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New data on Noctuoidea from the Canary Islands, Spain (Lepidoptera: Noctuoidea)

Authors:

Abstract

We describe three new species: Zebeeba orzolae Falck & Karsholt, sp. n. (Erebidae: Rivulinae), Nyctobrya vilfredi Falck & Karsholt, sp. n. (Noctuidae: Bryophilinae) and Agrotis corralejoi Falck & Karsholt, sp. n. (Noctuidae: Noctuinae). The unknown female of Mniotype loslobensis (Fischer, Saldaitis & Ivinskis, 2007) (Noctuidae: Hadeninae) and its genitalia are described and illustrated. Twelve species are recorded as new to the fauna of the Canary Islands: Epharmottomena eremophila (Rebel, 1895), Eublemma thermobasis Hampson, 1910, Eublemma candidana (Fabricius, 1794), Araeopteron ecphaea (Hampson, 1914), Gnamptonyx innexa (Walker, 1858) and Rhabdophera acrosticta (Püngeler, 1904) (Erebidae), Nycteola columbana (Turner, 1925) (Nolidae), Amyna axis Guenée, 1852, Polymixis aurora commixta (Rungs, 1943), Nonagria typhae (Thunberg, 1784), Mythimna languida (Walker, 1858) and Leucania zeae (Duponchel, 1828) (Noctuidae). Five species are, because of different errors, removed from the list of Lepidoptera found in the Canary Islands: Schrankia taenialis (Hübner, [1809]), Cosmia affinis (Linnaeus, 1767), Hecatera sancta (Staudinger, 1859), Agrotis boeticus (Boisduval, 1837) and A. aistleitneri Behounek & Speidel, 2009. The genus Gerarctia Hampson, 1905 is, based on personal information from H. Hacker, synonymised with Zebeeba Kirby, 1892, syn. n., and Gerarctia poliotis Hampson, 1905 is transferred to Zebeeba as Z. poliotis (Hampson, 1905), comb. n. DNA barcodes of Caradrina rebeli Staudinger, 1901 from different Canary Islands are compared, and it is concluded that the species occurs in two subspecies: C. rebeli rebeli Staudinger, 1901 and C. rebeli lanzarotensis Pinker, 1962.
145
New data on Noctuoidea from the Canary Islands, Spain
(Lepidoptera: Noctuoidea)
P. Falck & O. Karsholt
Abstract
We describe three new species: Zebeeba orzolae Falck & Karsholt, sp. n. (Erebidae: Rivulinae), Nyctobrya
vilfredi Falck & Karsholt, sp. n. (Noctuidae: Bryophilinae) and Agrotis corralejoi Falck & Karsholt, sp. n.
(Noctuidae: Noctuinae). The unknown female of Mniotype loslobensis (Fischer, Saldaitis & Ivinskis, 2007)
(Noctuidae: Hadeninae) and its genitalia are described and illustrated. Twelve species are recorded as new to the
fauna of the Canary Islands: Epharmottomena eremophila (Rebel, 1895), Eublemma thermobasis Hampson, 1910,
Eublemma candidana (Fabricius, 1794), Araeopteron ecphaea (Hampson, 1914), Gnamptonyx innexa (Walker,
1858) and Rhabdophera acrosticta (Püngeler, 1904) (Erebidae), Nycteola columbana (Turner, 1925) (Nolidae),
Amyna axis Guenée, 1852, Polymixis aurora commixta (Rungs, 1943), Nonagria typhae (Thunberg, 1784),
Mythimna languida (Walker, 1858) and Leucania zeae (Duponchel, 1828) (Noctuidae). Five species are, because of
different errors, removed from the list of Lepidoptera found in the Canary Islands: Schrankia taenialis (Hübner,
[1809]), Cosmia affinis (Linnaeus, 1767), Hecatera sancta (Staudinger, 1859), Agrotis boeticus (Boisduval, 1837)
and A. aistleitneri Behounek & Speidel, 2009. The genus Gerarctia Hampson, 1905 is, based on personal
information from H. Hacker, synonymised with Zebeeba Kirby, 1892, syn. n., and Gerarctia poliotis Hampson, 1905
is transferred to Zebeeba as Z. poliotis (Hampson, 1905), comb. n. DNA barcodes of Caradrina rebeli Staudinger,
1901 from different Canary Islands are compared, and it is concluded that the species occurs in two subspecies: C.
rebeli rebeli Staudinger, 1901 and C. rebeli lanzarotensis Pinker, 1962.
