Comments on the butterfly fauna (Papilionoidea) of Porto Santo (Madeira Archipelago): species list, distribution patterns, and butterfly-plant network

Abstract

Porto Santo (Madeira Archipelago) is a relatively old (11.1 to 14.3 Ma) and small volcanic island in the Atlantic Ocean. The main part of the island is characterised by semiarid climate and xeric vegetation, while the higher altitudes show subhumid conditions. So far, 11 butterfly species (Papilionoidea) have been detected on Porto Santo. The occurrence of Macroglossum stellatarum (Sphingidae) is published for the first time. The butterfly species of Porto Santo are commented and some comparisons with Madeira Island are discussed. The distribution of butterfly species on Porto Santo was studied during March 2017 (Papilionoidea; n = 204). Some observations of previous years were added (n = 21) and Macroglossum stellatarum (Sphingidae; n = 3) was also included. The butterfly-plant network was sampled during March 2017 (115 flower visits). Three species (Colias croceus, Pieris rapae and Vanessa cardui) show a wide occurrence on Porto Santo. As an exception, Vanessa vulcania prefers the subhumid regions. Leptotes pirithous, Macroglossum stellatarum, Pararge aegeria (which all newly colonised Porto Santo), and the migratory Danaus plexippus show a scattered distribution. On the whole, 115 butterfly-plant interactions could be recorded. The network shows more plant than butterfly species and is asymmetric: Seven butterfly species used 15 plant species. The butterfly and plant species were interconnected. The woody Echium species, E. nervosum (endemic to the Madeira Archipelago) and E. portosanctensis (endemic to Porto Santo), including hybrids, are key species as nectar resources for Vanessa atalanta, V. cardui, and V. vulcania. Vanessa vulcania could be found only within one grid in the north of the island at an altitude of 235 m a.s.l. Vanessa vulcania is probably a remnant of an earlier time period, where-due to lack of human impact-larger areas with microforests and in part Apollonias barbujana laurisilva existed.

Full text

ISSN 2183-279X (online edition) | doi: xx.xxxx/xxxxx-xxxx.xxxxxxxxx.x | Available online at: http://boletim.cm-funchal.pt
BOLETIM
MUSEU DE
HISTÓRIA NATURAL DO FUNCHAL
Vol. LXVIII (2018), Art. 349: 5-20
Received: 20 September 2018; Available online: 31 December 2018; Published: 31 December 2018
Comments on the buttery fauna (Papilionoidea) of Porto Santo
(Madeira Archipelago):
species list, distribution patterns, and buttery-plant network
By A. KrAtochwil
1 *
, A. SchwABe
2
& A. M. FrAnqu inho AguiAr
3
With 2 gures and 3 tables
1 Department of Biology/Chemistry, Ecology Section, University of Osnabrück, Barbarastr. 13, D-49069 Osnabrück, Germany.
* Corresponding author: anselm.kratochwil@biologie.uni-osnabrueck.de
2 Department of Biology, Technische Universität Darmstadt, Schnittspahnstr. 10, D- 64287 Darmstadt, Germany.
3 Entomology Lab, Laboratório de Qualidade Agrícola, Secretaria Regional de Agricultura e Pescas, Caminho Municipal dos Caboucos, 61, 9135-372
Camacha, Madeira, Portugal.
ABSTRACT: Porto Santo (Madeira Archipelago) is a relatively old (11.1 to 14.3 Ma) and small volcanic island in the
Atlantic Ocean. The main part of the island is characterised by semiarid climate and xeric vegetation, while the higher
altitudes show subhumid conditions. So far, 11 buttery species (Papilionoidea) have been detected on Porto Santo.
The occurrence of Macroglossum stellatarum (Sphingidae) is published for the rst time. The buttery species of Porto
Santo are commented and some comparisons with Madeira Island are discussed.
The distribution of buttery species on Porto Santo was studied during March 2017 (Papilionoidea; n = 204).
Some observations of previous years were added (n = 21) and Macroglossum stellatarum (Sphingidae; n = 3) was also
included. The buttery-plant network was sampled during March 2017 (115 ower visits).
Three species (Colias croceus, Pieris rapae and Vanessa cardui) show a wide occurrence on Porto Santo. As an
exception, Vanessa vulcania prefers the subhumid regions. Leptotes pirithous, Macroglossum stellatarum, Pararge
aegeria (which all newly colonised Porto Santo), and the migratory Danaus plexippus show a scattered distribution. On
the whole, 115 buttery-plant interactions could be recorded. The network shows more plant than buttery species
and is asymmetric: Seven buttery species used 15 plant species. The buttery and plant species were interconnected.
The woody Echium species, E. nervosum (endemic to the Madeira Archipelago) and E. portosanctensis (endemic to
Porto Santo), including hybrids, are key species as nectar resources for Vanessa atalanta, V. cardui, and V. vulcania.
Vanessa vulcania could be found only within one grid in the north of the island at an altitude of 235 m a.s.l. Vanessa
vulcania is probably a remnant of an earlier time period, where – due to lack of human impact – larger areas with
microforests and in part Apollonias barbujana laurisilva existed.
Keywords: Lepidoptera, Papilionoidea, Sphingidae, island biogeography, buttery diversity, Vanessa vulcania,
Madeira Archipelago, Porto Santo.

