mainland Asia Wide Oriental SE Asian mainland Himalaya + SE Asian mainland China/Taiwan Sundaland Lesser Sunda India/Sri Lanka Borneo Sumatra Java Philippines Sulawesi WEBER’S LINE Wide Melanesian New Guinea and Moluccas Moluccas New Guinea Bismarcks Solomons Fiji New Caledonia Australia Narosa: concinna gp 2101 SULAWESI Astygisa: vexillaria gp 119  Ectropidia 5219 21 Parantica: tityoides gp 1114  ▲ Achaea: serva gp 3 11111 ▲ Vitessa 1   21 1  21 22251111 WIDESPREAD ▲ Attacus + Coscinocera 11131 11211 ▲ Euploea: clade 21131 3111 1112   Graphium: sarpedon gp 111 1111 WIDE + SW PACIFIC Aegilia 11 1111 Eurema: tilaha + lacteola gps 12111  ▲ Paectes: cristatrix gp 21 111 ▲ Catochrysops 2 111 Lacera 21 11 ■ Petelia: medardaria gp 2111 1111 Parantica: crowleyi gp 11121151  Chasmina 111111 11111 Nadagarodes  11523 Neoscaptia 1 1636 Parantica: pumila gp  2111 OUTER Ophyx  21181211 Tellervo  1121 Psychonotis  1 122111 Bracca: bajularia gp 1 1121   Bracca: outgroup  6 MELANESIAN ARCS Udara: Perivaga  8 Callictita  9 Danis   19 Crasilogia  1713 INNER Polyacme  1411 Misalina 211111 1199 Graphium: agamemnon gp 1 1113 Catopyrops 1 122 Jamides: bochus gp 1 1111111 Polyura: pyrrhus gp 1 12111 Cerura: kandyia gp 1111 111 WIDE, ALLOPATRIC Teleclita: strigata gp 1111 11

50 60 70 80 90 100 PERCENTAGE SIMILARITY

Fig2 Continuation of the single-link dendrogram shown in Fig1 from the R-mode analysis of Lepidoptera taxa In the table of raw data the numbers of species of each taxon falling into each distributional category (listed at top) are indicated Symbols are used to facilitate cross-reference with the clusters recognised on the linkage diagram of Figs3 and 4 and discussed in the text

Cicadas These workers have recently published major biogeographic syntheses of this work (Boer, The majority of the data for the cicadas is pub- 1995a, b; Boer and Duffels, 1996a, b), that also lished by the group of researchers at the Univer- give comprehensive reference to the original sity of Amsterdam who have developed taxonomic monographs and distributional data phylogenies for virtually all cicada groups in Additional Oriental groups that extend into the Sulawesi and from the Moluccas eastwards archipelago in the Cryptotympana, revised 298 J( D( Holloway

Fig3 Linkage diagram for the Oriental cluster of Figs1 and 2 Links of 60-100% similarity are indicated by solid lines and those of 51-59% similarity by long dashed lines Clustering of taxa at a higher level than this is indicated by short dashed lines with arrows Large symbols indicate members of the clusters or groupings of taxa mentioned in the text: the same symbols indicate members of these groups in Figs1 and 2 Outlying taxa that do not fall into these groups are indicated by smaller solid circles Linkage with the Papuan subregion cluster is indicated by the encircled star Pattern in Lepidoptera and cicadas 299

l Misalina

Parantica: pumila gp Polyura: pyrrhus gp l l Psychonotis l l Tellervo Crasilogia l Polyacme l Bracca: Jamides: bajularia gp bochus gp l l l Ophyx

Graphium: l J agamemnon gp Danis Bracca: l l outgroup ORIENTAL CLUSTER Nadagarodes l l Callictita ll Neoscaptia Udara: l NEW GUINEA Perivaga Catopyrops MELANESIAN ARCHIPELAGO

Fig4 Linkage diagram for the Papuan cluster of Fig2 Conventions are as for Fig3

by Hayashi (1987a, b), Chremistica (Bregman, extend into the Papuan subregion (bounded to 1985) and Dundubia (Bloem and Duffels, 1976) the west by Weber’s Line of Faunal Balance after were also included the findings of Holloway and Jardine, 1968) The These data were initially analysed using the widespread groups are represented fairly evenly same distributional categories as for the butter- throughout the whole of the Indo-Australian flies, but cicada species were virtually unrepre- tropics and make a major contribution to the sented in the more widespread faunal elements faunas of Pacific archipelagos Within the more easterly archipelagos cicada Linkage diagrams for these two main clusters species are often restricted to one or two islands are shown in Figs 3 and 4, with links down to (including one or two endemic to very small is- 50% similarity being illustrated: clustering in of lands, excluded from the analysis), and they are outlying taxa above that level is also shown The also localised within the island of New Guinea widespread taxa (solid squares) form a moder- (Duffels and Boer, 1990) Therefore a second ately cohesive cluster distinct from a much analysis was performed on Papuan subregion larger grouping of taxa with much stronger rep- cicada genera using a much finer-grained set of resentation (species richness) in the Oriental re- distributional categories that are virtually those gion Within this larger grouping there is no identified in Boer (1995b) and combinations of definite substructure, but there is some polariza- them This finer-grained analysis provides an tion of groups (open and solid circles) across a opportunity to test objectively the coincidence continuum There are various small groupings of cicada groups as a whole over areas of ende- peripheral to these main clusters: a tight quartet mism recognised subjectively by Boer (1995b) of taxa with high endemism in Sulawesi (a com- parable situation in the Philippines is much more weakly defined); a loose association of Results: Lepidoptera taxa (solid triangles) with some affinity to the widespread cluster, but of a more Melanesian The dendrogram resulting from the single-link character cluster analysis, and the raw data from which it is derived, are shown in Figs 1 and 2 The dendrogram indicates a primary segregation of Oriental taxa: sympatric v( allopatric groups Papuan subregion (Melanesian archipelagos) groups from a combination of Oriental and The polarization of Oriental taxa is between taxa widespread ones Many of the Oriental groups (solid circles) that are particularly rich in species 300 J( D( Holloway