KEY WORDS: Lepidoptera, Erebidae, Nolidae, Noctuidae, new species, DNA barcodes, Canary Islands, Spain.
Nuevos datos sobre Noctuoidea de las Islas Canarias, España
(Lepidoptera: Noctuoidea)
Resumen
Describimos tres nuevas especies: Zebeeba orzolae Falck & Karsholt, sp. n. (Erebidae: Rivulinae), Nyctobrya
vilfredi Falck & Karsholt, sp. n. (Noctuidae: Bryophilinae) and Agrotis corralejoi Falck & Karsholt, sp. n.
(Noctuidae: Noctuinae). Se describe e ilustra la desconocida hembra y genitalia de Mniotype loslobensis (Fischer,
Saldaitis & Ivinskis, 2007) (Noctuidae: Hadeninae). Doce especies se registran como nuevas para las Islas Canarias:
Epharmottomena eremophila (Rebel, 1895), Eublemma thermobasis Hampson, 1910, Eublemma candidana
(Fabricius, 1794), Araeopteron ecphaea (Hampson, 1914), Gnamptonyx innexa (Walker, 1858) and Rhabdophera
acrosticta (Püngeler, 1904) (Erebidae), Nycteola columbana (Turner, 1925) (Nolidae), Amyna axis Guenée, 1852,
Polymixis aurora commixta (Rungs, 1943), Nonagria typhae (Thunberg, 1784), Mythimna languida (Walker, 1858)
y Leucania zeae (Duponchel, 1828) (Noctuidae). Cinco especies son, debido a diferentes errores, removidas de la
lista de Lepidoptera encontradas en las Islas Canarias: Schrankia taenialis (Hübner, [1809]), Cosmia affinis
(Linnaeus, 1767), Hecatera sancta (Staudinger, 1859), Agrotis boeticus (Boisduval, 1837) y A. aistleitneri
Behounek & Speidel, 2009. El género Gerarctia Hampson, 1905 es, basado sobre información personal de H.
Hacker, sinonimizado con Zebeeba Kirby, 1892, syn. n. y Gerarctia poliotis Hampson, 1905 es transferido a
Zebeeba como Z. poliotis (Hampson, 1905), comb. n. Se compara el ADN código de barras de Caradrina rebeli
SHILAP Revta. lepid., 50 (197) marzo 2022: 145-165 eISSN: 2340-4078 ISSN: 0300-5267
1901 de diferentes Islas Canarias y se concluye que la especie ocurre en dos subespecies: C. rebeli rebeli Staudinger,
1901 y C. rebeli lanzarotensis Pinker, 1962.
PALABRAS CLAVE: Lepidoptera, Erebidae, Nolidae, Noctuidae, nuevas especies, ADN código de barras, Islas
Canarias, España.
Introduction
Fieldwork undertaken in the Canary Islands since 2016 by the first author has revealed a number of
new and partly undescribed species of Lepidoptera for these islands. It may not be surprising that several
species of smaller, so-called Microlepidoptera have been overlooked by previous researchers, but an
increase of 15 % of the relatively large Pyraloidea species (FALCK et al., 2019) shows that the
Lepidoptera fauna of the Canary Islands is probably not as well investigated as one might have expected.
The Noctuoidea is a mega-diverse group of larger moths. During the last decade, family level
classification has undergone a number of changes (ZAHIRI et al., 2010, 2012, 2013).In the most
recent list to include Canary Island Lepidoptera VIVES MORENO (2014) listed 2 species of
Notodontidae, 2 Nolidae, 29 Erebidae, 1 Euteliidae and 99 Noctuidae. Here we describe 1 new
Erebidae and 2 new Noctuidae. Moreover, we record 6 species of Erebidae, 1 Nolidae and 5 Noctuidae
as new to the fauna of the Canary Islands, and we suggest that five species are removed from the list of
Canary Islands, for various reasons.
