Habitat, isolation and the evolution of Madeiran landsnails

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1 BiologicalJoumal of the Linnean Society (1996), 59: With 2 figures Habitat, isolation and the evolution of Madeiran landsnails L. M. COOK The Manchester Museum, University ifmanchester, Manchester, M139PL Received 8 December 1995, acceptedfir publication 2 February 1996 The Madeiran archipelago consists of Madeira itself, Porto Santo and the Deserta islands. On Madeira, the forest and the coastal floral associations are so different that their faunas are effectively isolated and have undergone largely independent development. There are different faunal associations on the eastern peninsula and in the SE coastal region, which may have been separated from each other in the past. On Porto Santo, western and eastern hills have different faunas. Most observations on the fauna are compatible with evolution by allopatric speciation, consequent upon isolation on different islands or mountains, as opposed to parapatric or sympatric processes following disruptive selection. Some cases where the taxonomy is difficult to unravel may, however, indicate parapatric speciation; examples belong to the genera Discula and Heterostoma (Helicidae) and Amphorella (Ferussaciidae). Most evidence relating to species composition in communities is compatible with a balance of random immigration and extinction, rather than selective interaction, allowing clusters of similar sympatric species to accumulate. However, this impression may indicate that test procedures are insufficiently sensitive to detect interactions, and detailed ecological studies are required. Questions about speciation and distribution would be clarified if dates of divergence were established. <C 1996 The Linnean Society of London ADDITIONAL KEY WORDS: - island fauna - land snails - speciation. CONTENTS Introduction. The setting Endemism on the Madeiras Molluscan evolution Speciation Faunal assemblages Conclusions Prospect Acknowledgements References INTRODUCTION The Madeiran islands have a rich molluscan fauna with a high frequency of endemic species. There are over 250 species and subspecies, of which 74% are endemic (Walden, 1984a, 216 species in Walden's 1983 list). The foundations for the systematics of the land molluscs were laid by Albers (1854), Lowe (1831, 1852, 1854), Castello de Paiva (1867) and Wollaston (1878). Lowe and Wollaston (particularly the /96/ $ The Linnean Society of London

2 458 L. M. COOK latter) were meticulous in recording the provenance of their material and in questioning the correctness ofunexpected location data, providing the framework for the biogeography as well as the taxonomy. In order to extend the geographical information, a series of quantitative studies of patterns of distribution has been carried out on Madeira (Cook,]ack & Pettitt, 1972; Cook, Cameron & Lace, 1990; Cameron & Cook, in prep.), Porto Santo (Cameron, Cook & Hallows, 1996) and to some extent the Deserta islands (Cook et al., 1972, 1990). The fossil history of the fauna at a particular place has also been studied (Cook, Goodfriend & Cameron, 1993; Goodfriend etal., 1996). Although some aspects, such as the fossil deposits on Porto Santo and the distributions on the Desertas need further attention, we now have a good picture of species distributions on the archipelago. Islands have always been thought ofas test beds for theories ofevolution; what can now be said about the evolution of the Madeiran snails which was not apparent to the nineteenth century pioneers? THE SETTING The Madeiran archipelago is in the Atlantic ocean, 900 km south-west of Portugal and 800 km west of Morocco (Fig. 1). It consists of the island of Madeira, 58 km at its largest dimension and rising to an altitude of 1860 m, and two other clusters of islands. Porto Santo is 40 km to the north-east and rises to 520 m. The ocean bed separating it from Madeira drops to 2600 m. About 10km to the south-east of Madeira are the three Deserta islands. They are 480 m high at the highest point, and are linked to Madeira by a submarine ridge falling to no more than 140 m along its crest. It is therefore possible for Madeira and the Desertas to have been connected as a result of sea level fall during a glacial phase, whereas Porto Santo could never have been so attached. Porto Santo is on a large shelf covered by shallow water, from which a number of islets also emerge. At one time it was probably a much larger island, and the deposits on it containing fossil snails were formed from consolidated sand derived from extensive beaches which existed at times of lowered sea level. Madeira appears to have been created by four main phases of volcanic activity. The first dates from between 20 and 12Myr and the fourth from 0.75Myr. (Zbyszewski, 1972; Mitchell-Thorne, 1985; reviewed by Cook et al., 1990). The eastern peninsula is formed ofrocks between 4 and 0.74 Myr in age. Along the south coast of the island there is a series of small volcanic cones which are younger than this but not recent. The Desertas are the same age as the peninsula, while the youngest rocks on Porto Santo are at least 12Myr old. Porto Santo may therefore have been a source for colonizations of Madeira after periods of vulcanism. To the north of this group is a series of sea mounts, some of which have been islands (Pastouret etal., 1980). They may have formed a route by which the Madeiras were colonized from the Iberian peninsula. ENDEMISM ON THE MADEIRAS To put the molluscan radiation in context, it is worth noting briefly the patterns in other groups. The islands show a range of diversity and endemism which is broadly related to ability to disperse. There are 1100 species ofplants (Hansen, 1969;

