Patterns of larval hostplant usage among hawkmoths (, ) from La Réunion, with a comparison of the Mascarenes with other regions of the world.

Marc At t i é 1, Ian J. Ki t c h i n g 2 & Jacques Ve s l o t 3

Résumé.— Modèles d’utilisation des plantes-hôtes par les larves de sphinx (Lepidoptera, Sphingidae) à la Réunion, avec une comparaison des Mascareignes avec d’autres régions du monde.— Les espèces de sphingides de la Réunion sont en grande partie originaire de Maurice et de Madagascar d’où elles ont migré avec probablement des préadaptations écologiques, notamment des préférences pour des plantes-hôtes. Dans la présente étude on cherche à savoir si les préférences alimentaires sont conservées ou bien si elles résultent d’adaptations à des contraintes locales. À l’aide d’Analyses Factorielle des Correspondances (AFC), on a comparé le spectre de plantes-hôtes de six espèces polyphages à large distribution (Acherontia atropos, convolvuli, fulvinotata, nerii, celerio, H. gracilis) et on constate qu’elles utilisent généralement le spectre de familles de plantes-hôtes disponible, les restrictions observées correspondant le plus souvent à l’absence de familles botaniques à la Réunion plutôt qu’à une spécialisation alimentaire. On peut noter cependant que a été trouvé sur plusieurs genres d’Apocynacées (, , Pachy- podium, ) mais semble, à la Réunion, peu fréquent sur Thevetia peruviana, une espèce com- mune appartenant à la même famille. De même Coelonia fulvinotata a été observé sur camara, une exotique invasive, alors que d’autres plantes-hôtes potentielles sont présentes sur l’île. L’analyse factorielle des données sur les plantes-hôtes se rapportant aux espèces distribuées dans les 11 genres de sphinx présents à la Réunion a permis de constater que les principales associations avec les plantes-hôtes sont conservées dans l’aire de distribution de ces espèces, bien que des spécificités alimentaires puissent exister localement. Les associations montrent clairement l’existence d’un conservatisme taxinomique (Acherontia, Agrius et Coelonia sont associés de préférence avec des Euasterids I) ou sont de nature phytochimique (Nephele densoi est associé à deux familles lactifères appartenant à des sous-classes botaniques distinctes).

Summary.— The hawkmoth fauna of La Réunion is largely the result of colonization from and Madagascar. These will have arrived with preadapted ecologies, including larval hostplant preferences. Here we investigate these preferences to determine whether they have been conserved or have adapted to local constraints. Using Factorial Correspondence Analysis (FCA), we compare the hostplant spectra of six widespread, polyphagous species (Acherontia atropos, , Coelonia fulvinotata, Daphnis nerii, , H. gracilis) and show that generally they utilize the full spectrum of available families, any restriction reflecting absence of particular families from La Réunion rather than specialization. However, although Daphnis nerii is found on several (Nerium, Ochrosia, Pachypodium, Tabernaemontana) on La Réunion, it seems to avoid the abundant species, Thevetia peruviana. Also, Coelonia fulvinotata seems to have developed an association with the invasive alien, , even though several other potential recorded hostplants also occur on the island. In some instances, absence of records in the region may also be due to a lack of field observations. We also analysed the known global host ranges of all species of the 11 sphingid genera on La Réunion. We found principal hostplant associations are conserved across geographical regions, although there may be local preferences, and that these associations may be taxonomic (Acherontia, Agrius and Coelonia are preferentially associated with Euasterids I) or phytochemical (Nephele densoi, associated with two taxonomically distinct lactiferous families).

1 corresponding author. 10 chemin des Vacoas, 97490, Sainte-Clotilde, La Réunion, . E-mail: marc.attie@ gmail.com 2 department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK. E-mail: [email protected] 3 cemagref, UR Hydrobiologie, route de Cézanne CS 40061, F-13182 Aix-en-Provence Cedex 5. E-mail: [email protected]

Rev. Écol. (Terre Vie), vol. 65, 2010.

­– 3 ­– Most hawkmoths (Sphingidae) occurring on the islands in the (e.g. Acher- ontia atropos, A. convolvuli, Basiothia medea, Coelonia fulvinotata, C. solani, Daphnis nerii, Euchloron medea, Hippotion celerio and H. gracilis) have morphological adaptations that aid dispersal over large distances, especially when assisted by favourable weather conditions such as strong winds. This provides a partial explanation of the broad geographical distributions of many of these species, which can encompass whole continents or even the entire Old World. The sphingid fauna of La Réunion, an island that is only 3 myr old (Chamalaun & McDou- gall, 1966), appears to have developed in large part through dispersal and colonization from Mauritius (itself only 8 myr old) (McDougall & Chamalaun, 1969) and Madagascar, which are respectively only 180 and 700 km distant. Various intrinsic and extrinsic factors interact to determine and maintain -hostplant relationships (Lewinsohn & Roslin, 2008). Intrinsic factors, such as the adaptive resources of (Moran, 1988), tend to optimize the host range of the phytophagous insects, and oper- ate within the phylogenetic context of historical biological constraints (Mitter & Farrel, 1991). However, extrinsic factors, such as which actually occur within the insect’s geographi- cal and ecological range, also play a prominent part (Dethier, 1954; Ehrlich & Raven, 1964; Agrawal et al., 2006), and Beck & Kitching (2007) demonstrated that larval diet breadth was the best predictor of range size, highlighting further the close association between these two factors. Climatic factors, human activity, genetic constitution and interspecies competition can have fundamental, if generalized, roles in determining the distributions of sphingid species (Pittaway, 1993), and habitat disturbance may affect the biodiversity of hawkmoth assem- blages, as shown by Beck et al. (2006) in Southeast . Studies of spatial models of plant-insect interactions have made rapid advances in recent years (Farell & Mitter, 1994; Thompson, 1994; Mopper, 1998; Mopper et al., 2000; De Jong & Nielsen, 2002). It is now well known that phytophagous insects play a significant part in determining plant distributions and evolutionary interactions are used to explain the radiative adaptations of plants and insects; for example, insect speciation in relation to the morphologi- cal and chemical variability of plants (McEvoy, 2002; Thompson, 1994). Insect adaptations to local biotopes are important components of the evolutionary process that lead to hostplant spe- cialization (Gotthard et al., 2004). The degree of preference for different plants is determined in part by the plant stimuli received by the insect (Courtney et al., 1989) and such preferences have surely played a part in the evolution of sphingids and their larval hostplant choices. The study of trophic relationships of hawkmoths on the isolated oceanic island of La Réunion is particularly interesting because the entire sphingid fauna would seem to have originated by natural dispersal from the pre-existing neighbouring islands of Madagascar and Mauritius. As a result, these hawkmoth species will have arrived with preadapted ecologies, including larval hostplant preferences. How, then, did these new arrivals persist under the new environmental constraints? Did they conserve their ancestral ecological characteristics (niche conservatism) (Wiens & Graham, 2005), the hostplant specificities of their geographical sources, or did they change to adapt to the new local conditions on La Réunion? When viewed on a global scale, many widespread sphingid species appear to be highly polyphagous, but on a local scale they may not, in fact, show the same larval hostplant spectrum in all parts of their ranges. Instead, they may adapt their trophic relationships to local conditions so that the pat- terns on large continental areas differ from those on smaller islands. A comparison of hostplant use between narrowly endemic and widespread species of hawkmoths, in both continental areas and on islands such as La Réunion, can provide insights into general ecological patterns. In addition, it has been shown that taxonomically related insects often feed on taxonomically related plants, as demonstrated by Ehrlich & Raven (1964) for different families of butterflies. Does such taxonomic conservatism occur in Sphingidae, at the species and/or generic level, and at what geographical scale? Is a particular species of Sphingidae, for example, conserva- tive in its larval hostplant choice on small islands but more polyphagous on continental-sized areas? In the present paper, we attempt to answer these questions by analysing the host range of the Sphingidae of La Réunion in the different regions in which they occur worldwide, includ- ing islands and continental areas, and by studying at a global scale the host range of all species in those genera present on La Réunion for which larval hostplant data exist.

­– 4 ­– Materials and methods

Sphingidae of the Indian Ocean Madagascar is characterized by a moderate-sized sphingid fauna of some 60 species (Griveaud, 1959; Carcasson, 1968). On the Comoro Islands, Turlin (1996) recorded 38 species and subspecies. Diversity in the , is low but not poor considering the young age and restricted area of the archipelago: 13 (Vinson, 1938) or 12 species (Viette, 1992, 1996) on Mauritius, there being uncertainty regarding the presence of Coelonia solani solani (Boisduval 1833) and the possible extinction of the endemic soror Rothschild & Jordan 1903, and 15 species on La Réunion. Mauritius and La Réunion have closely similar Sphingidae faunas, although C. solani is apparently absent from Mauritius but has been recorded in La Réunion (Attié & Morel, 1997), likewise Nephele densoi (Keferstein 1870) and biguttata (Walker 1856). There are also several diurnally-active endemic species: M. soror and trochilus (Guérin-Méneville 1843) on Mauritius, and M. milvus (Boisduval 1833) and C. apus (Boisduval 1833) on La Réunion (Boisduval, 1833; Guénée, 1862; Viette, 1957, 1992, 1996; Guillermet, 2006). The other species present on La Réunion also occur to Madagascar and Comoro Islands and some have broader distributions (Tab. I). The status of (Cramer 1779) in the Mascarenes is reviewed in this study.

Table I Geographical distribution of the 15 species of hawkmoths recorded on La Réunion Macroglossinae

Regions Acherontia atropos Acherontia Agrius convolvuli Coelonia fulvinotata Coelonia solani Basiohia medea Cephonodes apus Daphnis nerii megaera lacordairei Euchloron Hippotion celerio Hippotio gracilis Hyles biguttata aesalon Macroglossum Nephele densoi Nephele oenopion La Réunion + + + + + + + + + + + + + + + Mauritius + + + + + + + + + + Madagascar + + + + + + + + + + + + + Comoro Islands + + + + + + + + + African mainland + + + + + + + Western Palaearctic region + + + +

Ho s t p l a n t r e c o r d s On La Réunion, one of us (M.A.) undertook intensive fieldwork in indigenous forests, secondary vegetation and gardens between 1995 and 2002, mostly during the warm and wet season from October to March (Tab. II). Most field records date from this period but a few derive from subsequent years. The study sites were of two types: (A) “undisturbed habitats” that have essentially native vegetation; and (B) “disturbed habitats”, including cultivated areas and secondary vegetation that has been invaded by exotic plants. Both indigenous and exotic plants along and adjacent to pathways were examined and hawkmoth eggs and larvae collected. damage and larval frass often provided indications of the presence of hawkmoth nearby. Larvae were reared to adult stage in the laboratory using the hostplant upon which they were found. This permitted the identities of those for which the immature stages were previously unknown (e.g. Macroglossum milvus) to be confirmed. Hawkmoth and plant nomenclature follows Kitching & Cadiou (2000) and the Flore des Mascareignes (1976-2009) respectively. The system proposed by the Angiosperm Phylogeny Group (henceforth APG) (2003) was used as the best current hypothesis of the higher classification of plants. Hostplant associations were primarily literature records extracted from the HOSTS database maintained by the Natural History Museum, London, U.K. (Robinson et al., 2001), supplemented by records from literature and web sources known not to have yet been incorporated into that resource. Very few associations have been reported in the literature from the Indian Ocean region, particularly from native plants. With regard to the Mascarenes, data for Mauritius are

­– 5 ­– sourced primarily from Vinson (1938), and for La Réunion from Boisduval (1833), Guenée (1862), Guillermet & Guillermet (1986) and Viette (1957, 1992, 1996). Hostplant records for Madagascar are scarce, Griveaud (1959) adding only a single reliable record (Daphnis nerii on Cataranthus) to those from the older literature. Those records listed by

Table II Sites on La Réunion searched for eggs and larvae of sphingids by MA or communicated directly to him, with altitude and vegetation type (see text for explanation). (+: immature stages found, M: data from Muséum d’Histoire Naturelle of La Réunion (Parnaudeau, pers. comm., 2003), *: died in prepupation)

Type Altitude of

Sites (m) vegetation atropos Acherontia Agrius convolvuli Cephonodes apus Coelonia fulvinotata Coelonia solani Daphnis nerii Hippotion celerio Hippotion gracilis Hyles biguttata milvus Macroglossum aesaelon Macroglossum Nephele densoi Bébour 1350 A + + Bois-Court 950 B + + + Cap-Noir 1200 A + + Cilaos 1000 B Col des Boeufs 2000 B + Colorado 650 B + + + Forêt Dugain 800-950 A + Grand-Bassin 800-1000 B + + + Grande-Ravine 50 B + La Bretagne 250-300 B + + + La Montagne 300 B + Le Brûlé (Saint-Denis) 800 B + Mafate B + Mare-Longue (Nature Reserve) 270 A +* + Plaine d’Affouches 900-1200 A + + Plaine des Chicots 1200-1400 A + Plaine des Fougères 1450 A + + Ravine Bernica 50 B + Ravine de la Chaloupe St-Leu 400 B + Ravine de la Grande Chaloupe 70 B + Roche-Verre-Bouteille 1150 A Sainte-Clotilde 70 B + + + Saint-François 500 B + Saint-Gilles les Hauts 400 B + Takamaka 700-750 A + Tampon 700-800 B + + M M Vallée-Heureuse 790 A +

