BR.J.ENT.NAT.HIST..12: 1999 69 THOMAS VERNON WOLLASTON AND THE MADEIRAN BUTTERFLY FAUNA—A RE-APPRAISAL Michael A. Salmon Avon Lodge, Woodgreen. New Forest, Hampshire SP6 2AU. AndrewWakeham-Dawson MillLaine Farm. Offliam. Lewes, East Sussex BN73QB. Abstract. In 1847, the British entomologist T. V. Wollaston visited Madeira and commenced a study of the invertebrate fauna of that island. He returned several timesandhisworkculminated in thepublication ofInsecta Maderensiain 1854.Two years later he published On the Variation ofSpecies with especial reference to the Insects, a text which clearly anticipated On the Origin ofSpeciesbymeans ofNatural Selection by his friend Charles Darwin. Although Wollaston made only a small collection of Madeiran butterflies, he is remembered by the endemic subspecies of Pieris brassicae L. that bears his name. The present authors visited Madeira on a number of occasions during the period May 1997-August 1998 to survey Lepidoptera in the Parque Ecologico do Funchal, part of a long-term project to provideinformation fora fieldguideto the Lepidoptera ofthepark. Thispaperisan extension ofthat survey and presents data on all the Madeiran butterflies. Introduction 'The island being resorted to by so many invalids,' wrote Professor Henry Moseley, on visiting Madeira in 1872 (Moseley, 1880), 'the cemetery forms a conspicuous feature in the scenery.' With this statement the scene is set, for almost 150 years ago the young T. V. Wollaston (1822-1878) Fig. 1, an ailing consumptive of25 summers, first set sail for Madeira in the hope that constant sunlight and an equableclimate might result in acure forhis tuberculosis. However, the truth is that few such individuals survived long. In travelling south, Wollaston was merely following that long line of Victorian invalids despatched overseas by doctors who were, themselves, unable to offer any effective treatment. Indeed, there was Uttle hope of a cure, for the days of definitive anti-tuberculous medication were yet to come, butWollaston, initially at least, did ratherwell. Hewas so taken bythe beauty ofthe island, the friendliness ofits inhabitants and the starthng mountain scenery, that he returned on a number ofoccasions. While there he seized every opportunity to further his interest in entomology, and three prolonged visits to that island, undertakenatdifferen—tperiodsoftheyear, supplied thebasis forthemostimportant of his written works the monumental Insecta Maderensia (1854). This quarto volume of677 pages was illustrated with exquisite coloured drawings by Professor J. O. Westwood FRS, and describes Madeiran insects in minute detail together with other works (e.g. Wollaston, 1858) (Fig. 2). All that happened a very long time ago, and since then a not inconsiderable number ofother naturalists have also made their way to the North Atlantic Islands of Macaronesia and some of them have studied the butterfly fauna of Madeira. Indeed, for a country with less than 20 butterfly species Madeira would appear to have received a great deal ofattention. Many ofthese later authors, however, wrote little more about the entomology of Madeira and its neighbouring islands, Porto Santo and the Ilhas do Desertas, than short annotated lists ofthe species recorded. 70 BR.J.ENT.NAT.HIST., 12: 1999 andasfarastheColeopterawereconcerned, suchpapershaveoftenturnedouttobe no more than extensions of Wollaston's original works. So, was Wollaston an important figure and, ifso, what did he actually achieve? WOLLASTON'S EARLY LIFE Thomas Vernon Wollaston was born on 9 March, 1822, at Scotter, Lincolnshire, the youngest son of Major Wollaston ofShenton Hall, Nuneaton. He was directly related to both William Wollaston (1659-1724), author of the Religion ofNature Delineated, and Dr William Hyde Wollaston FRS (1766 1828), the celebrated chemist whofirstdiscovered themetallicelementsrhodium and palladium. Educated at the Grammar School, Bury St Edmunds, and Jesus College, Cambridge, he took his BA in 1845, and his MA in 1849. It was while at Cambridge that Wollaston first developed an interest in entomology, stimulated by the teaching and example ofhis mentors. Professor C. C. Babington FRS and, in