Boletim do Museu de História Natural do Funchal No. LXVIII, Art. 3496
RESUMO: Porto Santo (Arquipélago da Madeira) é uma pequena ilha vulcânica relativamente antiga (11.1 a 14.3
Ma) situada no Oceano Atlântico. A parte principal da ilha é caracterizada por ter um clima semiárido e vegetação
xérica, enquanto as altitudes mais altas mostram condições sub-húmidas. Até agora, 11 espécies de borboletas
(Papilionoidea) foram detectadas no Porto Santo. A ocorrência de Macroglossum stellatarum (Sphingidae) é publicada
pela primeira vez. As espécies de borboletas do Porto Santo são comentadas e algumas comparações com a ilha da
Madeira são discutidas. A distribuição das espécies de borboletas no Porto Santo foi estudada durante março de
2017 (Papilionoidea; n = 204). Algumas observações de anos anteriores foram adicionadas (n = 21) e Macroglossum
stellatarum (Sphingidae; n = 3) também foi incluído. A rede de plantas de borboletas foi amostrada durante março
de 2017 (115 visitas de ores). Três espécies (Colias croceus, Pieris rapae e Vanessa cardui) são bastante frequentes
no Porto Santo. Como exceção, Vanessa vulcania prefere as regiões sub-úmidas. Leptotes pirithous, Macroglossum
stellatarum, Pararge aegeria (todas introduções recentes no Porto Santo), e a espécie migratória Danaus plexippus
migratórias mostram uma distribuição mais dispersa. No total, foram registradas 115 interações borboleta-planta. A
rede inclui mais plantas do que espécies de borboletas e é assimétrica: sete espécies de borboletas usaram 15 espécies
de plantas. As espécies de borboletas e plantas estavam interligadas. As espécies lenhosas do género Echium, E.
nervosum (endémica do arquipélago da Madeira) e E. portosanctensis (endémica do Porto Santo), incluindo os seus
híbridos, são espécies-chave como recursos de néctar para Vanessa atalanta, V. cardui, e V. vulcania. Vanessa vulcania
só foi encontrada dentro de uma grade no norte da ilha a uma altitude de 235 m. Vanessa vulcania é provavelmente
um remanescente de um período de tempo anterior, em que – devido à falta de impacto humano – existiam áreas
maiores com microorestas, assim como Laurissilva do Barbusano.
Palavras-chave: Lepidoptera, Papilionoidea, Sphingidae, biogeograa de ilhas, diversidade de borboletas, Vanessa
vulcania, Arquipélago da Madeira, Porto Santo.
INTRODUCTION
The rst list of Papilionoidea of Madeira Island
(seven species) was published by
Felder
(1862), based
on specimens collected during the Novara Expedition
(1857-1859), the rst broad-scale scientic mission of the
Austrian Imperial Navy (
Basch-ritter
, 2008). The collector
was the Austrian zoologist Georg von Frauenfeld (1807-
1873), though he did not visit Porto Santo.
In nine expeditions, the British entomologist
Thomas Vernon Wollaston (1822-1878) collected much
more intensively from the Madeira Archipelago (
Machado
,
2006), including Madeira Island, the Desertas Islands, and
Porto Santo. His main interest was the study of Madeiran
Coleoptera (
Wollaston
, 1854), but also Hymenoptera (e.g.,
wild bees;
KratochWil
, 2018) and Lepidoptera (
Karsholt
,
2000). For generations, this immense material served as
a base for further analyses, which resulted in numerous
publications and species descriptions.
BaKer
(1891) analysed
the Papilionoidea (11 species) collected by Wollaston
and commented the list (e.g., abundancy, morphological
features, dierences to mainland forms), but without
further information (e.g., on which island they had been
collected). As a rule, Wollaston had labelled or colour-
marked the specimens according to the islands where
they were collected, or (in most cases) had numbered
and listed them in books (
Machado
, 2006). However,
BaKer
(1891) obviously did not consider localities as important.
Furthermore, most of the details had been documented
by Wollaston in lists, but some of them were hard to
nd or lost.
BaKer
(1891) was the rst who mentioned
Macroglossum stellatarum in relation to Madeira Island.
reBel
(1917) listed 13 Papilionoidea species for the
Madeira Archipelago without further comments and
without notes of the occurrences on dierent islands;
reBel
(1940) listed 11 species (without Pieris rapae and Vanessa
atalanta, mentioned in
reBel
, 1917). A more detailed list by
Martin
(1941) refers only to Madeira Island.
The rst list of the Papilionoidea of Porto Santo was
published by
carvalho
(1983). In 1980 and 1981, he observed
four species in one locality (Achada de Baixo): Pieris rapae,
Colias croceus, Lampides boeticus, and Vanessa cardui. His
list also included Vanessa indica occidentalis, cited by
BaKer
(1891), but the Sphingid species Macroglossum stellatarum
was not mentioned.
oWen
et al. (1987) listed 14 species for
Madeira Island.
A comprehensive overview of the butteries of
the Madeira Archipelago was given by
Meyer
(1993).