­– 6 ­– Desegaulx de Nolet (1984) for the Indian Ocean region are certainly unoriginal secondary citations that mostly cannot be precisely assigned to a particular country and thus cannot be used in a comparative analysis of diet among the regions. Hawkmoth-plant associations observed on La Réunion, and more broadly in the Mascarenes (La Réunion, Mauritius), are compared with those of other countries and regions in which the hostplants of those genera of Sphingidae that occur on La Réunion have been particularly well studied: (Moulds, 1981, 1984), Ivory Coast (Vuattoux, 1978; Vuattoux et al., 1989), South Africa (Pinhey, 1975; Robertson, 2004), East Africa (Le Pelley, 1959; Sevastopulo, 1949- 1975, 1980), Europe (Marktanner, 1976), and the western Palaearctic region (Pittaway, 1993). All sources on this list are also incorporated into HOSTS (Robinson et al., 2001). In the process of compiling the hostplant data, we encountered numerous anomalous records. For example, some hostplants reported for Acherontia atropos in Europe by Marktanner (1976), such as Anethum, Daucus, Brassica, Beta and Nerium, are unusual. The first four are all garden crops and could refer to wandering prepupal larvae that had actually fed on other plants growing among the crops. The Nerium record initially appeared doubtful but Westwood 1847 has also been recorded on this plant in (Beller & Bhenchitr, 1936), as well as on serpentina (Apocynaceae) in (Harvir Singh et al., 1984). In Europe, A. atropos is not autochthonal, but migrates north from Africa (Forster & Wohlfahrt, 1960; Pittaway, 1993). Thus, some European hostplant records may be simply opportunistic oviposition on plants in an unfamiliar flora that the larvae just happen to be capable of digesting. Many such errors may occur but go undetected because the larvae die at an early stage of development (Janz, 2003). Other atypical records include those of Hippotion celerio in Ivory Coast on Moraceae (Ficus) and Euphorbiaceae (Manihot), which Vuattoux et al. (1989) themselves noted as “associations non conformes”. It is unlikely that these records, which have never been observed in other countries, were simple errors but nor is it clear to what degree the larvae successfully developed on these plants, as no survivorship figures were given. Although some of these dubious associations may be erroneous, we nevertheless included them in our analyses because, in the absence of evidence to the contrary, they could represent regional dietary specializations. We did, however, correct some clearly erroneous literature records. For example, the association between Macroglossum milvus and foetida on Mauritius reported by Vinson (1938) certainly refers not to M. milvus, which is strictly endemic to La Réunion, but to M. aesalon or to M. soror, a Mauritian endemic that may now be extinct (Viette, 1992). The larvae of “Coelonia fulvinotata” briefly described by Griveaud (1959) are A. atropos, because the anal horn is described as “recourbée et tuberculée”, a feature that distinguishes A. atropos from C. fulvinotata (Kitching, 2003). Griveaud (1959) gave four host records for this misidentification: Dahlias, Duranta, and Meliaceae. The first two records originate from South Africa (Fawcett, 1901) not Madagascar. The third was derived from Denso (1944), who actually said explicitly that this was a family on which he had not found larvae of C. fulvinotata. He only recorded C. fulvinotata larvae from a low growing, heavily cut, red-flowered Meliaceae that was not identified further. Thus, all the records listed by Griveaud (1959), other than the undetermined red-flowered Meliaceae, are erroneous associations for C. fulvinotata on Madagascar. This demonstrates the need to determine the original sources of records and not rely uncritically on secondary citations. Finally, confusion between C. fulvinotata and C. solani explains why the larvae and hostplants of C. fulvinotata have often been reported in literature as those of C. solani. The hostplants for C. solani on Mauritius cited by Boisduval (1833) (), Vinson (1938) (Nicotiana, Ehretia, Nuxia) and Mamet & Williams (1993) (Solanum) all refer to C. fulvinotata (Attié & Morel, 1997; M. Attié, unpublished rearing data). Viette (1996) questioned the presence of C. solani on Mauritius. We found adults of only C. fulvinotata in the collection of the Mauritius Sugar Industry Research Institute (J.R. Williams collection, Le Réduit, Mauritius) and there were no specimens of C. solani in the collection of the Ministry of Agriculture (Le Réduit, Mauritius). These observations argue strongly that C. solani does not occur on Mauritius.

Factorial Correspondence Analysis (FCA) We here define a record as an association between a species of hawkmoth and a plant , irrespective of the number of species of hostplant genus involved. Plant records at the species level can provide important information about the relationship between specialized species of hawkmoths and their hostplants. However, for generalist species associated with numerous hostplants in different genera or families, we consider analysis at the levels of plant genus to be sufficiently precise. Furthermore, if records were analysed at the level of hostplant species, the disparity in numbers of records between cultivated areas and natural vegetation, and between well-known and poorly known , becomes greater. Furthermore, entomologists, rather than botanists, made the majority of plant identifications and thus identification to plant species is the factor with the greatest degree of uncertainty in the data. Analysis at the level of plant genus greatly reduces the potential for erroneous identifications. Factorial correspondence analysis (FCA) is a factor analysis designed to explore two-way frequency tables by representing the distances between rows and the distances between columns in a space of lower dimension. We first use FCA to investigate association frequencies by region and by species of Sphingidae. The species distributions and availability of hostplant records by region led us to select the following six species of Sphingidae: Acherontia atropos, Agrius convolvuli, Coelonia fulvinotata, Daphnis nerii, Hippotion celerio and H. gracilis. Hippotion gracilis was for a long time treated as a junior of H. eson. However, Eitschberger (2006) showed them to be two separate species that are sympatric throughout most of the African mainland. However, in the absence of voucher material, there is no way of allocating pre-2006 hostplant records between the two species. Thus, they are here treated as a single unit and referred to as H. gracilis for convenience (and because it seems that H. gracilis may be the only one of the two present on La Réunion). Records from La Réunion were incorporated into a Mascarenes data set that also included records from Mauritius. Likewise, records from Europe were incorporated into a more general western Palaearctic region data set. Data from Ivory Coast, South Africa, East Africa and Australia were sufficiently great to be analysed independently. However, the comparisons between regions are not entirely equivalent for these six species of hawkmoths, for not all

­– 7 ­– Table III Hawkmoth species in those genera found on La Réunion for which hostplants have been recorded. Codes are the abbreviations used for selected species in Fig. 3. * indicates the 41 species analysed in the second FCA

SA1* Acherontia atropos SH41* SM81* Macroglossum fritzei SA2* Acherontia lachesis SHY42 Hyles annei SM82 SA3* Acherontia styx SHY43 Hyles apocyni SM83 Macroglossum heliophila SAG4* SHY44 Hyles biguttata SM84 Macroglossum hirundo SAG5* Agrius convolvuli SHY45* Hyles calida SM85 Macroglossum insipida SAG6 Agrius cordiae SHY46 Hyles centralasiae SM86 Macroglossum mediovitta SB7 Basiothia aureata SHY47 Hyles chamyla SM87 Macroglossum micacea SB8 Basiothia charis SHY48 Hyles chuvilini SM88 Macroglossum milvus SB9* Basiothia medea SHY49 Hyles costata SM89* Macroglossum neotroglodytus SB10 Basiothia schenki SHY50 Hyles cretica SM90 Macroglossum nycteris SC11 Cephonodes apus SHY51 Hyles dahlii SM91* Macroglossum obscura SC12* Cephonodes armatus SHY52 SM92 Macroglossum particolor SC13* SHY53* Hyles euphorbiarum SM93 Macroglossum passalus SC14* Cephonodes kingii SHY54* SM94 Macroglossum poecilum SC15* Cephonodes picus SHY55 SM95* Macroglossum prometheus SC16 Cephonodes tamsi SHY56* SM96* Macroglossum pyrrhosticta SC17 Cephonodes xanthus SHY57* SM97 Macroglossum regulus SCO18* Coelonia fulvinotata SHY58 Hyles livornicoides SM98 Macroglossum sitiene SCO19 Coelonia solani SHY59 Hyles nervosa SM99* Macroglossum stellatarum SD20* SHY60 Hyles nicaea SM100 Macroglossum stigma SD21 Daphnis minima SHY61 Hyles perkinsi SM101 Macroglossum tenebrosa SD22* Daphnis nerii SHY62 Hyles robertsi SM102* Macroglossum trochilus SD23* Daphnis placida SHY63 Hyles sammuti SM103 Macroglossum vacillans SD24 Daphnis protrudens SHY64 Hyles siehei SM104 Macroglossum variegatum SD25 Daphnis torenia SHY65 SM105 Macroglossum vicinum SE26* Euchloron megaera SHY66 SN106* Nephele accentifera SH27 Hippotion aurora SHY67 Hyles wilsoni SN107* Nephele aequivalens SH28 Hippotion balsaminae SHY68 Hyles zygophylli SN108 Nephele argentifera SH29* Hippotion boerhaviae SM69 Macroglossum aesalon SN109 Nephele bipartita SH30 Hippotion brennus SM70 Macroglossum affictitia SN110* Nephele comma SH31* Hippotion celerio SM71 Macroglossum alcedo SN111 Nephele densoi SH32 Hippotion echeclus SM72 Macroglossum alluaudi SN112 Nephele discifera SH33* Hippotion gracilis SM73 Macroglossum aquila SN113 Nephele funebris SH34 Hippotion geryon SM74 Macroglossum assimilis SN114 Nephele hespera SH35* Hippotion osiris SM75* SN115 Nephele oenopion SH36 Hippotion rafflesii SM76 Macroglossum bifasciata SN116* Nephele peneus SH37 Hippotion rosae SM77 Macroglossum bombylans SN117* Nephele rosae SH38 Hippotion roseipennis SM78* SN118 Nephele subvaria SH39* SM79 Macroglossum divergens SN119 Nephele vau SH40* Hippotion scrofa SM80 Macroglossum dohertyi

­– 8 ­– are present in each of the studied regions. All species selected from the Mascarenes are present in Ivory Coast, South Africa and East Africa. In this last region, however, Agrius convolvuli associations have been recorded only at plant family level and therefore this species was excluded from comparisons involving East Africa. Acherontia atropos, C. fulvinotata, D. nerii and H. gracilis are absent from Australia, where among the studied species only A. convolvuli and H. celerio have been recorded. Coelonia fulvinotata and H. gracilis are absent from the western Palaearctic region. We then used FCA to investigate association frequencies within the plant families used by the 41 hawkmoth species from around the world that belong to the 11 genera present on La Réunion (Acherontia, Agrius, Basiothia, Cephonodes, Coelonia, Daphnis, Euchloron, Hippotion, Hyles, Macroglossum and Nephele). Only those hawkmoth species with at least three records of host genera were included because rare species bring associations too specific to be analysed (Tab. III, species indicated by * were used in the FCA). Likewise, those plant families with only one or two recorded genera were also removed. Correspondence analysis is sensitive to sampling effort but it is always difficult to know how this affects the resulting patterns, especially when there is no information to quantify sampling effort. However, at the plant genus level, the selection of not too low marginal frequencies for both rows and columns, and, at the plant family level, the increased marginal frequencies of aggregated data should lessen this risk. All statistical analyses were undertaken using the R statistical package (R Development Core Team, 2006).

Results

Patterns of host use on La Réunion Searches of 27 sites in different habitats on La Réunion resulted in the collection of larvae or eggs of 12 hawkmoth species of the subfamilies Sphinginae (Acherontia atropos, Agrius convolvuli, Coelonia fulvinotata and C. solani) and Macroglossinae (Cephonodes apus, Daph- nis nerii, Hippotion celerio, H. gracilis, Hyles biguttata, Macroglossum aesalon, M. milvus and Nephele densoi) (Tab. II). A total of 61 larval hostplants was recorded, of which 19 asso- ciations are new (Annex 1). The percentage of hostplant genera on the plant families and the percentage of endemic, indigenous and exotic associations for 12 species of hawkmoths are summarized in Table IV.