Material and methods
Most of the specimens were attracted to an 8 watts super actinic light. Genitalia were dissected
following ROBINSON (1976). Whole specimens were photographed with a Canon EOS 700D camera
equipped with a Canon EF 100 mm objective. The genitalia slides were photographed using a Soptop
CX40T Trinocular microscope in conjunction with a Toup Tek P10500A-E3 / E3ISPM05000KPA-E3 /
5.0MP USB3 camera.
DNA samples were prepared from dried legs according to the prescribed standards and processed
at the Canadian Centre for DNA Barcoding (CCDB, Biodiversity Institute of Ontario, University of
Guelph) to obtain the 658 base-pair long barcode fragment of the mitochondrial COI gene (cytochrome
c oxidase I). Intra- and interspecific distances of DNA barcode fragment were calculated using analytic
tools of BOLD with the Kimura 2-parameter model of nucleotide substitution. Neighbour-joining trees
were constructed using MEGA 6 (TAMURA et al., 2013) under the Kimura 2-parameter model for
nucleotide substitutions with the closest European species in BOLD as outgroups. Genetic clusters are
presented with their barcode index number (BIN; cf. RATMNASINGHAM & HERBERT, 2013).
We examined the morphology from all species and the DNA Barcode from new and cryptic species.
Abbreviations used
GP Genitalia preparation
PF Collection of Per Falck, Neksø, Denmark
MNCN Collection of Antonio Vives, Museo Nacional de Ciencias Naturales, Madrid, Spain
NHMUK Natural History Museum, London, United Kingdom
ZMUC Zoological Museum, Natural History Museum of Denmark, Copenhagen, Denmark
Results
EREBIDAE
RIVULINAE
The genus Zebeeba Kirby, 1892 = Gerarctia Hampson, 1905, syn. n., is originally a South African
P. FALCK & O. KARSHOLT
146 SHILAP Revta. lepid., 50 (197) marzo 2022
genus and includes, besides the Mediterranean species Z. falsalis (Herrich-Schäffer, 1839), several East
African and a number of South African species, several of which are still undescribed (HACKER,
2021: 80 and pers.com. 15-II-2021). Until now, about 20 species have been described.
Zebeeba poliotis (Hampson, 1905), comb. n.
Ingura poliotis Hampson, 1905. Ann. Mag. Nat. Hist., (7) 15(89): 449
Zebeeba orzolae Falck & Karsholt, sp. n. (Fig. 1)
Holotype 1: SPAIN, LANZAROTE, Mojón Blanco, Orzolá, 20 m, 21-X-10-XI-2019, leg. P. Falck,
genitalia slide 3473PF, DNA sample Lepid Phyl 0601PF/CILEP0600-20 (PF).
Description: Adult. Wingspan 14.5 mm. Labial palp porrect, segment 2 whitish grey with grey
scale tuft, segment 3 grey, very short. Antenna pale grey, with indistinct grey rings, very shortly ciliate.
Head and neck pale grey mottled with darker grey. Tegula and thorax grey. Forewing ground colour
whitish grey mottled with grey, at dorsum before tornus an indistinct large reddish-brown spot,
reniform stigma creamy white bordered distally by dark grey, postmedian fascia indistinct dark grey not
reaching dorsum, apical streak indistinct, dark grey. Hindwing grey, paler towards base. Fringe
yellowish white.
Male genitalia (Figs 29, 29a): Uncus long, relatively broad, pointed. Valva short, narrow and
knob-shaped. Saccus very broad, anterior margin concave. Phallus large, longer than valva, distally
slightly bent.
Female genitalia: Unknown.
Diagnosis: Z. orzolae resembles Z. poliotis (Fig. 2), the only other Zebeeba species known from
the Canary Islands (Spain). Males can be distinguished by the shortly ciliate antenna, bipectinate in Z.
poliotis, otherwise it can be distinguished by the whitish coloured forewing without wing pattern in the
inner half and the smaller wingspan. Z. orzolae also resembles some South African species e. g. Z.
mediorufa (Hampson, 1910) and Z. fuscipars (Hampson, 1910) (HACKER, 2021: 601). It can be
distinguished by the lack of wing pattern in the inner half of forewing. In the genitalia the very short,
narrow and knob-shaped valva and the broad concave saccus are characteristic. The genitalia of Z.
poliotis are figured by PINKER (1965: pls. 23).