PERATOPHYGA Mainland Asia New Guinea (outgroup: 6 species) Sundaland, Philippines, Sulawesi Sundaland to S Moluccas Borneo, Malaya, Sumatra Sulawesi Borneo Vanuatu Borneo B Bismarcks Sulawesi Solomons Borneo, Malaya, Sumatra New Guinea Mainland SE Asia Bismarcks Borneo, Sumatra BRACCA Sulawesi Borneo S Moluccas Borneo, Sumatra Sulawesi Borneo, Malaya Sulawesi S Moluccas E Sulawesi Sulawesi Malaya, Sumatra, Borneo NE Himalaya, Malaya, Borneo Sumatra, Borneo Java, Sumatra Fig5 Cladogram with areas for the ennomine geometrid Philippines, Sangihe genus Peratophyga, with groupings as justified by Holloway Philippines (1993), exemplifying the sympatric Oriental type of genus N and S Moluccas Australia Bismarcks Australia, S Moluccas N and S Moluccas in the mainland Asian and Sundanian distribu- Kai tional categories, and those (open circles) that S Moluccas, New Guinea, Bismarcks are less well represented on the Asian mainland Solomons but have higher endemism in the Philippines Sundaland, Philippines and Sulawesi and are more likely than the other IDEA Sundaland group to be represented in the Melanesian archi- Philippines pelagos The first group of taxa has some ende- Sulawesi S Moluccas mism in Borneo, less in the Philippines and N Moluccas, New Guinea Sulawesi, and often shows a high degree of Sulawesi sympatry in mainland or Sundanian areas Those Sri Lanka, S India in the second group have a more even distribu- Mainland Asia tion of their species amongst the distributional Mainland Asia, Sundaland categories and exhibit greater allopatry, and Sundaland may therefore offer a better chance of detecting Fig 6 Cladograms with areas for the ennomine geometrid compatibility of area relationships in their genus Bracca and the danaine genus Idea, from cladistic structure For convenience in the text Holloway (1991) The latter and the emolliens clade (E) of Bracca exemplify the allopatric Oriental type of genus The following, these groups will be referred to as bajularia clade of Bracca (B) exemplifies the Melanesian sympatric versus allopatric Oriental groups archipelago type of genus in the Papuan subregion cluster, The ennomine genus Peratophyga illustrates the Bracca outgroup being in the New Guinea subcluster the sympatric group (Fig 5) Relationships within the genus are as described by Holloway (1993) The genus is particularly rich in various Sundanian categories, including Bornean Melanesian, as well as island endemics endemics: ten out of the fifteen species occur in Holloway (1982, 1987b) grouped these together Borneo, and the clades recognised within the subjectively with the Teleclita strigata and Cerura genus are also both sympatric and allopatric in kandyia groups, but these are segregated in the character In the eurypylus group of the swal- analysis by virtue of representation in Australia lowtail butterfly genus Graphium (Saigusa et al( and greater allopatry in the western part of their 1977), the component species are generally range Holloway suggested that, as there were much more widespread, but again the majority no obvious characters to indicate a strongly are represented in Borneo dichotomous cladistic structure within any of Within the allopatric group, different degrees them, and the islands grouped by the more of complexity are shown In Tanaorhinus, the widespread elements largely reflected modern Phalera grotei group and Byrsia there is very geography, these patterns probably resulted little overlap amongst an array of species that from relatively recent episodes of rapid ancestral fall into distributional categories of intermediate dispersal, followed by vicariance over the character, e(g(, mainland Oriental, Sundanian, islands in modern juxtaposition, perhaps in Pattern in Lepidoptera and cicadas 301 response to Pleistocene climatic changes AVITTA Indo-Australian tropics More complex patterns are exhibited by the Borneo Eurema sari + candida group, Besida, Erconholda, Mainland Asia, Sundaland the Bracca emolliens group and Idea, though in NE Himalaya, Borneo none of these is there any strong evidence of Sundaland commonality of pattern, merely that of trends In Borneo, Sumatra Sri Lanka, S India the Eurema example a Melanesian sister-pair is NE Himalaya, Sundaland sister to an Oriental complex, with one Andamans allopatric trio overlapping the composite range Philippines of the unresolved remainder extensively Besida Lesser Sundas Sulawesi, Moluccas is sister to a widespread Melanesian genus Moluccas, New Guinea, Australia, (added to the data), but has two Sundanian spe- Bismarcks New Caledonia cies and a Philippines-Sulawesi sister-pair Solomons, Vanuatu Erconholda also exhibits the Sulawesi-Philip- Fig7 Cladogram with areas for the noctuid genus Avitta, pines relationship but its interaction with exemplifying a widespread Oriental cluster genus From Sundanian areas is different (Holloway, 1987b) Holloway (1984) Phylogenetic hypotheses for the Bracca group and Idea are shown in Fig 6 In both, there can be seen a sort of progression from west to east within the cladogram, the earliest branches be- vide into species groups that fall within some of ing the more easterly species However, that for the other clusters in the analysis For example, Idea ‘commences’ in India and mainland Asia in the noctuid genus Avitta (Fig 7), allopatric and ‘terminates’ in New Guinea, and that for the and sympatric Oriental clades occur as sister- emolliens group of Bracca ‘commences’ in Sun- groups daland and ‘terminates’ in the Solomons, with The looser association of widespread taxa earlier branches Melanesian rather than conti- (solid triangles) consists of less species-rich nental Asian Even within the area of overlap of groups where the proportion of Melanesian and the two cladograms, that for the Bracca group is Pacific endemics is equal to, or greater than that more species-rich, has a similar outlying taxon of widespread taxa, with more allopatry overall in Sulawesi, but lacks corresponding representa- This character is also seen in the Graphium tion of Sulawesi further up the sequence sarpedon clade and Aegilia pair Distributions of Nevertheless, this general west to east pro- species of Paectes in the ‘solid triangle’ group gressive pattern of areas is seen also in a Brooks and of Aegilia were mapped by Holloway (1985) Parsimony Analysis for Lepidoptera where areas The widespread groups just discussed are lacking a particular clade were coded ‘?’ (unin- generally considered to be biogeographically formative) rather than zero (primitively absent) uninformative, an assertion that is generally true when data for that clade were tabulated for much of their distribution through the Indo- (Holloway, 1991), and in general area cladograms Australian archipelago, but the mobility of these for (Schuh and Stonedahl, 1986) and groups has also led to them being relatively well butterflies (Vane-Wright, 1990), also discussed represented in remoter Pacific archipelagos and illustrated by Holloway (1991) This pattern where they often show a much higher degree of will be discussed further below endemism There will undoubtedly be a strong stochastic element in any pattern in this ende- mism but within this, as indicated earlier, some Groups of widespread taxa geological signal may be observed For exam- ple, patterns involving New Caledonia, Vanuatu, The major widespread cluster contains groups Rotuma, Samoa and Fiji may reflect changes in (solid squares) with a high proportion of species the relative positions of these island groups in in the most widespread category (through the the past (Holloway 1979: pp 211-213, 1983; Indo-Australian tropics) and in other wide- Duffels, 1988) spread categories Endemism in the Oriental re- gion tends to be low except sometimes in the Philippines and Sulawesi, but the groups in- Sulawesi taxa clude a number of Melanesian archipelago endemics Several are species-rich genera that, The small cluster of taxa with high endemism in when subjected to cladistic analysis, may subdi- Sulawesi consists mostly of those with extensive 302 J( D( Holloway radiations of species within Sulawesi, with a few POLYACME New Guinea species from these radiations occurring in some New Guinea Australia, New Guinea, S Moluccas cases further east in the Moluccas or New New Guinea Guinea, and with a sister-relationship to Orien- Australia tal taxa to the west These could be considered New Guinea further examples of west-to-east progressive New Caledonia patterns An exception within this cluster is the CRASILOGIA group New Guinea tityoides group of Parantica where affinities are Australia with the Lesser Sundas and Banda island arcs New Guinea generally New Guinea New Guinea New Guinea Australia Papuan subregion taxa: New Guinea and New Guinea, S Moluccas Melanesian archipelago groups Australia New Caledonia Groups in the Papuan subregion cluster segre- New Guinea gate into New Guinea and Melanesian archi- New Guinea pelago components, a segregation supported by Fig8 Cladograms with areas for two geometrid genera ex- emplifying the New Guinea subcluster of the Papuan the linkage diagram (Fig 4), where the two subregion cluster: Polyacme (); Crasilogia with groups remain separate except for linkage Papuanticlea (upper and lower clades respectively; through the noctuid genus Ophyx Members of Larentiinae) From Holloway (1984) the New Guinea cluster are virtually restricted to that island Species in these groups outside New Guinea tend to be in Australia and New Caledo- nia or in the Moluccas Any representation fur- therein), but, given current hypotheses of the ther afield in the Melanesian archipelagos is by geological complexity of the area, this is some- species in the widespread Melanesian category, thing of a misnomer (Polhemus, 1996; Polhemus presumably relatively dispersive Examples from and Polhemus, 1998 this volume), and Inner and the Geometridae, the genera Polyacme and Outer Melanesian archipelagos might be prefer- Crasilogia (including Papuanticlea), are shown able terminology However, the cicada data sug- in Fig 8 gest, along with geological evidence, that the The Melanesian archipelago groups have their situation is far more complex (Boer, 1995b), and species evenly distributed from the Moluccas this primary segregation is merely a ‘coarse fo- through New Guinea into the archipelagos to cus’ pattern the east This distribution type is exemplified by There is also the possibility that the apparent the bajularia clade of Bracca (Fig 6) A phylo- segregation of the two groupings has arisen genetic hypothesis for Ophyx, the genus inter- through operation of factors other than Earth mediate between the two groups, is shown in history: some genera may have responded to Fig 9 It segregates into two clades, one that growth of the land area of New Guinea by would probably fall within the New Guinea speciation; others, perhaps more adapted eco- cluster if coded separately, and the other an logically to archipelagic conditions, including unusual combination of a small Melanesian ar- having greater powers of dispersal, may have chipelago group that is sister to an Australia/ speciated extensively only in the Melanesian is- New Caledonia pair land groups For example, the geometrid genus Associated with the Melanesian cluster is a Nadagarodes, placed in the Melanesian group pairing of the lycaenid butterfly genus Catopyrops here, has a cladistic structure (Holloway, 1984) with the agamemnon group of Graphium, where with weak segregation within it of New Guinea several species endemic to various Melanesian and Melanesian clades, but also with indication archipelagos are associated with species in more of much interchange between New Guinea, the widespread categories The case of Graphium Moluccas and the Solomons The sister-genus will be revisited later was suggested to be Probithia, falling into one These distinct New Guinea and Melanesian of the more widespread categories among the patterns are often cited as evidence for the con- Oriental groupings, extending from India to Fiji vergence of two distinct geological systems, (discussed further by Holloway, 1991) sometimes termed Inner and Outer Melanesian Two factors may strengthen a geological, arcs (Holloway, 1984; Boer, 1995b; references rather than ecological, interpretation of this di- Pattern in Lepidoptera and cicadas 303