3 MADEIRAN SNAIL FAUNA 459 Sjogren, 1972). Colonizing species would have arrived as seeds, floating rafts or spores, but 11% of the present list is endemic. Some endemics are dominant components ofthe montane subtropical rain forest which clothes the higher slopes of Madeira. Others are adapted to survive on the dry rocky terrain existing on all the islands nearer to sea level. There are no mammals definitely native to the islands except bats, all others having been introduced by man in the last 500 years. A single endemic lizard species is abundant and shows colour and biometrical variation between islands (Cook, 1979; Bischoff, Osenegg & Mayer, 1989). Of 27 resident breeding land bird species, one is endemic to Madeira and two to Madeira and the Canary Islands; eleven species have Madeiran subspecies and four have subspecies shared with the Canaries (Hounsome, 1993 and pers. commun.) Where subspecific rank is not considered to have been reached there are sometimes biometrical differences from continental individuals. Some species which might have been expected, such as the tits (Paridae), are absent, while species that are on the islands tend to have broader niches and less sexual dimorphism than their continental neighbours (Hounsome, 1993). There are 12 species of butterflies, of which one is endemic and four are local subspecies (Swash & Askew, 1982; Owen & Smith, 1993). There are 58 families of Coleoptera, with about 875 species (817 on Madeira itself); about 260 species (30%) are endemic (Baez, 1993). Few ofthese beetle families contain endemic genera and most endemic genera are monotypic; some, however, have developed species swarms. In the Curculionidae, Laparocerus has 23 endemic species, and Acalles, 7 Atlantic Ocean Madeiras.,', Canaries '".. o. at? D N.Africa 20km Figure I. Map of the Madeiran archipelago, showing its location in the Atlantic Ocean and positions of the three island groups. On Madeira the approximate limit of forest is shown. On Porto Santo 200 m contours indicate position of eastern and western mountains. The 200 m hydrographic contour is indicated.

4 460 L. M. COOK species. In the Colydiidae Tarphius contains 21 species while in the Carabidae, Trechus is almost entirely confined to Madeira with 20 species. Machado (1992) provides data for the Carabidae from which the overall picture can be seen. There are 31 genera comprising 46 non-endemic species, three monotypic endemic genera, and 12 genera consisting of67 species, 47 of which are endemic. To take one example from the beetles, which influenced Wollaston (1856), the genus Tarphius (Colydiidae) consists mostly ofsmall wingless species living under rotting timber in the mountains of Madeira. Outside the damp forests they hardly occur. There is one other species on Porto Santo, and 26 others spread over other Atlantic islands, Mediterranean Europe and north Africa (Dajoz, 1977). At least some ofthese live on epiphytes and are more active runners than the dead wood species, the difference in habitat probably making them more subject to accidental passive dispersal. Wheater (1993) compared similarity of species composition in beetle samples, using his own data and that of Erber & Hinterseher (1988). The greatest separation was between the high lands of Madeira and a group including the Madeiran peninsula and the other islands, reflecting the effect of the vegetational difference between the montane woodlands and the dryer scrub and grass habitats wherever it occurred. Baez (1993) gives the following values for endemism in other groups ofarthropods: Araneae (41%), Dermaptera (46%), Acari (56%), Trichoptera (57%), Diplopoda (59%). Of over 50 species of millipedes (Diplopoda), 31 are endemic (Enghoff, 1992; ].G. Blower, pers. comm.). Some European genera are missing but the genus Cylindroiulus has 25 species which have evolved to occupy a much wider range of niches than European species of the genus. A similar pattern is found in the 46 terrestrial isopod species; few genera, but a multiplicity of species in those which are present, with cases of similar but distinct species on Madeira and Porto Santo (Vandel, 1961). MOLLUSCAN EVOLUTION Similar patterns are found in the Mollusca. Table 1 gives numbers of species in families for north-west Europe, Britain, the Madeiras and Turkey, to illustrate a region adjacent to Europe. The European fauna has a relatively even spread, with six families out of 27 having 19 or more species. Britain has a fauna with a similar distribution of species between families as the European one. The Madeiras not only have twice as many species as Britain, but many fewer families and more helicids than any of the other regions. Species proliferation has occurred in two other families besides the Helicidae, the Pupillidae and the Ferussaciidae, which are relatively poorly represented elsewhere. We have examined the extent to which species are localized within islands and looked for evidence ofcompetition, which might limit the diversity at a given locality or lead to selection for character displacement between taxa. In order to relate this information to the general question of evolution it is necessary first to define more specific questions. These revolve round two issues, namely (a) whether speciation is primarily a consequence of isolation or is driven by disruptive selection, and (b) whether the community assembled in a given habitat results from accidental colonization balanced by accidental loss or from competition for available niches. In either case, the question is whether the basic catalyst which seeds the ongoing sequence of events is a random occurrence or a selective process.

