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Biogeographical observations on the PhilippinePapilionoidea (Lepidoptera)Tommaso Racheli a & Maurizio Biondi aa Dipartimento di Biologia Animale e dell'Uomo (Zoologia) , Università ≪LaSapienza≫ , viale dell'Università 32, Roma, I‐00185, ItalyPublished online: 28 Jan 2009.

To cite this article: Tommaso Racheli & Maurizio Biondi (1989) Biogeographical observations on the PhilippinePapilionoidea (Lepidoptera), Bolletino di zoologia, 56:4, 333-347, DOI: 10.1080/11250008909355660

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Boll. Zool. 56: 333-347 (1989)

Biogeographical observations on thePhilippine Papilionoidea (Lepidoptera)

TOMMASO RACHELIMAURIZIO BIONDIDipartimento di Biologia Animale e dell'Uomo (Zoologia),Università «La Sapienza»,viale dell'Università 32, I-00185 Roma (Italy)

ABSTRACT

The Papilionoidea of the Philippines and Palawan were studiedwith different statistical methods to show distribution and faunisticsimilarities. The results can be summarized as follows: a) Palawanis significantly different from the Philippines, while the similaritiesare virtually undistinguishable from Borneo and Malaysia; b) thehigh percentage of taxa endemic to the Philippines and the lowfaunistic similarity with the surrounding areas of Malaysia, Borneoand Taiwan demonstrate that the butterflies of the Philippines arewell characterized; c) the total number of species on each island issignificantly correlated with the island area and less influenced bythe distance from the source-pools; d) the total number of endemictaxa on each island is highly correlated with the island area; e) thereare different behaviours in the relation species-area in each family.

KEY WORDS: Island biogeography; Lepidoptera; Philippines.

ACKNOWLEDGEMENTS

The idea of the present paper was crystallized during a trip tothe Philippines of T. Racheli with his late friend P. Brignoli. Theauthors wish to thank the Trustees of the British Museum of Na-tural History, London, for access to Philippine specimens' underthe care of the Staff of the Rhopalocera section. Papers, informa-tion and criticism were received from P. Ackery, M. Audley-Charles, J. C. Balouet, A. C. Cassidy, J. P. Duffels, L. R. Heaney, J.D. Holloway, R. I. Vane-Wright and A. Zilli. N. Falchi drew thetables and graphics. Our sincere thanks for their valuable support.

(Accepted 22 May 1989)

INTRODUCTION

The Philippines constitute a fringing archipelago ex-tending in a N-S direction for nearly 2000 km andwithin the triangle Palawan-Lu2on-Mindanao they em-brace more than 7000 islands (Fig. 1).

Wallace (1860) in his classic paper on the definitionof the limits of the Australian and Indian regionsplaced the Philippine islands in the latter region butcommenting that «... they are in some respects ofdoubtful location...)). Later, he placed the Philippinesdefinitively in the Oriental region (Wallace, 1876).Since then a vast number of reports have shed morelight on the biotic differences between the two regions,without however clarifying the position of the Philip-pines. Wallace's line and the other lines drawn for theArchipelago are discussed in many studies based ondifferent theoretical and methodological approaches(Raven, 1935; Scrivenor, 1943; Mayr, 1944; Holloway& Jardine, 1968; Lincoln, 1975; Simpson, 1977; Whit-more 1981, 1987; Keast, 1983). Perhaps the moreenlightening contribution is that of Simpson (1977)who commenting upon the number of «lines», citesHolloway & Jardine's (1968) paper, stating that:«From their figures and maps, it appears that supportfor almost any of the seven lines enumerated above canbe found, and also evidence against any of them andfor and against still others».

Wallace's line does not seem to constitute a realbarrier when insects are considered (Gressitt, 1961).Corbet (1943) reported that the Large Sunda islandsshare only a few of 28 genera of lycaenids with thePhilippines, and hence he favours Huxley's line.

The purpose of the present paper is to analyze thedistribution of some groups of Lepidoptera in the ma-jor islands of the Philippines in comparison with Tai-wan, Borneo, Palawan and Malaysia. Sulawesi and theSunda Islands have not been included in the analysisbecause distributional data of Rhopalocera are incom-plete.

MATERIAL AND METHODS

A total of 330 species of Papilionoidea belonging to the Papi-lionidae, Pieridae, Danaidae, Nymphalidae and Satyridae were in-cluded in this study. Lycaenidae were omitted because of scarcetaxonomic and distributional data. The data are based on an ex-haustive review of early reports and of recent faunistic and syste-matic papers on the butterflies of the Philippines, especially thoseby Yata (1981), Aoki et al. (1982), Tsukada & Nishiyama (1982)and Ackery & Vane-Wright (1984). Other sources were specimensin the collections of the British Museum (Nat. Hist.) London,unpublished data from various contributors and material collectedin the Philippines by T. Racheli during 1983.