OPHYX New Caledonia tremely widespread components One is the Australia largely Melanesian agamemnon group referred Solomons Solomons to above It is sister to the other pair One of New Guinea, S Moluccas these, the eurypylus group, has already been Bismarcks mentioned as an example of the sympatric Ori- New Guinea, Bismarcks, ?Australia ental type The other, the sarpedon group, was New Guinea also mentioned as one of the more widespread New Guinea New Guinea group that has a relatively high proportion of New Guinea more localised species in the eastern part of its New Guinea range: this is primarily a lineage distributed New Guinea, Bismarcks through the Moluccas, New Guinea, Australia New Guinea, Bismarcks and New Caledonia, a group, therefore, of weak New Guinea, S Moluccas, Bismarcks N Moluccas New Guinea cluster character New Guinea In Bracca (Fig 6), the Melanesian bajularia group New Guinea is segregated from the emolliens group in an New Guinea unresolved quadrichotomy with two other spe- Fig9 Cladogram with areas for the noctuid genus Ophyx, cies, one widespread, one endemic to Sulawesi intermediate between the New Guinea and Melanesian ar- The emolliens clade itself has already been dis- chipelago subclusters of the Papuan subregion cluster From Holloway (1984) cussed as illustrating a west to east progressive structure in a sister-relationship to a trio of spe- cies endemic to Sulawesi This whole complex is probably sister to a clade of six species used chotomy in pattern The first is the extent to as an outgroup in the analysis by Holloway which putative Melanesian groups still show (1991) This clade is restricted to New Guinea some restriction to the more recently accreted and falls within the New Guinea group in the Melanesian terranes in New Guinea, such as the analysis northern ranges of mountains The second is the Holloway (1984) discussed four closely existence of sister-pairs of groups, one of each related genera of lithosiine arctiid Two of these distribution type There are few clear-cut exam- were included in the analysis One falls into the ples of this in the Lepidoptera groups in this Melanesian cluster (Neoscaptia) and the other, analysis, the best being the Danis-Psychonotis Byrsia, has already been noted as an allopatric sister-pair in the Lycaenidae, though no phylo- widespread group of low diversity The genetic analysis of this pairing has been under- remaining two genera are Scaptesyle, probably a taken The situation in Ophyx is another possi- sympatric Oriental group, and Damias, with ble example; Holloway (1984) also illustrated over thirty species restricted mostly to New possible sister-groups within the arctiid genus Guinea, but with three species in the southern Cyana where the Melanesian component does Moluccas (Seram, ) The genus has its also show restriction to northern New Guinea greatest richness in the southeastern peninsula of New Guinea and is relatively weakly represented in the northern structures Combinations of patterns In the nymphalid butterfly genus Parantica, the pumila group is of Melanesian character, but Both Melanesian and New Guinea patterns may the genus has no New Guinea components, the also be represented amongst a number of pumila group being sister to the tityoides and closely related lineages, but not in a strict sister- crowleyi groups, the former already mentioned relationship There are a number of examples for its high endemism in Sulawesi, and the latter amongst the Lepidoptera groups included in the having high endemism in the Philippines as well analysis, though sometimes the data were not as a rather widespread Oriental character This good enough to permit all lineages to be in- complex is sister to the aglea group which has a cluded They are drawn from a number of differ- wide Indo-Australian distribution to as far east as ent families the Solomons (see cladogram in Holloway The genus Graphium has (1984) from Ackery and Vane-Wright, 1984) already been mentioned, as has the geometrid In another powerfully flighted nymphalid ge- genus Bracca The analysis of Graphium by nus, Polyura (Smiles, 1982), the pyrrhus group, Saigusa et al( (1977) revealed three major line- weakly associated with the Melanesian cluster in ages Each of these includes one or more ex- the analysis, is in an unresolved triplet with a 304 J( D( Holloway