5 MADE1RAN SNAIL FAUNA 461 Speciation It is convenient to distinguish auopatric, parapatric and sympatric patterns of species formation. If speciation is allopatric it is initiated by random geographical isolation, even if strong selection is involved in future divergence. Parapatric speciation, where species are formed by breakage of genetic continuity at a boundary between differently adapted races, is initiated by disruptive selection which increases fitness by decreasing gene flow. Sympatric speciation mimics one or other of these processes on a miniature scale; if it should arise through chromosome reorganization of some kind, then a random genetic discontinuity has occurred, while host- or habitat-specificity models imply disruptive selection commencing at the start ofthe process (Fig. 2). Evolutionary radiation on the Madeiran islands unquestionably began with allopatric radiation. The islands have existed for a maximum of about 15Myr with several subsequent phases of extreme volcanic activity on Madeira. The fauna is related to that of Europe, rather than north Africa, 30% belonging to, or closely related to, genera present in the European Tertiary (Walden, 1984a,b). At least 20 colonizations were involved (Cameron & Cook, 1992), and affinities with extinct and extant European genera suggest that they arrived at a wide range oftimes (Cameron, TABLE 1. Distribution of species between families for Mollusca in north-west Europe, Britain, Madeira and Turkey. Data for north-west Europe and Britain from Kerney & Cameron (1979), Madeira from Walden (1983), Turkey from Schutt (1993). NA - data not available Family NWEurope Britain Madeira Turkey Cyclophoridae Pomatiidae Aciculidae 8 I 0 2 Ellobiidae Succineidae Cochlicopidae Pyramidulidae I Vertiginidae Orculidae Chondrinidae Pupillidae Valloniidae Pleurodiscidae Enidae Endodontidae Arionidae NA Vitrinidae Zonitidae Milacidae NA Limacidae NA Euconulidae Ferussaciidae Subulinidae Clausiliidae Testacellidae NA Bradybaenidae Helicidae Families (27) Species

6 462 L. M. COOK Original species undergoes divergence which is I (a) flopatric ---. geographic l or (b) ~ympatric ---. chromosomalj or I (c) Sympatric ~ disruptive or selection (d) ~arapatric I Figure 2. Patterns of species formation as discussed in the text. divergence ---. reproductive isolationt incompatibility I two new species Cook & Hallows, 1996). Having arrived fortuitously, they established themselves in the absence ofspecialist predators or parasites and with no other competitors except for colonizers which may have arrived before. Why, then, was there a subsequent proliferation of endemic species to the point that, given their size, the islands are among the most species-rich on earth for molluscs? One contributory factor is that three groups ofislands are involved, Madeira, the Desertas and Porto Santo which, to different extents are isolated from each other. Allopatric, or vicariance, speciation took place between them. The helicid genus Leptaxis, consisting of species 15mm or more in diameter, is a case in point. The most common species on Madeira is L. undata. On the Desertas there is a very similar species, L. vulcania, and on Porto Santo, L. nivosa, which differs somewhat from that pair. All live in similar ecological conditions and the most reasonable interpretation is that they have diverged in isolation on the separate island groups. Indeed, L. vulcania is sometimes considered to be a subspecies ofl. undata. Madeira also has L. furoa and L. membranacea, however, which are found in damper highland conditions. There is a second species, L. leonina, recognized from the Desertas. This is found on the southernmost island, but also at the south end of Deserta Grande, where it coexists with L. vulcania. Differentiation, to the level of specific status, therefore also occurs within islands. Another species, L. erubescens, occurs on all three groups, although very locally on Porto Santo. The existence of allopatric species on different islands suggests that the water barriers between them provide very effective isolation, but one species has been able to colonize all three groups. Despite such anomalies, there are numerous examples from other families and genera which can be accounted for most easily by the distribution of the islands. Within islands there are geographical patterns of faunal composition. On Madeira, with its high and humid interior and dry fringing coastal strip, the ecological differences are so great as to lead to development of distinct moist woodland (the Clethro-Laurion of Sjogren, 1972) and dry 'Mediterranean' floral associations (the Aeonio-Lytanthion of Sjogren). Some snail species, such as pupillids in the genus Leiosiyla and vitrinids in the genus Phenacolimax, only frequent the damper wooded conditions, while others, such as many ofthe helicids, are very successful in the coastal conditions, and may be derived from colonizing ancestors adapted to similar conditions. These two associations have evolved, and endemic species have

