A pictorial key for identification of the hoverflies

(Diptera: Syrphidae) of the Madeira Archipelago

Carla Rego‡, John Smit§, António Franquinho Aguiar|, Délia Cravo|, Andreia Penado‡, Mário Boieiro¶
‡ Centre for Ecology, Evolution and Environmental Changes, Azorean Biodiversity Group, Faculty of Sciences, University of
Lisbon, Lisboa, Portugal
§ European Invertebrate Survey - the Netherlands / Naturalis Biodiversity Center, Leiden, Netherlands
| Laboratório de Qualidade Agrícola, Secretaria Regional de Agricultura e Desenvolvimento Rural, Camacha, Madeira,
Portugal
¶ Centre for Ecology, Evolution and Environmental Changes, Azorean Biodiversity Group, University of Azores, Angra do
Heroísmo, Portugal

Corresponding author: Carla Rego ([email protected])

Academic editor: Torsten Dikow

Abstract

Background

Syrphid flies are important ecological indicators and provide crucial ecosystem services,
being important pollinators and biological control agents of insect pests. These charismatic
insects are conspicuous and, due to their size and colourful patterns, are relatively easy to
identify. However, the lack of user-friendly literature (e.g. photographic guides) for most
areas may hamper its wider selection as a study group in biodiversity and ecological
studies. The syrphid fauna of Madeira Archipelago comprises 26 species, including four
endemics (Eumerus hispidus Smit, Aguiar & Wakeham-Dawson, 2004; Melanostoma
wollastoni Wakeham-Dawson, Aguiar, Smit, McCullough & Wyatt, 2004; Myathropa usta,
Wollaston, 1858 and Xanthandrus babyssa, Walker, 1849), but, despite the current good
taxonomic knowledge on this group, information on species distribution, ecology and
conservation is still lacking. Here, we provide a pictorial key to the adult hoverflies of
Madeira Archipelago highlighting diagnostic characteristics and present photographs of
both males and females (in dorsal and lateral views) in colour plates. The key and plates
will help researchers to differentiate these species, thus encouraging the use of this insect
group in future bioindication studies. In addition, this study also aims to engage a broader
audience of non-experts in improving the knowledge on the distribution and ecology of
Madeira syrphids.

©
.
New information

We provide a checklist for the hoverflies of Madeira Archipelago and a pictorial key to help
on species identification.

Keywords
Flower flies, Macaronesia, Madeira endemics, photographic guide, species identification,
syrphids, taxonomic key

Introduction
Syrphids, commonly known as hoverflies or flower flies, belong to a large family of flies
(Diptera: Syrphidae) with over 6,000 known species (Rotheray 1993Evenhuis and Pape
2021). These flies are conspicuous and easy to distinguish from other insects due to their
colour patterns, morphology and flying behaviour (Ball and Morris 2015). Adult hoverflies
feed on honeydew, pollen and nectar and are amongst the most important flower visiting
insects in many ecosystems (Wackers et al. 2008, Inouye et al. 2015, Doyle et al. 2020).
Larvae, on the other hand, exploit a wide variety of food resources, including fungal fruiting
bodies (mycophagous), dung, tree sap, nests of social insects, decaying vegetation and
wood (saprophagous), whereas other larvae mine the leaves and stems of plants
(phytophagous) or predate other insects (zoophagous), including leafhoppers, coccids and
aphids (Rotheray 1993). For these reasons, hoverflies are considered important pollinators
and biological control agents of insect pests, providing crucial ecosystem services (
Ankersmit et al. 1986, Nelson et al. 2012, Dunn et al. 2020, Pekas et al. 2020). More
recently, the role of hoverflies as ecological indicators has also been stressed in many
studies since they are easy to sample and identify, their life cycle is well-known and the
larvae from different species have distinct environmental requirements for their
development (Sommaggio 1999, Sommaggio and Burgio 2014, Ball and Morris 2015,
Dunn et al. 2020).

The syrphid fauna of Madeira has been studied since the mid-nineteenth century by
several authors who contributed to a better understanding of species diversity and
distribution in this Archipelago (Walker 1849, Wollaston 1858, Loew 1860, Schiner 1868,
Thomson 1869, Bigot 1884, Osten-Sacken 1884, Becker 1908, Becker 1921, Frey 1939,
Frey 1949). Frey (1949) updated the syrphid fauna checklist and reported 21 species for
Madeira Archipelago. During this first century of reports on the Madeiran syrphid fauna, the
number of recorded species increased considerably, showing a similar pattern to most of
the other groups of terrestrial arthropods in the Archipelago (Borges et al. 2008). More
recently, other authors made significant contributions to the knowledge of this group of flies
in Madeira Archipelago (Gomes and Baez 1990, Barkemeyer 1999, Pita and Gomes 2003,
Smit et al. 2004, Wakeham-Dawson et al. 2004, Aguiar et al. 2005, Pita et al. 2009). For