Kratochwil et al., Comments on the buttery fauna of Porto Santo2018 7
He distinguished the dierent islands of the Madeira
Archipelago and found the following species numbers
without vagrants: Madeira Island, 14 species; Porto Santo,
ve species; Deserta Grande, Bugio, Selvagem Grande,
and Selvagem Pequena, one species each.
oWen & sMith
(1993) analysed the buttery fauna of Macaronesia and
stated the same species numbers for Madeira Island and
Porto Santo as
Meyer
(1993).
An updated list of the butteries of the Madeira
Archipelago was presented by
aguiar & Karsholt
(2006).
The rst distribution maps of the endemic butteries
(Papilionoidea) of Madeira Island were produced by
WaKehaM-daWson
et al. (2002a), but distribution maps of all
species were not available.
WaKehaM-daWson
et al. (2002b)
also provided some further notes of the butteries of the
Madeira Archipelago, and
WaKehaM-daWson
et al. (2004)
presented the last published checklist of Madeira Island
(13 species) and Porto Santo (10 species).
Although numerous studies have already focused
on the butteries of Porto Santo, their distribution patterns
and networks with owering plants have not been studied
until now.
An overview of wild bees (Hymenoptera, Anthophila)
on Porto Santo has been recently published by
KratochWil
& schWaBe
(2018), which proves the existence of a wide
distribution of most wild-bee species – except for the
bumblebee Bombus terrestris lusitanicus, which prefers
the subhumid regions – and demonstrates the signicant
importance of the Echium species for ower-visiting bees.
The following topics will be presented in this study:
(1) a commented checklist of buttery species of
Porto Santo;
(2) a rst overview of the distribution patterns of
11 buttery species occurring on Porto Santo and their
nectar resources;
(3) the structure of the buttery-plant network in
Porto Santo.
Physico-geographical factors
Porto Santo is a relatively old (11.1 to 14.3 Ma) and
small volcanic island (area 42 km2) within the Madeira
Archipelago (
geldMacher
et al., 2000), with latest volcanic
activities occurring about eight million years ago (
cooK
,
2008). The island is separated from Madeira Island by an
ocean bed more than 2000 m deep.
Geologically Porto Santo has a volcanic base, and the
summits (e.g., Pico do Facho 517 m a.s.l., Pico da Gandaia
499 m a.s.l., Pico Branco 451 m a.s.l.) are characterised
mainly by trachyte and basaltic structures. More than a
third of the island is covered by quaternary sediments,
mainly between the northeastern and southwestern
mountain areas. Calcareous sand from the Pleistocene
period formed eolianites, which is solidied with more
than 50 m vertical thickness in part. The southern coast is
dominated by a long sandy beach, while the other parts of
the coast are characterised by steep clis. Perennial water
currents are not present, but some perennial springs exist
(
Faust-lichtenBerger
, 1988).
The bioclimate was classied as Mediterranean
xeric oceanic by
rivas-Martínez
(2009) with a pronounced
summer aridity. Precipitation values are very low from
June to August (376 mm/a), while mean temperature
varies between 15.7 °C (February) and 22.8 °C (August).
There are only few habitat types, and for the last
500 years there has been a strong anthropogenic impact.
There are about 446 phanerogamic plant species (nine
endemic species for Porto Santo, 29 endemic species for
the Madeira Archipelago, and 19 endemic species for
Macaronesia). All in all, 286 plant species are (probably)
native, and 103 species are (probably) introduced (
JardiM &
Menezes de sequeira
, 2008;
JardiM & Menezes de sequeira
, 2011;
Jones
et al., 2014;
KratochWil & schWaBe
, 2018).
The island was cleared to a great extent from woody
vegetation (original vegetation: mainly dry microforests of
Mayteno umbellatae-Oleo maderensis sigmetum). From
the end of the 18th century, aorestations were made with
Pinus pinaster and Pinus halepensis. Most habitats belong
to the Olea maderensis series (Mayteno umbellatae-Oleo
maderensis sigmetum);
capelo
et al. (2004, 2005). The
Echium species are the main nectar sources for many
butteries. The natural distribution of E. nervosum is
concentrated in the southern part, while E. portosanctensis
occurs naturally in the subhumid zone (
carvalho
et al., 2010).
Echium nervosum can also be found in plantings near the
roads, while E. portosanctensis was planted in the northern
part (hybrids exist). Small elements of the subhumidic
series are still present around the main summits and on
the north-facing slopes (
capelo
et al., 2004).
Extended sand habitats occur especially in the
southeastern part of the island and are relatively rich
in Lotus glaucus. There are also drift walls with Cakile
maritima, as well as halonitrophytic rocky and sandy
sites with Matthiola maderensis, Mesembryanthemum
crystallinum, and Senecio incrassatus. The island has
a very rich ruderal ora and vegetation, now mainly
characterising fallow land. Ruderal plant species such as
Asphodelus stulosus, Convolvulus althaeoides, Rapistrum
rugosum, and Sonchus oleraceus play a role as nectar
resources for butteries.

Boletim do Museu de História Natural do Funchal No. LXVIII, Art. 3498
METHODS
Buttery database
We observed the butteries of Porto Santo mainly
during one visit from 19 to 31 March 2017 (A . Kratochwil and
A. Schwabe). We recorded the distribution pattern of the
butteries and their interactions with plants. Qualitative
observations from the dry spring of 2012 (16-20 March)
preceded the 2017 study. Due to the presence of masses
of owers after the relatively wet winter in 2016/2017 (see
KratochWil & schWaBe
, 2018), the time period in March 2017
was very good for buttery monitoring.
Our approach covered the main habitat types. All in
all, we monitored 45 sites in the northern and southern
parts of the island – most localities with a full xeric, some
localities with a subhumid mesic-xeric bioclimate. The
number of observation sites reected the proportion of
the climatic types on the island. Most of these sites were
identical with the wild-bee sampling sites (
KratochWil &
schWaBe
, 2018). We were not able to observe butteries
in the at central part of the island (airport, golf course)
because access to the area was not permitted. The GPS
data of all the collected or observed buttery species were
recorded with a Garmin Oregon 700.
We recorded ten buttery species and 204 buttery
individuals (Table 1), including 115 ower visits. Fourteen
further observations by A. F. Aguiar were included in
the database (six species from nine localities, and ve
ower visits on three plant species). Additionally, seven
specimens collected on Porto Santo were found in the
collections of the Museu de História Natural do Funchal
(Madeira, Portugal).
Data analysis
The localities were plotted on a grid (1 km x 1 km)
based on the military map of the Madeira Archipelago
(2004). Interaction networks were analysed using the
bipartite package (
dorMann
et al., 2008;
r core teaM
, 2018).
RESULTS AND DISCUSSION
Commented checklist of the buttery species of
Porto Santo
We were able to analyse the distribution pattern of 11
species (Table 2, Fig. 1), which are commented below. The
Sphingid Macroglossum stellatarum (three observations,
two ower visits observed) was included in the analysis.
We observed the following species
(in alphabetical order):
Colias croceus (Georoy in Fourcroy, 1785)
This Palaearctic migrating species – widespread
in Europe except the North (occurring in southern
Scandinavia, Baltic states; see
tolMan & leWington
, 1997),
but common in Central and Southern Europe, and with
further distribution in North Africa and Southwest Asia
(
tshiKolovets
, 2011) – is native to the Madeira Archipelago
and the Canary Islands (
tolMan & leWington
, 1997).
On Madeira Island, C. croceus was very common
on open sites (
salMon & WaKehaM-daWson
, 1999), and in
July and August 1998, was the most abundant buttery
on the island (
salMon & WaKehaM-daWson
, 1999). The main
distribution is concentrated in dry coastal areas and near
human settlements (
Meyer
, 1993). Colias croceus occurs
at all altitudes from sea level up to 1,800 m a.s.l. (
Meyer &
hellers
, 1990;
Kudrna
, 1997;
salMon & WaKehaM-daWson
, 1999).
sWash & asKeW
(1982) reported that C. croceus is the most
abundant Madeiran buttery at altitudes above 1,600 m
a.s.l. Colias croceus was also recorded on Porto Santo (
Meyer
& hellers
, 1990;
aguiar & Karsholt
, 2006).
It is suggested that this species (probably) breeds on
Madeira Island (
vieira
, 1999;
WaKehaM-daWson & aguiar
, 2003;
WaKehaM-daWson
et al., 2004). The ratio between breeding
and migrant individuals is unknown, but it seems that
vagrants are in minority.
salMon & WaKehaM-daWson
(1999) present dierent
varieties (forms) of C. cro ceus from the Madeira Archipelago:
var. cremonae Verity, 1911; var. dawsoni Bollow, 1930; var.
geisleri Bryk, 1923; var. radiata Nitsche, 1932; and var.
faillae Stefanelli, 1900.
Fuchs
(1993) also described the
high variability of colour forms of C. croceus from Faial
(Azores). Such wing colour patterns, based on phenotypic
plasticity, may be produced physiologically in response
to environmental stress (e.g., temperature, general stress
response) and may be genetically xed in a population
(
otaKi
et al., 2010). This is shown particularly in the genus
Vanessa (summarised in
hiJaMa
et al., 2002).
aguiar & Karsholt
(2006) reported that var. helice
(
hüBner
, 1879) is rather frequently seen on Porto Santo (5-
10% of all populations;
tolMan & leWington
, 1997).
In the Museu de História Natural do Funchal three
females of the form ‘helice’ are deposited, collected in May
1942 (unknown collector, determined by J. T. Smit in 1998),
and also two females of the same form, collected on 30
March 1988 by F. Zino (determined by J. T. Smit in 1998).