A matrix for binary data (absence/presence) was compiled forthe species occurring on each Philippine island (Luzon = LUZ;Mindoro = MRO; Panay = PAN; Samar = SAM; Negros = NEG;Leyte = LEY; Cebu = CEB; Bohol = BOH; Mindanao = MIN) andPalawan (PAL).

a) A first similarity analysis was carried out using the coefficient

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334 T. RACHELI, M. BIONDI

Fig. 1 - Map of the Philippines.

of Baroni Urbani & Buser (1976). This was more suitable for thepresent approach and more indicative than other overall similarityindices (Biondi, 1988). The equation of this index reads as follows:S** = (Vad + a) / (Vad + a + b + c), where a = number of OTUs(Operational Taxonomic Units) in common between the two is-lands to be compared; b = number of OTUs present in the first butnot in the second island; c = the number of OTUs present in thesecond but not in the first island; d = the number of OTUs absentin both islands compared but present in the others considered.

b) A second similarity analysis was carried out using the re-semblance equation of Preston (1962). This index has been specifi-cally proposed for studies of insular biogeography. It is stressed thatthe results obtained by this index take into account indirectly thedifference among the island areas. Its equation is: a " ( l / z ) + b"(1/z) = 1, where a = the proportion of the joint fauna foundin one primary area, b = the proportion of the joint fauna found inthe other and z = the dissimilarity index. To compare the dendro-grams, the similarity index s = 1 — z was used.

The similarity values obtained by both analyses were subjected tocluster anal> sis according to the flexible method of Lance & Williams(see Sneath & Sokal, 1973), using the parameter «beta» = — 0.25.

The same statistical procedures have been used for endemic taxa(N = 138) and for each single family. Here endemism is consideredas every identifiable taxonomic unit proper to the Philippine islands+ Palawan or part of this complex.

c) To evaluate faunistic affinities, the Papilionoidea faunas ofthe Philippines and Palawan and those of Borneo, Malaysia andTaiwan were compared using the same procedures described in a).For comparison in the latter case the principal coordinates analysis(PCO) was used (see Gower, 1966; Digby & Kempton, 1987).

d) A multiple regression analysis was performed using the for-ward method with a significance level set at 51. As dependentvariables the total number of species and endemics on each islandwere compared respectively with physical parameters such as alti-tude, area, average distance of each island from mainland Asia,Borneo, Mindanao and Luzon (Table I). Elevations and distancesconsidered were obtained from Operational Navigation Charts(Defense Mapping Agency Aerospace Center, St. Louis Air ForceStation, Missouri). Surfaces are according to Heaney (1984, 1986)and Lawlor (1986). Each parameter has been lognormally consi-dered (base e).

RESULTS

The similarity analysis with Baroni Urbani &. Buser'sindex and cluster analysis and PCO performed on thetotal number of Papilionoidea (see Figs. 2a, b, 4a)demonstrates that: 1) Palawan is well apart from thePhilippines because the similarity is high with Borneoand Malaysia, both of which act as a source of itsfauna, and less with Taiwan. 2) The Philippines aredivided into two main clusters: one including Luzonand Mindanao, another all the remaining islands. Be-cause Luzon and Mindanao are the largest islands, theyhave a high number of species, most of which have awide distribution. The similarity is due largely to thecommon presence of wide-range species in the twoislands. 3) The similarity analysis on the endemics ofLuzon and Mindanao (Fig. 3b) with respect to all theremaining islands of the archipelago, demonstrates thatthe geographical factor affects the distribution of thesetaxa. In fact, the northern islands (Mindoro, Panay,Samar and Negros) have a higher faunistic affinity withLuzon, while the southern ones (Cebu, Leyte and Bo-hol) are more similar to Mindanao. The same analysiswas performed considering the Papilionoidea in toto(Fig. 3a), but it did not provide any additional data.

The dendrograms (Fig. 4c, d, e, f, g) obtained bythe analysis (Baroni Urbani & Buser's index + clusteranalysis) on each single family show that the distinctive-ness of Palawan from the rest of the Philippines isfairly constant, except for Pieridae and Danaidae (Fig.4d, e) which include many high vagile species. In fact,in the case of these two families, Palawan shows asignificative affinity to Luzon and Mindanao. This isbecause also Palawan, which on the basis of our resultsbehaves as a landbridge, has a considerable number ofspecies with a high percentage of wide range elements(80.461). The affinity of Palawan to Cebu in the Pieri-dae is probably ascribable to the extinction of someendemic species following high deforestation. Interest-ingly, based on Satyridae (Fig. 4g), the islands are dis-tributed into two well-defined clusters: a south-oriental(Mindanao, Bohol, Leyte and Samar) and a north-

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BIOGEOGRAPHICAL OBSERVATIONS ON THE PHILIPPINE PAPILIONOIDEA 335

TABLE I - Total number of species and endemics on each island and physical parameters.