particular, the Revd Hamlet Clark (1823-1867) (Fig. 3), a fellow undergraduate, who was elected Vice President ofthe Entomological Society of London in 1864. During his early years at Cambridge, Wollaston, with Hamlet Clark and the Revd J. F. Dawson organized collecting trips to the fens and S—uffolk breckland. In 1843, Wollaston published his—first entomological paper a short account of the beetles found near Launceston and it appears that from then onwards he was to devote most ofhis hfe to a continuous study of entomology and natural history. Under the guidance of Professor Babington and Hamlet Clark, Wollaston was soon elected a Fellow of the Cambridge Philosophical Society and then, in 1847, to Fellowship of the Linnean Society. Although his main interest was in the Coleoptera, Wollaston studied many other orders of insects as well as making studies of the botany of Madeira and its land shells. At least two species ofLepidoptera are named after him. There is Httle doubt that Wollaston's work on the North Atlantic Islands did much to stimulate others who were interested in the various endemic forms that inhabited them. What he achieved was to make people think. Duringhis undergraduatedays, Wollaston wasopposed to the theoryofevolution but later, as a friend ofCharles Darwin, he came to accept that some forms which could hardly be regarded otherwise than as individual species might have a recognizable derivative origin. Wollaston admired Darwin for his accurate field observations, but his own observations convinced him that there was no evidence that one specieseverevolved into another. Heconcluded that specieswereseparately createdaccordingto somedivineplan, thedetailsofwhichcouldbeelucidated by the study ofnatural history. He insisted that his thesis stemmed not from religious belief but from observation and application of the inductive scientific method (Cook, 1995). In 1856, he published On the Variation ofSpecies with especialreference to the Insects, a text which clearly anticipated Darwin's The Origin ofSpecies by means of Natural Selection {\S59). Wollaston dedicated thework to his friend: "Whoseresearches, in variousparts of theworld, haveadded somuch toourknowledgeofZoologicalgeography." Darwin, in turn, sent him a complimentary copy of the Origin ofSpecies. In spite of their conflicting views Wollaston and Darwin remained good friends, although the former'scriticism ofTheOriginofSpeciesclearly strained therelationship. Wollaston was quite unable to accept the central theme ofDarwin's message (Anon, 1860). "A cold shuddering comes over us at what we are compelled to regard as a glorious non sequitur, and that, too, from premiseswhichwecannot admit!" Darwin responded to this in a letter to his son William: "There was a ratherfierce attack on it in Annalsof — BR J. ENT. NAT. HIST., 12: 1999 Nat. HistorybymyfriendWollaston. ... ByJovetheBookhasmaderowenough,&I shd[sic]nowliketogetonquietlywithmywork."Although Darwinwouldrathernot have become involved he was provoked into retaliation. In July 1860, he wrote to Professor Westwood, who was attempting to raise money to purchase Wollaston's collectionsofMadeiranlandandfreshwatershells: "Ihavespentsomuchmoneylately that I am not willing to subscribe forthepurchase ofWollaston'scollections foryour Museum."' (Burkhardt, et al. 1993). In the introductions to his various works on the fauna of the Atlantic islands Wollaston put forward his own hypothesis that theendemicColeopteracould not be satisfactorily referred to any geographical area now existing, but rather to some Atlantic region of which they were the sole representatives of modern times. This view remained unchallenged until the latter half of the present century when a number ofauthors, including Mayr (1969), Bush (1975 and 1982), and Cain (1984), individuallyaddressed the twin problems ofevolution and speciation. More recently, Owen and Smith (1993a), who visited the North Atlantic Islands on a number of occasions, reviewed more than 300 publications on the butterfly fauna of Macaronesia, and postulated that this fauna consisted offour elements: a.) endemic taxa related to existing ones on the neighbouring continents; b.) relict