Comparison of generalists and specialists on La Réunion Of the generalist species, Acherontia atropos is the most polyphagous, being recorded from 24 genera in 11 plant families (Tab. IV). Euasterids I account for the large majority of these, and 67 % are from order , of which , and are the most frequently observed larval hosts in La Réunion. The other host families of A. atro- pos are represented by single genera classified in orders assigned to Eurosids I (Cucurbitaceae and ) or, for the first time, Annonaceae in Magnoliids. Hippotion gracilis is the next most generalist species on La Réunion, being recorded from 12 genera in eight hostplant fami- lies (Tab. IV), mostly (Basal Monocots) and Vitaceae (). If we consider all the regions in which they occur, Agrius convolvuli, Coelonia fulvino- tata, Daphnis nerii and Hippotion celerio are generalists with a complex host use. However, on La Réunion, the first two of these species have been recorded on only one hostplant family and D. nerii and H. celerio on only two. D. nerii was recorded on five genera of Apocynaceae and one Moraceae. Apocynaceae thus seems to be the preferred hostplant family for this spe- cies on the island. The abundance of cultivated Apocynaceae, especially Nerium oleander, on La Réunion may preferentially attract ovipositing D. nerii despite being well rep- resented (and recorded as a hostplant on Mauritius and elsewhere). It is also noteworthy that another apocynaceous genus, Thevetia, has never been recorded as a hostplant on La Réunion, despite records from both East Africa and the western Palaearctic region and T. peruviana being commonly cultivated on the island. The other three species present few hostplant records on La Réunion. Agrius convolvuli was observed only on a species of but it probably also feeds on other cultivated species of not searched in this study. Its larvae are well known to cause damage to sweet (Ipomoea batatas), on which Vinson (1938) reported this to be a minor on Mauritius, and other crops in India and (Bell & Scott, 1937). Likewise, C. fulvinotata has a diverse host range in the African regions but was recorded on La Réunion only from Lantana camara (Verbenaceae). Hippotion celerio

­– 9 ­– Table IV Percentage of hostplant genera in plant families for 12 species of hawkmoths on La Réunion and percentage of endemic, indigenous and exotic associations according species of Sphingidae

Subclasses Orders Plant families atropos Acherontia Agrius convolvuli Coelonia fulvinotata Coelonia solani Cephonodes apus Daphnis nerii Hippotion celerio Hippotion gracilis Hyles biguttata milvus Macroglossum aesalon Macroglossum Nephele densoi Magnoliids Magnoliales Annonaceae 4.2 Basal Alismatales Araceae 33.3 Monocots 8.3 Core 50

Geraniales Geraniaceae 8.3 Rosids Onagraceae 8.3 Unplaced Vitaceae 50 16.7 Rosids Cucurbitales Cucurbitaceae 4.2 Eurosids I Fabales Fabaceae 4.2 Moraceae 20 100

Balsaminaceae 8.3 100 Unplaced Boraginaceae 8.3 Euasterids I Apocynaceae 80 8.3 33.3 Rubiaceae 4.2 100 8.3 66.7 100

Acanthaceae 4.2 Euasterids I Bignoniaceae 20.8 Lamiales Oleaceae 12.5 Stilbaceae 4.2 Verbenaceae 25 100 100

Convolvulaceae 100 Solanaceae 8.3

Endemics 10.7 0 0 100 100 33.3 0 15.4 0 75 0 0

% plant Indigenous 7.1 0 0 0 0 0 50 30.8 100 25 0 100 Exotics 82.1 100 100 0 0 66.7 50 53.8 0 0 100 0 Associations 28 1 1 1 2 6 2 13 1 4 1 1

Number of Genera 24 1 1 1 2 5 2 12 1 3 1 1

Families 11 1 1 1 1 2 2 8 1 2 1 1

­– 10 ­– was less frequently encountered than H. gracilis and recorded only on (Polygonaceae) in montane habitats up to 2000 m altitude, and on (Vitaceae). Of the specialist species (those restricted globally to one or two host families; Tab. IV), Cephonodes apus was found only on two species of Rubiaceae (Gentianales), as were also the two species of Macroglossum, M. aesalon and M. milvus. The latter species was additionally found on Loganiaceae (also Gentianales). Rubiaceae have also been cited as hostplants of Basiothia medea in South Africa (Pinhey, 1975), but on La Réunion the species has only been recorded from a single adult captured in the montane rain forest of Bébour (MHN, Saint-Denis, La Réunion) and its hostplant remains unknown. B. medea may be migratory from Mauritius or Madagascar. The larva of Coelonia solani described by Attié & Morel (1997) feeding on Cleroden- drum (Verbenaceae) is the only record from La Réunion, and the hostplants from Madagas- car given by Griveaud (1959) refer to C. fulvinotata. Hyles biguttata and Nephele densoi were observed very infrequently. H. biguttata was only found as eggs on a species of Eri- caceae, and just a single larva of Nephele densoi was found on Ficus reflexa (Moraceae) growing in relict semi-dry habitat of the island. On Madagascar, the latter species had previously been recorded from this host (as F. melleri; Paulian & Viette, 1956), and also on Banian (F. benghalensis; Guillermet & Guillermet, 1986) and Nerium (Apocynaceae; Desegaulx de Nolet, 1984). Both N. densoi and D. nerii seem to be specialized on host families with abundant latex.

Comparison of La Réunion endemics and widely distributed species For those species with very broad distributions, incorporating Africa and often significant parts of the rest of the Old World (A. atropos, A. convolvuli, D. nerii, H. celerio), we observed few associations with the endemic plants of La Réunion. A. atropos was observed primarily on exotic plants (82 % of the recorded associations, Tab. IV), but was also recorded on a number of endemic and indigenous plants. For example, the species was infrequently found on Nuxia verticillata (Stilbaceae), which is common in the native forests, and slightly more frequently in semi-dry habitats on the indigenous species of (O. lancea, O. europaea var. africana). Larvae have also been recorded on cultivated plants of Tarenna borbonica (Rubiaceae) and Clerodendrum heterophyllum (Verbenaceae), both of which are endemic to La Réunion and Mauritius. Agrius convolvuli was recorded on only one exotic Convolvulaceae, but the species prob- ably feeds on other species of Ipomoea. Daphnis nerii larvae are most often observed on exotic plants (66.7 % of associations). However, they were found on endemic plants, such as the threatened Tabernaemontana persicariaefolia (Apocynaceae), a small of relict semi-dry habitat that is sometimes entirely defoliated and the fruits also eaten (M. Attié, pers. obs.), and , a scarce tree that occurs in submontane forest. Daphnis nerii and A. atropos are the best-known species in the larval stage on the island. Larvae of both may be commonly observed on garden plants at low altitude throughout the year. The widely dis- tributed Hippotion celerio was recorded on cultivated Vitis (Vitaceae) and frequently on the indigenous Rumex abyssinicus (Polygonaceae), which is common in open habitats at middle and high altitudes. Coelonia fulvinotata and Hippotion gracilis (Mascarenes, Madagascar, Comoro Islands and African region) are essentially associated with exotic plants. Coelonia fulvi- notata was recorded only on the exotic invasive, Lantana camara (Verbenaceae), and the native hostplant of C. fulvinotata on La Réunion remains unknown, begging the question of what native plant did this moth feed prior to the introduction of Lantana by man? Nuxia verticillata, mentioned above as a hostplant of A. atropos on La Réunion, was also noted on Mauritius in association with larvae of C. fulvinotata, which Mamet & Williams (1993) confused with those of C. solani. This indigenous plant might be an original reunionese hostplant of C. fulvinotata before Lantana camara became naturalized on the island. How- ever, although Guillermet & Guillermet (1986) stated that C. fulvinotata was a common species in the south of the island, and larvae have been also recorded on L. camara in Ivory

­– 11 ­– Coast (Vuattoux, 1978; Vuattoux et al., 1989), on La Réunion larvae were observed only after several years of searching in a particular biotope where L. camara was growing in the understorey of a montane secondary forest habitat or in the relict semi-dry forest. The abundance and the chemical attractiveness of Lantana certainly will play an important role in the feeding behavior of C. fulvinotata without excluding the possibility of other associa- tions. In addition, a larva was observed on Grande Comore (Ngazidja) feeding on Cananga odorata (Annonaceae) (M. Attié, pers. obs., 2004). This plant is very abundant and widely cultivated near the natural vegetation that is the habitat of C. fulvinotata on the Comoro Islands, where it may thus be another preferred hostplant. However, on La Réunion, C. ful- vinotata seems rather to be closely associated with forest habitat and thus will rarely come into contact with C. odorata, which occurs only as an ornamental plant in gardens and along recently cultivated roadsides in towns. Hippotion gracilis, which was listed by Guenée (1862), Vinson (1938), Viette (1996) and Guillermet (2006) as H. eson, is found throughout subsaharan Africa (where it is sympatric with H. eson), Seychelles, Comoro Islands, Mada- gascar, Mauritius and La Réunion (I.J. Kitching, unpubl. data). This species has 13 associa- tions with both exotic (53.8 %) and endemic and indigenous plants (respectively 15.4 and 30.8 %). Among these latter associations are three : annulata, endemic to La Réunion and Mauritius; the two indigenous plants, C. rotundifolia (Vitaceae), often found on cliffs, and Danais fragrans (Rubiaceae) in semi-dry forest; and Alocasia senderiana (Araceae) in wet secondary habitat. Commonly found in gardens, the two species of Hip- potion (H. celerio and H. gracilis) also occur on indigenous and endemic plants in relictual indigenous vegetation or in secondary vegetation. Four species with more regional distributions, Coelonia solani, Hyles biguttata, Nephele densoi and Macroglossum aesalon aesalon, also mostly feed on exotic plants, but with some records from indigenous and endemic plants. Coelonia solani was recorded in La Réunion only on Clerodendrum heterophyllum (Verbenaceae) in a semi-dry habitat (Attié & Morel, 1997). This family has only one endemic on La Réunion and Mauritius (C. heterophyllum) but is strongly represented by 24 exotic species (Flore des Mascareignes, 1976-2009). Hyles biguttata is associated with Agarista salicifolia, an indigenous Ericaceae (Anderes, 1989) that occurs on La Réunion, Mauritius, Madagascar, and throughout Africa (Flore des Mascareignes, 1976-2009). Observations of H. biguttata on this plant on La Réunion at 650-950 m altitude, and records on Madagascar above 1000 m (Griveaud, 1959; Desegaulx de Nolet, 1984), sug- gest this species is confined to submontane and montane habitats.Macroglossum aesalon aesa- lon was recorded in only one locality on the island, on the exotic Rubiaceae, . Nephele densoi was recorded on the indigenous Ficus reflexa but has also recorded on the exotics, F. bengahalensis (Moraceae) (Guillermet & Guillermet, 1986) and Nerium oleander (Apocynaceae) (Desegaulx de Nolet, 1984). The endemic species Cephonodes apus and Macroglossum milvus, adults of which are diurnally active, are closely associated with native forests and larvae have been recorded only on endemic and indigenous plants. Oviposition by C. apus was observed once in semi-dry for- est on the common Rubiaceae, Antirhea borbonica, and larvae were also found on a cultivated plant of the endemic coffee, Coffea mauritiana (Rubiaceae). Boisduval (1833) noted the rarity of this moth (cited as Cephonodes hylas but certainly referable to C. apus as C. hylas is absent from La Réunion according to Viette (1996)) and since then, its frequency does not seem to have increased. Adults of C. apus were very rarely observed during the searches. In contrast, Macroglossum milvus is a relatively common species in lowland rain forest, submontane semi-dry forest, and submontane and montane rain forests (Tab. II). Larvae were frequently observed on the indigenous plant, Danais fragrans, and on Chassalia corallioides (Rubiaceae) and Geniostoma borbonicum (Loganiaceae), two plants endemic to La Réunion that are relatively common in native forests. The endemic moths, Macroglossum milvus and Cephonodes apus, and the indigenous species, Hyles biguttata, are all strictly associated with native plants in native forests, or in or near relict forests, in contrast to the other macroglossine species, which are observed princi- pally in gardens, cultivated areas and secondary vegetation.

­– 12 ­– Comparison of host ranges on the Mascarenes and other regions Host range may reflect different feeding behaviors in a species according to region, with perhaps differences being manifest between islands and continental areas. The first two com- ponents derived from factorial correspondence analysis of the frequencies of plant genera, both by moth species and region, for the six selected hawkmoth species (A. atropos, A. convolvuli, C. fulvinotata, D. nerii, H. celerio and H. gracilis), are shown in Figure 1. The first component, which separates Hippotion celerio (Hc) widely from the other species, also strongly discrimi- nates Australia (AU) from the other regions (EA, IC, IO, SA), as well as the western Palaearc- tic region (WP), though to a lesser extent. These regions are indeed associated with particular hostplant profiles of Hippotion celerio. The second component widely separates Acherontia atropos (Aa) and Coelonia fulvinotata (Cf), and discriminates the western Palaearctic region and, though less clearly, the Mascarenes from the African regions. This is a consequence of a broad host range in Acherontia atropos combined with a rather smaller number of associa- tions in Coelonia fulvinotata in the Mascarenes and the absence of the latter species from the western Palaearctic region.

WP 0.6 Aa 0.4

MS 0.2 Comp2 -0.0 Ac Hc IC -0.2 Dn Hg SA EA AU -0.4

Cf -0.6

-2.0 -1.5 -1.0 -0.5 0.0 0.5 1.0

Comp1

Figure 1.— First factorial plan of the FCA of the 181 larval hostplants at plant genus frequency table for six selected polyphagous hawkmoth species (Aa: Acherontia atropos, Ac: Agrius convolvuli, Cf: Coelonia fulvinotata, Dn: Daphnis nerii, Hc: Hippotion celerio, Hg: H. gracilis) and by region (AU: Australia, EA: East Africa, IC: Ivory Coast, MS: Mascarenes, SA: South Africa, WP: western Palaearctic) (58.1 and 29.5 % of total inertia).

The results (Fig. 1, Tab. V) allow us to recognize two groups of species. First are those species in which the host ranges observed in the Mascarenes and other regions are diverse. On the Mascarenes, Acherontia atropos shows a greater range of hostplant genera than in East Africa, South Africa and Ivory Coast (Tab. V, Fig. 1). In contrast, its host range in the western Palaearctic region is much greater, with almost twice the number of families being repre- sented. These differences may be partly the result of climatic and floristic differences among the regions, but will also be affected by the different levels of recording effort applied in each region. In addition, the greater host range of A. atropos in the western Palaearctic region might reflect more generalist oviposition preferences in females migrating from tropical and sub- tropical to more temperate habitats. Hippotion gracilis shows a similar range to that observed in other countries, although it must be remembered that for the African records it is not yet possible to distinguish those hostplant records that refer to H. gracilis and those that refer to

­– 13 ­– H. eson (Tab. V). Further work is necessary to determine the precise hostplant spectra of these two recently distinguished species. Then there are the other species in which the host range on the Mascarenes is often smaller than in other regions where they occur, being restricted to one or a few families. On the Mascarenes, Agrius convolvuli, Coelonia fulvinotata, Daphnis nerii and Hippotion celerio have all been recorded on only 1-3 host families, which contrasts strongly with observations from the other studied regions in which they occur (Tab. V).