A C . D H J K M N O

PALLUZMROPANSAMNEGLEYCEBBOHMIN

1047102312401522152214221516157616381728

503087133206233295247294563

4887125362331383101471831150

3491113779960858733671737737609

11785108171973513429721312297136704421386499078

2086293025858501349176524381013792

2954

8126810645814

38413035112

2812224116

152917612610161231

62117121710129920

34784529442433363183

32271919201513181827

273717612717211141

2126612141218111023

71644315321624443372

23332822301725172041

17418711374108679611192204

A: average distance (Km) from continent; B: average distance (Km) from Luzon; C: average distance (Km) from Mindanao; D: averagedistance (Km) from Borneo; E: area (Km); F: altitude (m); G: number of endemic species of Papilionidae; H: idem, of Pieridae; I: idem,of Danaidae; J: idem, of Nymphalidae; K: idem, of Satyridae; L: total number of endemic species; M: number of species of Papilionidae; N:idem, of Pieridae; O: idem, of Danaidae; P: idem, of Nymphalidae; Q: idem, of Satyridae; R: total number of species.

1.0 .5

PAL"

MAL"

BOR-

TAI-

LUZ-

MIN-

MRO-

CEB-

SAM"

LEY"

BOH-

PAN"

NEG"

AXIS I

B

MM LEVLUZ SAM

MRO BOHCEB

— ; PANNEG

Fig. 2 - A: Similarity analysis among the Philippine islands and their main source areas. B: Similarity analysis among the Philippine islandsand their main sources areas performed with principal coordinates analysis (PCO).

western faunistic block (Cebu, Negros, Panay, Luzon,Mindoro and Palawan). This situation represents therelatively low vagility of the species belonging to thisfamily, and the two faunistic blocks may reflect differ-ent ways of colonization.

The dendrograms obtained with Preston's indexconfirm the faunistic diversity of Palawan from thePhilippines proper (Fig. 5). Substantial differences inthe intra-archipelago similarity emerge from a compari-son of Preston's index and the Baroni Urbani & Buser'sindex. In fact, the similarity between Luzon and Min-

danao is remarkably lower with Preston's index, andthere is a geographic trend in the shaping of the clus-ters. The differences between the two analyses are dueto the distinct mathematic nature of the two indices.Baroni Urbani & Buser's index is an overall similarityindex of general application for Boolean matriceswhere the parameters considered are the total numberof positive (1-1) and negative (0-0) matches and mis-matches (1-0; 0-1) (Baroni Urbani & Buser, 1976).Preston's index is based on much more complexmathematic presuppositions and, as quoted above, is

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336 T. RACHELI, M. BIONDI

3 .3-E" .2-

.1 •

0 -

•7 »

.6 S

OUJ

Z

Xo

B1

.9

.8

.7 m3

.6 5T

O

Z

Fig. 3 - Similarity within the Philippines with respect to Luzon andMindanao. A: total number of species. B: endemic species.

specifically suitable for insularity topics. The calculus ismore difficult than that of the other index, and requiresa computer or Preston's table (Preston, 1962, p. 419)to solve the resemblance equation. When Preston's tableis used the value of the index is somewhat approxi-mate. This explains the different results obtained withthe two analyses. In particular, the position of Luzonand Mindanao in the dendrograms could be due to thegreat difference of the island areas, which influencesPreston's index.

Holloway and Vane-Wright (personal communica-tion) found that the LUZ + MIN connection given bythe similarity analysis using Baroni Urbani & Buser'sindex was spurious because of the inadequacy of themethod. However, the dendrogram shown in Figure4a supports the actual objective sharing out of endemictaxa and the Pleistocenic eastern track along the Vi-sayas that connects the two major islands, as was pre-dicted but not corroborated by data on mammals(Heaney, 1985). The distributions of species such asPachliopta phegem (Hopffer, 1866), Pachliopta mariae(Semper, 1878), Graphium idaeoides (Hewitson, 1885),Eurema sarilata (Semper, 1891), Euploea tobleri Semper,1878, Ptychandra schadenbergi (Semper, 1887), Ptjchandraleytensis Banks, Holloway & Barlow, 1976, tread in aneastern track, while Pachlioptaphlegon (Felder & Felder,

1864), Tirumala ishmoides Moore, 1883, Larippa pata(Moore, 1858), Bassarona piratica (Semper, 1888) in awestern track.

The similar climatic indices of Luzon and Mindanao(Whitmore, 1975) may have influenced the present-day distribution of types of vegetation and hence of thebutterflies; furthermore these islands have constitutedPleistocene rain forest refugia. Delias themis Hewitson,1861, Delias nuydaorum Schroeder, 1975, Delias baracasaSemper, 1890, Appias aegis (Felder & Felder, 1861),Pantoporia epira (Felder & Felder, 1863) may representrefugial entities in Luzon and Mindanao.

Therefore, both similarity indices, even if they cangive different results, are equally useful for a morecomplete interpretation of biogeographic events.