endemic descendants ofan extinct Saharan fauna; c.) recent (mostly post-fourteenth century) immigrants from Europe and East Africa; d.) cosmopolitan migrants. They suggested that an unknown but substantial number of species, mainly from Laurisilva which covered much of the island of Madeira, had probably become extinct since theislandswerecolonized by Europeansin thefourteenthcentury. They were able to show that the number ofspecies and the number ofendemic forms on each ofthe North Atlantic Islands were positively correlated with vegetation density and negatively correlated with distance from the nearest continent. Wollaston in Madeira Twoyearsaftergraduation, duringtheautumnof1847, Wollastondevelopedearly symptoms of severe pulmonary disease. His doctors advised him to seek a milder — chmate from the harsh British winter months ahead advice that led him to convalesce in Madeira during the winter of 1847-1848. It was here that over thenext few years, in spite ofappalling ill health, he took every possible opportunity to study the entomology and natural history ofthe island. The result was the accumulation of amassivecollection ofColeoptera, Lepidoptera and otherordersofinsects, aswell as the first scientific collection of Madeiran land shells. On his return to England, Wollaston agreed to continue convalescence but this time in the West Country. He left his elegant town house in Park Lane, London; purchased property in Devon; married theyoungestdaughterofaclosefriend;wrotea slimvolumeofpoems Lyra Devoniensis (1868); only to die a few years later, at the age of 57, of pulmonary haemorrhage, theconsequenceoftuberculosis. Hewasburied atTeignmouth, Devon. Madeira was not the only island that Wollaston visited. He seemed fascinated by islands, and visited, among others, the Canaries, the Cape Verdes, St Helena and Lundy. At each place he collected assiduously, his illness permitting, and produced detailed lists ofspecies which included many that were new to science. In St Helena hespecializedintheCossonidae(Coleoptera), collectingfromthatislandnolessthan 54 separate speci—es, and this—interest ultimately led to the description of 255 new cossonid species world-wide which was approximately 180 more than all other coleopterists had achieved. Looking back to those far offtimes one is immediately struck by the strength of purpose that Wollaston showed while engaged in these — 72 BR.J.ENT.NAT.HIST.,12: 1999 Studies. His illness was at times so severe that he suffered an almost unremitting battle to keep going. He once wrote "The constant warfare between physical incapacity and will was a curious feature, even at Madeira, where halfmy work was actually written in bed, and when suffering more or less from bleeding ofthe lungs; orelse while sittingin achair in thegarden, baskingin the sunshine." But Wollaston wasobviouslymadeofstern stuff. Intheprefaceto Insecta Maderensiahewrote, "In May of 1850,. . . having procured a tent, I again set sail for the island,—prepared to take up my abode, during the hotter period, in districts as yet but imperfectly explored; and by applying myselfin good earnest (at elevations, moreover, difficult ofaccessexcept at that pecuharseason), I conceived that I should beinaposition, at the close ofmy third sojourn, to attempt a more lengthened and systematic treatise than I had at the beginning ventured to contemplate". During the 1850s, the expatriate community in Funchal was sizeable, (averaging around 500 adults in the summer months and 700 in winter (Nash, 1990), and Wollaston found within it a number ofdevoted friends. One ofthese, John Gray, a wealthy business man and amateur coleopterist, invited him to join his yacht, the Miranda, for excursions to remote parts of the island, and later to the Canaries archipelago and Cape Verde Islands. Among his companions on these trips was Richard Thomas Lowe, another Cambridge man and Chaplain to the Enghsh Church in Funchal. Revd Lowe was a keen naturalist, particularly interested in botany as well as the Mollusca. Unfortunately his practice ofattempting to convert expatriate parishioners to the ways ofthe Roman Church led him into deep waters. When news of