Co m par i s o n o f p l a n t f a m i l y u s e pat t e r n s in t h e Ma s c ar e n e s w i t h t h o s e f o u n d t h r o u g h o u t t h e r e s t o f t h e ra n g e o f s i x s e l e c t e d m o t h s p e c i e s Hostplant selection by female hawkmoths depends first and foremost on plant availability, itself a function of the distributional ranges and other biological and ecological attributes of the plants. But it also depends to some extent on the behavioural plasticity and adaptability of the moths, that is, their ability to utilize suboptimal or novel hostplants when their preferred hosts are unavailable. This combination of extrinsic and intrinsic factors can result in the develop- ment among the species of regional dietary preferences. A fuller analysis of host use by region in the six selected hawkmoth species, at the level of plant genus, is presented in Table V. Acherontia atropos and Coelonia fulvinotata were recorded respectively on 29 and 13 families of dicots (Tab. VI) with good representation of each on Solanaceae and Verbenaceae in all regions (Tab. V). Similarly, Bignoniaceae is an important family for these two moths, except in Ivory Coast for A. atropos, probably due to a lack of field observations, and in the western Palaearctic region and in the Mascarenes for C. fulvinotata (the species is absent from the former region). On the Mascarenes the diet of C. ful- vinotata is restricted on Boraginaceae, Solanaceae, Stilbaceae and Verbenaceae. In the western Palaearctic region, the paucity of records of A. atropos on Verbenaceae and Bignoniaceae is attributable to the poor representation of these plant families in this area, both of which have essentially tropical and subtropical distributions (Heywood, 1985). Indeed, the Flora Europaea Database (1998) lists only 11 species of Verbenaceae in Europe, of which only five are native, and no species of Bignoniaceae. All western Palaearctic records of A. atropos on Bignoniaceae have been from introduced garden plants, such as Catalpa and . Agrius convolvuli was recorded principally on Convolvulaceae in most regions but the small number of plant genera recorded and the absence of data on the frequency of records does not allow us to test the importance of the other host families, such as in Ivory Coast or Asteraceae in East Africa and the western Palaearctic region. Daphnis nerii was recorded principally on Apocynaceae but Rubiaceae is also well represented in all regions except the western Palaearctic (Tab. V). The Old World species, Hippotion celerio and H. gracilis show a diverse but essentially common range of both monocots and dicots (Tab. VI). Hippotion celerio could thus show some hostplant specialization according to region. This species was most often recorded on Polygo- naceae and Vitaceae, although the first family was unrecorded in Ivory Coast and South Africa and the second unrecorded in East Africa and Ivory Coast. In South Africa, most records were on Vitaceae, whereas in Australia, records were more diverse, with a predominance of Araceae, Vitaceae and Polygonaceae. In the Mascarenes, all records are from Polygonaceae, Rubiaceae and Vitaceae (Tab. V). No Fabaceae were recorded in the Mascarenes or western Palaearctic region, whereas associations between H. celerio and this family (Acacia in East and South Africa, Afzelia in Ivory Coast) seem to be typical of the African regions. In Ivory Coast, the rather unusual families Convolvulaceae and Nyctaginaceae have been also reported. Hippotion gracilis has host families in common with H. celerio and has been predominantly recorded on Araceae, Vitaceae and Rubiaceae in all regions studied and no regional dietary preferences are apparent on the Mascarenes (Tab. V). Barplots at the level of plant superorder show that each of the six selected species of hawkmoth has a diet strongly restricted to a particular higher group of plants (Fig. 2). Acher- ontia atropos, Agrius convolvuli, C. fulvinotata and D. nerii are essentially associated with Euasterids I (EA1), whereas H. celerio and H. gracilis show a more diversified diet, including Monocots, Asterids and Rosids. Hippotion celerio and H. gracilis are well recorded in different

­– 14 ­–

b of families of b N

7 8 6 4 4 1 2 1 3 8 3 8 3 11 22

b of genera of b N 4 8 2 3 1 5 8 5 aphnis nerii,

12 14 13 25 38 12 14

D aprifoliaceae C ipsacales D

------0 0 0 0

7.9

steraceae A sterales A - - - -

0 0 0 0 0 0 0 25 20

2.6

14.3

piaceae A piales A

------0 0 0 0

E uasterids II 5.3

olanaceae S oelonia fulvinotata, ------

8 40

8.3 C 12.5 14.3 16.7 14.3 30.8 15.8

onvolvulaceae C - - - - -

0 0 0 25 60

7.2 100 100 S olanales 37.5 66.7

erbenaceae V

0 0 0 0 0 75 20 24

2.6 12.5 16.7 21.4 16.7 35.7 15.4

tilbaceae S

- - - -

------

0 0 0 4 20

crophulariaceae S ------grius convolvuli,

A leaceae O ------

0 0 0 12

8.3 8.3 7.1 15.4 15.8

amiaceae E uasterids I L

0 0 0 0 0 0 0 0

8.3 7.1 7.7 2.6 37.5 14.3 16.7 Bignoniaceae ------

0 0

25 20

2.6 L amiales

12.5 33.3 14.3 23.1 entianales G ubiaceae R ------

0 0 0 0 0 4

cherontia atropos, 12.5 entianales G pocynaceae A

A

V ------

nplaced U Boraginaceae

------

0 0 0 0 8 0 0

20

7.2

ricales E sterids a b le A

- - - -

- - - - - 0 0 0 0 0

T 25

apindales S apindaceae S

------

apindales S utaceae R

------0 0 0 0

5.3

alvales M alvaceae M ------

0 0 0 0 0 E urosids II

33.3 osaceae R

------0 0 0

7.7 7.9

oraceae according to region H ippotion celerio and . gracilis according M

------

R osales

abales F abaceae F

E urosids I

------4 0 8 0 0

nagraceae

O ------

elastomataceae M ------0 0 0 0 0

25

M yrtales

R osids nplaced U itaceae V

------

aryophyllales C Polygonaceae

------

- - - 0 0 0 0 0 20

aryophyllales C yctaginaceae N C ore ------

E udicots

onocots M Basal lismatales A raceae A ------A frica A frica A frica O rders A frica A frica A frica R egions S ubclasses Palaearctic Palaearctic

Sphinginae axa T E ast I vory C oast S outh M ascarenes W E ast I vory C oast S outh A ustralia M ascarenes W E ast I vory C oast S outh M ascarenes Acherontia atropos Acherontia Agrius convolvuli Agrius C. fulvinotata C. Percentage Percentage of hostplant genera in principal plant families for the six species,

­– 15 ­–

b of families of b N

2 6 3 3 2 5 6 4 3 9 6 8 6 8 10

b of genera of b N 8 7 5 5 6 8 3 7 8 aphnis nerii,

12 16 12 19 10 14

D aprifoliaceae C ipsacales D

------

steraceae A sterales A

------

piaceae A piales A

------E uasterids II

olanaceae S oelonia fulvinotata, ------

C 16.7

onvolvulaceae C ------

0 0 0 0

5.3 S olanales 16.7

erbenaceae V

------

tilbaceae S

------

crophulariaceae S ------grius convolvuli,

16.7

A leaceae O ------

12.5

amiaceae E uasterids I L

------Bignoniaceae ------

5.3

L amiales

) entianales G ubiaceae R

0 0 0 0

20

8.3 5.3 7.1

cherontia atropos, 14.3 33.3 33.3 28.6 12.5 12.5 12.5 entianales G pocynaceae A A

------

75 80

5.3 nplaced U Boraginaceae 66.7 57.2 62.5 ( continued

------

V ricales E Balsaminaceae sterids A ------

20

5.3 7.1

14.3 12.5 12.5 apindales S apindaceae S ------a b le 0 0 0 0

14.3 12.5 T apindales S utaceae R

------

alvales M alvaceae M

------E urosids II

20 6.2

osaceae R

------

oraceae according to region H ippotion celerio and . gracilis according M ------

0 0 0 0

R osales

14.3 16.7 abales F abaceae F E urosids I ------

0 0 0 0

6.2 16.7 12.5

nagraceae O ------

10

14.3 16.7 14.3 12.5

elastomataceae M ------

20

M yrtales

R osids nplaced U itaceae V - - - - -

0 0

21 20 25

33.3 16.7 14.3 14.3 62.5 aryophyllales C Polygonaceae ------

0 0 20

8.3 15.8 33.3

aryophyllales C yctaginaceae N C ore ------

0 0 11 20 17 10

8.3 7.1 E udicots 12.5

onocots M Basal lismatales A raceae A - - - - - 0 0 0 0 21 10 25 28.6 35.7 12.5 A frica A frica A frica A frica A frica A frica Orders Regions Palaearctic Subclasses

Palaearctic

E ast I vory C oast S outh M ascarenes W Australia E ast I vory C oast S outh M ascarenes

axa T E ast I vory C oast S outh M ascarenes W Daphnis nerii Daphnis Hippotion celerio Hippotion H. gracilis H. Macroglossinae Percentage Percentage of hostplant genera in principal plant families for the six species,

­– 16 ­–

Table VI

b of families of b N Percentage of hostplant genera in principal plant families for Acherontia atropos, Coelonia fulvinotata, Agrius convolvuli, 2 6 3 3 2 5 6 4 3 9 6 8 6 8

10 Daphnis nerii, Hippotion celerio and H. gracilis, recorded in Ivory Coast, East Africa, South Africa, Australia, the Mascarenes

b of genera of b N and western Palaearctic region 8 7 5 5 6 8 3 7 8 aphnis nerii,

12 16 12 19 10 14

D aprifoliaceae C ipsacales D

------

steraceae A sterales A

------

piaceae A piales A E urosids I E uasterids II ------E uasterids II Basal Core

Subclasses Monocots Eudicots Rosids Euasterids I

olanaceae S oelonia fulvinotata, ------

C 16.7

onvolvulaceae C ------

0 0 0 0

5.3 S olanales A sterales 16.7

erbenaceae V

A lismatales D ioscoreales C aryophyllales C aryophyllales U nplaced M yrtales F abales U nplaced G entianales G entianales

------Orders Lamiales Solanales

tilbaceae S

------

crophulariaceae S ------grius convolvuli,

16.7

A leaceae O ------

- Nb of Nb of 12.5

Araceae Dioscoreaceae Nyctaginaceae Polygonaceae Vitaceae Onagraceae Fabaceae Boraginaceae Apocynaceae Rubiaceae Bignoniaceae Lamiaceae Oleaceae Verbenaceae Convolvulaceae Solanaceae Asteraceae amiaceae L Taxa genera families E uasterids I ------

- Acherontia atropos 0 0 0 0 0 0 4.2 2.8 1,4 2.8 11.1 5.6 8.3 12.5 1.4 8.3 4.2 72 29 Bignoniaceae Coelonia fulvinotata 0 0 0 0 0 0 0 7.1 0 0 25 10.7 3.6 21.4 3.6 7.1 3.6 28 13 ------

5.3

L amiales

) entianales G ubiaceae R Agrius convolvuli 0 0 0 5.9 0 0 0 0 0 5.9 0 17.6 0 5.9 35.3 0 11.8 17 9

0 0 0 0

20

8.3 5.3 7.1

cherontia atropos, 14.3 33.3 33.3 28.6 12.5 12.5 12.5 entianales G pocynaceae A Daphnis nerii 0 0 0 0 3 0 3 0 60.6 18.2 0 0 0 0 0 0 0 33 9 A

------

75 80

5.3 nplaced U Boraginaceae 66.7 57.2 62.5 Hippotion celerio 10.3 0 7.7 7.7 17.9 5.1 5.1 0 2.6 7.7 2.6 0 2.6 0 2.6 2.6 0 39 21 ( continued

------

V ricales E Balsaminaceae sterids A Hippotion gracilis 25 7.1 7.1 0 14.3 10.7 0 0 0 17.9 0 0 0 0 0 0 0 28 11 ------

20

5.3 7.1

14.3 12.5 12.5 apindales S apindaceae S ------a b le 0 0 0 0

14.3 12.5 T apindales S utaceae R

------

alvales M alvaceae M

------E urosids II

20

6.2 osaceae R regions on Asterids (EA1) and Rosids (ROS), with additional associations of H. celerio with

------

- Core Eudicots (CED) and H. gracilis with Basal Monocots (BMO).

oraceae according to region H ippotion celerio and . gracilis according M ------

0 0 0 0

R osales

14.3 16.7 abales F abaceae F Extended analysis at the level of moth genus E urosids I ------0 0 0 0 6.2 16.7

12.5 Finally, we extend the analysis to include all species of those hawkmoth genera occur- nagraceae O ------

10

16.7 14.3 14.3 12.5 ring on La Réunion in a preliminary investigation of taxonomic conservatism of their larval

elastomataceae M hostplants on a global scale. The results of the FCA analysis at the level of plant family of ------

20

M yrtales

R osids the 41 species of hawkmoths belonging to the 11 genera present on La Réunion (Tab. I), for nplaced U itaceae V - - - -

- which there are at least three host genus records, are shown in Figure 3. The cluster dendro- 0 0

21 20 25

33.3 16.7 14.3 14.3 62.5 aryophyllales Polygonaceae C gram (Fig. 4) and the FCA allows the discrimination of five groups: A) Acherontia, Coelonia ------0 0

20 8.3

15.8

33.3 and Agrius; B) Basiothia, Cephonodes, Macroglossum and some species of Hippotion (SH29,

aryophyllales yctaginaceae N C SH39, SH40), one species of Hyles (SHY45) and one species of Daphnis (SD20), all charac- C ore ------0 0 11 20 17 10

8.3 7.1 E udicots

12.5 terized by affinities with the Rubiaceae; C) Daphnis and Nephele species, mainly associated

onocots M lismatales A raceae A Basal Basal with Apocynaceae; D) the remaining Hippotion species and Euchloron megaera (SE26); and - - - - - 0 0 0 0 21 10 25 28.6 35.7 12.5 E) the remaining Hyles species. However, it should be noted that for most moth genera only a proportion of the included species was available for analysis (Tab. VII) and so the results may change when the hostplants of these further species are discovered. A frica A frica A frica A frica A frica

A frica Kitching (2003) demonstrated that Acherontia and Coelonia are sister taxa and that Agrius Orders Regions Palaearctic Subclasses

Palaearctic (plus the East African genus, Callosphingia) was sister to these two within a monophyletic E ast I vory C oast S outh M ascarenes W Australia E ast I vory C oast S outh M ascarenes

axa T E ast I vory C oast S outh M ascarenes W Daphnis nerii Daphnis Hippotion celerio Hippotion H. gracilis H.