7 MADElRAN SNAIL FAUNA 463 developed, virtually in isolation from each other because of the extreme habitat differences brought about by difference in altitude. On Porto Santo, Cameron, Cook & Hallows (1996) have argued that although no more than 5 km apart, the different mountain groups have effectively been isolated for much ofthe time. When sea levels were higher there would have been incursions which reduced the connection between east and west, and when sea levels were lower large areas of sand on the shallow coastal shelf would have been exposed, giving rise to the sand drift found between the mountains. Few species have been able to adapt to the sandy conditions, the main evolutionary thrust being miniature allopatric radiations on isolated hills. The refuges themselves differ in size and height, and thus in the protection they afford against natural fluctuations in climate or sandblow. This is probably why the western refuges are depauperate relative to the larger and more interconnected hills of the east. The eastern peninsula of Madeira has a fauna with a different composition from that of the south coast of the island. It has several species of its own and some woodland species. There are also taxa nearly confined to it on Madeira, which are abundant on the Desertas and Porto Santo. As a result, there is a faunal discontinuity which does not correspond to any present-day physical barrier or marked habitat change (Cook et al., 1990), and a historical explanation may be sought. There is evidence that the strip ofhigh land running north-south across the eastern peninsula to the east ofthe town ofmachico is less than 0.75 Myr old while land to either side is earlier and may be up to 4 Myr old (Walker, 1975). If that is so, it could have isolated the fauna to the east. In view of their faunal affinities, it is noteworthy that the peninsula and the Deserta islands were possibly connected at times of extreme low sea level. Volcanic action and sea level change could have provided contact and isolation at different times so as to permit both recruitment and allopatric divergence. An alternative possibility is that an ecological barrier has been present, separating the two coastal faunas. The Aeonio-Lytanthion is primarily a field layer alliance of low growing herbs and grasses interspersed with low shrubs, while the Clethro Laurion forest has a dominant tree or shrub layer. Sjogren (1972) suggests that until human colonization the latter association extended at least 100 m lower than it does now, citing nineteenth century observers in support. As is often the case, human interference may have extended the field layer alliance, which favours the dry facies molluscs. Ifat some stage the forested area extended downwards by 200 m, the south coast habitats could have been split into isolated sections. Both ecological and geological processes may therefore have been responsible for allopatric species divergence in the snails. Ifthe habitat ofa species consists ofdifferent subunits in which divergent selection occurs, and the average displacement of individuals is small compared with the extent ofthe subunits, then in principal disruptive selection could result in speciation. This would be parapatric if the subunits are geographically arranged and abut onto each other, or sympatric if the subunits are niches within the same territory. We do not need to distinguish further between these two processes, or other variants of them, because no situation will be discussed where the presumed speciation process is strictly sympatric. There has been much argument about the likelihood of nonallopatric speciation. It would be necessary for the subunits, each with their own controlling factors to maintain individuals with alternative genotypes long enough for selection for isolating mechanisms to be effective. Different commentators have taken

8 464 L. M. COOK different standpoints on the likelihood of non-allopatric speciation (with qualifications: Bush, 1975 and White, 1978 for; Mayr, 1963 and Barton & Hewitt, 1985 against), while Maynard Smith (1966) analysed conditions for sympatric speciation, without commitment to one view or the other. Evidence of selection for increased sexual isolation has been scarce in the literature, although recently, Noor (1995) presented data favouring it in two Drosophila species. Distribution of species may also shed light on the frequencies of these processes (Lynch, 1989). Land molluscs are good candidates for parapatric speciation because of their very low mobility. In the most complete study available of a genus of snails on islands, that of Pastula in the Society Islands, Johnson, Murray & Clarke (1993) concluded that each of the coexisting species is an important part of the biotic environment of the others, and that the effect of congeners is stronger than any other known environmental effect. In some cases there is also hybridization, or possibly incipient divergence, between species. This makes it possible that disruptive selection in situations where there is genetic continuity between the evolving elements, has been of major importance. The evidence for primary division as opposed to secondary contact is not conclusive, however, and the interaction effects could have arisen after genetic isolation. On the Madeiran islands, there are several examples of variation which should be considered in this context. The early investigators were conscious of a need to distinguish between species and variants within species (Wollaston, 1856, 1878) and as a result, tended to present a very orderly picture of the taxonomy. Nevertheless, there were cases, such as that of Discula polymorpha where Wollaston's (1878) treatment shows that difficulties arose. We examined the shell morphology of this species on the Madeiran peninsula (Cook etal., 1990). Over much of the length ofthe transect studied, variation is clinal, the shell getting slightly larger and flatter and the sculpture more tuberculate, rather than ridged, as one moves east. The peninsula ends in two islets, almost connected to the mainland. On these there is a sharp change in shape and sculpture to a high-spired tuberculate form (called subspecies agostinhoensis by Walden, 1983). On the two Desertan islands Ilheu Chao and Deserta Grande there are other combinations of size, spire angle and sculpture. These different dusters of characters are allopatric. The population on the very end of the peninsula, however, is high-spired, like those of the eastern islets but with sculpture like individuals from further west on the mainland. The change in the morphological characters appears to be associated with the slight break between mainland and islets but to occur at a different place for the different characters considered. Such noncoincidence of character changes has been found, for example, in Cepaea (Cain and Currey, 1963) and in Partula (Murray, 1972; Johnson et al., 1993), and may indicate a transition between differently adapted genomes. On Porto Santo, Discula attrita, D. calcigena and D. puluinata are very similar to each other. They exhibit stabilized combinations of the characters which show clinal and short distance variation in D. polymorpha on Madeira, and have separate distributions which meet each other (Cameron, Cook & Hallows, 1996). Although the first is a mountain species, being almost completely restricted to the Pico da Ana Ferreira, the other two live on the low land. If parapatric speciation occurs, they are suitable candidates. The helicid Heterostoma paupercula is another case where the evolutionary history is unclear. The small snails which were given this name by Lowe (1831) differ from all other helicids on the islands, but show so much variation in shell morphology and genitalia that they were later given three specific names in two genera (Mandahl Barth, 1984; Walden, 1983). A taxonomic interpretation would stand up ifdifferent