2
instance, both Smit et al. (2004) and Wakeham-Dawson et al. 2004 described new
endemic species, clarified the identity of ambiguous taxa and provided a thorough revision
of Madeiran Syrphidae nomenclature. The current knowledge on the taxonomic diversity of
Madeiran syrphids was updated by Smit (2008) and it was included in a comprehensive
reference work on Madeira Archipelago biodiversity (Borges et al. 2008). Smit (2008) listed
26 species of hoverflies including four endemics (Eumerus hispidus Smit, Aguiar &
Wakeham-Dawson, 2004; Melanostoma wollastoni Wakeham-Dawson, Aguiar, Smit,
McCullough & Wyatt, 2004; Myathropa usta, Wollaston, 1858 and Xanthandrus babyssa,
Walker, 1849) and provided general information on species distribution in the Archipelago.
Despite the current good taxonomic knowledge on Madeiran syrphids, there is still a
significant gap regarding species distribution, ecology and conservation.

During the last decades, there has been a growing interest in biodiversity conservation by
the general public that has extended to several charismatic invertebrate groups, such as
dragonflies and butterflies. In oceanic islands, like Madeira, invertebrate conservation
needs to be fostered by engaging researchers, decision-makers and common citizens in
knowing, valuing, protecting and making public the unique diversity of life forms of these
ecosystems. This interplay is urgent since the biodiversity of oceanic islands worldwide is
under threat due to various factors (e.g. land-use change, invasive species, climate
change) and, jointly with significant declines in endemic species abundance, many human-
driven extinctions have been documented in these unique ecosystems, including in
Madeira Archipelago (Goodfriend et al. 1994, Gardiner 2003, Fontaine et al. 2007, Régnier
et al. 2009, Rando et al. 2012, Terzopoulou et al. 2015). Halting biodiversity loss in oceanic
islands is mandatory and, to accomplish this goal, a multidisciplinary strategy needs to be
implemented. This strategy should include monitoring programmes targeting specific
invertebrate groups and the use of expeditious, user-friendly and reliable techniques (
Borges et al. 2018).

Here, we aim to provide a user-friendly pictorial key for the identification of Madeira’s
hoverflies, a charismatic bioindicator and ecologically-important insect group. The key was
designed for use by non-experts and, altogether with the photos of male and female
specimens of all known species occurring in the Archipelago, aims to engage a diverse
audience in improving current knowledge on these conspicuous flies.

Materials and methods

Study area
Madeira Archipelago is located in the Atlantic Ocean, nearly 600 km from the African coast
(Morocco) and 450 km north from the Canary Islands, between latitudes 32°24′ and
33°07′N and longitudes 16°16′ and 17°16′W. The Archipelago is formed by three groups of
volcanic islands and islets: Madeira, Porto Santo and the Desertas Islands (Boieiro et al.
2015). Madeira is the largest island of the group (~ 740 km2) and is characterised by a
rugged topography with a steep coastline combined with deep ravines, high peaks and an
altitudinal plateau in the central part of the Island (the highest mountaintop is Pico Ruivo at

3
1862 m). Madeira harbours various habitats, from coastal dry areas to humid laurel forests
(Laurisilva) and heathland at higher altitudes, that support a diverse fauna and flora (
Menezes et al. 2011, Boieiro et al. 2013, Boieiro et al. 2015). Porto Santo, the second
largest island, lies ~ 40 km NE of Madeira, has several islets surrounding it, all included in
a network of protected areas (Alves et al. 2015). Natural vegetation cover was severely
destroyed by human activities following colonisation and, currently, is mostly composed of
herbaceous plants which are scarce in some areas. Further, in some mountaintops, pine
forests were planted to mitigate soil erosion effects. The Desertas Islands is a group of
three islands (Ilhéu Chão, Deserta Grande and Bugio) that include a Nature Reserve. Both
Bugio and Deserta Grande (the largest ones) are crest-like islands with steep slopes and
have large areas deprived of vegetation or are covered by herbaceous plants while Ilhéu
Chão, with a flat surface, has a well-preserved herbaceous vegetation cover (Menezes et
al. 2005).

Laboratory work
Specimens, both males and females, from all the known species reported to Madeira
Archipelago were studied under a stereomicroscope. Most of the specimen’s images were
taken with a Leica M125 motorised stereomicroscope, equipped with a IC80 HD digital
camera and LAS-Leica Application Suite 3.8 Software. For image stacking, we used the
LAS Module “Multifocus” and post-processed the images in ®Adobe Photoshop CC. We
also used a Canon 7D digital slr camera with a Canon EF 100 mm 2.8 L Is USM macro
lens to capture the habitus of some specimens. The study specimens are deposited in the
entomological collections of the Laboratório de Qualidade Agrícola (ICLAM) (Madeira,
Portugal) and Naturalis Biodiversity Center, Natural (RMNH) (Leiden, the Netherlands).