Solomons endemic and the sympatric Oriental exception being the Raiateana group of genera delphis group However, this triplet is sister to a (Solomons eastwards) that may be related to the single New Caledonian endemic Oriental Cryptotympana (Duffels, 1988, pp 80- Thus, though some gross pattern emerges 81) also included in the analysis Most of the from the distributions of Lepidoptera groups, Chlorocystini groups (asterisks in Fig 10) fall that may represent geological signal, the phylo- within the New Guinea cluster, the exception genetic structures within such groups, and be- being one group of the genus Baeturia Two of tween them when they come together in larger the four Cosmopsaltriaria groups (daggers) fall groupings, exhibit no strong pattern, indicative within the Melanesian cluster, the others being of process (e(g(, dispersal and speciation) that is the outlying Aceropyga + Moana group and the probably of a highly stochastic nature As stated New Guinea cluster genus Cosmopsaltria, which in the introduction, it is rare to find Lepidoptera bears a sister-relationship to the other Papuan groups that range from India into the Pacific and subregion groups (Duffels, 1983) Both tribes yet show the maximal endemicity of their taxa have relatives in Sulawesi, the Chlorocystini be- throughout that range advocated by Page and ing represented by their sister-tribe (Boer, 1995), Lydeard (1994) for biogeographical analysis the Prasiini (double asterisks) The Chlorocystini also have a number of small genera endemic to forests along the eastern seaboard of Australia Results: cicadas from Cape York to southern New South Wales, a feature roughly paralleled by some New Guinea The analysis of cicadas using the same distribu- groups in the butterfly analysis The significance tional categories as for the Lepidoptera yielded of this will be assessed in the final, general dis- similar clusters, but with virtually no overlap cussion between Oriental and Papuan subregion The areas of endemism recognised for the groups, the link between the two being at al- finer grained analysis of cicada groups are most zero percentage similarity (Fig 10) Indeed, somewhat more numerous than those identified the range of similarity values registered is con- by Boer (1995b), but are generally similar, par- siderably more extreme than that for the Lepi- ticularly within New Guinea (Fig 11) Species in doptera each group were either endemic to these or rep- The Oriental cluster shows more definite in- resented in various pairs, triplets or more of ternal structure, with a tight trio of groups virtu- them, yielding a total of 50 distributional catego- ally restricted to Sulawesi and three pairings of ries into which all could be assigned The results groups centred on the Philippines, on Sunda- of the analysis are shown in Fig 12 both in the land and mainland Asia, and on the Banda arcs form of a dendrogram and a minimum spanning This is probably a reflection of the much smaller tree sample of Oriental groups than for the Lepidop- The dendrogram reveals no strong clustering tera, for amongst these pairings there is little structure, and similarity levels are generally low, commonality of area relationships One of the not much exceeding 50% The spanning tree Philippines groups is associated with Sulawesi confirms this, with the groups falling more or and the other with Sundaland One member of less into a chain from groups in New Guinea the Banda arc pair has its centre of richness at and the Moluccas through to the Melanesian ar- the western end of the geological structure, in chipelago groups The spanning tree does bring Sumatra, and the other is more diverse at the together as neighbours the groups in the sub- eastern end, in the Lesser Sundas and around clusters of the coarse-grained analysis These : a further example of a Banda arc pattern groupings do not bear very close relationship to is illustrated for a group of Ptilomera (water- the subjective assignation of whole groups to strider) species by Polhemus (1996) ancestral areas of endemism by Boer (1995b) The Papuan subregion groups are, like those He referred Cosmopsaltria to central New Guinea, for the Lepidoptera, clearly segregated into New Baeturia as a whole, Guineapsaltria and Mira- Guinea and Melanesian archipelago clusters, the bilopsaltria to northern New Guinea, Thaum- former relatively tight, the latter less so, with astopsaltria, Papuapsaltria and Gymnotympana one outlier, Aceropyga + Moana, that has high to the Papuan peninsula, and Aedeastria to the endemism from the Bismarck Islands eastwards, Bird’s Head These assignations need not neces- but particularly in Fiji (Aceropyga) sarily be incorrect, but the analysis here does As mentioned earlier, these cicada groups fall suggest that stochastic processes such as disper- mainly within two major tribal groupings, the sal have tended to blur any clear patterns that Pattern in Lepidoptera and cicadas 305 SE Asian mainland Widespread mainland Asia India + Sri Lanka Malaya Borneo Sumatra Java Malaya + Sumatra Sumatra + Borneo Lesser Sundas Timor Lesser Sunda + Wallacea Philippines Sulawesi WEBER’S LINE Moluccas New Guinea + Bismarcks, Moluccas Australia (+ New Guinea) New Guinea Bismarcks Solomons Vanuatu Fiji Tonga/Samoa Fiji, Tonga, Samoa Papuapsaltria *  18 Baeturia: viridis gp *  7 Baeturia: exhausta gp *  5 Baeturia: loriae gp *  5 Mirabilopsaltria *  15 Baeturia: nasuta gp *  1101 Cosmopsaltria ✝   22 18 Baeturia: con + gutt gps *  36 NEW GUINEA Aedeastria *  37 MELANESIAN Gymnotympana *  32213 ARCHIPELAGOS Guineapsaltria *  114 Thaumastopsaltria *  115 Raiateana gp   41 Inflatopyga ✝   6 Baeturia: bloetei gp * 1 228111 Diceropyga ✝   511546 OUTER Aceropyga + Moana ✝   21192 Lembeja: foliata gp ** 13 SULAWESI Prasia ** 7  WALLACEA Dilobopyga ✝  301 Cryptotympana: accipiter gp  73  PHILIPPINES Chremistica: tridentigera gp 21314  Cryptotympana: recta gp 21121  SUNDANIAN/ASIAN Dundubia: terpsichore gp 1121  Cryptotympana: acuta gp 11133  BANDA ARCS Cryptotympana: diomedea gp 311 