Macroglossinae . Acherontia and Coelonia are principally associated with Bignoniaceae, Verben-

Percentage Percentage of hostplant genera in principal plant families for the six species, aceae and Lamiaceae, in the order Lamiales (Tab. VIII). In general, host use at the plant family

­– 17 ­– Acherontia atropos Agrius convolvuli Coelonia fulvinotata

Western Palaearctic South Africa Indian Ocean Ivory Coast East Africa Australia

Daphnis nerii Hippotion celerio Hippotion gracilis

Figure 2.— Distributions of larval hostplant genera by superorder by region (Labels. AST: Asterids; BED: Basal Eudicots; BMO: Basal Monocots; CED: Core Eudicots; EA1: Euasterids I; EA2: Euasterids II; ER1: Erosids I; ER2: Erosids II; ROS: Rosids).

SE26

1 SH33 SH35 SHY53 SHY54 SHY5 6 SH31 SH29 SH39 SM99 SHY5 7 SH40 SM75 SC14SC15

0 SM89 SHY45 SM85 SAG4 SB9 SAG5 SC13 SM78 SA1 SA2 B SCO18 SD20 SA3

Comp2 -1 A

SD22

SN106

-2 SD23 C SN107

SN116

-1.0 -0.5 0.00.5 1.0 Figure 3a.— Column coordinates in the first factorial plan of the FCA of the hostplant genus frequency table for 41 selected species of hawkmoths (for codes see Table III) and for not too rare hostplant family (23.5 and 21.2 % of total inertia).

­– 18 ­– 0

1. SHY57 E SHY56 SHY54

SHY53 .5

SM99 SM85 SM89 0 0 SC15 0. SA1 SD20 SM78 SC14 SN116 SN107 SN106 SH40 SHY45 SD22 SAG4 SC13 SM84 SA3 SA2 Comp4 SD23 SH29 SB9 SAG5 SH39 -0.5 SCO18 SH31

-1.0 SH35

SH33

D -1.5

SE26

-1.5 -1.0 -0.5 0.00.5 1.0

Comp3

Figure 3b.— Column coordinates in the second factorial plan (16.9 and 10.8 % of total inertia) (for codes see Tab. III). 0

1. Ar c Nyc Vit Pol Lil Ona Ar a .5 Am a Ros

Cap Rub 0 0 As t Scr 0.

Con Sol Ole Big

Axis2 Mal -0.5 Lam Fab Ac a Ver

Bor -1.0 -1.5 -2.0

Apo

-1.0 -0.5 0.00.5 1.0

Axis1

Figure 3c.— Row coordinates in the first factorial plan (Aca: , Ama: Amaranthaceae, Apo: Apocynaceae, Arc: Araceae, Ara: Araliaceae, Ast: Asteraceae, Big: Bignoniaceae, Bor: Boraginaceae, Cap: Caprifoliaceae, Con: Convolvulaceae, Fab: Fabaceae, Lam: Lamiaceae, Lil: Liliaceae, Mal: , Nyc: Nyctaginaceae, Ole: Oleaceae, Ona: Onagraceae, Pol: Polygonaceae, Ros: , Rub: Rubiaceae, Scr: Scrophulariaceae, Sol: Solanaceae, Ver: Verbenaceae, Vit: Vitaceae). ­– 19 ­– 0 2.

Lil 1.5

Ros

1.0 Ama

Ona Cap .5 4 Pol Ast Axis Mal

0 0 Ole 0. Nyc Rub Scr Fab Apo Sol Ver Aca Bor Lam -0.5 Con Big

-1.0 Ara Vit

-1.5 Ar c

-1.0 -0.5 0.00.5

Axis3

Figure 3d.— Row coordinates in the second factorial plan (for abbreviations see Fig. 3c).

Figure 4.— UPGMA cluster dendrogram of hawkmoth species according to the plant genera recorded in the areas of their distribution.

­– 20 ­– Table VII Global number of species in the 12 hawkmoth genera occurring on La Réunion and the percentage studied

Genera No. species Nb of studied % species Acherontia 3 3 100 Agrius 6 3 50 Basiothia 5 4 80 Cephonodes 17 7 41 Coelonia 3 2 67 Daphnis 10 6 60 Euchloron 1 1 100 Hippotion 37 15 41 Hyles 29 27 93 Macroglossum 84 37 44 Nephele 22 14 64

level in Coelonia and Acherontia is very similar; among the major Acherontia host families, only Solanaceae and Fabaceae were rarely or not recorded respectively for Coelonia. Agrius is associated primarily with Convolvulaceae (Solanales), although Lamiaceae (Lamiales) is also a major hostplant family in Ivory Coast (Vuattoux et al., 1989). Nevertheless, at the level of plant order, Agrius conforms to the pattern observed in Acherontia and Coelonia (Figs. 3 & 4; group A). Among the Macroglossinae, Basiothia, Cephonodes and Macroglossum are essentially Rubiaceae feeders, with over 60 % of all records being from this family (group B) (Tab. VIII, Figs. 3 & 4). Daphnis (six species studied of 10 known) is also associated with Rubiaceae (as Daphnis hypothous (SD20)) (group B), but the most frequently recorded host family is Apocy- naceae (group C). Both Rubiaceae and Apocynaceae are members of the order Gentianales. Lamiales, Solanales and Gentianales are all closely related and currently classified in an unre- solved trichotomy within Euasterids I. Moreover, the clade formed by these orders comprises the majority of Euasterids I (APG, 2003). Nephele shows a similar pattern to that observed in Daphnis, but with additional host elements (group C). The fourteen (of 22) species for which hostplant records are available can be divided into three groups (Tab. IX). The first includes N. oenopion and N. rosae, restricted to Rubiaceae. The second group comprises species restricted to Apocynaceae: N. aequivalens, N. argentifera, N. bipartita, N. comma, N. funebris, N. peneus and N. vau. Nephele aequiva- lens has also been recorded on Fabaceae and Sapotaceae (MacNulty, 1970) but both records are of larvae reared in captivity and may not represent natural hosts. Likewise, the record of N. comma on Cassia (Fabaceae) by Grei (1990) may be a typographical error for Car- issa (Apocynaceae). The third group consists of two species, N. accentifera and N. densoi, in which the Apocynaceae association is supplemented by records on Moraceae and Rubiaceae. Moraceae belongs to the order Rosales in subclass Eurosids I, and thus is distantly related to the orders comprising Euasterids I. However, like Apocynaceae, Moraceae have sticky latex and the moth-plant association here may be one determined not by phylogeny but rather by plant chemistry. A similar pattern is shown by the New World hawkmoth genera Isognathus and Erinnyis (Macroglossinae, ), both of which are also associated with Apocy- naceae but in which many species are most frequently recorded on another lactiferous Eurosids I family, Euphorbiaceae (see also below under Hyles). The species of Hyles analysed in the FCA present a diet different from that of the other hawkmoth genera (Fig. 3b; group E), although Hyles calida shows affinities with Rubiaceae

­– 21 ­–

and 7 1 4 36 23 13 14 17 36 46 15 M acroglossum Nb of families H yles, 4 51 13 49 33 64 43 25 151

100 123

Nb of genera

II uasterids E sterales A

H ippotion, Asteraceae 6 0 0 3 0 0 2 0 0

7.8 4.9 Solanaceae

0 0 0 2 0 0

7.9 7.8 6.1 1.6 2.4 Convolvulaceae E uchloron,

0 0 1 0 0

2.6 7.7 6.1 1.6 S olanales 0.8 17.6 Verbenaceae 0 0 0 0 7.9 7.8

7.7 4.1 1.6 2.3 18.2

D aphnis, Stilbaceae

0 0 0 0 0 0 0 0 0

0.7 6.1 Scrophulariaceae

2 2 0 3 0 0 3 0 0

7.7 1.6 Oleaceae C ephonodes,

6 0 0 2 3 0 2 0 0

L amilaes 4.7 2.4 Lamiaceae E uasterids I 0 0 0 0 0 0 0 0

5.3 5.9 9.1

C oelonia, Bignoniaceae VIII

2 0 2 0 0 2 0 0 0

9.9

27.3 entianales G Rubiaceae 0 0 a b le

25 16 32 2.6 3.9 8.1

53.8 69.4 65.1 T entianales G Apocynaceae

0 0 2 0 1 56

1.3 0.0 0.8 2.3

39.1

nplaced A grius, Basiothia, U Boraginaceae

0 0 0 0 0 0 0 0

2.6 6.1 1.6

sterids A ricales E Balsaminaceae

0 2 0 0 0 0 1 0

7.7 0.8 2.3

osales R recorded in different regions of the world regions in different N ephele recorded Rosaceae A cherontia,

2 0 0 0 0 0 0 0

2.6 5.7 2.3

abales F Fabaceae

0 0 0 3 0 8

4.1 6.3 5.7 E urosids I

10.6 13.7 yrtales M Onagraceae

0 0 0 0 0 0 4 0

1.6 6.5 2.3

nplaced U Vitaceae

R osids 0 2 0 0 3 0 0

10 1.6 1.6 100

udicots E ore C aryophyllales C Nyctaginaceae

0 2 0 0 0 0 0 5 0 0

3.3

iliales L Liliaceae

0 0 0 0 0 0 0 0 0 0

4.9

lismatales A Araceae Basal 0 0 0 0 0 0 0 0 14 3.9 7.7 M onocots Orders Genera Subclasses Acherontia Agrius Basiothia Cephonodes Coelonia Daphnis Euchloron Hippotion Hyles Macroglossum Nephele Percentage of hostplant genera Percentage in principal plant families for

­– 22 ­– Table IX Percentage of hostplant genera in plant families for 14 species of Nephele

Subclasses C ore E udicots Eurosids I Euasterids I

Orders C aryophyllales F abales R osales G entianales G entianales

Nb of Nb of

Taxa N yctaginaceae F abaceae M oraceae A pocynaceae R ubiaceae genera families

Nephele accentifera 0 0 11.1 55.6 33.3 9 3 Nephele aequivalens 0 20 0 80 0 5 2 Nephele argentifera 0 0 0 100 0 2 1 Nehele bipartita 50 0 0 50 0 2 2 Nephele comma 0 20 0 80 0 5 2 Nephele densoi 0 0 33.3 66.7 0 3 2 Nephele discifera 0 0 0 100 0 1 1 Nephele funebris 50 0 0 50 0 2 2 Nephele hespera 0 0 0 100 0 1 1 Nephele oenopion 0 0 0 0 100 3 1 Nephele peneus 0 20 0 80 0 5 2 Nephele rosae 0 0 0 0 100 7 1 Nephele subvaria 0 0 0 100 0 1 1 Nephele vau 0 0 0 100 0 1 1

(group B). An extended analysis (Tab. X) of 27 species of Hyles (29 species known) shows that they can be separated into four groups (Fig. 5). The first group consists of a small number of species associated with Asterids, especially Euasterids I, although records are few. Hyles apocyni is restricted to Apocynum (Apocynaceae) in SW Tajikistan. Hyles biguttata (the puta- tive sister group of H. livornicoides from Australia; Hundsdoerfer et al., 2009) was recorded on Gentianaceae (Gentianales) and Acanthaceae (Lamiales), both in Euasterids I, and also Ericaceae, which are placed in the Basal Asterids. The second group comprises those species restricted to Rosids, especially Eurosids I: H. chuvilini, H. costata, H. cretica, H. nervosa, H. robertsi and H. sammuti on Euphorbiaceae; H. dahlii also on Onagraceae; and H. tithy- mali also on Vitaceae. The third group comprises those species with Asterid and Rosid asso- ciations. Among this group, one sub-group centred on the species preferentially associated with Rosids: H. euphorbiae, H. euphorbiarum, H. gallii, H. hippophaes (the only species recorded on Thymelaeaceae (Thy); Tab. X), H. lineata (with 30 host families), H. livornica (with 29 host families), H. livornicoides, H. nicaea (virtually restricted to the plant genus

­– 23 ­– 6 2 1 1 1 1 1 5 5 3 1 6 3 15 N b of families 9 2 1 1 1 1 2 9 6 3 1 8 4 24