Identification keys

Key to the hoverflies of the Madeira Archipelago

The taxonomic key to the adult stages of hoverfly species of the Madeira Archipelago relies on
pictorial information to ease interpretation of characters and includes information on morphological
differences between males and females. Additionally, photos of male and female specimens of all
species (in dorsal and lateral views) are presented in colour plates. In the dichotomous key, the
couplet leads present one or more morphological characteristics indicated with symbols (e.g.
asterisks for one lead and Roman numerals for the other) which help to easily identify those
characteristics in the associated figures. The terminology of morphological characters used in this
key follows Thompson (1999). Abdominal tergites and sternites are abbreviated with a ‘t’ or ‘s’,
respectively.

Face entirely yellow (i); scutellum

1 always yellow, clearly lighter than 2
scutum (ii) (Fig. 1)

4
 Face dark or yellow with median
dark stripe, sometimes obscured
– by dense pollinosity (*); scutellum 15
never yellow, sometimes orange-
brown (**) (Fig. 1)

Very large species, over 18 mm,

with hornet-like appearance;
scutum with yellow markings on
anterior half and orange-red
2 colouration on posterior half; Milesia crabroniformis
metafemur with a small tooth
apicoventrally (i); wings with a
yellow tinge along the costa (ii)
(Fig. 2)

Smaller species, at most 15 mm,

never with a hornet-like
appearance; if yellow markings
present on scutum, then restricted
– to lateral margins and scutum 3
never with orange-red markings;
metafemur never with a small
tooth apicoventrally; wings without
yellow tinge along costa

Thoracic pleura with distinct yellow

3 markings (i); scutum with distinct 4
yellow bands laterally (ii) (Fig. 3)

Thoracic pleura without yellow

markings (*); scutum without
– 6
distinct yellow bands laterally (**)
(Fig. 3)

Abdomen distinctly margined (i);

4 male with tooth-like protuberance Ischiodon aegyptius
on metatrochanter (ii) (Fig. 4)

Abdomen not margined (*); male

– without tooth-like protuberance on 5
metatrochanter (Fig. 4)

5
 Scutum with yellow lateral band
restricted to the anterior part of
wing base (i); abdomen of males
5 Sphaerophoria rueppelli
about as long as wings when
folded; smaller species: 5-8 mm
(Fig. 5)

Scutum with yellow lateral band

uninterrupted, continuing
posteriorly of wing base (*);
– Sphaerophoria scripta
abdomen of males clearly longer
than wings when folded (**); larger
species: 7-12 mm (Fig. 5)

Abdomen more or less parallel-

6 sided, as broad as scutum (i) (Fig. 7
6)

Abdomen clearly broadening and

– oval shaped, clearly broader than 8
scutum (*) (Fig. 6)

Abdomen with ‘double bands’ on

7 Episyrphus balteatus
t3, t4 (i) (Fig. 7)

Abdomen without ‘double bands’,

with oblique yellow spots (*),
– Meliscaeva auricollis
sometimes connected to form
bands (Fig. 7)

Wing vein R4+5 in basal half of

cell r4+5 almost parallel to M,
curving upwards in apical part (i);
8 eyes pilose (ii); frons distinctly 9
swollen, more obvious in males
(iii); larger species: 10-15 mm (Fig.
8)

Wing vein R4+5 more or less

straight, converging from vein M
from the base of cell r4+5 (*); eyes
– 11
bare, except S. torvus (**); frons
not swollen (***); smaller species:
7-13 mm (Fig. 8)

6
 T3 and t4 with slender yellow or
white lunulate maculae, clearly
constricted in the middle
9 10
(i), yellow or white markings
covering less than half the length
of t3 (Fig. 9)

T3 and t4 with larger yellow

maculae, which are at most slightly
– constricted in the middle (+), Scaeva albomaculata
yellow markings covering more
than half the length of t3 (Fig. 9)

Abdominal spots yellow in live

specimens; spots on t3 with hind
10 edges curved, their outer and inner Scaeva selenitica
corners equally close to anterior
edge of the tergite (*) (Fig. 9)

Abdominal spots almost white in

live specimens; spots on t3 with
hind edges straight and oblique,
– Scaeva pyrastri
their outer corners distinctly further
removed from anterior edge of
tergite than inner corner (ii) (Fig. 9)