To obtain further information on the faunistic affini-ties of the insular complex, a similarity analysis thatconsiders only the endemisms in the sense previouslydefined has been carried out to exclude an eventuallevelling effect caused by the elements of wide distribu-tion. The results are as follows (Figs. 4b and 5b): 1)the separation of Palawan from the Philippines is em-phasized even more. 2) As far as the Philippines areconcerned, the dendrogram obtained by Baroni Urbani& Buser's index contains two clusters, the former con-sisting of Luzon, Mindoro and Mindanao, the latter allthe remaining islands (Fig. 4b). When Preston's indexwas used, the dendrogram shows a geographical trend:the south-eastern and the north-western islands, re-spectively (Fig. 5b).

The regression analysis shows a significant correla-tion between the number of species and island area(Fig. 6a, b). No correlation was obtained with altitudeor distance from mainland or from large islands, withthe exception that the Pieridae are significantly corre-lated with altitude. The coefficients of correlation ob-tained with the regression analysis for area and altitudeare reported in Table II: The total number of the Papi-lionoidea reveals a coefficient of correlation r = 0.64(level of significance 4.32). Considering endemics, thecoefficient of correlation r equals 0.78 (level of signifi-cance 0.822). Different behaviour is shown in eachfamily because the faunistic data are not exhaustive andalso because of the different ecology of these taxa.Moreover, the species-area relationship is, particularlyfor archipelagos, often based on simplistic conjectures,while many other important parameters are not exa-mined, e.g., the influence of the interaction amongislands. Hence, islands within an archipelago can host arelatively higher number of species than islands ofequivalent area but isolated. This can be the case ofCebu and Bohol. These two islands, the innermost inthe archipelago, show the highest values for speciesdensity and the lowest values for the endemism rate(Table III). This is due to the influence of the nearbyislands: therefore Cebu and Bohol have the typical roleof «transit areas» with a behaviour analogous to con-tinental islands. The lower value of species density for

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BIOGEOGRAPHICAL OBSERVATIONS ON THE PHILIPPINE PAPILIONOIDEA 337

1.0

PAL -

L U Z -

M I N -

MRO-

CEB-

S A M -

LEY-

BOH-

PAN-

NEG-

P A L -

L U Z -

MIN -

SAM-

LEY -

BOH-

CEB -

MRO

PAN

NEG

B

P A L -

C E B -

L U Z -

M I N -

M R O -

NEG-

P A N -

S A M -

L E Y -

BOH-

PAL-

LUZ -

MIN -

MRO-

SAM-

BOH-

PAN-

LEY -

NEG-

CEB-

PAL-

LUZ-

MRO-

PAN-

CEB-

NEG-

SAM-

LEY -

BOH-

MIN -

Fig. 4 - Similarity analysis within the Philippines performed with Baroni Urbani & Buser's index and cluster analysis. A: total number ofPapilionoidea; B: endemic Papilionoidea; C: Papilionidae; D: Pieridae; E: Danaidae; F: Nymphalidae; G: Satyridae.

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338 T. RACHELI, M. BIONDI

NEG-

PAN-

MIN-

BOH-

LEY-

SAM-

CEB-

MRO-

LUZ-

PAL-

1.0

L E Y -

SAM—

CEB —

MIN-1

BOH—'

PAN—

MRO—l

LUZ—•

NEG-l

PAL-"

1.0

C

.8

PAN-

CEB-

LEY-

SAM-

BOH-

MIN-

NEG-

MRO-

LUZ-

PAL-

— '

J

PAN

SAM

MRO

BOH-

MINi

NEGj

LEY'

CEB-

LUZ-

PAL-

— '

hIH

MIN-

BOH-

LEY-

SAM-

NEG-

CEB-

PAN-

MRO-

LUZ-

PAL-

Fig. 5 - Similarity analysis within the Philippines performed with Preston's index and cluster analysis. A: total number of Papilionoidea; B:endemic Papilionoidea; C: Papilionidae; D: Pieridae; E: Danaidae; F: Nymphalidae; G: Satyridae.

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BIOGEOGRAPHICAL OBSERVATIONS ON THE PHILIPPINE PAPILIONOIDEA 339

D

5.5

COJJ

</>

u.ocrLUB

| 4 . 5

4

-

•

CEB•

BOH

A

PALD

MRO ^ ^ - ^ * ^ ^LEY Q ^ < * * ^

a ^^*^^^-***'^ SAM

^ ^ ^ D

NEG

D

f i l l

MIN•LUZ ^ ^

TABLE III - Endemism rates and density in each island. Density (DENS)= (number of species / area) X 100.

8.5 9.5 10AREA (log)

10.5 11.5

Fig. 6 - Species area relation. A: total number of species (y =0.216 x + 2.672); B: endemic species (y = 0.278 x + 1.020).

the outer island of Samar (DENS = 0.551) supportsthis view. The endemism rate (ER): number of ende-mics/total number of species in each island (Table III)shows somewhat homogeneous values (0.324 < ER^ 0.417) for the Philippines, while the value is consi-derably lower (ER(PAL) = 0.195) for Palawan. Thisunderlines the continental features of the latter island.