N b of genera

II uasterids E oodeniaceae G

0 0 0 0 0 0 0 0 0 0 0 0 0

11.1 II uasterids E steraceae A

0 0 0 0 0 0 0 0 0 0 0 0 0 0

A sterales

olanales S olanaceae S

0 0 0 0 0 0 0 0 0 0 0 0

16.7 12.5 crophulariaceae S

0 0 0 0 0 0 0 0 0 0 0 0 0 0

leaceae O

0 0 0 0 0 0 0 0 0 0 0 0 0

4.2

canthaceae A L amiales

0 0 0 0 0 0 0 0 0 0 0 0 0

33.3 ubiaceae R

0 0 0 0 0 0 0 0 0 E uasterids I 0 0

55.6 12.5 entianaceae G

0 0 0 0 0 0 0 0 0 0 0 0 0

33.3 pocynaceae A

0 0 G entianales 0 0 0 0 0 0 0 0 0

100 100 16.7 riclaes E sterids A ricaceae E

0 0 0 0 0 0 0 0 0 0 0 0 0

33.3 apindales S utaceae R

0 0 0 0 0 0 0 0 0 0 0 0 0

11.1 alvales M hymelaeaceae II T

0 0 0 0 0 0 0 0 0 0 0 0 0

E urosids 16.7 osaceae R

0 0 0 0 0 0 0 0 0 0 0 0 0

12.5 laeagnaceae E

0 0 0 0 0 0 0 0 0 0 0 0 0

R osales

33.3 alpighiales M uphorbiaceae a b le X E T

0 0 0 0 0 0 0

50

8.3

100 100 100 22.2 12.5 agales F agaceae F 0 0 0 0 0 0 0 0 0 0 0 0

4.2

11.1 E urosids I abales F abaceae F

0 0 0 0 0 0 0 0 0 0 0 0

11.1 11.1 nplaced U Zygophyllaceae

0 0 0 0 0 0 0 0 0 0 0 0 0 0

nagraceae O

0 0 0 0 0 0 0 0 0

50

22.2 16.7 20.8 37.5 yrtaceae M

0 0 0 0 0 0 0 0 0 0 0 0 0

M yrtales

11.1 nplaced U itaceae R osids V

0 0 0 0 0 0 0 0 0 0 0

25

4.2

11.1 yctaginaceae Percentage of hostplant genera in principal plant families for 27 species H yles Percentage N

0 0 0 0 0 0 0 0 0 0 0 0 50

12.5

0 0 0 0 0 0 0 0 0 0 0 0

4.2 12.5 Polygonaceae

0 0 0 0 0 0 0 0 0 0

25

4.2

12.5 22.2 maranthaceae A

C ore E udicots C aryophyllales

0 0 0 0 0 0 0 0 0 0 0 0 0 0

anthorrhoeaceae X sparagales A

0 0 0 0 0 0 0 0 0 0 0 0 0 0

iliaceae L iliales L Basal 0 0 0 0 0 0 0 0 0 0 0 0 0 100 M onocots axa T O rders S ubclasses Hyles euphorbiarum Hyles euphorbiae Hyles dahlii Hyles cretica Hyles costata Hyles chuvilini Hyles chamyla Hyles centralasiae Hyles calida Hyles hippophaes Hyles biguttata Hyles gallii Hyles apocyni Hyles annei

­– 24 ­– 2 2 2 1 1 2 2 1 5 2 5 29 30 N b of families 3 3 3 1 1 3 2 1 5 2 6 59 57

N b of genera

II uasterids E oodeniaceae G

0 0 0 0 0 0 0 0 0 0 0 0 0

II uasterids E steraceae A

0 0 0 0 0 0 0 0 0

6.8 5.3 A sterales

olanales S olanaceae S

0 0 0 0 0 0 0 0 0 0 0

1.7 3.5

crophulariaceae S

0 0 0 0 0 0 0 0 0 0

50 3.4 1.8

leaceae O

0 0 0 0 0 0 0 0 0 0

20 3.4 1.8

canthaceae A

L amiales

0 0 0 0 0 0 0 0 0 0 0 0 0

ubiaceae R H yles

0 0 0 0 0 0 0 0 E uasterids I

6.8 3.5

33.3 66.7 33.3 entianaceae G

0 0 0 0 0 0 0 0 0 0 0 0 0

pocynaceae A

0 0 0 G entianales 0 0 0 0 0 0 0 0 0 0

riclaes E sterids A ricaceae E

0 0 0 0 0 0 0 0 0 0 0

1.7 1.8

apindales S utaceae R

0 0 0 0 0 0 0 0 0 0 0 0

16.7 alvales M hymelaeaceae II T )

0 0 0 0 0 0 0 0 0 0 0 0 0

E urosids osaceae R

0 0 0 0 0 0 0 0 0 0 0

5.1 8.8

laeagnaceae E

0 0 0 0 0 0 0 0 0 0 0 0

R osales 1.7

alpighiales M uphorbiaceae E

0 0 0 0

50

1.7 1.8

100 100 100

66.7 33.3 16.7 agales F agaceae a b le X ( continued F T 0 0 0 0 0 0 0 0 0 0 0 0 0

E urosids I abales F abaceae F

0 0 0 0 0 0 0 0 0 0

20

6.8

16.7 nplaced U Zygophyllaceae

0 0 0 0 0 0 0 0 0

20 50 3.4 1.8

nagraceae O

0 0 0 0 0 0 0 0 0 0

5.1

66.7 10.5 yrtaceae M

0 0 0 0 0 0 0 0 0 0 0

1.7 M yrtales

16.7 nplaced U itaceae R osids V

0 0 0 0 0 0 0 0 0

20

3.4 3.5

33.3 yctaginaceae Percentage of hostplant genera in principal plant families for 27 species Percentage N

0 0 0 0 0 0 0 0 0 0 7

20 1.7 Portulacaceae

0 0 0 0 0 0 0 0 0 0 0

1.7 3.5 Polygonaceae

0 0 0 0 0 0 0 0 0 0 0 7

10.2 maranthaceae A C ore E udicots C aryophyllales

0 0 0 0 0 0 0 0 0 0 0 7

1.7

anthorrhoeaceae X sparagales A

0 0 0 0 0 0 0 0 0 0 0 0

33.3 iliaceae L iliales L Basal 0 0 0 0 0 0 0 0 0 50 8.5 3.5 M onocots 66.7 axa T O rders S ubclasses Hyles wilsoni Hyles vespertilio Hyles tithymali Hyles siehei Hyles sammuti Hyles robertsi Hyles perkinsi Hyles nicaea Hyles nervosa Hyles livornicoides Hyles zygophylli Hyles livornica Hyles lineata

­– 25 ­– Figure 5.— Barplots of Hyles and superorder associations.

Euphorbia (Euphorbiaceae) (Hundsdoerfer et al., 2005a, b) but also recorded on the Scro- phulariaceae genus, (Wilkinson, 1956) (thus 50 % with Eup (ER1) and 50 % with Scr (EAI)), H. vespertilio principally on Onagraceae, and H. wilsoni on different families of Rosids (Euphorbiaceae, Fabaceae, Myrtaceae and Rutaceae) and on Asterids (Rubiaceae with 33.2 % of records). A second sub-group includes species preferentially associated with Asterids: H. calida (SHY45) (group B) (Fig. 3a) and H. perkinsi preferentially associated with Rubiaceae. The fourth group includes species with Monocot associations. One sub- group includes species restricted to monocots: H. centralasiae and H. siehei are associated with Eremurus (Liliaceae, Liliales), the larvae of which eat the flowers and young fruits not the , and a second sub-group includes a species recorded on monocots and Rosids: H. zygophylli. Species of Hippotion (15 species studied on 37 known) are preferentially associated with five plant families that are classified in quite different orders and informal groups: Araceae (Alismatales, Basal Monocots), Rubiaceae (Gentianales, Euasterids I), Vitaceae (unplaced to order in Basal Rosids), Onagraceae (Myrtales, Basal Eurosids) and Nyctagi- naceae (Caryophyllales, Basal Core Eudicots) (Tab. XI, Fig. 6). However, it is important to note that, with the exception of H. celerio and H. gracilis, no species has been recorded on all these families. According to their host use, the species of Hippotion can be separated into three groups. The first comprises those apparently restricted to one host family, each from different subclasses: H. aurora on Nyctaginaceae (Core Eudicots), H. rafflesii on Balsaminaceae (Asterids), H. rosae and H. roseipennis on Vitaceae (Rosids) and H. geryon on Araceae (Basal monocots) with one record also from Cissus (Vitaceae). A second group of species comprises Asterids/Rosids associations: H. rosetta (SH39) and H. scrofa (SH40) preferentially associated with Asterids and particularly Rubiaceae (group B) (Fig. 3a). The third group contains polyphagous species recorded from Monocot/Asterid/Rosid associa- tions: with species preferentially associated with Asterids, such as H. boerhaviae (group B) and H. brennus on Rubiaceae, and H. echeclus on Pedaliaceae and Rubiaceae; species preferentially associated on Monocots and Asterids, such as H. balsaminae, H. velox and

­– 26 ­– Table XI Percentage of hostplant genera in principal plant families for 15 species of Hippotion

Core Eurosids Eurosids

Subclasses Basal M onocots Basal M onocots C ommelinids Eudicots Rosids I II Asterids Euasterids I E uasterids II

Orders A lismatales D ioscoreales C ommelinales C aryophyllales C aryophyllales D illeniales U nplaced M yrtales C ucurbitales F abales Brassicales S apindales Ericales G entianales Lamiales S olanales A sterales

itaceae Nb of Nb of

Taxa A raceae D ioscoreaceae Pontederiaceae N yctaginaceae Polygonaceae D illeniaceae V O nagraceae C ucurbitaceae F abaceae Brassicaceae R utaceae Balsaminaceae T heaceae R ubiaceae O leaceae Pedaliaceae S crophulariaceae C onvolvulaceae A steraceae genera families Hippotion aurora 0 0 0 100 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 Hippotion 40 0 0 0 0 0 0 20 0 0 0 0 20 0 0 0 0 0 20 0 5 4 balsaminae Hippotion 5.6 5.6 0 11.1 0 0 5.6 0 5.6 5.6 0 0 5.6 5.6 33.3 0 0 0 5.6 0 18 12 boerhaviae Hippotion brennus 25 0 0 0 0 25 0 0 0 0 0 0 0 0 50 0 0 0 0 0 4 3 Hippotion celerio 16.1 1.6 0 4.8 6.5 1.6 12.9 4.8 0 0 0 0 1.6 0 12.9 1.6 0 4.8 3.2 0 62 24 Hippotion echeclus 0 0 20 20 0 0 0 20 0 0 0 0 0 0 20 0 20 0 0 0 5 5 Hippotion gracilis 25.7 5.7 0 5.7 0 5.7 14.3 8.6 0 0 0 0 2.9 0 14.3 0 0 0 2.9 0 35 14 Hippotion geryon 100 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 Hippotion osiris 22.7 0 0 0 13.6 0 22.7 9.1 0 0 0 0 4.5 0 13.6 0 0 0 4.5 0 22 9 Hippotion rafflesii 0 0 0 0 0 0 0 0 0 0 0 0 100 0 0 0 0 0 0 0 1 1 Hippotion rosae 0 0 0 0 0 0 100 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 Hippotion 0 0 0 0 0 0 100 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 roseipennis Hippotion rosetta 0 0 0 12.5 0 0 12.5 0 0 0 0 0 0 0 62.5 0 0 0 12.5 0 8 4 Hippotion scrofa 0 0 0 0 5.3 0 15.810.5 0 0 0 5.3 5.3 0 31.6 5.3 0 0 5.3 10.5 19 10 Hippotion velox 33.3 0 0 16.7 0 0 0 0 0 0 16.7 0 0 0 16.2 0 0 0 16.7 0 6 5

H. gracilis (group D) (Fig. 3b), on Araceae (Monocots), Balsaminaceae (H. balsaminae/H. gracilis), Convolvulaceae (H. balsaminae/H. velox/H. gracilis) and Rubiaceae (H. velox/H. gracilis) (Asterids) (group D); species associated indifferently into Monocots, Asterids and Rosids, such as H. celerio (group D); and species preferentially associated with Rosids, such as H. osiris on Vitaceae and Onagraceae. Euchloron, with only a single species, E. megaera, appears to be linked exclusively to Vitaceae (Cissus, Parthenocissus (Pinhey, 1975), (Kroon, 1999) and Vitis (Vuat- toux et al., 1989)), unplaced within the Rosids.

­– 27 ­– Figure 6.— Barplots of Hippotion and superorder associations.

Discussion

Regional diet The prevalence on La Réunion of records from a reduced number of plant families seems to suggest a regionally distinct diet. However, it is surprising, for example, that A. atropos was not found on Bignoniaceae in Ivory Coast when both it and C. fulvinotata have been recorded on Millingtonia, and Tecoma in East Africa, and Bignonia in South Africa. The only records of Sphinginae on Bignoniaceae in Ivory Coast are for C. fulvinotata on Kigelia (Vuattoux et al., 1989), although other genera of this family, such as Markhamia, Newbouldia, Spathodea and Stereospermum, do occur (Mensbruge, 1966). There is a similar absence of H. celerio records in Ivory Coast on Araceae and Polygonaceae, whereas these associations are represented in other African regions, suggestive of a lack of field observations rather than a regionally distinct hostplant selection. In contrast, it is notable that the host range of Coelo- nia fulvinotata on Mauritius and the three African regions is larger than that on La Réunion, where it has only been recorded on the invasive Verbenaceae, Lantana. This species seems to prefer the abundant L. camara in secondary forest and hostplant selection appears to change with habitat quality. Indeed, on the Mascarenes, the question remains: why has C. fulvinotata been recorded on Mauritius principally in open and cultivated areas but on La Réunion only in secondary forest?