Scutum pollinose and dull (i);

abdomen with relatively slender
11 yellow bands on t3-t4 (ii); ventral 12
calypter with long erect pili on
dorsal surface (iii) (Fig. 10)

Scutum pollinose, but clearly

shining (*); abdomen typically with
yellow spots on t3-t4, sometimes
– connected to form bands 13
(**); ventral calypter lacking long
erect pili on dorsal surface (***)
(Fig. 10)

Eyes pilose, sparse and short in

females (i); wing cell BM entirely
12 Syrphus torvus
covered by microtrichia (ii) (Fig. 11
)

Eyes bare (*); wing cell BM basal

– Syrphus vitripennis
¼ bare (**) (Fig. 11)

7
 Face in frontal view at least as
broad as one eye (i); femora at
13 14
least partially black at the base (ii)
(Fig. 12)

Face in frontal view

clearly narrower than one eye (*);
femora entirely yellow (**);
– Eupeodes nuba
abdomen normally with broad
yellow maculae, sometimes
connected to form bands (Fig. 12)

Scutellum predominantly yellow

pilose (i); abdominal maculae
reaching lateral margins of
14 tergites, normally with spots, but Eupeodes corollae
frequently connected to form
bands (ii); male with larger
genitalia (iii) (Fig. 13)

Scutellum predominantly black

pilose (*); abdominal maculae not
reaching lateral margins of
– tergites, usually with spots, only Eupeodes luniger
rarely connected to form bands
(**); male with smaller genitalia
(***) (Fig. 13)

Wing vein R4+5 with a strong dip

in the cell below (i); larger species
(10-16 mm), sometimes with
15 16
metallic bronze luster, but often
with a bee-like appearance (Fig.
14)

Wing vein R4+5 without a strong

dip in the cell below (*); smaller
– 19
species (4-12 mm), never with a
bee-like appearance (Fig. 14)

Eyes spotted (i). Entire body

16 largely with metallic bronze luster Eristalinus aeneus
(Fig. 15)

Eyes never spotted, but either

– striped or concolorous. Body 17
without bronze luster

8
17 Eyes striped (*) (Fig. 15) Eristalinus taeniops

Eyes concolorous, without stripes

– 18
(+) (Fig. 15)

Eyes with bands of pili (i); wing cell

18 Eristalis tenax
R1 closed (ii) (Fig. 16)

Eyes without bands of pili; wing

– Myathropa usta
cell R1 open (**) (Fig. 16)

Face in profile with a facial

19 20
tubercule (i) (Fig. 17)

Face in profile more or less

straight (*), sometimes mouth-
– 23
edge clearly protruding (**) (Fig.
17)

Face entirely black (i); abdomen

either entirely dark or with orange-
20 21
yellow spots; larger species: 5-12
mm (Fig. 18)

Face creamy yellow with a black

facial stripe (*); abdomen black or
– Paragus mundus
partially red; very small species:
4-6 mm (Fig. 18)

Abdomen oval, clearly broader

than scutum (i); female with
abdomen entirely black or with
21 very small, rounded spots (i); male Xanthandrus babyssa
with broad orange-yellow spots on
t3 and t4 (ii), which are sometimes
connected (Fig. 19)

Abdomen slender and parallel-

– sided, as broad as scutum (*) (Fig. 22
19)

Abdomen entirely black in males

(**), at most with reduced orange
22 Melanostoma wollastoni
markings in the female (*); larger
species: 7-10 mm (Fig. 19)

9
 Abdomen with clear orange
markings, triangular on t3 and t4 in
– females (+), rectangular in males Melanostoma mellinum
(++); smaller species: 5-8 mm (Fig.
19)

Abdomen clearly petiolate, t2

23 constricted in basal half (i); smaller Neoascia podagrica
species: 5-6 mm (Fig. 20)

Abdomen never that clearly

petiolate, t2 never constricted in
– basal half, abdomen more or less 24
parallel-sided. Larger species:
6-13 mm (Fig. 20)

Abdomen with a broad orange

24 band (*); larger species: 10-13 mm Xylota segnis
(Fig. 20)

Abdomen without orange; smaller

– 25
species 6-10 mm (+) (Fig. 20)

Thoracic pleura not dusted (i) and

scutum with a pair of longitudinal
pollinose vittae, almost reaching
25 Eumerus hispidus
scutellum (ii); legs entirely black
(iii); abdomen with pollinose spots
on t2-t4 (iiii) (Fig. 21)

Thoracic pleura heavily dusted,

continuing on the frontal half of the
lateral side of scutum (*); legs
– bicoloured (**); abdomen with Syritta pipiens
yellow spots on the lateral sides of
t2 and t3, t4 with dusted
areas (Fig. 21)

Analysis
The hoverfly species of Madeira Archipelago
The syrphid fauna of Madeira Archipelago comprises 26 species, all considered to be
native to these islands (Table 1). Four of them are endemics (Eumerus hispidus,
Melanostoma wollastoni, Myathropa usta and Xanthandrus babyssa) which occur mostly in
native laurel forests (Laurisilva) and altitudinal heathlands in Madeira Island (except for E.