these heretic practices came to the ears of parliament. Lord Palmerston was provoked to announce that there was a 'scandal in Madeira.' He dispatched anenvoy, theArchbishopofCapeTown, to Funchalat once, hopingthat mediationmightput anend to these 'vilepractices.' TheArchbishop, however, failed in his mission and Palmerston ordered Lowe's immediate resignation, an order that Lowejustasquickly rejected. Through allthisWollastoncontinued to support Lowe and, together, they organized furthercollectingtripsto remoteand inaccessibleparts oftheisland. WewonderiftheArchbishop ofCapeTown was persuaded to develop an interest in the natural history ofMadeira. Wollaston was totally captivated by the beauty of Madeira although, in 1854, journeys to the more central parts ofthe island must have been extremely difficult. Travel on horseback, tramping for hours over ill-defined rocky tracks into the evergreen Laurisilva, orattemptingto land on the more remoteparts ofthecoastHne were all fraught with danger, especially to a man so physically unfit but Wollaston revelled in thechallenge and readily accepted thedifficulties that beset him alongthe way. Hemaywell havebeenmindfulofthewordso—fCharles Kingsley(1819-1875), a fellow naturalist and author of The Water Babies indeed, these two would appear to have had much in common. "The naturalist," wrote Kingsley, should be "strong in body; able to haul a dredge, climb a rock, turn a boulder, walk all day, uncertain where he shall eat or rest; ready to face sun and rain, wind and frost, and to eat or drink thankfully anything, however coarse or meagre . . ." On his trips, Wollaston was usually accompanied by hiswife, a knowledgeable lepidopterist, and Portuguese porters who were responsible for bringing daily supplies. He was able to describe in glowing terms the joys of camping out in wildest Madeira. There is something amazingly luxurious in betaking oneself to Tent-hfe, after months of confinement and annoyance (it may be entirely, partially it must be) in the heat and noise of Funchal. We are th—en perhaps more than ever open to the favourable impressions of an alpine existence; and who can adequately tell the ecstasy ofa first encampment on these invigorating hills! To turn out, morning after morning, in the solemn BR.J.ENT.NAT. HIST.. 12: 1999 73 — Stillness of aerial forests, where not a sound is heard, save ever and anon a woodman's axe in some far-off tributary ravine, or a stray bird hymning forth its matin song to the ascending sun; to feel the cool influence ofthe early dawn on the upward swa—rd, and to mark the thin clouds offleecy snow uniting gradually into a solid bank, affording glimpses the while, as they join and separate, of the fair creation stretched out beneath; to smell the damp, cold vapour rising from the deep defiles around us, where vegetation is still rampant on primaeval rocks and new generations oftrees are springing up, untouched by man, from thedecayingcarcases ofthe old ones; to listen in the still, calm evening air to the humming ofthe insect world (the most active tenants ofthese elevated tracts); and to mark, as the daylight wanes, the unnumbered orbs of night stealing one by one on to the wide arch of — heaven, as brilliant as they were on the first evening of their birth; are the lofty enjoyments, all, which the intellectual mind can grasp in these transcendent heights.' In 1890. Mrs Wollaston presented George Baker with the greater part ofher late husband's collection ofMadeiran Lepidoptera. It contained 11 butterfly species and amuch largernumberofmoths. Wollaston's originalaccount(1858)consists oflittle more than short descriptive paragraphs, in Latin, ofthe various species, but Baker (1891) transcribed these, added notes of his own, and put forward his own theory that humidity ofclimate might be largely responsible for variation in certain species (see also Stainton, 1859). From July 20-August 3, 1998, the au—thors visit—ed Madeira under the auspices of the Madeiran scientific funding body CITMA to survey Lepidoptera in the Parque Ecologico do Funchal. This was part ofa long-term project started in May 1997, to provide information for a field guide to the Lepidoptera