Kratochwil et al., Comments on the buttery fauna of Porto Santo2018 9
Table 1 – List of buttery species of Porto Santo (MA = Madeira Archipelago, CI = Canary Islands), number of observations and number of
ower visits in this study.
Tab le 2 – Numbers of observed individuals, localities, and grids of 11 species. * = including data from
carvalho
(1983).
Tab le 3 – Visited plant species and number of observations in March 2017.
Family Species Status Number
individuals
Number flower
visits
Data from the authors and
Museu de História Natural do Funchal
Pieridae Pieris rapae (Linnaeus, 1758) native/introduced? 16 2
Pieridae Colias croceus (Geoffroy in Fourcroy, 1785) native 82 23
Lycaenidae Leptotes pirithous (Linnaeus, 1758) introduced? 12 2
Lycaenidae Lampides boeticus (Linnaeus, 1767) native 4 2
Nymphalidae Danaus plexippus (Linnaeus, 1758) native/introduced? 6 1
Nymphalidae Vanessa atalanta (Linnaeus, 1758) native 19 18
Nymphalidae Vanessa cardui (Linnaeus, 1758) native 68 58
Nymphalidae Vanessa vulcania (Godart, 1819) endemic (MA, CI) 10 10
Nymphalidae Pararge aegeria aegeria (Linnaeus, 1758) introduced 5 2
Sphingidae Macroglossum stellatarum (Linnaeus, 1758) native 3 2
Data from other observers
Nymphalidae Hypolimnas missipus (Linnaeus, 1764) migrant - -
Lycaenidae Lycaena phlaeas phlaeoides (Staudinger, 1901) endemic (MA) - -
Number
of grids Localities Individuals
Colias croceus 19 38 77
Danaus plexippus 4 4 4
Hypolimnas misippus 2 2 3
Lampides boeticus 3* 4* 4
Leptotes pirithous 5 6 12
Macroglossum stellatarum 3 3 3
Pararge aegeria 4 4 5
Pieris rapae 11 13 14
Vanessa atalanta 3 3 19
Vanessa cardui 914 68
Vanessa vulcania 1 2 10
Number
flower visits
Echium nervosum (including hybrids) 86
Lotus glaucus 6
Asphodelus fistulosus 5
Leontodon taraxacoides 3
Cakile maritima 2
Convolvulus althaeoides 2
Matthiola maderensis 2
Brassica nigra 1
Echium portosanctensis 1
Galactites tomentosa 1
Mesembryanthemum crystallinum 1
Oxalis pes-caprae 1
Rapistrum rugosum 1
Senecio incrassatus 1
Sonchus oleraceus 1

Boletim do Museu de História Natural do Funchal No. LXVIII, Art. 34910
Fig. 1 – Distribution patterns of 11 buttery species from Porto Santo.