10
hispidus which occurs in coastal and drier habitats of different islands) (Smit et al. 2004,
Smit 2008). There are significant differences on the syrphid fauna between island groups
since 25 species were recorded in Madeira, 14 in Porto Santo and seven in Desertas
Islands. Differences in island area, altitude and habitat heterogeneity are the main drivers
of species richness differences between islands, with some species being restricted to
native forest areas in Madeira Island. We believe that the syrphid species inventory of
Madeira Archipelago is near complete, but additional sampling should be carried out in
Porto Santo and Desertas since these smaller islands were less sampled than Madeira
proper. Furthermore, taking in consideration the increase in trade and tourism to Madeira in
recent decades, it is expected that new species will arrive at the Archipelago; therefore, it is
important to implement a monitoring scheme for early detection of introduced species in
the short term.

Acknowledgements
We acknowledge the financial support from Fundação para a Ciência e a Tecnologia (FCT,
Portugal) through project PTDC/BIA-BIC/1013/2014, UIDB/00329/2020-2024 and contract
DL57/2016/CP1375/CT0001 to MB. We also thank Instituto das Florestas e da
Conservação da Natureza (IFCN) for allowing the collection of specimens in Madeira
Archipelago.

References

• Aguiar AM, Smit JT, Wakeham-Dawson A (2005) Eristalinus taeniops (Wiedemann,

1818) (Diptera: Syrphidae) new to Madeira Island, Portugal. Dipterists Digest 12: 20‑21.
• Alves F, Rodrigues J, Montes R, Menezes D, Oliveira P, Silva V (2015) Islets of Porto
Santo: a treasure to be preserved. Serviço do Parque Natural da Madeira, 98 pp.
• Ankersmit GW, Dijkman H, Keuning NJ, Mertens H, Sins A, Tacoma H (1986)
Episyrphus balteatus as a predator of the aphid Sitobion avenae on winter wheat.
Entomologia Experimentalis et Applicata 42: 271‑277. https://doi.org/10.1111/j.
1570-7458.1986.tb01032.x
• Ball S, Morris R (2015) Britain’s Hoverflies. Princeton University Press, Princeton, 312
pp. https://doi.org/10.1515/9781400866021
• Barkemeyer W (1999) Zur Kenntnis der Syrphidenfauna des Madeira-Archipels.
TenDenZen Supplement: 115‑134.
• Becker T (1908) Dipteren der Insel Madeira. Mitteilungen aus dem Zoologischen
Museum in Berlin 4: 181‑206.
• Becker T (1921) Neue Dipteren Meiner Sammlung. Mitteilungen aus dem Zoologischen
Museum in Berlin 10: 1‑93.
• Bigot JM (1884) Diptères nouveaux ou peu connus. 24e partie. Syrphidi (2e partie).
Annales de la Société Entomologique de France 32: 73‑116.
• Boieiro M, Aguiar AF, Aguiar CA, Borges PA, Cardoso P, Crespo L, Menezes D, Pereira
P, Rego C, Silva I, Silva PM, Serrano AR (2013) Madeira, the biodiversity pearl - valuing