9 MADElRAN SNAIL FAUNA 465 characters assorted together, but they do not appear to do so (Lace, 1992; Cook & Lace 1993; Cameron, Cook & Gao,1996). On Porto Santo, individuals from sandy locations tend to be larger and paler than those from rocky locations. In other species which have been studied, such variation is sometimes non-genetic (e.g, Baur, 1988). We conclude on the present evidence that adaptive variation takes place within a single species in response to local ecological conditions. Where genetic, it is aided by very low mobility. A Madeiran example from another family concerns Amphorella tomatellina (Ferussaciidae). Walden (1983) recognized two forms, the widespread A. t. tomatellina and A. t. minor, which occurs on the peninsula. This was distinguished by Wollaston (1878) but judged by Walden to be an ecophenotype. Measurements given by Cook etal. (1990) show it to be well separated in size from individuals in other samples, but that individuals from the north and south coasts of Madeira also differ significantly from each other in size and shape. Very close to these is another species, A. mitrformis, named by Lowe (1852), but thought by Wollaston (1878) to be an elongated form of A. tomatellina. This occurs on the Desertas and Madeira, sometimes at the same sites as apparent A. t. tomatellina. Different populations of this supposed taxon have different degrees of development of the columellar lamellae, so that other features besides size and shape are involved. The situation is unresolved, and while that is so, there remains the possibility that we are seeing local parapatric speciation. Differentiation over a very small scale occurs in molluscs in other island studies (e.g. Hadfield & Miller, 1989). Faunal assemblages There are two opposed views on the kind of community assemblage we should expect in any given habitat. At one extreme, it may be the product of a balance of accidental colonization and loss ofspecies, with no interaction between them. At the other, there may be competition for niches, the precise nature ofwhich is defined by the characteristics of the colonizing species. We have examined these possibilities in two ways, by statistical tests for species interaction and by looking at the distribution of size and shape of species within families. Evenness of species abundance distributions is a measure of the nearness the distribution gets to having equal numbers ofindividuals in each species. Examination of the fossil sequence (Goodfriend et al., 1993, 1994) and ofhabitat (Cook et al. 1990, Cameron, Cook & Hallows, 1996) shows that although one or two of the nonendemic species are indigenous most have been introduced after human colonization. Cook (1984) compared the evenness of samples which have only endemic species with those which include non-endemics. If relative abundance is determined by species interaction and competition the distributions should not differ, whereas if the newly incoming species simply add on to the existing fauna, the evenness should go up. In the samples compared it went up. If the test is not too insensitive, then either the species do not interact noticeably and replacement is not taking place, or else the balance has not had time to develop in the 500 years during which there has been an influx ofnon-endemics due to human activities. Similarly, disturbed sites on the north coast of Madeira failed to show evidence of competitive interaction between endemics and non-endemics (Cameron & Cook, in prep). Of course, if it

10 466 L. M. COOK takes more than halfa millennium for a balance to start to develop then in a situation where frequent introductions occur the random model effectively applies. Pulmonate snails as a whole have a characteristic size and shape distribution, tending to be either high-spired or disc-shaped (Cain, 1977). This is very apparent when shell height (H) is plotted on diameter (D). For a community with a range of sizes, average values of H on D for species tend to follow a line greater than 45 or one less than 45, with few equidimensional examples. If sizes and shapes are related to available niches, then there should be a minimum distance in the HID space between species occupying the same habitat. If dimensions do matter in this way, then recently mixed populations are likely to display smaller average distances than ones which have evolved for a long time. A comparison between samples with endemics only and those which are mixed endemic, non-endemic showed no difference in average distance (Cook, 1984), so that again there is no evidence of interaction. Results from the statistical tests therefore do not disagree with the random model. The other way of examining the relation of species within communities via their HID distributions is to determine whether species from an available family fill in the HID space occupied elsewhere by another family, which is missing from the fauna (Cameron & Cook, 1989). Compared with N.W. Europe there is a deficiency of large, high-spired species, due to the absence of Chondrinidae, Orculidae and Enidae and poor representation of Clausiliidae. To some extent their place has been taken in the HID space by Ferussaciidae, which grow to a larger size than in Europe. Madeiran ferussaciids, however, differ from species of similar shape in European families in being more likely to burrow and less likely to climb on vertical surfaces. The scatter of disc-shaped species in the HID space is dominated by Helicidae in Madeira, much less so in Europe, and the helicids extend to a smaller range of size. However, both small low-spired and large high-spired species are less well represented in Madeira then in Europe and seemingly the Madeiran fauna has failed to fill these apparent niches. This may be because they have had too little time to adapt, or because the direct response route is unavailable since it means passing through an adaptive trough, or simply because the niche argument does not apply. It is probable that a community is partly structured by competition and partly not. A particular case on Madeira which seems to indicate competition is replacement of Caseolus bowdichianus by the introduced 17zeba pisana, which occurred on the eastern peninsula at the time offirst human occupation (Goodfriend et al., 1994).Johnson et al. (1993) conclude that evolution in the genus Partula on Moorea cannot be understood unless the competitive relationships ofthe different species are taken into account. At a higher level oftaxonomic distinctness, Baur (1988) found that variation in size in Chondrina clienta was affected by presence and density ofbalea peroersa. These examples remind us that detailed long-term ecological and behavioral studies are required to resolve the issue. The types of test described here may simply be too general to detect interactions at the level at which they occur. CONCLUSIONS We now have a much more detailed picture of the patterns of distribution of species than was available to the founders of the study of the Madeiran snail fauna,