02040 60 80 100 PERCENTAGE SIMILARITY Fig10 Single-link dendrogram and data for the R-mode analysis of cicada taxa using the same distributional categories as for the Lepidoptera Chlorocystini groups are indicated by *, Prasiini groups by ** and Cosmopsaltriaria groups by ✝

may have existed, though on a much more lo- cated in bold for New Guinea groups in the list calised scale than for the Lepidoptera This is following The list includes the genus or species seen most strongly in Baeturia itself where, group, area (or areas in ambiguous cases) of even if the overall ‘weight’ of the genus is in endemism suggested by Boer, followed by the northern New Guinea, some groups (nasuta, highest few links (percentage similarity in brack- loriae) approach Cosmopsaltria in their repre- ets) with other taxa in order of similarity sentation in central New Guinea, and another Gymnotympana (Papuan peninsula): Papua- (bloetei) has a distribution resembling that of the psaltria (51); Mirabilopsaltria (56); Cosmo- Melanesian groups of the Cosmopsaltriaria psaltria (44); Guineapsaltria, Baeturia loriae The correlation of the analysis with ancestral group (43) areas of endemism categories of Boer can be Guineapsaltria (Papuan peninsula): Mirabilo- assessed by examining the extent to which taxa psaltria (45); Gymnotympana (43); Thaum- in each category have high links primarily with astopsaltria (40) other taxa in that category Such links are indi- Thaumastopsaltria (Papuan peninsula): 306 J( D( Holloway

Fig11 Areas of endemism recognised for the finer R-mode analysis of cicada patterns, after Boer (1995b) The solid lines delimit the areas of endemism of Boer; broken lines indicate further subdivision of these for this analysis

Guineapsaltria (40); Baeturia nasuta group There is some general affinity amongst (38), Papuapsaltria (35) Papuan peninsula groups on the one hand and Papuapsaltria (Papuan peninsula): Baeturia Northern New Guinea ones on the other, loriae group (56); Mirabilopsaltria (52); though Mirabilopsaltria of the latter also has Gymnotympana (51); Cosmopsaltria (47) links with the former, and Cosmopsaltria has its Baeturia loriae group (Papuan peninsula, major links with members of both groups Central New Guinea): Baeturia nasuta group (59); Papuapsaltria (56); Gymnotympana (43); Cosmopsaltria (41) Discussion Cosmopsaltria (Central New Guinea): Mirabilopsaltria (59); Papuapsaltria (47); Given this plethora of pattern variation in both Baeturia nasuta group (46); Gymnotympana, insect groups, are there any basic themes within Baeturia exhausta group (44) it that might have a geological basis, to which Baeturia nasuta group (Central New Guinea, further detail may be added through considering Northern New Guinea): Baeturia loriae group fine-grained pattern in more localised and hope- (59); Cosmopsaltria (46); Papuapsaltria (42) fully more informative groups such as cicadas? Mirabilopsaltria (Northern New Guinea): Analyses for both groups do recognise a Baeturia exhausta group (73); Cosmopsaltria number of fairly gross areas of endemism within (59); Papuapsaltria (52); Gymnotympana, the Indo-Australian archipelago In the Oriental Guineapsaltria (45) region there is some segregation of Sundaland, Baeturia exhausta group (Northern New Philippines and Sulawesi groups, with some in- Guinea): Mirabilopsaltria (73); Cosmopsaltria teraction between Sulawesi and the Philippines, (44) Sulawesi and the Moluccas and New Guinea, Baeturia viridis group (Northern New and Sulawesi and the Lesser Sundas The cicada Guinea): Mirabilopsaltria, Papuapsaltria (40); patterns, with finer detail, also include definite Baeturia nasuta group (37) Banda arc groups Some of the smaller sub- Baeturia conviva + guttulinervis group groups of Lembeja, not included in the analysis, (Maluku, Northern New Guinea): Aedeastria (20) also feature Sulawesi and the Lesser Sundas Aedeastria (Bird’s Head): Baeturia exhausta (Jong, 1985, 1986, 1987) This variety of associa- group, Mirabilopsaltria (34); Cosmopsaltria (31) tions between Sulawesi and neighbouring areas Pattern in Lepidoptera and cicadas 307

1 Papuapsaltria 3 ■ 2 Baeturia loriae group 2 3 Baeturia nasuta group ■ 4 Baeturia exhausta group 1 ■ ■ 7 5 Mirabilopsaltria 6 Cosmopsaltria ■ 6 7 Gymnotympana ■ ■ 5 ■8 8 Guineapsaltria 10 ■ 9 Thaumastopsaltria 4 10 Baeturia viridis group ■ 9 11 Aedeastria ■11 12 Diceropyga 13 Baeturia bloetei group 14 Baeturia con + gutt groups ■ 12 14 ■ 15 Aceropyga + Moana 16 Inflatopyga 15 ■ ■13 17 Raiateana group ■17 02040 60 80 100 PERCENTAGE SIMILARITY ■ 16