Importance of host use and dispersal abilities of species There is certainly a relationship between the dispersal abilities of hawkmoths (and thus their capacity to colonize new areas) and evolution of their host use. Beck & Kitching (2007) found that larval diet breadth was the best predictor of range size and inter-island dispersion, and confirmed the importance of niche breadth on the geographical ranges of species. Invert- ing this relationship implies that the breadth of larval diet can be predicted to some extent from a knowledge of a species’ range, with more widespread species feeding on a broader range of plants. In the present study, we distinguished three patterns. The first, exemplified by

­– 28 ­– Acherontia atropos, Hyles lineata and H. livornica, includes species that have been reported on numerous host families (see Tabs. VI & X) and have very broad distributions. The second pat- tern, exemplified by Agrius convolvuli, is of a widely distributed moth that uses subsets of its total range of hostplant families in different parts of its distribution (although Convolvulaceae features in all regions and is the preferred hostplant family). These contrast with the third pat- tern, exemplified by many species of Hyles, which have both restricted distributions and are specialists. With regard to the endemic species of La Réunion Island, Macroglossum milvus and Cephonodes apus are clearly specialized and have a conservative diet. However, it is inter- esting to note that speciation in Macroglossum and Cephonodes has not been accompanied by a similar diversification in hostplants, most have maintained species of Rubiaceae or closely related plant families as hosts, whereas Acherontia or Agrius with few species (respectively 3 and 6) have a broad range of hostplants in different plant families (Tab. VIII). A low dispersal ability and adaptation to local biotopes, together with high hostplant availability, could explain the high levels of speciation within, and the broad distributions of, Macroglossum and Cepho- nodes. The diversity and the abundance of Rubiaceae and genetic constraints may explain why the species in these genera preserve a restricted link with Rubiaceae. In contrast, the ability of A. atropos and H. celerio, for example, to feed on a broad range of hostplant families may contribute significantly to their capacity for vagrancy, favoured by their morphology, which contributes to high dispersal abilities.

Taxonomic analysis versus phylogenetic indices It is now accepted that there are numerous potential pitfalls in simple comparisons of larval host plant range and diversity. Higher taxonomic categories have different limits in different taxonomic groups and thus are not comparable across different organisms. Thus, the phylogenetic relationships among the plants and among the insects should be taken into consideration to take account of the constraints imposed by the phylogenetic histories of both groups of organisms. Several phylogenetic indices have been proposed (Symons & Beccaloni, 1999; Beccaloni & Symons, 2000; Weiblen et al., 2006) but while they bring advantages in some areas, they also have their own constraints that restrict their applicability under other circumstances. So although we considered applying phylogenetic indices to our data, the task of compiling species-level phylogenies for all the plants involved across the six regions we studied was too great and in the absence of sufficiently complete data to make analyses mean- ingful, we refrained from doing so.

Conclusion

The different factors which play a role in the host range The differences observed in hostplant ranges between regions for a given species of hawk- moth are the result of a complex interaction of intrinsic and extrinsic factors. Hostplant use in Sphingidae is dependent upon environmental factors, such as climatic and floristic differences, as well as levels of interspecific competition, predation and parasitism, which act to constrain the species ecologically. However, the comprehensiveness of our understanding of host ranges is also significantly influenced by the level of applied recording effort. For example, the pau- city of records of Agrius convolvuli on La Réunion may be the result of a lack of observa- tions in cultivated areas (only native forests, natural secondary vegetation and gardens were surveyed in this study and there are few literature records from cultivated areas), whereas the larger host range in Ivory Coast, including Convolvulaceae and Lamiaceae, was the result, in part, of an intensive recording effort in cultivated areas. Genetic constraints also play a role in hostplant selection by Sphingidae, and the fre- quency of an association is undoubtedly correlated with the chemical attractiveness of the hostplants and the capacity of larvae to elude the plants’ defensive mechanisms. This study did not quantify the effects of hostplant chemistry on larvae and so it was not possible to determine

­– 29 ­– differential preferences among the recorded hostplants at the level of moth species. In addi- tion, hostplant attractiveness to insects is usually determined by the use of choice tests under laboratory conditions, but the degree, to which such results are applicable to natural conditions where such factors as the abundance and diversity of available plants will exert an influence on the oviposition behavior of females, is unclear. On La Réunion, we observed Coelonia fulvi- notata only on Lantana camara (Verbenaceae), an alien plant in the Mascarenes. Likewise, on the Comoro Islands, we recorded C. fulvinotata only on the widely cultivated plant, Cananga odorata (Annonaceae). In both cases, it is possible that female C. fulvinotata lay eggs on the most available plant. Dietary differences may have other origins. For example, the greater host range of A. atro- pos in the western Palaearctic region might reflect more generalist oviposition preferences in females migrating from tropical and subtropical to more temperate habitats. Subtle interactions between extrinsic and intrinsic factors may induce new constraints that lead to new responses from female moths. In the case of A. atropos, these interactions may result in females in tem- perate areas choosing to oviposit on atypical hostplant families when the usual tropical or subtropical hostplants are unavailable.

Taxonomic conservatism and chemical specialisation Our analyses show that hawkmoth species preserve their principal hostplant family pref- erences across regions, although regions may also have their own particular associations, such as Lamiaceae for C. fulvinotata in Ivory Coast. However, when a plant family is prominent in a region, we may ask whether the association really is a regional specificity or is simply due to a lack of records from other regions? For example, A. atropos was recorded on Bignoniaceae in East and South Africa but these records for this family are lacking in Ivory Coast despite that family being well represented and frequent in that country. A clear taxonomic conservatism was demonstrated for Macroglossum, Cephonodes and Basiothia species, which are preferentially associated with Rubiaceae, and for the species of Daphnis, which are preferentially associated with Apocynaceae and Rubiaceae. Similarly, Acherontia, Agrius and Coelonia are preferentially associated with plant families belonging to Euasterids I. Moth-plant associations may not be determined only by plant but also by plant chemistry as, for example, in the case of Nephele densoi, which is associated with Apocy- naceae (Euasterids) and Moraceae (Rosids), two taxa not closely related but lactiferous families, reflecting a similar pattern found in the New World hawkmoth genera,Isognathus and Erinnyis, which are associated with lactiferous Apocynaceae (Euasterids) and Euphorbiaceae (Rosids) (Robinson et al., 2001). Those species of Daphnis principally associated with Apocynaceae (Asterids) also utilize other lactiferous plant families both within and outside the Asterids. For example, D. nerii is principally associated with Apocynaceae and Rubiaceae (Asterids), but has been reported on Sapotaceae (Asterids) and Moraceae (Rosids). Such species appear to show chemical as well as taxonomic conservatism but the relative contributions of each, and their impact on the evolution of hostplant choice in Sphingidae, has yet to be determined.

Acknowledgements

We are grateful to G. Morel for his field assistance and suggestions, to R. Parnaudeau (Muséum d’Histoire Naturelle de Saint-Denis) for data localities, and to T. Monnier for A. atropos records from Bois-Court.

Note added in proof

Some additional hostplants were published by Martiré & Rochat (2008) but too late to be included in our study. The new records are as follows: Agrius convolvuli on Merremia umbellata (Convolvulaceae), Coelonia fulvinotata on floribunda (Oleaceae), Daphnis nerii on F. floribunda, Hyles biguttata on Agarista buxifolia (Ericaceae), and Nephele oenopion oenopion on Mussaenda arcuata (Rubiaceae). Furthermore, MA finally discovered eggs of Daphnis nerii on Thevetia peruviana on 09. VIII.2009 at La Bretagne. However, they proved to be parasitized by a species of Microhymenoptera and the palnt was devoid of larvae.

­– 30 ­– References

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­– 33 ­– Annex 1

Hostplants recorded on La Réunion for Acherontia atropos, Agrius convolvuli, Cephonodes apus, Coelonia fulvi- notata, C. solani, Daphnis nerii, Hippotion celerio, H. gracilis, Hyles biguttata, Macroglossum aesalon, M. milvus, Nephele densoi and N. oenopion oenopion with localities and references. Other hostplants are recorded for the first time.

Locality Taxa Plant family Hostplants Status (altitude m) References Sphinginae Acherontia atropos Verbenaceae Clerodendrum heterophyllum Endemic: La Chaudron (50) Attié (1999) (Poiret) R. Br. Réunion and Mauritius Verbenaceae Clerodendrum thomsonae Exotic Sainte-Clotilde (30) Attié (1999) Balf. Verbenaceae L. = Exotic Sainte-Clotilde D. repens L. Verbenaceae Lantana camara L. Exotic Cap-Noir (1200), Attié (1999); Colorado (650), Attié et al. (2005); Grand-Bassin (1000), Williams (1952) Sainte-Clotilde, Tam- [for Mauritius] pon (600), Bois-Court (1350), Mafate, all the island (waste lands) Verbenaceae volubilis L. Exotic Sainte-Clotilde Attié (1999) Verbenaceae Stachytarpheta urticifolia Exotic Colorado Attié (1999) (Salisb.) Verbenaceae Verbena officinalis L. Exotic Tampon Oleaceae Jasminum sp. Exotic Saint-Gilles les Hauts (400) Oleaceae Ligustrum ovalifolium Hassk. Exotic Bois-Court Attié (1999) Oleaceae Ligustrum robustum Blume Exotic Ravine des Cabris Attié et al. (2005) var. walkeri (Decaisne) Mansf. Oleaceae Olea europaea L. subsp. Indigenous Grand-Bassin Attié (1999) africana (Miller) Oleaceae Olea lancea Lam. Indigenous Grand-Bassin Attié (1999) Bignoniaceae Jacaranda mimosifolia D. Exotic Bois-Court Don Bignoniaceae jasminoides Exotic Chaudron (Lindl.) K. Schum. Bignoniaceae Spathodea campanulata Exotic Chaudron Attié (1999) Beauv. Bignoniaceae pallida (Lindley) Exotic Chaudron Attié (1999) Miers Bignoniaceae Tecoma stans (L.) Juss. ex Exotic Saint-Gilles (10) Attié (1999) Benth & Hook. Rubiaceae Tarenna borbonica (E. G. & Endemic: La Chaudron Attié (1999) A. Henderson) Réunion and Mauritius Stilbaceae Nuxia verticillata Comm. ex Endemic: La Bébour Attié (1999) Lam. Réunion and Mauritius Boraginaceae Cordia macrostachya (Jacq.) Exotic Sainte-Clotilde Williams (1948) Roem. & Schult. = C. curas- [Mauritius]; savica (Jacq.) Roem. & Attié (1999) Schult.

­– 34 ­– Boraginaceae Ehretia acuminata R. Br. Exotic Chaudron Vinson (1938) [Mauritius]; Attié (1999) Acanthaceae lupulina Lindl. Exotic Chaudron Annonaceae Cananga odorata (Lam.) Exotic Saint-Denis (Grand- Hook. f. & Thomson Canal) (150) Fabaceae Wisteria sinensis (Sims) Exotic Chaudron Attié (1999) Sweet. Cucurbitaceae Momordica charantia L. Exotic Saint-Paul, La Vinson (1938) Bretagne (250) [Mauritius] Solanaceae Physalis peruviana L. Exotic La Réunion Solanaceae Solanum auriculatum Ait. Exotic Bois Court Solanaceae Solanum melongena L. Exotic Saint-Pierre Mamet (1955) (Bassin-Martin) [Mauritius]; Attié (1999) Solanaceae Solanum tuberosum L. Exotic Guillermet & Guill- ermet (1986)

Agrius convolvuli Convolvulaceae Ipomoea purpurea (L.) Roth. Exotic Chaudron Coelonia fulvinotata Verbenaceae Lantana camara L. Invasive exotic Roche-Verre-Bouteille Attié (1999) (1150) Bois-Court Coelonia solani Verbenaceae Clerodendrum heterophyllum Endemic: La Ravine de la Chaloupe Attié & Morel (Poiret) R. Br. Réunion and (200) (1997) Mauritius Macroglossinae Cephonodes apus Rubiaceae Coffea mauritiana Lam. Endemic: La La Bretagne (300) Réunion and Mauritius Rubiaceae Antirhea borbonica Gmel. Endemic: La Grand-Bassin Attié (1999) Réunion and Mauritius Daphnis nerii Apocynaceae Nerium oleander L. Exotic Chaudron, La Desegaulx de Nolet Bretagne, Tampon (1984); Attié (1999) Apocynaceae Ochrosia borbonica J. F Endemic: La Tampon Gmel. Réunion and Mauritius Apocynaceae Pachypodium sp. Exotic Montagne (250) Apocynaceae Tabernaemontana divari- Exotic Saint-Gilles cata (L.) R. Br. Ex Roem. & Schult. = T. coronaria (Jacq.) Willd. = Ervatamia coronaria (Jacq.) Stapf. Apocynaceae Tabernaemontana persicari- Endemic: La Grande-Ravine (70), Attié (1999) aefolia Jacq. Réunion and Ravine de la Chaloupe, Mauritius Chaudron Ravine de la Grande- Chaloupe (20) Moraceae Ficus benjamina L. Exotic La Bretagne (250)