Kratochwil et al., Comments on the buttery fauna of Porto Santo2018 11
If we use the form ‘helice’ in the broader sense for
all bright forms (white to yellowish wing colour; ‘alba’
polymorphism), four of the specimens we observed have to
be assigned to this form. So far, the literature has reported
the form ‘helice’ only for female butteries (
lorKovic &
herMan
, 1961;
liMeri & Morehouse
, 2016;
liMeri
, 2017).
Our observations demonstrate that C. croceus is also
quite common throughout Porto Santo from sea level
up to the highest mountains (19 grids, 38 localities; Fig.
1). Colias croceus could be detected on the owers of ten
plant species: Asphodelus stulosus (Xanthorrhoeaceae)
n = 5; Brassica nigra (Brassicaceae) n = 1; Convolvulus
althaeoides (Convolvulaceae) n = 2; Echium nervosum
(Boraginaceae) n = 1; Galactites tomentosa (Asteraceae) n =
1; Leontodon taraxacoides (Asteraceae) n = 3; Lotus glaucus
(Fabaceae) n = 6; Matthiola maderensis (Brassicaceae) n =
2; Mesembryanthemum crystallinum (Aizoaceae) n = 1; and
Oxalis pes-caprae (Oxalidaceae) n = 1.
Danaus plexippus (Linnaeus, 1758)
This species, originally native to North and Central
America, nowadays has – by migration events or man-made
introductions – a presence in the warmer zones (southern
Portugal, southern Spain, India, Papua New Guinea, other
East Indian islands, Australia, New Zealand, Hawaii, and
southern Peru to Canada;
tolMan & leWington
, 2008). There
are migratory and non-migratory populations (
FreedMan
et al., 2018). Danaus plexippus is classied as a migratory
species in Madeira, Porto Santo, and Canary Islands (
oWen
& sMith
, 1993). According to
aguiar & Karsholt
(2006), this
species has occurred periodically on Madeira Island since
1889, and was seen for the rst time in Porto Santo in 1955
(
pereira
, 1989). The exact date of arrival is unknown, and not
recorded by
BaKer
(1891) nor
cocKerell
(1923). A successful
colonisation was possible because the species is able to
use introduced plant species as larval food (
oWen & sMith
,
1989). Local populations established in the Canary Islands
in 1880 or 1887 (
higgins & riley
, 1970;
leestMans
, 1975;
Báez
,
1998), and the species was also spotted on Madeira Island
(
sousa
, 1986, 1991) and Porto Santo (
gardner & classey
, 1960;
vieira
, 1999). The establishment on Madeira and Porto
Santo is believed to have happened in August 1980 (
aguiar
& Karsholt
, 2006).
salMon & WaKehaM-daWson
(1999) and
shoWler
(2001) hypothesised an intended introduction.
During the last two decades, D. plexippus has also become
a resident species in the Azores (
neves
et al., 20 01).
In the Museu de História Natural do Funchal one
female, collected on 14 June 1970 (unknown collector)
and one male, collected in September 1957 (unknown
collector), are deposited; both were determined by J. T.
Smit in 1998.
One ower visit of this species was observed on
Bougainvillea by A. F. Aguiar. On Porto Santo, D. plexippus
was detected on four grids and in four localities (Fig. 1)
ying at high speed with directed movement.
Lampides boeticus (Linnaeus, 1767)
This seasonal migratory species is widely distributed
all over the world in southern regions with higher
temperatures, and in subtropic and tropic zones: Southern
Europe (reaching occasionally to Germany and the British
Isles), North Africa, subtropical and dryer regions in South
and Southeast Asia, Australia, New Zealand, and Hawaii. In
Macaronesia L. boeticus is distributed on the Canary Islands,
the Azores (
Felder
, 1862;
leestMans
, 1974), and Madeira
Island (
reBel
, 1940;
Meyer & hellers
, 1990;
Meyer
, 1993;
tolMan
& leWington
, 1997;
tshiKolovets
, 2011). On Madeira Island the
species was recorded by
BaKer
(1881),
sWash & asKeW
(1982),
and
salMon & WaKehaM-daWson
(1999), and classied both
as common by
oWen
et al. (1987) and as not common by
WaKehaM-daWson
et al. (2004).
carvalho
(1983) reported this species in two localities
on Porto Santo (Achada de Baixo, 18 September 1980, and
Farrobo, near the airport, 17 September 1980). The species
was observed in two further localities (Vila Baleira, 16
July 2001; road to Fonte da Areia, 6 July 2017). Lampides
boeticus was detected in three grids (Fig. 1) and one ower
visit on Lotus glaucus was observed.
WaKehaM-daWson &
aguiar
(2003) documented egg laying on Lotus glaucus on
Porto Santo (in July 2002).
Leptotes pirithous (Linnaeus, 1767)
The range of L. pirithous spans from North Africa,
Southern and Central Europe, Turkey, the Middle East,
Saudi Arabia, and Central Asia to India (
tolMan & leWington
,
1997). The species is also present in the Canary Islands
(
tshiKolovets
, 2011), where the rst detection occurred in
1998 (
aguiar & Karsholt
, 2006). Probably originating from
the Canary Islands, L. pirithous was introduced to the
Madeira Archipelago. Records of L. pirithous on Madeira
Island and Porto Santo were reported in 2001 by
hall &
russell
(2001),
WaKehaM-daWson
et al. (2002b), and
aguiar
et
al. (2002). One ower visit of this species was observed
on Rosmarinus ocinalis by A. F. Aguiar.
WaKehaM-daWson
et al. (2002b) reported observations of this species ying
around Lotus glaucus in the sand dunes of Calheta (Porto
Santo). On Porto Santo, L. pirithous was detected on ve
grids and in six localities (Fig. 1).

Boletim do Museu de História Natural do Funchal No. LXVIII, Art. 34912
Macroglossum stellatarum (Linnaeus, 1758)
This migratory species has a permanent distribution
in the southern Palaearctic region including North Africa,
the Near East, and Pakistan, reaching eastwards to China
and Japan. The northern areas are populated only in
warmer months.
The rst record of M. stellatarum was reported by
BaKer
(1891), who characterised this species as common for
Madeira Island. Further detections were reported by
reBel
(1940, 1940b) and
Martin
(1941). Macroglossum stellatarum
was also detected on Selvagem Grande (
aguiar & Karsholt
,
2006). On Porto Santo, M. stellatarum was found on three
grids and three localities (Fig. 1), only on owers of Echium
nervosum (Boraginaceae) n = 2.
Pararge aegeria aegeria (Linnaeus, 1758)
The nominal species is distributed in Maghreb,
the Iberian Peninsula, the Balearic Islands, southern
and southwestern France, southwestern Switzerland,
Peloponnesus, Corsica, Sardinia, Sicily, southern Italy,
Crete, Lesbos, Samos, Kos, Karpathos, Cyprus, southern
Turkey, and the Near East (
tshiKolovets
, 2011). Madeira was
recently colonised – the rst specimen was observed
by the lepidopterist Hoegh-Guldberg in 1967 on
Madeira Island (
oehMig
, 1983). Numerous detections of
this introduced species were documented in the mid-
seventies (
higgins
, 1977;
oehMig
, 1977, 1982;
oWen
et al., 1987;
aguiar & Karsholt
, 2006). In 1981, P. a. aegeria was at least
as common as P. xiphia on Madeira Island (
sWash & asKeW
,
1981). Nowadays, P. a. aegeria is one of the more common
butteries on Madeira Island living in lower elevations
mainly in residential areas, disturbed habitats, cultivation
sites, and gardens (
lace & Jones
, 1984;
oWen
et al., 1987;
shreeve & sMith
, 1992;
Fernández-ruBio & garcia-Barros
, 1995;
oWen
et al., 2008). It was not seen on Porto Santo in 1981
(
sWash & asKeW
, 1982), but a single specimen was detected
in 1998 (
vieira
, 1999).
On Porto Santo, P. a. aegeria was detected on four
grids and in four localities (Fig. 1) and could be observed
on two plant species: Cakile maritima (Brassicaceae) n = 1
and Senecio incrassatus (Asteraceae) n = 1. Territorial and
mating behaviours were noted.
Pieris rapae (Linnaeus, 1758)
This species has a trans-Palaearctic, North-
African distribution including the Arabian Peninsula and
subtropical Asia. It was introduced and is now resident
in North America and Australia (
tshiKolovets
, 2011), and
it is less frequent in northern Fennoscandia (
tolMan &
leWington
, 1997). The rst indications for an occurrence on
Madeira Island came from
reBel
(1917), but in his checklist
the taxon P. rapae is characterised by a question mark. In a
footnote he referred to Otto Sterz, who observed P. rapae
on Madeira Island in March 1909. A voucher specimen
does not exist.
P. rapae is a recent introduction on Madeira Island
and has become one of the most abundant buttery
species (
aguiar & Karsholt
, 2006). According to
WolFF
(1975),
a specimen of P. rapae from Madeira Island is deposited
in the Museu de História Natural do Funchal, Madeira,
collected on 15 December 1971.
WolFF
(1975) documented
a single observation in 1973, but occurrence in substantial
numbers started in 1974. Three years later, P. rapae was
widespread on the island and extremely common. The
altitudinal distribution of this buttery species spans from
sea level up to more than 1,500 m a.s.l., and it is found
mainly on coastal sites and at lower altitudes (human
settlements and agricultural areas).
A colonisation with established populations on
Madeira Island was conrmed by, e.g.,
oehMig
(1977),
sWash
& asKeW
(1982),
lace & Jones
(1984),
sousa
(1986), and
Meyer
& hellers
(1990).
While
Meyer
(1993) hypothesised an introduction with
vegetables from the mainland,
WolFF
(1975) is convinced
that numerous specimens were transported from Portugal
to Madeira Island during special weather periods. He
pointed out a particularly favourable meteorological
condition for air transport in 1974. Furthermore,
WolFF
(1975) stated that the specimens of Madeira must be
classied as form ‘metra’ (
stephens
, 1824), which also occurs
in Portugal.
The rst record on Porto Santo was reported by
carvalho
(1983) at Achada de Baixo on 17 September 1981;
see also
oWen & sMith
(1993) and
aguiar & Karsholt
(2006).
Meyer & hellers
(1990) observed the species on Porto Santo,
too, and it was also found on Deserta Grande (
sWash &
asKeW
, 1982).
P. rapae is quite common throughout Porto Santo
(11 grids, 13 localities; Fig. 1) and could be detected on the
owers of two plant species: Cakile maritima (Brassicaceae)
n = 1; and Sonchus oleraceus (Asteraceae) n = 1.
Vanessa atalanta (Linnaeus, 1758)
This migratory buttery species is widespread
throughout North America, down south to Guatemala,
Cuba, Hispaniola, the Atlantic islands, North Africa,
Europe, Ireland, Britain, southern Fennoscandia, the
Mediterranean islands, Pakistan, Kashmir, and north of the