Fig12 Single-link dendrogram (left) for the finer R-mode analysis of cicada patterns, with the linkage diagram to which it is related (right) In the linkage diagram heavy solid lines indicate links at 50% and above, light lines those at 40-49% Broken lines indicate taxa clustering in (minimum spanning tree) at a lower level of similarity Heavy arrows indicate the division between the New Guinea and Outer Melanesian Archipelago groups of Fig10

is of interest, and may suggest a composite ori- Cosmopsaltria itself, are distributed mainly gin for the current island, discussed briefly be- amongst the Melanesian archipelagos Duffels low In the Papuan region there are two much (1990) proposed a possible phylogeny for the clearer patterns: the New Guinea groups and Cosmopsaltriaria group within Sulawesi, and re- Melanesian archipelago groups, the former of- viewed the localisation of the species within the ten with Australian associations Cicada groups island The group is represented by sister-genera, sharing these patterns have very much more Dilobopyga and Brachylobopyga Dilobopyga has species richness than do Lepidoptera groups, three major species groups Most of the species but there are fewer of them When this species are localised within the island, and some sub- richness occurs in larger islands, such as New groups within the major groups are concen- Guinea and Sulawesi, the cicada taxa concerned trated in one or other of the geographical com- are often localised within the island, and may ponents The genus is largely restricted to the reveal something of any geologically divergent north, centre and eastern peninsulas Apart from histories among the components In the Lepi- an island-wide species, only two species are lo- doptera, such internal localisation, if it ever oc- calised in southwest Sulawesi, a sister-pair, one curred, has usually been blurred by subsequent on the island of Saleyer There is one species in movements of the species, leading to a high de- the southern Moluccas Brachylobopyga has gree of sympatry (Holloway, 1991) only a pair of species, one central and another in Boer (1995b) focused mainly on the similari- the southwest So, as for the Lembeja foliata ties between the biogeography of the Prasiini + group of the Prasiini, discussed below, the Chlorocystini and the Cosmopsaltriaria cicada greatest diversity is in the north and east of the groups, but the differences, particularly in rela- island, though with richness also in the centre tion to Sulawesi and the composite nature of the Within both groups there are indications of vi- Papuan subregion, may be as interesting It is cariance between western and eastern groups worth considering these groups in a little more of species detail (Fig 13), though the reader is recom- The tribe Prasiini is strongly centred on mended also to consult Boer (1995b), and Boer Sulawesi Its sister-group, the tribe Chlorocystini, and Duffels (1996a, b) occurs from the Sula Islands east to Samoa and As mentioned earlier, the Papuan subregion northeastern Australia as discussed earlier The Cosmopsaltriaria groups, with the exception of sister relationship of the two together is thought 308 J( D( Holloway

East Asia/Ryukyu Meimuna

Muda SUNDALAND + Ryukyu

Prasia L harderi group LESSER SUNDAS L fatiloqua group Prasiini L parvula group SULAWESI Brachylobopyga L foliata group N + E Dilobopyga L robusta group NEW GUINEA Jacatra SUMATRA, JAVA Arfaka Bird’s Aedeastria Head Rhadinopyga Solomons Inflatopyga Thaumastopsaltria PP Maluku/Bismarck/Solomons Diceropyga Mirabilopsaltria NNG Cystopsaltria, Cystosoma AUS Bismarck/Solomons/Samoa Moana Fiji Aceropyga Papuapsaltria PP+CNG Cosmopsaltria

Guineapsaltria NNG Cosmopsaltriaria Chlorocysta, Owra, Glaucopsaltria AUS Gymnotympana PP Chlorocystini Venustria Aus Scottotympana, NNG Baeturia Maluku/Bismarck/Solomons/Samoa bloetei group

Fig13 Diagrams of phylogenetic hypotheses for the Prasiini and Chlorocystini cicadas (left) and the Cosmopsaltriaria (right) from Boer (1995b), indicating major instances of overlap in their distributions Areas listed to the left and right are possible Inner and Outer Melanesian archipelago ancestral ranges for each group Central blocked areas indicate areas of overlap that are ambiguous, possibly indicating early dispersal between the two systems or colonisation of intermediately situated areas Broken lines with arrows suggest instances where more recent dispersal may have occurred The foci of endemism within the Chlorocystini follow those recognised by Boer (1995b) The repeated (paralogous) three-area pattern involving Australia is indicated by heavier lines and bold type: the double line to Chlorocysta, Owra and Glaucopsaltria represents a single branch on the cladogram, as the last genus is sister to the first two From Holloway (1997)

to be the Sundanian genus Muda (Fig 13) spathulata (Duffels, 1990) – an extended Philip- Among the Prasiini, the genus Prasia itself is pines-Sulawesi-Lesser Sunda pattern Other endemic to Sulawesi, with seven species (Jong, groups of Lembeja include one in New Guinea, 1985) The genus Arfaka is found in the Bird’s Obi and that is sister to a major Head (Vogelkop) area of New Guinea, and a group endemic to Sulawesi Whilst the species group of taxa currently in Lembeja is restricted of Prasia form an allopatric array distributed to the Lesser Sunda Islands throughout the main island, those of the The Prasiini are completed by sister-genera, Lembeja foliata group are concentrated in the Jacatra, found in Java and Sumatra and thus the north and east of the island, also Sangihe Island only exclusively Sundanian member of the (Jong, 1987), whereas those of the fatiloqua and group, and Lembeja sensu stricto that also con- parvula groups are found in the extreme north tains groups endemic to Sulawesi, but ranges and the extreme southwest (Jong, 1986) There widely from northern Borneo and Mindanao to are two sister-pairs that span these extremes the Lesser Sundas, New Guinea and northern The Lesser Sunda species appear to form an in- Queensland This extensive range is exhibited terrelated group, some still undescribed Twenty by one group, the fatiloqua group (Jong, 1987; species of Lembeja occur in Sulawesi (Duffels, Boer, 1995b) that is found in northern Borneo, 1990) Mindanao, Sulawesi, Sumba, , south Phylogenetic and biogeographic analyses of New Guinea and north Queensland The con- the Chlorocystini by Boer (1995a, b) indicate a nection with northern Borneo appears to have thrice repeated pattern involving Australia as sis- been via Mindanao rather than direct from ter area to a pair of areas of endemism in New Sulawesi – a distribution also shown by Ayesha Guinea: the southeast Papuan peninsula and the Pattern in Lepidoptera and cicadas 309