11
 the native habitats and endemic life forms. Sociedade Portuguesa de Entomologia 80
pp.
• Boieiro M, Aguiar A, Rego C, Borges P, Serrano AR (2015) The biodiversity of terrestrial
arthropods in Madeira and Selvagens archipelagos. Revista Ibero Diversidad
Entomológica @ccesible 6B: 1‑20.
• Borges PA, Cardoso P, Kreft H, Whittaker RJ, Fattorini S, Emerson BC, Gil A, Gillespie
RG, Matthews TJ, Santos AM, Steinbauer MJ, Thebaud C, Ah-Peng C, Amorim IR,
Aranda SC, Arroz AM, Azevedo JM, Boieiro M, Borda-de-Agua L, Carvalho JC, Elias
RB, Fernandez-Palacios JM, Florencio M, Gonzalez-Mancebo JM, Heaney LR, Hortal J,
Kueffer C, Lequette B, Martin-Esquivel JL, Lopez H, Lamelas-Lopez L, Marcelino J,
Nunes R, Oromi P, Patino J, Perez AJ, Rego C, Ribeiro SP, Rigal F, Rodrigues P,
Rominger AJ, Santos-Reis M, Schaefer H, Sergio C, Serrano AR, Sim-Sim M,
Stephenson PJ, Soares AO, Strasberg D, Vanderporten A, Vieira V, Gabriel R (2018) A
Global Island Monitoring Scheme (GIMS) for the long-term coordinated survey and
monitoring of forest biota across islands. Biodiversity and Conservation 27: 2567‑2586.
https://doi.org/10.1007/s10531-018-1553-7
• Borges PAV, Abreu C, Aguiar AMF, Carvalho P, Jardim R, Melo I, Oliveira P, Sérgio C,
Serrano ARM, Vieira P (Eds) (2008) A list of the terrestrial fungi, flora and fauna of
Madeira and Selvagens archipelagos. Direcção Regional do Ambiente da Madeira and
Universidade dos Açores, Funchal and Angra do Heroísmo. [ISBN 978-989-95790-0-2]
• Doyle T, Hawkes WLS, Massy R, Powney GD, Menz MHM, KR W (2020) Pollination by
hoverflies in the Anthropocene. Proceedings of the Royal Society of London B 287
(20200508). https://doi.org/10.1098/rspb.2020.0508
• Dunn L, Lequerica M, Reid CR, Latty T (2020) Dual ecosystem services of syrphid flies
(Diptera: Syrphidae): pollinators and biological control agents. Pest Management
Science 76: 1973‑1979. https://doi.org/10.1002/ps.5807
• Evenhuis NL, Pape T (Eds) (2021) Systema Dipterorum, version 3.5. http://diptera.org/.
Accessed on: 2022-1-05.
• Fontaine B, Bouchet P, Van Achterberg K, Alonso-Zarazaga MA, et al. (2007) The
European union’s 2010 target: Putting rare species in focus. Biological Conservation
139: 167‑185. https://doi.org/10.1016/j.biocon.2007.06.012
• Frey R (1939) Die Arthropodenfauna von Madeira nach den Ergebnissen der Reise von
Prof. Dr. O. Lundblad Juli-August 1935. XIX. Diptera Brachycera. Arkiv för Zoologi 31A
(20): 1‑18.
• Frey R (1949) Die Dipterenfauna der Inseln Madeira. Commentationes Biologicae 8
(16): 1‑47.
• Gardiner BO (2003) The possible cause of extinction of Pieris brassicae wollastoni
Butler (Lepidoptera: Pieridae). Entomologist's Gazette 54: 267‑268.
• Gomes A, Baez M (1990) Contribución al conocimiento de los sírfidos del archipiélago
de Madeira (Diptera, Syrphidae). Vieraea 19: 339‑345.
• Goodfriend GA, Cameron RA, Cook LM (1994) Fossil evidence of recent human impact
on the snail fauna of Madeira. Journal of Biogeography 21: 309‑320. https://doi.org/
10.2307/2845532
• Inouye DW, Larson BM, Ssymank A, Kevan PG (2015) Flies and flowers III: Ecology of
foraging and pollination. Journal of Pollination Ecology 16 (16): 115‑133. https://doi.org/
10.26786/1920-7603