Kratochwil et al., Comments on the buttery fauna of Porto Santo2018 13
Himalayas to the Amur river (
Field
, 1971;
tolMan & leWington
,
1997;
tshiKolovets
, 2011). The American and Old World
populations were characterised as separate subspecies
(
vane-Wright & hughes
, 2007).
Wollaston collected three specimens (Madeira
Island, The Mount, without date), which is recorded by
BaKer
(1891) and
reBel
(1917).
sWash & asKeW
(1982) stated
that V. atalanta is never numerous on Madeira Island, in
contrast to the observations of
WaKehaM-daWson
et al. (2004)
which classied the occurrence as relatively common. The
preferred habitats are human settlements with gardens
(
lace & Jones
, 1984;
oWen
et al., 1987;
WaKehaM-daWson
et al.,
2004). Vanessa atalanta was also recorded on Porto Santo
and Deserta Grande (
aguiar & Karsholt
, 2006; see also
Meyer
& hellers
, 1990).
On Porto Santo, V. atalanta is restricted to sites with
high abundancies of Echium owers (3 grids, 3 localities;
Fig. 1) and could be detected only on the owers of Echium
nervosum (Boraginaceae) n = 18.
Vanessa cardui (Linnaeus, 1758)
This cosmopolitan migratory species (except for
Australia and New Zealand) is widely distributed and a
permanent resident of North Africa and the Mediterranean
zone (
tolMan & leWington
, 2008). Vanessa cardui was also
recorded in Venezuela (
tshiKolovets
, 2011). The species
is resident on the Madeira Archipelago (Madeira Island,
Porto Santo, Deserta Grande, Selvagem Grande and
Selvagem Pequena) (
aguiar & Karsholt
, 2006). On Madeira
Island, the species is common from 50 to 400 m a.s.l. in
cultivated terraces and coastal sites (
WaKehaM-daWson
et al.,
2004), and, similar to Colias croceus, also concentrated on
drier sites (abundant in Ponta de São Lourenço;
oWen
et al.,
1987), and near human settlements (
Meyer
, 1993). Vanessa
cardui is also quite common throughout Porto Santo from
sea level up to the highest mountains (9 grids, 14 localities;
Fig. 1), but not so common as Colias croceus.
There is evidence that this species breeds on
Madeira Island but appears to be also a migrant (
oWen
,
1989;
salMon & WaKehaM-daWson
, 1999). Breeding in winter
was observed by
oWen
(1987).
On 18 January 1921,
cocKerell
(1923) collected an
aberration of V. cardui on Porto Santo, deposited in the
Oxford University Museum of Natural History (OUMNH).
Such similar stress-induced colour-pattern modications
were considered as phenotypic plasticity in V. cardui (
otaKi
,
2007).
V. cardui could be detected on the owers of
three plant species, with high preference of Echium
(Boraginaceae): E. nervosum n = 55; E. portosanctensis n = 1;
and Rapistrum rugosum (Brassicaceae) n = 2.
Vanessa vulcania (Godart, 1819)
In the literature this species was classied as a
subspecies of V. indica Herbst, 1797 (e.g.,
Field
, 1971;
tolMan
& leWington
, 2008) or a species of its own (e.g.,
Bernardi
,
1961;
leestMans
, 1975b, 1978;
tshiKolovets
, 2011). Synonyms
are Pyrameis (?) vulcania Godart, 1819, and Pyrameis
callirhoe var. occidentalis Felder, 1862.
leestMans
(1978)
demonstrated the clear separation between V. vulcania
and V. indica.
V. vulcania is a Macaronesian endemic species,
occurring in the Madeira Archipelago and the Canary
Islands (
BaKer
, 1891;
Meyer & hellers
, 1990;
tolMan & leWington
,
1997;
WaKehaM-daWson & aguiar
, 2003;
vane-Wright & hughes
,
2007;
tshiKolovets
, 2011). Within the Madeira Archipelago, V.
vulcania was detected on Madeira Island, Porto Santo, and
Deserta Grande (
Meyer & hellers
, 1990;
aguiar & Karsholt
,
2006).
There were also occasional detections (natural or
introductions) of V. vulcania in Germany (
gerisch
, 1975,
1978;
reinhardt & gerisch
, 1982), Western and Northern
Europe (
opheiM
, 1960), and the Iberian Peninsula (
Fernández-
vidal
, 1989;
Fernández-ruBio
, 1991) with no establishment.
One of the rst records came from
Felder
(1862): A
male was collected during the Novara Expedition (1857-
1859) by Georg von Frauenfeld on Madeira Island and
described as Pyrameis Callirhoë Hübner, var. occidentalis
Felder, 1862. Alfred Russel Wallace (1889) cited Wollaston
(
Wollaston
, 1856) and noted that specimens from Porto
Santo are much smaller and darker than those of Madeira
(cited also in
salMon & WaKehaM-daWson
, 1999).
reBel
(1940)
mentioned that the specimens from Madeira Island are
not dierent from those of the Canary Islands.
V. vulcania is widely distributed on Madeira Island
from sea level to altitudes up to 1,000 m a.s.l., but it is not
common (
WaKehaM-daWson & aguiar
, 2002;
WaKehaM-daWson
et al., 2004). In contrast,
oWen & sMith
(1993) noted that V.
vulcania is well established and common, more abundant
than V. atalanta.
sWash & asKeW
(1982) reported frequent
observations, mostly at low and medium altitudes in
cultivated areas (see also
oWen
et al., 1987).
shapiro
(1992a) suggests a possible introduction by
Portuguese traders; however, this is not the opinion of
salMon & WaKehaM-daWson
(1999).
shapiro
(1992a, b) noted a
habitat preference for laurisilva forests and hypothesised
that V. vulcania could be also considered as a relict species
of former broadleaved forests.