11 MADEIRAN SNAIL FAUNA 467 and a better understanding ofthe possible nature ofadaptation and ofthe process of evolution as a whole. The majority of speciation appears to be allopatric and consequent upon the isolation, topography and disturbed geological history of the islands. There are some instances, however, which may represent selectively induced parapatric speciation. On the longer time scale, species from the available families fill some gaps in the HID distribution usually occupied by families which are accidentally missing, but in the shorter term we have not yet found evidence for interactions between species. On the whole, communities appear to be random assemblages of available and suitable species, or else the interaction effects take a very long time to come about. Wollaston (1856, 1878) refused to accept that speciation occurred, on the basis, he said, of his scientific observations (see Cook, 1995). He raised two objections. One was that we do not see speciation occurring. The other was that if it did occur through natural selection, then we would not expect to get associations of very similar sympatric species. Two points can be made about the first assertion. Even in a set ofislands with such a high species richness, species have come into being at a mean rate no faster than one per years, given the time scale and the number of colonizations (Cook, 1995). For comparison, Darwin's finches appear to have speciated at a rate ten times as fast (Grant, 1986). Species formation would not be witnessed often, and some of the variable taxa discussed above may in fact be evidence ofit. Secondly, in difficult cases, Wollaston (1878) tended to weight the evidence in his favour. As he says of Amphorella mitrifirmis, "... these abstract questions of 'species' and 'variety' are so difficult (indeed in many instances so impossible) to solve, that where any given form is sufficiently well defined to be easily recognisable, and is not connected with its nearest ally by decided intermediate links, I prefer for my own part (at any rate in those instances where it has already been enunciated and published) to accept it as specifically distinct." As to the coexistence of similar sympatric species, the reason is still unclear. It is not unaccountable, however, if there is frequent allopatric speciation, resulting from isolation ofterritories with similar habitats, and subsequent rejoining and overlap. If competition were strong, one or other of the new species would be replaced, but if, as seems to be the case, competition is weak or takes a long time to operate, then a build-up of similar sympatric species would take place. Overall, the evidence suggests that speciation is usually allopatric and competition weak. In considering why species should proliferate on isolated archipelagoes, compared with continental areas, it is notable that different types oforganisms have responded in similar ways. Although we might argue, for example, that some particular conditions on Madeira favour parapatric speciation in snails, that is hardly likely to apply to all the animal groups which have a paucity ofgenera but radiations within some ofthem. It is more reasonable to suggest that similar responses relate to a feature all groups share - a small probability ofarrival and common experience of a sequence of geographical isolating incidents brought about by vulcanism and sea level change. The endemic snail fauna is abundant in many parts of the islands, and has not suffered from human interference in the same way as, for example, endemic molluscs of the Pacific islands, perhaps because they are not so specialist in their requirements (Cameron, 1995). Although study of the Madeiran fossil bed suggests that a few species became extinct at the time ofcolonization by man, others may have increased

12 468 L. M. COOK their extent (Goodfriend et al., 1994). However, some species have extremely limited ranges within a single island, and could be totally eliminated by even a small building project; for that reason they are listed by the LU.C.N. as vulnerable (Wells et al., 1983). The distinctive helicid Actinella laciniosa occurs only at the north end of the small northern Deserta island, and the equally striking Dtscula turricula has a similar restricted range on the Porto Santan offshore islet of Cima. A varied community, described in a transect of the Cabo do Garajao by Cook et al. (1972), is now under a tourist development, and in general the southern coastal fauna can be regarded as threatened. Removal of sand from the Madeiran fossil bed for building purposes revealed the stratification, but at the same time destroyed much of it (Goodfriend et al., 1996). Measures exist to protect the Madeiran forest and the Deserta islands as national parks. Development of the drier low-lying parts of Madeira and Porto Santo for housing, industry and tourism could have severe effects, however, and planning should take the indigenous flora and fauna into account. PROSPECT With respect to molluscan evolution three approaches are now needed. One is to study the ecology of suitably chosen species in detail and to measure degree of niche overlap. Only then can we properly assess the importance of species interactions. Another is to take suitable pairs or groups of species and use molecular methods to measure their divergence (Johnson et al., 1986; Simon, 1991). Evidence obtained in this way may then be enlarged by construction of cladistic trees, giving order of emergence ofspecies. An indication of the timing and sequence ofsplitting should be obtained, which would help to explain why patterns of distribution differ between families and genera. The succession of colonizers may have arrived at any time over 10-15Myr while the fossil record extends for at most a million years. We can speculate about the phylogenetic processes involved, but the actual evolutionary paths followed in different groups have a strong historical component. The details we now have are like entries from a diary with most ofthe pages torn out; new directions ofenquiry could fill some critical gaps. ACKNOWLEDGEMENTS I am grateful to R.A.D. Cameron and all other colleagues working on the Madeiran islands for stimulating discussions. Work referred to has been funded by N.E.R.C., The Royal Society and The Linnean Society and the National Geographic Society. REFERENCES AlbersJC Malacographia Maderensia sive enumeratio molluscorum quae in insulis Maderae etportus Sancti autvive extant autfossilis repenuntur. Berlin: Reimer. Barton NH, Hewitt GM Analysis of hybrid zones. AnnualReview 'IfEcology and Systematics 16: Baez M Origin and affinities of the fauna of Madeira. Boletim do Museu Municipal do Funchal Suppl. 2: 9-40.