12
• Loew H (1860) Die Dipteren-fauna Südafrika’s. Erste Abtheilung Abhandlungen des
Naturwissenschaftlichen Vereins für Sachsen und Thüringen in Halle (1858-1861) 2:
57‑402.
• Menezes D, Freitas I, Gouveia L, Oliveira P, Pires R, S F (2005) The Desertas islands.
Serviço do Parque Natural da Madeira, Funchal, Portugal, 94 pp.
• Menezes D, Freitas I, Domingues M, Oliveira P (2011) Madeira-Natural paradise.
Serviço do Parque Natural da Madeira, 127 pp.
• Nelson EH, Hogg BN, Mills NJ, Daane KM (2012) Syrphid flies suppress lettuce aphids.
BioControl 57: 819‑826. https://doi.org/10.1007/s10526-012-9457-z
• Osten-Sacken CR (1884) List of the Diptera of the Island of Madeira, so far as they are
mentioned in entomological literature. Entomologist’s monthly Magazine 21: 32‑34.
https://doi.org/10.5962/bhl.part.22005
• Pekas A, Craecker ID, Boonen S, Wackers FL, Moerkens R (2020) One stone; two
birds: concurrent pest control and pollination services provided by aphidophagous
hoverflies. Biological Control 149: 104328. https://doi.org/10.1016/j.biocontrol.
2020.104328
• Pita MT, Gomes A (2003) Notes on the syrphid fauna of Madeira Archipelago and the
Salvage Islands (Diptera, Syrphidae). Vieraea 31: 33‑38.
• Pita MT, Silva I, Menezes D, Gomes A (2009) On the syrphid fauna of the Madeira
Archipelago and the Salvage Islands, with some first records from Deserta Grande and
Selvagem Grande (Diptera: Syrphidae). Boletín Sociedad Entomológica Aragonesa 44:
425‑433.
• Rando JC, Pieper H, Alcover JA, Olson SL (2012) A new species of extinct fossil scops
owl (Aves: Strigiformes: Strigidae: Otus) from the Archipelago of Madeira (North Atlantic
Ocean). Zootaxa 3182: 29‑42. https://doi.org/10.11646/zootaxa.3182.1.3
• Régnier C, Fontaine B, Bouchet P (2009) Not knowing, not recording, not listing:
numerous unnoticed mollusk extinctions. Conservation Biology 23: 1214‑1221. https://
doi.org/10.1111/j.1523-1739.2009.01245.x
• Rotheray GE (1993) Colour guide to hoverfly larvae (Diptera, Syrphidae). Dipterists
Digest 9: 1‑156.
• Schiner (1868) Diptera. Reise der österreichischen Fregatte Novara um die Erde in den
Jahren 1857, 1858, 1859. Zoologischer Theil 2 (1): 1‑338.
• Smit J (2008) Syrphidae. In: Borges PAV, Abreu C, Aguiar AMF, Carvalho P, Jardim R,
Melo I, Oliveira P, Sérgio C, Serrano ARM, P V (Eds) A list of the terrestrial fungi, flora
and fauna of Madeira and Selvagens archipelagos. Direcção Regional do Ambiente da
Madeira and Universidade dos Açores, Funchal and Angra do Heroísmo,, 337–338. pp.
• Smit JT, Aguiar AM, Wakeham-Dawson A (2004) The hoverflies (Diptera, Syrphidae) of
the Madeiran Archipelago, Portugal. Dipterists Digest 11: 47‑82.
• Sommaggio D (1999) Syrphidae: can they be used as environmental bioindicators?
Agriculture, Ecosystems and Environment 74: 343‑356. https://doi.org/10.1016/
S0167-8809(99)00042-0
• Sommaggio D, Burgio G (2014) The use of Syrphidae as functional bioindicator to
compare vineyards with different managements. Bulletin of Insectology 67: 147‑156.
• Terzopoulou S, Rigal F, Whittaker RJ, Borges PA, Triantis KA (2015) Drivers of
extinction: the case of Azorean beetles. Biology Letters 11: 1‑4. https://doi.org/10.1098/
rsbl.2015.0273

13
• Thompson FC (1999) A key to the genera of the flower flies (Diptera: Syrphidae) of the
neotropical region including descriptions of new genera and species and a glossary of
taxonomic terms. Contributions on Entomology International 3: 319‑378.
• Thomson CG (1869) 6. Diptera, Species nova descripsit. Kongliga Svenska Fregatten
Eugenies Resa Omkring Jorden 2 (1): 443‑614.
• Wackers F, Rijn PCJ, GE H (2008) Honeydew as a food source for natural enemies:
making the best of a bad meal? Biological Control 45: 176‑184. https://doi.org/10.1016/
j.biocontrol.2008.01.007
• Wakeham-Dawson A, Aguiar AM, Smit J, Mccullough A, Wyatt N (2004) Melanostoma
wollastoni sp. n. (Diptera, Syrphidae) from Madeira, Portugal. Dipterists Digest 10:
89‑94.
• Walker F (1849) List of the specimens of the dipterous insects in the collection of the
British Museum, part 3. The British Museum (Natural History), London. 485‑687.
• Wollaston TV (1858) Brief diagnostic characters of undescribed Madeiran Insects.
Annals and Magazine of Natural History 1: 113‑125. https://doi.org/
10.1080/00222935808696882

14
Figure 1.
Differences in face and scutellum colouration patterns.

15
Figure 2.
Small apicoventral tooth in metafemur and yellow tinge along the costa.

16
Figure 3.
Differences in scutum and thoracic pleura colouration.

17
Figure 4.
Differences in abdomen margin and metatrochanter protuberance.

18
Figure 5.
Differences in scutum colouration and abdomen morphology.

19
Figure 6.
Differences in abdomen morphology.

20
Figure 7.
Differences in abdomen colouration patterns.

21
Figure 8.
Differences in eye pilosity, frons and vein R4+5 morphology.

22
Figure 9.
Differences in colouration pattern of t3 and t4.

23
Figure 10.
Differences in scutum shine, colouration pattern of t3-t4 and in the presence of long erect pili
on dorsal surface of ventral calypter.