Molecular diagnosis: We obtained DNA barcode fragments of 601 bp. Barcode Index Number
(BIN) BOLD: AEE6226.
Biology: Early stages unknown, but the larva probably feeds on Asparagaceae (HACKER, 2021:
80). The specimen was attracted to an 8 watts super actinic light in a sandy area near the coast.
Distribution: Known only with certainty from the northern part of the island of Lanzarote, but
probably also from Fuerteventura - see remarks. The species is probably endemic to the Canary Islands
(Spain).
Etymology: The species is named after the small town Orzolá placed nearby the type-locality.
Remarks: HACKER & SCHMITZ (1996: 180) mention a Noctuidae species from Fuerteventura
“Auf dieser Insel konnte jedoch eine kleinere, Phycitinae-ähnliche Noktuide nachgewiesen werden, die
habituell poliotis ähnelt, jedoch einen anderen Fühler- und Genitalbau aufweist und vermutlich eine
unbeschriebene Art darstellt”, the specimen is also figured (Tafel M: 1). It has not been possible to
study this specimen, but it most likely belongs to Z. orzolae.
HYPENODINAE
Schrankia taenialis (Hübner, [1809])
Pyralis taenialis Hübner, [1809]. Samml. Eur. Schmett.: pl. 23, fig. 151
The species is mentioned by REBEL & ROGENHOFER (1894: 67) from The Canary Islands (La
Palma) on the basis of two specimens “Zwei geflogene, auffallend kleine (Exp. 14-16 mm) und
schmalflügelige Exemplare durch Prof. Simony am 26 August 1889 an einer mit Juncus und
NEW DATA ON NOCTUOIDEA FROM THE CANARY ISLANDS, SPAIN
SHILAP Revta. lepid., 50 (197) marzo 2022 147
Petroselinum bewachsenen feuchten Stelle…”. The specimens are most likely confused with S.
costaestrigalis (Stephens, 1834), a quite common species from Gran Canaria, La Palma and Tenerife.
HACKER & SCHMITZ (1996: 170) also doubted the La Palma record of S. taenialis, and it is not
mentioned by BÁEZ et al. (2010: 312). The species should be removed from the list of Lepidoptera
occurring in the Canary Islands (VIVES MORENO, 2014: 614).
CALPINAE
Epharmottomena eremophila (Rebel, 1895) (Fig. 3)
Armada eremophila Rebel, 1895. Verh. Zool.-bot. Ges. Wien, 45: 350
Material examined: SPAIN, FUERTEVENTURA, Corralejo, 10 m, 1 0, 1-27-XI-2017, leg. P. Falck;
Caldereta, 120 m, 1 1, 27-II-19-III-2018, leg. P. Falck (PF). New to the Canary Islands.
Distribution: North Africa and the Middle East, Ghana (FREINA & BEHOUNEK, 1996: 16).
Biology: Early-stage unknown. The adult flies during winter.
Remarks: E. eremophila is probably a migrant from Africa.
BOLETOBIINAE
Eublemma thermobasis Hampson, 1910 (Fig. 4)
Eublemma thermobasis Hampson, 1910. Cat. Lep. Phal., 10: 135, pl. 152, fig. 27
Material examined: SPAIN, GRAN CANARIA, Puerto Rico, 50 m, 1 1, 11-24-VI-2018, leg. P. Falck;
San Filipe, 25 m, 2 11, 24-X-13-XI-2020, leg. P. Falck; Lanzarote, Mojón Blanco, Orzolá, 20 m, 1 0,
21-X-10-XI-2019, leg. P. Falck, DNA sample Lepid Phyl 0603PF/CILEP602-20; Tenerife, El Médano,
15 m, 12 11, 4 00, 1-20-III-2017, genitalia slide 2499PF, same data but 6 11, 1 0, 18-XI-8-XII-2018,
same data but 6 11, 1 0, 21-V-3-VI-2019, leg. P. Falck, DNA sample Lepid Phyl 0602PF/CILEP601-
20 (all PF). New to the Canary Islands.
Distribution: Morocco (one record from 1994), East Africa from Sudan to Ethiopia and the
Arabian Peninsula from Yemen to Jordan (HACKER, 2019: 335). The species is considered to be
resident in the Canary Islands (Spain).