Tmesisternus New Guinea Tmesisternopsis Sulawesi (+ New Guinea) Elaidius New Guinea TMESISTERNINI Sphingnotus Moluccas to Solomons Sepicana, Pascoea New Guinea Buprestomorpha, Arrhenotoides ENICODINI New Caledonia Blapsilon, Epiblapsilon Trigonoptera New Guinea Temnosternopsis Temnosternus Australia (Queensland) Karadinia, Falsapolia HOMONOEINI Sri Lanka; Sundaland to Philippines & Solomons Tribe separated from ENICODINI New Zealand BUMETOPIINI Philippines to Solomons and Queensland CRINOTARSINI Philippines; New Guinea to Fiji Fig14 Relationships within the Tmesisternini tribal complex of the cerambycid suggested by Gressitt (1984), with distributions also included

central northern part of New Guinea This pat- Hence, despite considerable disparity in the tern is indicated by thick lines in Fig 13 In con- geography of these two tribal groupings out- trast to the Cosmopsaltriaria, only one of three of side Sulawesi, they show at least partial conver- the northern New Guinea components shows a gence in their geography within the island The wide extension into neighbouring archipelagos, higher taxonomic diversity of the Prasiini is as the genus Baeturia with a species group that much in the Banda arcs (Sumatra to Lesser ranges from the Moluccas to Samoa Its Papuan Sundas) as in Sulawesi, and also extends to the peninsula sister-group also shows some exten- eastern and southern periphery of New Guinea, sion, to the northern Moluccas and again to Aus- so the component of convergence with the tralia In one of the other trios there is unique Cosmopsaltriaria may postdate the initial devel- representation of the New Guinea Bird’s Head opment and radiation of ancestral lineages of Thus, whilst the Chlorocystini have their main the tribe There is a further overlap of representation in Australia – at a generic level Cosmopsaltriaria, Prasiini and Chlorocystini in only, as species richness is an order of magni- the New Guinea Bird’s Head, with each contrib- tude lower than in New Guinea: the relevance of uting a genus endemic to, or centred on, that this when considering area relationships is de- area: Rhadinopyga, Arfaka and Aedeastria re- batable (Nelson and Ladiges, 1996) – and south- spectively (Fig 13) There may also be an early eastern and northern New Guinea, the north-south separation of the biota of Sulawesi, Cosmopsaltriaria have their main representation with the weak east-west one mentioned above in archipelagos from the Bismarck Islands to Fiji developing later (see also Holloway, 1997) and Samoa in a sister-relationship to a grouping Similar northern versus southern patterns can in the central and southeastern mountains of be seen in a number of weevil and longhorn New Guinea (Fig 13) The archipelagic group groups (Gressitt, 1956, 1982) The overlaps principally with the Chlorocystini in the Pachyrhynchini weevils are a northern group Bird’s Head and Papuan peninsula, and is also Both northern and southern groups are seen in represented in the Moluccas The sister-group to the longhorn (Cerambycidae) Tmesisternini this complex is in Sulawesi, discussed above, and tribal complex The northern groups are rarely the sister genus to the whole subtribe is sug- represented on Sulawesi, if at all But, unlike gested to be Meimuna, an east Asian genus ex- the Cosmopsaltriaria, all are represented in the tending no further south than the Ryukyu Is- Philippines, the weevil tribe being extremely lands Thus the main feature in common be- diverse there Another good example of a tween these two tribal groupings is rich repre- northern group ranging from the Philippines to sentation in Sulawesi Otherwise they appear to Fiji excluding Sulawesi is the frog genus have developed roughly in parallel, the Platymantis (Allison, 1996) Morley (1998, this Chlorocystini + Prasiini along a more southerly volume) suggests the flora of the eastern archi- axis than the Cosmopsaltriaria, alone of the two pelagos could have been derived from floras in involving Sundaland and the Lesser Sundas the Philippines and Sulawesi, an observation 310 J( D( Holloway that is not inconsistent with such northern and geographic pattern with hypotheses of changes southern patterns The two lineages of Aporosa in geography due to plate tectonic processes discussed by Schot (1998, this volume) also thus is difficult: the paper by Boer (1995b) rep- show parallels resents the most significant attempt to date We Gressitt (1984) had begun to investigate the still need a methodology for pattern analysis that Tmesisternini complex in detail just before his does not introduce the sort of constraints that untimely death In this work he gave an outline the past focus on a hypothetical unique set of of a phylogenetic hypothesis for the complex area relationships has threatened to do A com- This is summarised in Fig 14 The northern plementary combination of R-mode and Q-mode groups referred to above are the Bumetopiini approaches as first advocated by Holloway and and Crinotarsini The Tmesisternini are an es- Jardine (1968) may be one way forward sentially southern group, with a natural group of Michaux (1991, 1994, 1998 this volume) has four genera in Queensland, a genus in Sulawesi already made preliminary observations on the and three distinct groups in New Guinea One of relationship of panbiogeographic, R-mode types these also includes a genus, Sphingnotus, that, of pattern to the geology of the region The like the Baeturia bloetei group in the northern and southern types of distribution dis- Chlorocystini cicadas, is found in the outer cussed above perhaps correlate with his (1994) Melanesian archipelagos, but consists of only Outer Melanesian track (the former) and a com- three rather widespread species bination of his Inner Melanesian track with con- A novel feature of the beetle group relative to tinental Australia and his Inner Banda track, ex- the Chlorocystini is the inclusion of New Cal- cluding Borneo (the latter) edonia in both lineages of the Tmesisternini But there is a prime need for many more The sister-group of the Tmesisternini, the modern phylogenetic treatments of taxa from Homonoeini, ranges widely through the Indo- the region to add to our pattern sample and to Australian tropics, but the Tmesisternini + enable us to explore further the complementa- Homonoeini pairing is itself sister to the rity of cladistic (Q-mode) and panbiogeographic Enicodini, a tribe endemic to New Caledonia (R-mode) approaches to analysis of biogeo- The outgroups include a tribe endemic to New graphic pattern in relation to Earth history We Zealand as well as the two northern archipelagic also need a better understanding of biological distribution tribes mentioned above A partial process (dispersal, speciation, response to cli- explanation of these differences might be that matic factors) to facilitate interpretation of the the ancient history of the beetle group is Aus- patterns derived Sneath (1982) and Page and tralasian/Gondwanan, whereas that of the cica- Lydeard (1994) noted that the sort of geographi- das is Oriental, but this does not necessarily ex- cally progressive cladistic pattern described ear- plain why New Caledonia features within the lier, shown by so many Indo-Australian biologi- southern Queensland-New Guinea-Sulawesi (+ cal groups, could equally well arise through dis- Melanesian archipelagos) patterns in the beetles persal processes as vicariant ones: gradual bio- but not in that for the Chlorocystini + Prasiini logical dispersal through the archipelago from