24
Figure 11.
Differences in eye and wing cell BM pilosity.

25
Figure 12.
Differences in face morphology and femora colour.

26
Figure 13.
Differences in scutellum pilosity, abdominal colouration pattern and male genitalia.

27
Figure 14.
Differences in wing vein R4+5 morphology.

28
Figure 15.
Differences in eye colour patterns.

29
Figure 16.
Differences in eye pili bands and wing cell R1 morphology.

30
Figure 17.
Differences in face profile.

31
Figure 18.
Differences in face colour.

32
Figure 19.
Differences in abdomen morphology and colour patterns.

33
Figure 20.
Differences in abdomen morphology and colouration.

34
Figure 21.
Differences in leg colouration and body pollinosity.

35
a b

c d

Figure 22.
Episyrphus balteatus
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

36
a b

c d

Figure 23.
Eristalinus aeneus
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

37
a b

c d

Figure 24.
Eristalinus taeniops
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

38
a b

c d

Figure 25.
Eristalis tenax
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

39
a b

c d

Figure 26.
Eumerus hispidus
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

40
a b

c d

Figure 27.
Eupeodes corollae
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

41
a b

c d

Figure 28.
Eupeodes luniger
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

42
a b

c d

Figure 29.
Eupeodes nuba
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

43
a b

c d

Figure 30.
Ischiodon aegyptius
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

44
a b

c d

Figure 31.
Melanostoma mellinum
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

45
a b

c d

Figure 32.
Melanostoma wollastoni
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

46
a b

c d

Figure 33.
Meliscaeva auricollis
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

47
a b

c d

Figure 34.
Milesia crabroniformis
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

48
a b

c d

Figure 35.
Myathropa usta
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

49
a b

c d

Figure 36.
Neoascia podagrica
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

50
a b

c d

Figure 37.
Paragus mundus
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

51
a b

c d

Figure 38.
Scaeva albomaculata
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

52
a b

c d

Figure 39.
Scaeva pyrastri
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

53
a b

c d

Figure 40.
Scaeva selenitica
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

54
a b

c d

Figure 41.
Sphaerophoria rueppellii
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

55
a b

c d

Figure 42.
Sphaerophoria scripta
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

56
a b

c d

Figure 43.
Syritta pipiens
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

57
a b

c d

Figure 44.
Syrphus torvus
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

58
a b

c d

Figure 45.
Syrphus vitripennis
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

59
a b

c d

Figure 46.
Xanthandrus babyssa
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

60
a b

c d

Figure 47.
Xylota segnis
a: male in dorsal view
b: male in profile
c: female in dorsal view
d: female in profile

61
Table 1.
List of the Syrphidae species from Madeira, their distribution in the Archipelago (M – Madeira
Island, PS – Porto Santo Island and surrounding islets, D – Desertas Islands) and representative
photos of adult males and females (in dorsal and lateral views).

Species M PS D Photos

Episyrphus balteatus (De Geer, 1776) ● ● ● Fig. 22

Eristalinus aeneus (Scopoli, 1763) ● ● Fig. 23

Eristalinus taeniops (Wiedemann, 1818) ● Fig. 24

Eristalis tenax (Linnaeus, 1758) ● ● ● Fig. 25

Eumerus hispidus Smit et al., 2004 ● ● Fig. 26

Eupeodes corollae (Fabricius, 1794) ● ● ● Fig. 27

Eupeodes luniger (Meigen, 1822) ● ● ● Fig. 28

Eupeodes nuba (Wiedemann, 1830) ● Fig. 29

Ischiodon aegyptius (Wiedemann, 1830) ● ● ● Fig. 30

Melanostoma mellinum (Linnaeus, 1758) ● ● Fig. 31

Melanostoma wollastoni Wakeham-Dawson et al., 2004 ● Fig. 32

Meliscaeva auricollis (Meigen, 1822) ● Fig. 33

Milesia crabroniformis (Fabricius, 1775) ● Fig. 34

Myathropa usta (Wollaston, 1858) ● Fig. 35

Neoascia podagrica (Fabricius, 1775) ● Fig. 36

Paragus mundus Wollaston, 1858 ● ● Fig. 37

Scaeva albomaculata (Macquart, 1842) ● ● Fig. 38

Scaeva pyrastri (Linnaeus, 1758) ● ● ● Fig. 39

Scaeva selenitica (Meigen, 1822) ● Fig. 40

Sphaerophoria rueppellii (Wiedemann, 1830) ● ● Fig. 41

Sphaerophoria scripta (Linnaeus, 1758) ● ● Fig. 42

Syritta pipiens (Linnaeus, 1758) ● ● Fig. 43

Syrphus torvus Osten-Sacken, 1875 ● Fig. 44

Syrphus vitripennis Meigen, 1822 ● Fig. 45

Xanthandrus babyssa (Walker, 1849) ● Fig. 46

Xylota segnis (Linnaeus, 1758) ● Fig. 47

62