Biology: Unknown.
Remarks: We obtained DNA barcode from two specimens. Barcode Index Number (BIN) BOLD:
ACL6036.
Eublemma candidana (Fabricius, 1794) (Fig. 5)
Pyralis candidana Fabricius, 1794. Ent. Syst., 3(2): 245
Material examined: SPAIN, TENERIFE, La Cuesta, 1 0, III-1924, leg. H. P. Duurloo, genitalia slide
3818 M. Fibiger (ZMUC). New to the Canary Islands.
Distribution: Europe (Mediterranean countries), Turkey, Middle East, Caucasus, Trans-Caucasus
and Central Asia (FIBIGER et al., 2010: 68).
Biology: The larvae feed in April-May and again in July on flowers of Helichrysum (FIBIGER et
al., 2010: 68).
Remarks: The record of E. candidana from the Canary Islands is probably due to accidental
introduction or migration, as the species has not been found on the islands for almost a hundred
years.
The collector of the specimen, Hans Peter Duurloo was a Danish lepidopterist who travelled and
collected Lepidoptera in the Canary Island during 1924-1926 (HENRIKSEN, 1936: 413).
Araeopteron ecphaea (Hampson, 1914) (Fig. 6)
Araeoptera ecphaea Hampson, 1914. Ann. Nat. Hist., 13: 167
Material examined: SPAIN, GRAN CANARIA, Maspalomas, 10 m, 1 1, 1 0, 17-30-IX-2018, leg. P.
Falck (PF). New to the Canary Islands.
P. FALCK & O. KARSHOLT
148 SHILAP Revta. lepid., 50 (197) marzo 2022
Distribution: Central Africa, North Africa (Morocco, Tunisia), Middle East and Europe (the
southern Mediterranean from Greece to Spain) (FIBIGER et al., 2010: 52; HACKER, 2019: 243).
Biology: Early stages undescribed. E. ecphaea inhabits moist areas with Phragmites (FIBIGER et
al., 2010: 52).
Remarks: E. ecphaea is probably a resident of the Canary Islands.
EREBINAE
Gnamptonyx innexa (Walker, 1858) (Fig. 7)
Alamis innexa Walker, 1858. List Specimens lepid. Insects Colln Br. Mus., 15: 1797
Material examined: SPAIN, GRAN CANARIA, Carreteria, 455 m, 1 1, 8-20-VIII-2020, leg. P. Falck;
Pie de la Cuesta, 500 m, 1 1, 1 0, 21-VIII-4-IX-2020, leg. P. Falck (PF). New to the Canary Islands.
Distribution: Spain (FIBIGER et al., 2010: 233), Cape Verde Islands, North Africa, through the
Middle East to Afghanistan and India (FREINA & BEHOUNEK, 1996: 19; HACKER, 2016).
Biology: The larva feeds on Acacia species. It is continuously brooded, and the adults are
migratory (FIBIGER et al., op. cit.).
Remarks: G. innexa is probably a migrant from Africa.
Rhabdophera acrosticta (Püngeler, 1904) (Fig. 8)
Pericyma acrosticta Püngeler, 1904. Iris, 16: 290, pl. 6, fig. 6
Material examined: SPAIN, FUERTEVENTURA, Corralejo, 10 m, 1 0, 7-27-XI-2017, leg. P. Falck
(PF). New to the Canary Islands.
Distribution: From North Africa to the Middle. East Distribution map cf. HACKER (2016: 327).
In Europe one specimen from Malta (LEPIFORUM, 2021).
Biology: Early stages are probably undescribed.
Remarks: R. acrosticta is probably a migrant from Africa.
NOLIDAE
CHLOEPHORINAE
Nycteola columbana (Turner, 1925) (Fig. 9)
Sarrothripus columbana Turner, 1925. Entomologist’s Rec. J. Var., 37: 77
Material examined: SPAIN, FUERTEVENTURA, Corralejo, 10 m, 1 1, 7-27-XI-2017, leg. P. Falck,
genitalia slide 3472PF (PF). New to the Canary Islands.
Distribution: South Europe, Turkey, North Africa (Morocco, Algeria, Tunisia) and the Middle East
(FIBIGER et al., 2009: 138).
Biology: Early stages are probably undescribed. The larva feeds on Quercus.