Boletim do Museu de História Natural do Funchal No. LXVIII, Art. 34914
There are two dierent hypotheses of the origin of
V. vulcania:
(1)
leestMans
(1978) suggested that during glacial
maxima, the taxon of the V. indica complex could have
been distributed in the eremic zone of North Africa,
towards the east to Somalia and the Arabian Peninsula,
and from there to the Iranian Plateau, the Thar Desert
and the Tien Shan, and V. vulcania or its ancestor reached
the Canary Islands and Madeira Archipelago via southern
Morocco. In this scenario V. vulcania and V. indica indica
should be sister taxa (
leestMans
, 1978).
WahlBerg & ruBinoFF
(2011) note that the position of V. vulcania is not clear
within the V. indica complex, but it does appear to be
a separate lineage. This hypothesis is compatible with
the results on mitochondrial genetic variability within
the Hyles euphorbiae sensu lato lineage (Lepidoptera,
Sphingidae, Macroglossinae) in the western Palaearctic
(
hundsdoerFer
et al., 2010). Most haplotypes of H. tithymali
in Macaronesia, North Africa, and Yemen build a cluster of
similar haplotypes.
(2) Alternatively,
vane-Wright
(2007) discussed the
hypothesis of a North-American taxon reaching the
Atlantic islands from eastern North America, rather than
North Africa, while the Asian indica-group species reached
Asia from western North America via Beringia, a theory
supported by molecular ndings.
On Porto Santo, V. vulcania is restricted to subhumid
sites with high abundancies of Echium owers (1 grid, 2
localities; Fig. 1) and could be detected only on the owers
of Echium nervosum (Boraginaceae) n = 10.
There are additional data of other observers:
Hypolimnas misippus (Linnaeus, 1764)
The occurrence of this migrant nymphalid species
was recorded in
tennent
et al. (2013). Hypolimnas misippus
is widespread (Africa, Asia, Australia, West Indies, and
parts of Central and North America);
tolMan & leWington
(2008). This species is resident on the Cape Verde Islands,
occasionally reported in the Azores, the Canary Islands,
and the Madeira Archipelago (
tennent
et al., 2013). There
had been three detections in Porto Santo: both sexes, Pico
do Castelo, 16 November 2012; and one female at the golf
club, 18 November 2012, on owers of Lantana camara (2
grids, 2 localities; Fig. 1).
Lycaena phlaeas phlaeoides (Staudinger, 1901)
Lycaena phlaeas (Linnaeus, 1761) is a widespread
species with 22 subspecies in the Palaearctic, oriental,
and Ethiopian regions, and six subspecies in the North
American region (
Kohler
, 2007). Lycaena phlaeas phlaeoides
represents an endemic subspecies on the Madeira
Archipelago (
reBel
, 1940;
Bernardi
, 1961;
leestMans
, 1975;
Meyer & hellers
, 1990;
Meyer
, 1993;
tolMan & leWington
, 1997;
WaKehaM-daWson & aguiar
, 2003;
aguiar & Karsholt
, 2006;
tshiKolovets
, 2011).
Kohler
(2007) described the subspecies
as follows: ‘The rich brown, somewhat mottled colour
and jagged whitish postmedian band of the ventral hind
wing on this subspecies are distinctive’. This subspecies is
represented in the Madeira Archipelago also in dierent
forms: form ‘elea’ (Fabricus, 1798), form ‘coeruleapunctata’
(Rühl, 1895), form ‘obsoleta’ (Tutt, 1896), and form ‘radiata’
(Tutt, 1896); see also
salMon & WaKehaM-daWson
(1999).
cocKerell
(1923) detected a specimen without any dark
colours (ordinary form ‘phlaeas’). The subspecies rank was
not accepted, e.g., by
sWash & asKeW
(1982). According to
Meyer
(1993), the Madeiran populations are native.
L. phlaeas was mentioned by
Felder
(1862): A male
was collected during the Novara Expedition (1857-1859) by
Georg von Frauenfeld on Madeira Island. The distribution
of L. p. phlaeoides on Madeira Island was reported by
WaKehaM-daWson & aguiar
(2002) and
WaKehaM-daWson
et al.
(2002a) as widely spread at all altitudes up to 1,800 m a.s.l.
The occurrence on Porto Santo is mentioned by
WaKehaM-
daWson
et al. (2004).
Distribution patterns
The distribution patterns of 11 buttery species of
Porto Santo are summarised in Table 2 and Fig. 1. Colias
croceus shows the widest distribution on the island (19
grids), followed by Pieris rapae and Vanessa cardui (11
and nine grids). All other species are characterised by
a restricted distribution: Vanessa vulcania (one grid)
and V. atalanta (three grids) are concentrated on sites
with owering Echium; Leptotes pirithous (ve grids) and
Lampides boeticus (three grids) prefer hot, dry, and sunny
sites (
tshiKolovets
, 2011), in contrast to Pararge aegeria
(four grids), which use shadow sites with trees or bushes
(
tolMan & leWington
, 1997). The migrant species Danaus
plexippus and Hypolimnas misippus were detected on four
grids and two grids respectively, while the newly detected
Macroglossum stellatarum was found on three grids.
Flower visits and buttery-plant network
Table 3 shows the visited plant species in March
2017, which were used as nectar resources. Butteries
could be detected on 15 plant species. The key species
with the highest rates of visitation was Echium nervosum.