The highest ER values were obtained for Mindanao,Luzon and Leyte (ER(MIN) = 0.407; ER (LUZ) =0.417; ER (LEY) = 0.407). In the former two islands,these results can be attributed to the high altitude andlarge surface of Mindanao and Luzon, while Leyte hasa bridge effect with the two major islands of the Philip-pines.

The slope parameter (z) obtained for the Philippines(0.216) falls in the predicted range of values and in asituation of islands verging on oceanic characteristics.If Palawan is excluded, the z-value increases (0.225)

PALLUZMROSAMLEYPANNEGCEBBOHMIN

PDEA

0.190.420.400.390.410.360.340.320.340.41

PDAE

0.250.440.320.420.500.400.310.280.440.52

PIER

0.110.220.230.170.250.000.180.240.090.29

DANA

0.090.310.170.170.140.170.220.090.100.26

NYMP

0.210.450.390.400.370.370.420.360.360.43

SATY

0.260.640.610.54.0.570.590.480.530.450.49

DENS

1.480.171.160.551.500.540.702.562.380.21

thus supporting the hypothesis that Palawan is a land-bridge (see Heaney, 1986).

It has been suggested (Martin, 1981; Williamson,1981; Wright, 1981; Case & Cody, 1983; Lawlor,1986) that the species/area relationship and hence thez-value is biased both by the taxa under considerationand by the environmental diversity of the archipelagoitself. Therefore, we did not compare the slope para-meter obtained for the Philippines with others derivedfrom similar studies on Lepidoptera in different ar-chipelagos (Scott, 1972; Gross, 1975; Hockin, 1980;Miller, 1985).

CONCLUSIONS

Districts within the Philippines were defined bySemper (1892) who based his assumptions on the dis-tribution of the endemic Rhopalocera known at thattime. He recognized six districts: 1) Luzon; 2) Min-doro; 3) Palawan; 4) Panay, Negros, Cebu; 5) Samar,Leyte, Bohol; 6) Mindanao. According to Pagenstecher(1909), Palawan is a bridge between Borneo and thePhilippines, its rhopalocerous fauna being predomi-nantly Bornean. Schultze (1928) found that the faunaof Palawan was more related to that of Borneo thanthat of the Philippines.

Ackery & Vane-Wright (1984), on the basis of thedistribution of Danainae, subdivided the Philippines inseven zones: Palawan, Sulu Archipelago, Mindoro, Lu-;zon, Western Visayas (Negros, Cebu, Panay), EasternVisayas (Samar, Leyte, Bohol), Mindanao.

Comparisons between taxa endowed with differentdispersion ability are difficult to make. However, forbirds, Delacour & Mayr (1946) distinguished twomain provinces and three marginal districts. An East-ern Province with, besides small islands, Luzon, Min-danao, Samar, Leyte and Bohol and a Central Provincewith Panay, Negros and Cebu. The three districts in-

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TABLE II - Correlation coefficients from the regression analysis.

Total speciesEndemic species

Total speciesEndemic species

PDEA

0.64*0.78*

0.620.61

PDAE

0.510.64*

0.320.20

PIER

0.590.58

0.67*0.56

DANA

0.70*0.92*

0.450.66

NYMP

0.420.59

0.510.57

SATY

0.74*0.64*

0.650.48

* = significant at 51 /PDEA = Papilionoidea; PDAE = Papilionidae; PIER = Pieridae; DANA = Danaidae; NYMP = Nymphalidae; SATY = Satyridae.

elude Mindoro, the Babuyan groups and the Sulu ar-chipelago. Palawan is faunistically considered to belongto the Malaysian subregion. Diamond & Gilpin's(1983) data on Philippine birds are suggestive of astepping-stone model between the continent and thePhilippines. The distribution of the endemic hornbillsBuceros hydrocorax and Penelopides panini resembles therelations shown by endemic Papilionoidea (Fig. 4c).Very similar results were obtained for land-snails(Cooke, 1892), amphibians (Taylor, 1928; Inger,1954) and reptiles (Brown & Alcala, 1970). More re-cently, Heaney & Rabor (1982) and Heaney (1985,1986) have suggested, on the basis of terrestrial mam-malian fauna, four major regions based on richness,composition and degree of endemism, namely Pala-wan, Mindanao, Luzon and Negros.

To compare these results with those on butterflies,widespread species must be excluded and only theendemic taxa should be taken into consideration forvicariance purposes. Unfortunately only a few endemictaxa have been examined phylogenetically. The distri-bution of the Zethera complex (Vane-Wright & Smiles,1975) supports the views of Semper (1892) and corre-sponds fairly well with our results, particularly withthe endemic Papilionoidea (Figs. 4b, 5b).