13 MADEIRAN SNAIL FAUNA 469 Baur B Microgeographical variation in shell size of the land snail Chondrina clienta. Biological Joumal qf the Linnean Socie!>, 35: 247~259. Bischoff W, Osenegg K, Mayer W Untersuchungen zur subspezifischen Gliederung der Madeira Mauereidechse Podarcis dugesii (Milne-Edwards, 1829). Salamandra 25: Bush GL Modes of animal speciation. AnnualReview qf Ecology and ~stematics 6: Cain AJ Variation in the spire index of some coiled gastropod shells and its evolutionary significance. Philosophical Transaction qf the RoyalSocie!>, qf London, Series B 277: Cain AJ, CurreyJD Area effectsin Cepaea. Philosophical Transactions ofthe RoyalSociety qflondon, Series B 246: Cameron RAn Pattern of diversity in land snails: the effectsof environmental history. in: Van Bruggen AC, Wells SM, Kemperman CM, eds. Biodioersity and conservation of themollusca. Leiden: Backhuys Cameron RAn, Cook LM Shell size and shape in Madeiran land snails: do niches remain unfilled? BiologicalJoumal qf thelinnean Socie!>, 36: Cameron RAn, Cook LM The development of the diversity of the land snail fauna of the Madeiran archipelago. Biological Joumal ofthelinnean Socie!>, 46: Cameron RAn, Cook LM, Gao G Variation in snail species widespread on Porto Santo. Joumal of the Molluscan Studies 62: Cameron RAn, Cook LM, HallowsJ Land snails on Porto Santo: adaptive and non-adaptive radiation. Philosophical Transactions qf theroyalsociety oflondon, Series B 351: Castello de Paiva, Bario do (Antonio da Costa Paiva) Monographia molluscorum terrestriumfiuvialium lacustnum insularium Maderensium. Academic Press: Lisbon. Cook LM Variation in the Madeiran lizard Lacerta dugesii. Joumal cf Zoology 187: Cook LM The distribution ofland snails in eastern Madeira and the Desertas. In: Solem A, Van Bruggen A, eds. World-WuJe Snails. Leiden: EJ. Brill, Cook LM T. Vernon Wollaston and the "monstrous doctrine". Archives qf NaturalHistory 22: Cook LM, Lace LA Sex and genetic variation in a helicid snail. Heredity 70: Cook LM, Cameron RAn, Lace LA Land snails of eastern Madeira: speciation, persistence and colonization. Proceedings oftheroyal Socie!>, of London, Series B 239: Cook LM, Goodfriend GA, Cameron RAn Changes in the land snail fauna of eastern Madeira during the Quaternary. Philosophical Transactions qf theroyal Socie!>, of London, Series B 339: Cook LM,Jack T, Pettitt CWA The distribution ofland molluscs in the Madeiran Archipelago. Boletim domuseumunicipal do Funchal26: Dajoz R Coliopteres: Coiydiidae et Anommatidae paliarctiques. Paris: Masson. Enghoff H Macaronesian millipedes (Diplopoda) with emphasis on endemic species swarms on Madeira and the Canary Islands. BiologicalJoumal qf thelinnean Socie!>, 46: Erber D, Hinterseher W Contribution to the knowledge of Madeiran beetles. Boletim domuseu Municipal do Funchal40: Goodfriend GA, Cameron RAn, Cook LM Fossil evidence of recent human impact on the land snail fauna of Madeira. Joumal ofbiogeograpjry 21: Goodfriend GA, Cameron RAn, Cook LM, Courty M-A, FedoroffN, Kaufinan A, Livett E, TallisJ Quaternary eolianite sequence of Madeira: stratigraphy, chronology, and paleoenvironmental interpretation. Palaeogeography, Paleoclimatology, Paleoecology 120: Grant PR Ecology and evolution qf Darwin'sfinches. Princeton, NJ: Princeton University Press. Hadfield MG, Miller SE Demographic studies on Hawaii's endangered tree snails: Partulina proxima. Pacific Science 43: Hansen A Checklist of the vascular plants of the archipelago of Madeira. Boletim do Museu Municipal do Funchal24: Hounsorne MY Biometrics and origins of some Atlantic island birds. Boletim domuseumunicipal do Funchal Suppl. 2: Johnson MS, MurrayJ, Clarke B An electrophoretic analysis of phylogeny and evolutionary rates in the genus Partula from the Society Islands. Proceedings of the RoyalSocie!>, oflondon, Series B 227: Johnson MS, MurrayJ, Clarke B The ecological genetics and adaptive radiation of Partula on Moorea. Oifotd Survrys in Evolutionary Biology 9: Kerney MP, Cameron RAn A field guide to the land snails of Britain and north-west Europe. London: Collins. Lace LA Variation in the genitalia of the land snail Heterostoma paupercula (Lowe, 1831) (Helicidae) in Madeira. Biological Joumal qf thelinnean Socie!>, 46: Lowe RT Primitiae Faunae et florae Maderae et Portus Sancti. Transactions ofthecambridge Philosophical Socie!>, 4: 1-70 (Also published as a separate by Van Voorst, London, 1851). Lowe RT Brief diagnostic notices of new Madeiran land shells. AnnalsandMagazine ofnaturalhistory 9(2): , Lowe RT Catalogus molluscorum pneumonatorum insularum Maderensium: or a list of all the land and freshwater shells, recent and fossil, of the Madeiran Islands: arranged in groups according to their natural affinities; with diagnoses of the groups, and of the new or hitherto imperfectly defined species. Proceedings qf the Zoological Socie!>, qf London 22:

14 470 L. M. COOK LynchJD The gauge of speciation: on the frequencies of modes of speciation. In: Otte D, EndlerJA (eds.) Speciation and its consequences. Sunderland, MA: Sinauer, Machado A Monographia de los cardbtdos de las Islas Canarias (Insecta, Coleoptera). La Laguna: Instituto de Estudios Canarios. Mandabl-Bartb G Systematische Untersuchungen tiber die heliciden-fauna von Madeira. Abh. Senckenbergische Nauuforschende Gesellschoft 469: Maynard SmithJ Sympatric speciation. American Naturalist 100: Mayr E Animalspecies andevolution. Cambridge, MA: Harvard University Press. Mitchell-TboDll? RC Radiometric studies in macaronesia, Boletim do Museu Municipal do Funchal 37: Murray J Genetic dioersuy and natural selection. Edinburgh: Oliver & Boyd. Noor MA Speciation driven by natural selection in Drosophila. Nature, London 375: Owen DF, Smith DAS The origin and history of the butterfly fauna of the north Atlantic islands. Boletim domuseu Municipal do Funchal Supp!. 2: Patouret L, AuzendeJ-M, Le Lann A, OlivetJ-L Temoins des variations glacio-eustatiques du niveau marin et des mouvements tectoniques sur Ie bane de Gorringe (Atlantique du Nord-Est). Palaeogeography, Palaeoclimatology, Palaeoecology 32: Schutt H Tiakische Landschnecken. Wiesbaden: Hemmen. Sitnon C Organelle DNA studies of the evolution of recently derived species complexes in the Hawaiian islands. In: Dudley EC, ed. The uniry qf evolutionary biology. Portland, Oregon: Dioscorides Press, Sjogren E Vascular plant communities of Madeira. Boletim do Museu Municipal do Funchal26: Swash AR, Askew RR A survey of madeiran butterflies. Boletim do Museu Municipal do Funchal 34: Vandel A Les isopodes terrestres de l'archiepl maderien, Memoires dumuseum National d'histoire Naturelle. A22: Walden HW Systematic and biogeographical studies in the terrestrial Gastropoda of Madeira. With an annotated Check-list. Annales Zoologici Fennici 20: Walden HW. 1984a. The land mollusc fauna of Madeira in relation to other Atlantic islands and the Palaearctic region. In: Solem A, Van Bruggen AC, eds. World-Wuie Snails. Leiden: EJ. Brill, Walden HW. 1984b. On the origin, affinities, and evolution of the land Mollusca of the Mid-Atlantic islands, with special reference to Madeira. Boletim do Museu Municipal do Funchal36: Walker MD The petrology of Madeira and Porto Santo. Ph.D. thesis, Manchester: University of Manchester. Wells SM, Pyle RM, Collins NM. eds The IUCN intertebrate red data book. Gland, Switzerland: IUCN. Wbeater CPo Habitat similarities of Coleoptera from the Madeiran archipelago. Boletim domuseu Municipal do Funchal Supp!. 2: White l\gd Modes ofspeciation. San Francisco: Freeman. Wollaston TV On the variation of species with especial riference to theinsectai folloioed by an inquiry intothenature qf genera. London: Van Voorst. Wollaston TV Testacea Atlantica or the landand Freshwater Shells qf theazores, Madeira, Salvages, Canaries, Cape Verdes and SaintHelena. London: Reeve. Zbyszewsld G Levantamentos geologicos na parte oriental da ilha da Madeira e nas ilhas Desertas. Memonas academica das Ciencias, Lisboa 16:

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