ofthe Park, and to provide base-line information against which the effects of the attempted re- establishment of native vegetation (Laurisilva) on the butterfly fauna could be assessed (Wakeham-Dawson & Warren, 1998a). We were accompanied by entomological colleagues from the Natural History Museum (London) and the University ofWarwick. The authorities ofthe Parque Ecologico do Funchal put the Casa do Barreiro (970m.), one ofthe park houses used to accommodate visitors, at our disposal and we were able to visit most parts ofthe island to examine localities mentioned as specific in the literature. Although the butterflies ofMadeira are well documented, a number ofchanges, some highly significant, haveoccurred during the past thirty years. It is with this in mind that we have re-examined the status ofthe butterfly fauna as it appeared on this visit, and in relation to Madeira's Oceanic existence (MacArthur & Wilson. 1967). The Funchal Ecological Park The Parque Ecologico do Funchal was founded in 1994 as a centre for environmental education. It is situated above Funchal, the capital city ofMadeira, on the southern side ofthe island (Fig. 4). The Park covers an area ofabout lOkm-^; its lowest point is 500m. above sea level and it rises to over 1800m. near Pico de Arreiro. The range of elevation together with a number of steep gullies within the Park, allows the area to support an interesting assemblage of endemic plants and animals (Wakeham-Dawson & Warren, 1998b). One of the main aims of the Park authorities is to fell invasive Eucalyptus and Acacia, and replace them with Laurel trees. The hope is that native Laurel forest (Laurisilva) can be re-established. Laurisilva once covered most ofMadeira and is a relicofthe forests thatcovered southern Europe in theTertiaryPeriod (Press& Short, 1994). From sea-level (especially on the dry south side oftheisland) to c.300m. there BR.J.ENT.NAT.HIST.,12: 1999 Fig. 1. ThomasVernon WoUaston (1822-78). Fig. 2. Beetles sketched by T. V. WoUaston 1846. (Courtesy Mun. Museu Funchal). Fig. 3. Revd Hamlet Clark (1823-67). BR.J.ENT.NAT.HIST., 12: 1999 PICODOAREIRO (1810m.) 1000m. I RibeiroFrio- CentrodeRecepcao " (1125m.) CASADOBARREIRO (970m.) PARQUEECOL<5gICODOFUNCHAL Seixai ParqueEcologico SaoVicente 'doFuDChal Raba^al Encumeada RibeiraBrava FllNCHAL MADEIRA Fig. 4. Map ofthe Funchai Ecological Park (Parque Ecologico do Funchal). 76 BR.J.ENT.NAT HIST..12: 1999 was dry Laurisilva consisting ofApollonias harhujaua (Cav.) Bornm. with Euphorbia spp. andDragontrees{Dracaenadraco(L.) L.). From 300m. toabout 1200m.: humid Laurisilva ofLauras azorica (Seub.) Franco, etc., and from 1200m. to the summit of the island: Erica spp. (Tree Heathers) Laurisilva. Today, the scene is rather different. The humid Laurisilva is now restricted to the north and the dry Laurisilva is now almost totally replaced by urban development or agriculture (Press & Short, 1994). The humid Laurisilva is kept wet bycondensation ofwater from the frequent sea fogs that formoverthehigherregionsofthe island. Much ofthe original area ofLaurisilva on Madeira has been destroyed, and the Park's project may make a valuable contribution to the conservation of this important habitat type. According to Nash (1990) Joao Gon9alves Zarco, the Portuguese explorer who discovered Madeira in 1419, found the island covered with dense forest (in Portuguese, the word Madeira means wood). In orderto clear the ground for the first settlement he caused fire to be put to the trees. There is a local legend that the whole ofthe island burned for seven years but this is probably no more than a legend. Check-listofthe madeiran butterflies (arranged in the order that they are discussed below) Pieris rapae (L.) small white. Pierisbrassicaessp. wollastoniButler, madeiranlargewhite. (W) Endemic subspecies. Gonepteryx maderensis Felder madeiran brimstone. (W) Endemic species Colias crocea (Geoffroy) clouded yellow. (W) Hipparchia maderensis Baker, madeiran southern grayling. (W) Endemic species. Pararge xiphia Fab. madeiran speckled wood. (W) Endemic species. Pararge aegeria (L.) speckled wood Neohipparchia statilinus (Hufnagel) tree grayling. (US) Vanessa atalanta (L.) red admiral. (W) Vanessa indica ssp. vulccmia Godart. Indian red admiral. (W) Danausplexippus (L.) monarch or milkweed butterfly. Cynthia cardui (L.) painted lady. (W) Lycaenaphlaeas ssp. phlaeoides Staudinger. small copper. (W) Lampides boeticus (L.) long-tailed blue. (W) Colias hyale (L.) pale clouded yellow. (US) Cynthia virginiensis (Drury). american painted lady. (US) Issoria lathonia (L.). queen ofspain fritillary. (W) (OS) Hypolinmas misippus (L.) false plain tiger. (US) Danaus chrysippus (L.) plain tiger. (US) W = specimens in the T.V.WoUaston Collection (1847-1850) OS = occasional vagrant US = uncertain status Pieris rapae (L.) The small white is a relatively recent addition to the Madeiran fauna. Although there is the record 'ofa possible sighting' in 1909 (Swash & Askew, 1982), it was not until December 1971 thata specimenwasactuallycapturedand notuntil 1974thatthe species became definitely established. In July ofthat year a massive invasion occurred which Wolff(1975) suggested might have originated in Portugal and been driven by the prevailing Trade Wind towards Madeira. Why, Wolffasks, when similar climatic conditions must have occurred many times in the past, did sudden colonization not BR.J.ENT.NAT.HIST..12: 1999 77 occuruntil 1974?Thisquestionhasnotbeenanswered.WefoundtheSmallWhitetobe very common throughout the Parque Ecologico do Funchal (Table 1) and at the northernendoftheisland,inlandfromSeixal,andinthewoodedlaurel-coveredvalleys nearRaba9al. Itcouldbeseenflyingfromsea-leveltoanaltitudeofatleast 1300m. Itis also common on the island of Porto Santo. We observed females ovipositing on Cabbage (Brassica oleracea L.) and Chagas (Tropaeolum majiis L.) during July and August 1998. It has also been observed in Madeira laying on Coronopus didymus (L.) Sm. (Wakeham-Dawson, 1998). No named aberrations were observed. Pieris brassicae ssp. woUastoni Butler — Since Wollaston first collected this butt—erfly in mid-Victorian times and he collected a considerable series (Baker, 1891) it has continued to decline in numbers and must nowbe regarded as extinct. Thisdecline appears to be long-standing. Both Wollastonand Bakerthought this subspecieswaslittlemorethan amelanicvariation of Pieris brassicae L., forming a natural transition between P.brassicae and the endemic Canary Island P. cheirauthi Hiibner. Felder (1862), who published a list of the insects collected by scientific members of the frigate Novara, thought that this subspecies was P. cheirauthi, but Kudrna (1973), after critical examination of the genitalia of P. brassicae, cheirauthi and wollastoui, stated that there was sufficient evidence to consider wollastouias a subspecies ofP. brassicae. Themale genitalia are illustrated in Fig. 5a, for comparison with illustrations by Higgins (1975). Swash & Askew (1982) pose the question as to whether the decline in numbers of Pierisbrassicaessp. wollastouiand theincreasein P. rapaemight be inter-related; the decline ofthe former facilitating colonization by the latter, and the establishment of P. rapae possibly resulting in the final decline ofP. brassicae wollastoui. This seems unlikely, however, as both taxa co-exist throughout much of Europe. It therefore remains a subject for debate. Another possibility is that a parasite arrived on the island and destroyed the P. brassicae wollastouipopulation. In the United Kingdom 95% ofP. brassicae larvae are parasitized by the wasp Apauteles glomeratus L, and the population only survives because of the regular influx of migrants from the Continent (Cribb, 1983). Although Martin (1941) found P. brassicae wollastoui at Santo da Serra, Oehmig (1977)atPicoArreiroand Aguas Mensas, andGardner&Classey (1960)at Raba9al, Chao das Feiteras and Caramujo, de Worms (1964) observed only a single specimen in 1964, atMonte,c.700m. above Funchal. Sincethen therehavebeen onlyscattered sightings. Wolff(1975) reported a numberofspecimensin 1973 and 1974, and Swash & Askew (1982), various isolated sightings at Porto da Cruz, Ribeiro Frio, Funchal and SantaCruz. Theyconcluded that 'theonlysizeablepopulationlocatedwasin the valley leading to Faja da Nogueira where, in July and August, several specimens