Because our analysis was carried out mainly on theintra Philippine relations rather than with other neigh-bouring islands, no comparison can be made withother Wallacean islands. The importance of the islandof Sulawesi, its strange shape and central position inthe Indonesian archipelago, and the high number ofendemic taxa have long attracted the attention of na-turalists. As far as Lepidoptera are concerned, Hollo-way & Jardine (1968) and Holloway (1973) showedphenetically and phylogenetically that Sulawesi is relat-ed to the Philippines. More recently, Holloway (1987)used phylogenetic analysis to examine the Notodonti-dae and Limacodiidae of Sulawesi, and showed an as-sociation with the Philippines. This is congruent withthe hypothesis of the geological formation of Sulawesi, .

from convergent oriental and Australasian compo-nents, the former part has been isolated for a longertime from Borneo than it has from the Philippines(Audley-Charles, 1981, 1987). A Philippine-Sulawesilink has also been inferred by Kitching et al. (1987) onthe basis of the distribution and cladistics of Idea (Dan-ainae). The areas-species relationship (Irianjaya + Ma-luku, Sulawesi, Philippines) and (durvillei + idea, blan-chardi, electro.') strongly supports a Moluccas-Sulawesi-Philippines link. This link is supported by floweringplants (Balgooy, 1987), but not by mammals exceptbats (Musser, 1987).

Atrophaneura semperi (Felder & Felder, 1861) a Phi-lippine endemism, with a peculiar subspecies A. semperimelanotus (Staudinger, 1889) in Palawan and its sisterspecies A. kuehni (Honrath, 1886) in Sulawesi, maysuggest an ancient Philippine-Sulawesi link and a morerecent relation with Palawan.

The present results confirm that Palawan is faunisti-cally strictly oriental, even though it has acted and isacting as a two-way corridor for some taxa.,This issupported by Ptychandra talboti Hobby, 1940, found inBorneo with the rest of its representative only in thePhilippines and Palawan (Banks et al., 1976). Graphiummegarus (Westwood, 1845) and G. megaera (Staudinger,1888) are reciprocally exclusive of Sundaland and Pala-wan. Graphium stratocles (Felder & Felder, 186li) with anumber of populations scattered in the Philippines, oc-curs also in Palawan where its sister species, G. maca-reus (Godart, 1819) an oriental element, is at;its eas-tern limit. The Graphium antiphates complex has thesame pattern of distribution. The presence of-Philip-pine endemisms at species or superspecies level may bethe expression of a dispersal due to uplift of the seafloor by 160 m, which occurred during the "middlePleistocene, approx. 160 000 years ago (Heaney,1985). Euploea swainson (Godart, 1824) and- Euploeablossomae Schaus, 1929, western Philippine elements,support this hypothesis. :

The presence in north Luzon of strictly derived pa-

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leafctic species such as Papilio chikae Igarashi, 1965,and Papilio benguetanus Joicey & Talbot, 1923, may bedue to a sweepstake route via Taiwan, as opposed tothe «normal» south-north trend of typhoons as exem-plified by three Philippine danaids present in Taiwan,Danaus melanippus (Cramer, 1777), Euploea camaralze-man Butler, 1866, and Parantica luzonensis (Felder &Felder, 1863).

The papilionid genus Pachliopta comprehends a dozenspecies throughout its range from India to New Guineaand in the Philippines it has undergone radiation andspeciation with six endemics. The changes in watergaps during the last glaciations and the formation ofland bridges, other than over water dispersal, may ex-plain in many instances the actual distribution of but-terflies, as reported for non-volant mammals by Hean-ey (1986).

Vicariance and dispersal undoubtedly underlie thecomposition of the biotic scenario of the Philippines.Recent Pleistocenic phenomena rather than the com-plex geological history of the archipelago, have in-fluenced butterfly distribution as exemplified by thepresence of endemic or almost endemic genera such asZethera and Ptychandra, of sister and sibling species inPachliopta, Graphium, Delias, Danaus, Elymnias etc. andof the high number of endemics at species level. How-ever, ecological factors, differences in dispersal abili-ties of the considered taxa, habitat diversity, competi-tive displacement, particular niches of some species,arid reduction of natural habitats by man have allplayed a role in the distribution of Philippine butter-flies.

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Aoki T., Yamaguchi S., Uémura Y., 1982 - Satyridae, Libytheidae.In: E. Tsukada (ed.), Butterflies of the Southeast Asian islands,3. Plapac, Tokyo, 628 pp.

Audley-Charles M. G., 1981 - Geological history of the region ofWallace's line. In: T. C. Whitmore (ed.), Wallace's line and platetectonics. Clarendon Press, Oxford, pp. 24-35.

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Cooke A. H., 1892 - On the geographical distribution of the land-mollusca of the Philippines Is., and their relations to the Mol-lusca of the neighbouring groups. Proc. zool. Soc. London,1892: 447-469.

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Diamond J. M., Gilpin M. E., 1983 - Biogeographical umbilici andthe origin of the Philippine avifauna. Oikos, 41: 307-321.