­– 35 ­– Hippotion celerio Polygonaceae Rumex abyssinicus Jacq. Indigenous Grand-Bassin, Cap Attié (1999) Noir, Col des Boeufs (2000) Vitaceae Vitis vinifera L. Exotic Sainte-Clotilde Attié (1999) Hippotion gracilis Araceae Alocasia senderiana W. Bull Indigenous Saint-François (700) Attié (1999) Araceae Anthurium andreanum Linden Exotic Bois de Nèfles (St- Attié (1999) Denis), Sainte-Clotilde Araceae Syngonium sp. Exotic Saint-Denis Araceae (L.) Exotic Cilaos Desegaulx de Nolet Spreng. (1984) Vitaceae Cissus annulata Descoings Endemic: La Ravine Bernica (50) Attié (1999) Réunion and Mauritius Vitaceae Cissus rotundifolia (Forssk) Indigenous Ravine Bernica Vahl. Vitaceae Vitis vinifera L. Exotic Sainte-Clotilde Vinson (1938) [Mauritius] Rubiaceae Danais fragrans (Comm. ex. Indigenous Grand-Bassin Attié (1999) Lam.) Pers. Loganiaceae Geniostoma borbonicum Endemic: La Vallée-Heureuse (796) Attié (1999) Spreng Réunion Nyctaginaceae Boerhaavia coccinea Miller Indigenous Sainte-Clotilde Onagraceae Ludwigia octovalvis (Jacq.) Exotic Forêt Dugain (800) Raven Balsaminaceae walleriana Hook. f. Exotic Saint-François Geraniaceae Pelargonium roseum Willd. Exotic Bois Court Hyles biguttata Ericaceae Agarista salicifolia (Comm. Indigenous Colorado, Plaine Anderes (1989) ex Lam) G. Don d’Affouches (950) Macroglossum aesalon Rubiaceae Paederia foetida L. Exotic Brûlé (Saint-Denis) Macroglossum milvus Rubiaceae Chassalia corallioides (Cor- Endemic: La Plaine des Fougères, Attié (1999) dem.) Verdc. Réunion Plaine d’Affouches (1000), Grand-Bassin, Roche-Verre-Bouteille, Takamaka (600-796), La Fenêtre (Salazie) (1400) Rubiaceae Chassalia gaertneroides Endemic: La Plaine des Fougères, Attié (1999) (Cordem.) Verdc. Réunion Bébour Rubiaceae Danais fragrans (Comm. ex. Indigenous Colorado, Grand-Bas- Desegaulx de Nolet Lam.) Pers. sin, Cap-Noir, Vallée- (1984); Attié (1999) Heureuse Loganiaceae Geniostoma borbonicum Endemic: La Haut du Brûlé (Saint- Attié (1999) Spreng Réunion Denis) (980), Vallée- Heureuse Nephele densoi Moraceae Ficus reflexa Thunb. Indigenous Ravine de la Grande- Paulian & Viette Chaloupe (30) (1956) [Madagas- car]; Attié (1999)

­– 36 ­–

H c H c H c H c H c H c H c H c H c Australia Coelonia fulvi - Coelonia C f:

A a H c Western Palaearctic Agrius convolvuli , Agrius

H g H g H g H g Ivory Coast A c: H c g

H c H c H g H g East Africa HOSTS ustraliafrom literature or website records, with subsequent A

H g H g , atropos Acherontia South H c g Africa

A a H g H g H g H g H g H g Mascarenes Mascarenes A nnex 2 oast,western Palaearctic region and C Genera Cananga Alocasia Amorphophallus Anchomanes Anthurium Colocasia Cryptocoryne Syngonium Typhonium Zantedeschia Dioscorea Tacca Hibbertia Tetracera Beta Bougainvillea Commicarpus Emex Rheum éunion for the six species studied ( A a: studied species six the for R éunion vory I L a frica, A ast E Families A nnonaceae A raceae A raceae A raceae A raceae A raceae A raceae A raceae A raceae A raceae D ioscoreaceae D ioscoreaceae D illeniaceae D illeniaceae A maranthaceae N yctaginaceae N yctaginaceae N yctaginaceae N yctaginaceae Polygonaceae Polygonaceae outhand S ascarenes new records from records new M ascarenes ascarenes, M Order M agnoliales A lismatales A lismatales A lismatales A lismatales A lismatales A lismatales A lismatales A lismatales A lismatales D ioscoreales D ioscoreales D illeniales D illeniales C aryophyllales C aryophyllales C aryophyllales C aryophyllales C aryophyllales C aryophyllales C aryophyllales and in the in and I J K Subclasses M agnoliids Basal M onocots Basal M onocots Basal M onocots Basal M onocots Basal M onocots Basal M onocots Basal M onocots Basal M onocots Basal M onocots Basal M onocots Basal M onocots C ore E udicots C ore E udicots C ore E udicots C ore E udicots C ore E udicots C ore E udicots C ore E udicots C ore E udicots C ore E udicots ummaryof hostplant genera in the Classe Basal D icots M onocots M onocots M onocots M onocots M onocots M onocots M onocots M onocots M onocots M onocots M onocots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots additions or deletions by deletions or additions S ). Classification follows APG (2003). notata , D n: Daphnis nerii H c: Hippotion celerio g: gracilis ). Classification follows

­– 37 ­–

H c H c H c H c H c Australia

H c H c H c A a A a A c H D n H c Western Palaearctic

H c H c H g H g H g D n D n Ivory Coast

H c A c H g H g East Africa HOSTS c

H H c H c H c H g South H c g H c g Africa

H c H g A a H g A a H g H g H c g Mascarenes Mascarenes Genera Rumex Pelargonium Dissotis Jusiacea Ludwigia Cayratia Cissus Clematicissus Cyphostemma Leea Parthenocissus Rhoicissus Vitis Euonymus Euonymus Manihot Rinorea Acacia Afzelia Cajanus Lonchocarpus Vicia Wisteria itaceae itaceae itaceae itaceae itaceae itaceae itaceae itaceae iolaceae Families Polygonaceae G eraniaceae M elastomataceae O nagraceae O nagraceae O nagraceae O nagraceae V V V V V V V V C elastraceae E uphorbiaceae V F abaceae F abaceae F abaceae F abaceae F abaceae F abaceae R osids R osids R osids R osids R osids R osids R osids R osids Order C aryophyllales G eraniales M yrtales M yrtales M yrtales M yrtales M yrtales U nplaced U nplaced U nplaced U nplaced U nplaced U nplaced U nplaced U nplaced C elastrales M alpighiales M alpighiales F abales F abales F abales F abales F abales F abales Subclasses C ore E udicots R osids R osids R osids R osids R osids R osids R osids R osids R osids R osids R osids R osids R osids R osids E urosids I E urosids I E urosids I E urosids I E urosids I E urosids I E urosids I E urosids I E urosids I Classe E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots

­– 38 ­–

A c H c Australia

A a A a A a A a A a H c A a A a A a A a A a D n Western Palaearctic

H c D n D n D n Ivory Coast

H c H g D n East H c g Africa HOSTS

A a D n South H c g Africa

A a D n H g H b Mascarenes Mascarenes Genera Cannabis Ficus Fragaria Malus Pyrus Urtica Begonia Momordica Zygophyllum Brassica Abutilon Sterculia Citrus Ruta Cardiospermum Paullinia Philadelphus Impatiens Agarista Strophanthus Adenium Families C annabaceae M oraceae R osaceae R osaceae R osaceae R osaceae U rticaceae Begoniaceae C ucurbitaceae Zygophyllaceae Brassicacae M alvaceae M alvaceae M alvaceae R utaceae R utaceae S apindaceae S apindaceae H ydrangeaceae Balsaminaceae E ricaceae A pocynaceae A pocynaceae A pocynaceae Order R osales R osales R osales R osales R osales R osales R osales C ucurbitales C ucurbitales U np. E urosids I Brassicales M alvales M alvales M alvales S apindales S apindales S apindales S apindales C ornales E ricales E ricales G entianales G entianales G entianales Subclasses E urosids I E urosids I E urosids I E urosids I E urosids I E urosids I E urosids I E urosids I E urosids I E urosids I E urosids II E urosids II E urosids II E urosids II E urosids II E urosids II E urosids II E urosids II A sterids A sterids A sterids E uasterids I E uasterids I E uasterids I Classe E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots

­– 39 ­–

H c Australia

D n D n A a D n Western Palaearctic

D n D n D n D n D n D n D n D n D n H g Ivory Coast

D n D n D n D n D n D n D n D n D n East Africa HOSTS

D n D n D n D n D n D n H g South Africa

D n D n D n D n H g Mascarenes Mascarenes Genera Amsonia Asclepias Carissa Catharanthus Funtumia Holarrhena Hunteria Nerium Ochrosia Pachypodium Picralima Plumeria Rauvolfia Tabernaemontana Thevetia Vinca Voacanga Geniostoma Borreria Burttdavya Coffea Coprosma Families A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae A pocynaceae L oganiaceae L oganiaceae R ubiaceae R ubiaceae R ubiaceae R ubiaceae R ubiaceae Order G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales Subclasses E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I Classe E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots

­– 40 ­–

H c Australia

H c A a A a Western Palaearctic

C f A a D n H g Ivory Coast A c D n

C f C f C f D n D n H g East A a C f Africa HOSTS

C f C f D n South Africa

H c A a A a H g D n A a A a D n Mascarenes Mascarenes Genera Danais Hallea Mitracarpus Paedaria Pentas Rubia Tarenna Acanthus Asystasia Barleria Bignonia Catalpa Fernandoa Jacaranda Kigelia Markhamia Millingtonia Newbouldia Pandorea Families R ubiaceae R ubiaceae R ubiaceae R ubiaceae R ubiaceae R ubiaceae R ubiaceae R ubiaceae R ubiaceae R ubiaceae R ubiaceae R ubiaceae A canthaceae A canthaceae A canthaceae Bignoniaceae Bignoniaceae Bignoniaceae Bignoniaceae Bignoniaceae Bignoniaceae Bignoniaceae Bignoniaceae Bignoniaceae Order G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales G entianales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales Subclasses E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I Classe E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots

­– 41 ­–

H c Australia

A a A a A a A a A a A a A a A a H c A a H c Western Palaearctic A c

A c A c A a A a Ivory Coast A a f C a

C f a C f C f A A a A a A a East A a C f A Africa HOSTS

C f C f A a A a A a A a A a C f A a C f South Africa

A a A a A a A a A a A a Mascarenes Mascarenes Genera Podranea Spathodea Tabebuia Tecoma Aeschynanthus Basilicum Hoslundia Ocimum Pogostemon Pycnostachys Salvia Solenostemon Teucrium Fraxinus Jasminum Ligustrum Olea Phillyrea Syringa Sesamum Veronica Veronica Buddleja Paulownia Scrophularia Families Bignoniaceae Bignoniaceae Bignoniaceae Bignoniaceae G esneriaceae L amiaceae L amiaceae L amiaceae L amiaceae L amiaceae L amiaceae L amiaceae L amiaceae O leaceae O leaceae O leaceae O leaceae O leaceae O leaceae Pedaliaceae Plantaginaceae S crophulariaceae S crophulariaceae S crophulariaceae Order L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales Subclasses E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I Classe E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots

­– 42 ­–

A c A c H Australia c c

H c A a A c A A A a A a A a A a A a A a Western Palaearctic f C A c

C f C f a A c A c A a H c Ivory A a C f A a C f Coast A a C f A a C f A A a A c H c a A

C f a A A a East A a C f A a C f A Africa HOSTS f C

C f c A c A a A a A a C f A a C f A South A a C f A a C f Africa

C f A a A a A a A a A a A c A a A a C f A a C f A a C f Mascarenes Mascarenes Genera Verbascum Nuxia Aloysia Clerodendrum Duranta Gmelina Lantana Petrea Stachytarpheta Tectona Verbena Vitex Calystegia Ipomoea Lepistemon Merremia Xenostegia Atropa Lycium Nicotiana Physalis Solanum erbenaceae erbenaceae erbenaceae erbenaceae erbenaceae erbenaceae erbenaceae erbenaceae erbenaceae erbenaceae Families S crophulariaceae S tilbaceae V V V V V V V V V V C onvolvulaceae C onvolvulaceae C onvolvulaceae C onvolvulaceae C onvolvulaceae C onvolvulaceae S olanaceae S olanaceae S olanaceae S olanaceae S olanaceae S olanaceae Order L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales L amiales S olanales S olanales S olanales S olanales S olanales S olanales S olanales S olanales S olanales S olanales S olanales S olanales Subclasses E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I E uasterids I Classe E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots

­– 43 ­–

Australia

A a A c A a A a A a A a A a H c Western Palaearctic

A a A a A a Ivory Coast

A c East Africa HOSTS

C f C f South Africa

A a A a C f Mascarenes Mascarenes Genera Cordia Ehretia Anethum Daucus Cussonia Acanthospermum Chrysanthemum Conyza Dahlia Helianthus Laggera Lonicera Sambucus Symphoricarpos Families Boraginaceae Boraginaceae A piaceae A piaceae A raliaceae A steraceae A steraceae A steraceae A steraceae A steraceae A steraceae C aprifoliaceae C aprifoliaceae C aprifoliaceae Order U np. E uasterids I U np. E uasterids I A piales A piales A piales A sterales A sterales A sterales A sterales A sterales A sterales D ipsacales D ipsacales D ipsacales Subclasses E uasterids I E uasterids I E uasterids II E uasterids II E uasterids II E uasterids II E uasterids II E uasterids II E uasterids II E uasterids II E uasterids II E uasterids II E uasterids II E uasterids II Classe E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots E udicots

­– 44 ­–