The New Caledonian cicada fauna is entirely west to east or progressive vicariance of land Australian in affinity (Holloway, 1979, pp 234- areas through introduction of sea barriers in the 235) A more exclusively Australian, New Cal- same direction edonian and New Guinea pattern is seen in the Even if there is evidence of correlation be- plant genus Arytera (; Turner, tween biological and geological data in the 1995) structure of area relationships, geologists fre- The equation of gross pattern even between quently ask whether it is possible to date diver- such biogeographically informative groups as gence in biological lineages so as to permit di- the longhorn beetles and cicadas is thus not rect comparison with geological dates This is a without problems: the development of these topic where there is still much disagreement patterns may also involve stochastic processes, Holloway and Nielsen (1998), reviewing and this is not necessarily unexpected in com- cladistic biogeographic analyses of the Mediter- plex archipelagic situations where emergence ranean, noted that area relationships established and convergence of geological structures, and to be common to a number of biological groups change of relative positions, have been much and related to the post-Oligocene history of the more frequent than the sort of sundering and area by some authors were also manifest in a divergence that could lead to clear, repeated phylogenetic hypothesis based on molecular vicariant patterns Objective comparison of bio- data for another group that suggested the events Pattern in Lepidoptera and cicadas 311 concerned were much more recent Intermediate patterns include development of Dating of speciation events that relies on as- an area of endemism in the Philippines that sumption of a molecular clock and uses distance would appear to have had some interaction with data has been strongly criticized by cladistic a component of Sulawesi, such as its northern biogeographers (e(g(, Morrone and Carpenter, peninsula, development of patterns in the 1994), but should not be rejected entirely As Banda arcs that also included interaction with a Page and Lydeard (1994) indicated, the distance component of Sulawesi, probably the south- measures must be ultrametric for them to be western peninsula, and progressive breakdown clock-like (discussed also by Jardine et al(, 1969), of the segregation of the New Guinea areas of and care must be taken with the analyses If endemism from those of the Melanesian archi- these conditions are satisfied, such analyses pelagos The development of the Cosmopsaltria could well provide useful additional information pattern endemic to the New Guinea areas could to test concordance between biogeographic pat- have preceded considerably the expansion of tern and geological hypotheses (Page and the Chlorocystini into the outer archipelagos in Lydeard, 1994; Hedges et al , 1994) the form of the Baeturia bloetei group, and also The complexity of the Indo-Australian tropics the extension of Diceropyga of the Cosmopsaltriaria will undoubtedly require the sort of pragmatic into the Papuan peninsula This interpretation approach adopted by Funk and Wagner (1995) departs somewhat from that of Boer (1995b), in interpreting pattern in various plant and ani- though he also considered differential timing of mal groups that have speciated in the Hawaiian development of the various patterns The inter- archipelago They developed a series of hypo- action of structures between Sulawesi and New thetical patterns that might be expected under Guinea with the major foci of endemism is still different process assumptions (rate of evolution not clear Cicada patterns involving the Bird’s relative to dispersal, mode and time of colonisa- Head of New Guinea are complex, with interac- tion, direction of dispersals, etc ) in relation to tion seen with Sulawesi, New Guinea and current knowledge of the geological evolution Melanesian archipelago (Solomons) groups of the Hawaiian island chain, and compared (Fig 13) None of the islands comprising the cur- them with actual patterns The most frequent rent Moluccas appears to have acted as a strong pattern was of the progression type, with disper- centre of endemism, their faunas bearing a gen- sal from older to younger islands, this progres- eral relationship to neighbouring areas, though sion being in the form of clades or grades in more so with areas to the east than to the west relation to the islands, dependent on the relative (see also Jong, 1998 this volume) frequency of speciation relative to dispersal Patterns that are probably progressively more This last distinction might equally be applicable recent are the allopatric and widespread catego- to the differences observed within similar gross ries among Oriental taxa The former, where patterns in Lepidoptera and cicadas, leading to progressive, do not appear to discriminate sepa- much greater species-richness within each pat- rate areas of endemism, or composites in tern element of the latter, the paralogies of Nel- Sulawesi or New Guinea The species of the lat- son and Ladiges (1996) ter, more often than not, transcend the older ar- Is it possible, therefore, to represent the pat- eas of endemism and, through stochastic inter- terns discussed in this chapter in some sort of changes between islands, enable a strong signal sequence that might have some bearing on the from modern geography to be recaptured in geological history of the area? The basal group- scaling analyses such as in Holloway and ings in many of the more complex patterns in Jardine (1968) Whilst tending to obscure signals the Lepidoptera or cicadas are: Oriental- from the past, these groups may well be estab- Sundanian; tropical Australia with New Guinea lishing the foundations of signals that will be or New Guinea alone, possibly with a focus on detected in the distant future the Papuan peninsula and the Central and Sepik structures of the latter; outer Melanesian archi- pelagos, possibly also including some compo- Acknowledgements nents now seen as northern structures within New Guinea; Sulawesi, possibly a component of This chapter is based in part on a presentation the current island that was relatively isolated made in 1993 at the 13th meeting of the Willi from both Oriental and Papuan land areas at that Hennig Society in Copenhagen and in part on time Establishment of these patterns is therefore that made at the formative meeting for this book likely to have been relatively early in 1996 It owes much to interaction through the 312 J( D( Holloway years with Hans Duffels, Arnold de Boer, Dan tralian Systematic Botany 4: 117-128 Polhemus, Dick Vane-Wright, Rienk de Jong and Darlington, P J Jr 1957 Zoogeography: The Geographical Distribution of  John Wiley and Sons, New York others perplexed by the complexities encoun- Duffels, J P 1983 , phylogeny and tered in the biogeography of the Indo-Australian biogeography of the genus Cosmopsaltria, with remarks tropics, and therefore embracing the ‘heresy’ of on the historic biogeography of the subtribe searching beyond the goal of a unique structure Cosmopsaltriaria (Homoptera: ) Pacific for area relationship within the archipelago The Monographs 39: 1-127 Duffels, J P 1988 The cicadas of the Fiji, Samoa and Tonga coefficients for the analyses were calculated us- islands, their taxonomy and biogeography (Homoptera, ing software written by Gaden Robinson I am Cicadoidea) Entomonograph 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