Dickerson R. E., (ed.), 1928 - Distribution of life in the Philip-pines. Bureau of Science, Manila, 322 pp.

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Holloway J. D., 1973 - The taxonomy of four groups of butterfliesin relation to general patterns of butterfly distribution in theIndo-Australian area. Trans. R. ent. Soc. London, 125:125-176.

Holloway J. D., 1987 - Lepidoptera patterns involving Sulawesi:what do they indicate of past geography? In: T. C. Whitmore(ed.), Biogeographical evolution of the Malay Archipelago.Clarendon Press, Oxford, pp. 103-118.

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Whitmore T. C. (ed.), 1987 - Biogeographical evolution of theMalay Archipelago. Clarendon Press, Oxford, X-147 pp.

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APPENDIX - Papilionoidea of the Philippines and their distribution (presence/absence) amongst the islands.

~V\ A T1TT T/~\X T T r \ A T~*

PAPILIONIDAETroides rhadamantusTroides magellanusTrogonoptera trojanaPachliopta aristolochiaePachliopta atroposPachliopta phlegmPachliopta mariaePachliopta phegeusPachliopta schadenbergiAtrophaneura semperiAtrophaneura neptunusPapilio rumanzovia/lowiPapilio karnaPapilio chikaePapilio palinurusPapilio demotionPapilio benguetanusPapilio demoleusPapilio antonioPapilio helenusPapilio hipponousPapilio polytes alphenorChilasa osmanaChilasa paradoxaChilasa carolinensisChilasa clytiaGraphium codrusGraphium sarpedonGraphium sandawanum

PAL

10111000011110110101110101110

LUZ

11010110110101101101110001110

MRO

10010100010100100001110001110

SAM

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LEY

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PAN

10010110010100100001110000110

NEG

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CEB

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BOH

11010011010100100001110000110

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11010111010100100011111011111

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BIOGEOGRAPHICAL OBSERVATIONS ON THE PHILIPPINE PAPILIONOIDEA 343

APPENDIX - Continued.

PAL LUZ MRO SAM LEY PAN NEG CEB BOH MIN

Graphium aryclesGraphium dosonGraphium eurypylusGraphium agamemnonGraphium bathyclesGraphium dekssertiGraphium megaeraGraphium stratoclesGraphium macareusGraphium idaeoidesGraphium aristeusGraphium decolorGraphium euphratesLamproptera megesLamproptera curius

PIERIDAEh.eptosia ninaDelias themisDelias georginaDelias blancaDelias nuydaorumDelias henningiaDelias woodiDelias pasithoeDelias apoensisDelias levickiDelias diaphanaDelias baracasaDelias hypareteDelias mandayaPieris canidiaCepora boisduvalianaCepora aspasiaAppias lynddaAppias albinaAppias waltraudaeAppias nepheleAppias neroAppias indraAppias pandioneAppias remediosAppias libytheaAppias paulinaSaletara liberiaSaletara pandaSaletara nathaliaPhrissura aegisHebomoia glaucippeCatopsilia pyrantheCatopsilia pomonaCatopsilia scyllaCatopsilia florellaPareronia Valeria/boeberaPareronia phocaea/nishiyamaEurema alithaEurema harinaEurema adaEurema blandaEurema andersoniEurema simulatrixEurema sarilata

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(Continued)

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344 T. RACHELI, M. BIONDI

APPENDIX - Continued.

PAL LUZ MRO SAM LEY PAN NEG CEB BOH MIN

Eurema laetaEurema biuraiIxias darki

DANAINAEIdeopsis gauraIdeopsis juventaIdeopsis vulgarisIdea leuconoeIdea electraDanaus chrysippusDanaus melanippusDanaus affinisTirumala limniaceTirumala septentrionisTirumala ishmoidesTirumala choaspesTirumala hamataParantica aspasiaParantica vitrinaParantica luzonensisParantica phyleParantica dannattiParantica sitaParantica milagrosParantica davidiParantica schoenigiEuploea camaralzemanEuploea mulciberEuploea midamusEuploea phaenaretaEuploea euniceEuploea blossomaeEuploea tulliolusEuploea swainsonEuploea tobleriEuploea eyndhoviiEuploea SylvesterEuploea algea

NYMPHALIDAEAriadne merioneAriadne taeniataCupha ariasCupha erymanthisPhalanta phalantaPhalanta alcippeVagrans egistaArgynnis hyperbiusVindula erotaPaduca fas dataCirrochroa tycheCirrochroa menonesCirrochroa satellitaTerinos clarissaCethosia biblisCethosia hypseaCethosia luzonicaJunonia iphitaJunonia hedoniajunonia lemoniasJunonia orithyaJunonia almana

000

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BIOGEOGRAPHICAL OBSERVATIONS ON THE PHILIPPINE PAPILIONOIDEA 345

APPENDIX - Continued.