Kratochwil et al., Comments on the buttery fauna of Porto Santo2018 15
Most of the used plant species are endemic or native;
therefore, they were already present about 500 years ago,
before the beginning of the strong human impact. Most
of the endemic or native plant species are common and
widely distributed on Porto Santo. The plant species with
the highest buttery visitation rates is Echium nervosum,
endemic to the Madeira Archipelago, and distributed in
dry microforests, open rocky and dry habitats. Echium
nervosum is also planted on roadsites. Also visited by
butteries is Echium portosanctensis, endemic to Porto
Santo, growing in open rocky habitats of the subhumid
zone and planted locally on roadsides in the northern
part. Hybrids of these Echium species also exist.
A higher number of visits was observed on Lotus
glaucus (endemic to Macaronesia) in sandy habitats and
on Cakile maritima in driftline habitats. Butteries were
also observed on Matthiola maderensis, endemic to the
Madeira Archipelago and growing on coastal rocks and
clis, which are partly inuenced by salt-spray.
Apart from these endemic plant species, there is
a second group comprising native species with a wider
distribution in the Mediterranean region or in North
Africa that grow on Porto Santo, e.g., on roadsides and
fallows: Asphodelus stulosus, Brassica nigra, Convolvulus
althaeoides, Galactites tomentosa, Leontodon taraxacoides,
Rapistrum rugosum, and Sonchus oleraceus.
The bipartite graph of the buttery-plant network
of Porto Santo (Fig. 2) shows the ower-visiting buttery
species on the left-hand side and the visited plant species
on the right-hand side, both connected by interaction
links. The buttery-plant network is asymmetric: seven
buttery species versus 15 plant species.
The butterfly-plant network shows that the most
abundant butterfly species on flowers is Vanessa cardui
with a preference for Echium nervosum. Vanessa atalanta
and the endemic V. vulcania also show a preference
for the Echium species (E. nervosum, endemic for the
Madeira Archipelago, and Echium portosanctensis,
endemic for Porto Santo, including hybrids). In contrast,
Colias croceus has a diversified flower-visiting behaviour
(observed on ten plant species of different plant
families).
Fig. 2 – Bipartite graph of the buttery-plant network of Porto Santo.

Boletim do Museu de História Natural do Funchal No. LXVIII, Art. 34916
CONCLUSIONS
Porto Santo, a relatively old and small volcanic island
in the Atlantic Ocean, is a model area for the colonisation
of islands by insect species of dierent areageographical
origins. Regarding bee species, Porto Santo served as
an important colonisation source for Madeira Island
(
KratochWil & schWaBe
, 2018).
Porto Santo has only two buttery species which
are endemic to archipelagos of Macaronesia or parts of
it: Vanessa vulcania, endemic to the Madeira Archipelago
and the Canary Islands, which can be considered a relict
species on Porto Santo, and, at a subspecies level, Lycaena
phlaes phlaeoides (endemic to the Madeira Archipelago).
In contrast, many other buttery species are endemic
to Madeira Island: Hipparchia maderensis (Bethune-
Baker, 1891), Pararge xiphia (Fabricius, 1775), Gonepteryx
maderensis Felder, 1862, and Pieris wollastoni (Butler, 1886).
These species do not occur on Porto Santo, because there
are no habitat conditions of higher mountain zones. The
endemic butteries of Madeira Island prefer vegetation
mosaics of forest sites with forest edges and forest
islands (Pararge xiphia and Gonepteryx maderensis) or live
generally at higher altitudes (Hipparchia maderensis, 800-
1,800 m a.s.l.; Pieris wollastoni, 400-1,110 m a.s.l.). Whether
these species ever existed on Porto Santo is a question
that cannot be answered.
A possible exchange of buttery individuals
between Madeira Island and Porto Santo is facilitated
by the small distance of 45 km between the islands (e.g.,
in the case of Danaus plexippus, Vanessa atalanta, and
V. cardui, but also of other buttery species, e.g., Colias
croceus, which are migratory and good yers).
The buttery-plant network of Porto Santo shows
asymmetry: Seven buttery species used 15 dierent
plant species, but not as marked as the bee-plant network
with six wild-bee species and 27 dierent plant species
(
KratochWil & schWaBe
, 2018).
Like the wild-bee species, the woody Echium
species E. nervosum (endemic to the Madeira Archipelago)
and E. portosanctensis (endemic to Porto Santo), including
hybrids, are key species as resources for Vanessa atalanta,
V. cardui and V. vulcania.
V. vulcania is not common. This endemic buttery
species was found only within one grid in the north of
the island at an altitude of 235 m a.s.l. Vanessa vulcania
is a remnant of an earlier time period of the island, when
(due to lack of human impact) areas with microforests
and Apollonias barbujana laurisilva vegetation still
existed. Vanessa vulcania show a similar preference for
the subhumid zone as the bumblebee Bombus terrestris
lusitanicus (
KratochWil & schWaBe
, 2018).
ACKNOWLEDGEMENTS
We thank cordially Ysabel Gonçalves, checking the
collections of the Museu de História Natural do Funchal
(Madeira, Portugal).
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NOMENCLATURE
The nomenclature for plant species names follows
JardiM & sequeira
(2008). One plant species was newly
described in the year 2010: Echium portosanctensis
carvalho
,
pontes
,
Batista-Marques & JardiM
, 2010; see
carvalho
et al., (2010).