were seen nearthe riverand a few at greateraltitude up to about 1000m.' Two years later. Lace & Jones (1984) reported seeing the Large White at Boca da Risco on the northcoast, butOwenand Smith (1993a), whovisited theislandfivetimesduringthe 1990s, were unable to find either adults or larvae. One ofthe present authors (A.W- D), in six visits to Madeira since 1997, has been no more successful. In an extensive search during July 1998 we found no evidence that this butterfly still flies on Madeira. Resident entomologists have not seen this species in recent years and have been aware ofa decline in P. brassicae wollastoui that appears to have matched the increase in P. rapae (Antonio M. F. Aguiar, pers. comm.). BR.J.ENT.NAT HIST..12: 1999 Gonepteryx maderensis Felder This species (Plate 1, H), well known to Wollaston in the 1850s, is endemic to Madeira and formerly widely distributed and abundant. It was first described by Felder(1862)whothoughtitwasnomorethanavarietyofG. cleopatraL. Somethirty years later. Baker (1891) suggested that it formed a connecting link between the Mediterranean Gonepteryx cleopatra L. and Gonepteryx cleopatra ssp. cleobiile from the Canaries. More recently, Kudrna(1975) hasaccordedbothit and cleobiilespecific status. This action is supported by Brunton et al. (1996) who studied seven species of Gonepteryxunderultraviolet lighttoreveal theirultravioletreflectancepatterns. They wereabletoshowthat G.maderensisdiffered from otherspeciesinthat the ultraviolet pattern on the underside offemale specimens was typical ofGonepteryx males. The literature suggests that over the past 25 years the madeiran cleopatra has declined steadily in numbers. Kudrna (1975) described G. maderensis as local and uncommon at higher elevations in mountains; Gardner & Classey (1960) thought it might be restricted to certain northern parts of the island; while Swash & Askew (1982) reported sightings at Faja do Cedro Gordo, near Faja da Nogueira, and the Botanical Gardens at Ribeiro Frio. Manley & Allcard (1970) found this species at an altitude of 1200m. However, the Madeiran Cleopatra may in fact be more common than it appears, as it rarely comes down from the forest canopy. It is probably widespread but not common in March, May andJuly in areas ofLaurisilva between 400-1000m. InJuly 1998 weobserved small numbers flyingabout thelaurel forestat Chao da Ribeiro. This species was seen at Chao da Ribeiro near Seixal on the north coast in March (John Smit, pers. comm.) and again in May (W), and a female was seen ovipositing high up in the Laurisilva canopy on leaves of Rhamniis glandulosa Alton on 24.vii.1998. A malewas observed nectaringat Hypoclweris radicata L., and both sexes were observed feeding at Agapanthiis praecox Willd. flowers by the roadside near Ribeiro Frio and at several locations near the Encumeada Pass. A single male was seen in the Funchal Ecological Park on 30.vii.l998 (Table 1). This butterfly had probably flown south west from Ribeiro Frio with the prevailing wind (Duarte Camara, pers. comm.). This is encouraging because it suggests that the species may be able to re-colonise naturally in the Park, once Laurisilva is established. The male genitalia are illustrated in Fig. 5b. Colias crocea (Geoffroy) In July and August 1998, the Clouded Yellow was second only to the Madeiran Southern Grayling in abundance. It occurred at all altitudes from sea-level up to 1800m. and was often observed flying in the streets and gardens of Funchal. A femalewasobserved layingsingleeggson theuppersurfaceofthe leavesofTrifolium repens L. in shaded areas near small cabbage fields at Seixal. We also noticed that a PLATE MADEIRANBUTTERFLIES I A A: HipparchiamaderensisMale. B: HipparchiamaderensisFemale. C: Hipparctiia maderensisFemaleunderside.D; ParargeaegeriaMale. E: ParargeaegeriaMale Dwarf Var. F: Pararge xiphia Female. G: Pararge xiphia Female underside. H: Gonepteryx maderensis Male. I: Lycaena plilaeas ssp. phlaeoides var. caeruleopunctala. J; Vanessa indica vulcania. K: Colias crocea var. cremonae. L: Coliascroceavar. dawsoni.M: Coliascroceavar. geisleri. N: Coliascroceavar. radiata.O; ColiascroceamaleDwarfvar. P: Coliascroceavar. failiae.