PAL LUZ MRO SAM LEY PAN NEG CEB BOH MIN

Vanessa indicaVanessa dejeaniKaniska kanaceSymbrenthia annaSymbrenthia hypselisSymbrenthia hypatiaKhinopalpa poly niceYomo sabinaHypolimnas anomalaHypolimnas misippusHypolimnas bolinaDoleschallia bisaltideCyrestis cassanderCyrestis maenalisCyrestis niveaCyrestis kudratiChersonesia rahriaNeptis mindoranaNeptis duryodanaNeptis pampangaNeptis felisimilisNeptis sunicaNeptis cliniaNeptis nitetisNeptis cymelaNeptis vibusaNeptis iliraLasippa pataLasippa bellaLasippa illigerellaLasippa illigera'Lasippa ebusaPhaedyma columellaPantoporia damaPantoporia hordoniaPantoporia epiraPantoporia cyrillaAthyma alcameneAthyma magindanaAthyma maenasAthyma pravaraAthyma speciosaAthyma kasaAthyma gutamaAthyma mindanicaAthyma godmaniAthyma arayataAthyma epimethisAthyma cosmiaAthyma asuraAthyma jagoriAthyma larymnaModuza thespiasModuza pintuyanaModuza urdanetaModuza mataModuza jumaloniModuza nuydaiBassarona teutaBassarona piraticaLexias damalisLexias hikarugenzi

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00010011101111000100000100000001110010010000000100000000000000

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346 T. RACHELI, M. BIONDI

APPENDIX - Continued.

PAL LUZ MRO SAM LEY PAN NEG CEB BOH MIN

Lexias satrapesLexias panopusLexias dirteaDophla evelinaDichorragia peisistratusRohana parisatisRohana rheaEuripus halithersesProthoe franckAgatasa calydoniaPolyura athamasPolyura delphisPolyura schreiberCharaxes antonioCharaxes plateniCharaxes harmodiusCharaxes so/onCharaxes amycusCharaxes bernardusHestina dissimilisHestina waterstradtiModuza procrisl^ebadea marthaParthenos sylviaTanaecia arunaTanaeda calliphorusTanaecia leucotaeniaTanaecia susoni/borromeoiTanaecia hmvarthiTanaecia iapisTanaecia godartiiEuthalia lusiadaEuthalia anosiaEuthalia obsoletaEuthalia amandaEuthalia alphedaEuthalia moninaEuthalia mahadevaEuthalia acontheaEuthalia adoniaEuthalia djataEuthalia lubentinaBassarona dunya

LYBITHEIDAELybithea narinaLybithea geoffroyi

SATYRIDAEMelanitis atraxMelanitis ledaMelanitis phedimaMelanitis boisduvaliaElymnias pantheraElymnias daraElymnias kanekoiElymnias luteofasciataElymnias esacaElymnias congruensElymnias casiphonidesElymnias meliasElymnias sansoniElymnias beza

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BIOGEOGRAPHICAL OBSERVATIONS ON THE PHILIPPINE PAPILIONOIDEA 347

APPENDIX - Continued.

PAL LUZ MRO SAM LEY PAN NEG CEB BOH MIN

Lethe europaLethe chandkaLethe duraLethe dataensisNeorina lowiiMycalesis orseisMycalesis bisayaMycalesis tagalaMycalesis treadawayiMycalesis aramisMycalesis janardanaMycalesis perseusMycalesis mineusMycalesis felderiMycalesis itaMycalesis igoletaMycalesis kashiwaiiMycalesis georgiMycalesis fredericiMycalesis tamarauOrsotriaena medusRagadia magandaRagadia melindenaRagadia luzoniaRagadia tsukadaiRagadia crohonicaZethera hestioidesZethera thermaeaZethera pimpleaZethera musidesZethera musaPtychandra leucogynePtychatidra mindoranaPtychandra lorquiniiPtychandra schadenbergiPtychandra negrosensisPtychandra ohtaniiPtychandra mizunumaiPtychandra nakamotoiAcrophtalmia art em isAcrophtalmia letoA crophtalmia yamashitaiAcrophtalmia albofasciataYpthima stelleraYpthima pandocusYpthima sensilisFaunis lendsFaunis sapphoFaunis phaonFaunis stomphaxAmathusia phidippusAmathusia philippinaDiscophora oginaDiscophora philippinaDiscophora simplexDiscophora nechoDiscophora dodongDiscophora sondaicaZeuxidia amethystusZeuxidia semperiZeuxidia sibulana

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1100001100100111001010110101000101001000100110010011010000001

1000000100010010000010010000010101000000000110001010000010001

1100000100011011010010010000010100010000010110001011100010000

1000000100000101000010000000010101000000000110001010010001000

1100000100101010000010000001000001000000100110010010010000000

110000011

0111110001010100010001001101110101111110011010001101

Dow

nloa

ded

by [

Geb

ze Y

ukse

k T

ekno

loji

Ens

titïs

u ]

at 0

3:09

21

Dec

embe

r 20

14