Remarks: The genitalia of both sexes are figured by FIBIGER et al. (2009: figs 81, 245).
NOCTUIDAE
BAGISARINAE
Amyna axis Guenée, 1852 (Fig. 10)
Amyna axis Guenée, 1852. Hist. Nat. Insc. Spec. Gen. Lep. Noct., 5(1): 407
SPAIN, LAGOMERA, Valle Gran Rey, 1 larva, 25-II-2013, leg. J. Köhler (KÖHLER, 2014).
Material examined: SPAIN, LANZAROTE, Puerto del Carmen, 25 m, 1 1, 21-X-10-XI-2019, leg. P.
Falck; Mojón Blanco, Orzolá, 20 m, 3 11, 1 0, 21-X-10-XI-2019, leg. P. Falck; El Bosquecillo, 610 m,
4 11, 21-X-10-XI-2019, leg. P. Falck (all PF). New to the Canary Islands.
Distribution: A. axis is described from Tahiti and is widely distributed in the tropics around the
world. In Africa from the south to Sudan, Mauretania and Saudi-Arabia in the north.
Biology: The larva from La Gomera was found sitting on Rumex vesicarius L. without eating
NEW DATA ON NOCTUOIDEA FROM THE CANARY ISLANDS, SPAIN
SHILAP Revta. lepid., 50 (197) marzo 2022 149
(KÖHLER, 2014: 179).The larva is reported feeding from several plants e. g. Arachis hypogaea L.,
Beta vulgaris L., Celosia sp., Chenopodium album L. Bosc ex Moq., Corchorus capsularis L., Hibiscus
cannabinus L., Spinacia oleracea L. often as a pest (LEPIFORUM, 2008-2021).
Remarks: A. axis has probably become a resident of the Island of Lanzarote due to either
migration or accidental importation.
BRYOPHILINAE
The subgenus Nyctobrya Boursin, 1957 is treated in two relatively new articles BEHOUNEK &
SPEIDEL (2013) and FISCHER & FREINA (2014) with description of two new species. Hitherto, five
species and one subspecies are known: N. simonyi (Rogenhofer, 1889), N. simonyi debilis (Rebel,
1894), N. canaria (Alphéraky, 1890), N. maderensis (Bethune-Baker, 1891), N. pinkeri Behounek &
Speidel, 2013 and N. hierroana Fischer & Freina, 2014. Until recently it was assumed that the
subgenus was endemic to the Macaronesian region, but N. simonyi is now recorded from Morocco
(LEPIFORUM, 2008-2021).
Nyctobrya vilfredi Falck & Karsholt, sp. n. (Figs 11, 12)
Holotype 0: SPAIN, GRAN CANARIA, Ayacata, 1400 m, 17-30-IX-2018, leg. P. Falck, DNA sample
Lepid Phyl 0572PF/CILEP571-20 (ZMUC).
Paratypes: SPAIN, GRAN CANARIA, Los Tilos, 600 m, 1 0, 5-VIII-1979, leg. & P. Stadel Nielsen;
Barranco de la Virgen, Moya, 400 m, 1 0, 20-VII-1984, leg. P. Olsen, B. Skule & P. Stadel Nielsen;
Pinos de Galdar, 1200 m, 15 11, 46 00, 24-VII-1984, 14 11, 45 00, 31-VII-1984, genitalia slides
3653, 3836 M. Fibiger, leg. P. Olsen, B. Skule & P. Stadel Nielsen (all ZMUC); Ayacata, 1400 m, 1 1,
2 00, 17-30-IX-2018, genitalia slide 3294PF, DNA sample Lepid Phyl 0574PF/CILEP573-20, leg. P.
Falck (PF); Pie de la Cuesta, 500 m, 1 0, 17-30-IX-2018, genitalia slide 3300PF, DNA sample Lepid
Phyl 0573PF/CILEP572-20, leg. P. Falck (PF); Carreteria, 455 m, 14 11, 23 00, 8-20-VIII-2020, leg.
P. Falck, genitalia slides 3462PF, 3463PF, DNA sample Lepid Phyl 0758PF/CILEP757-20 (MNCN,
PF); Fontanales, 1100 m, 7 00, 8-20-VIII-2020, leg. P. Falck (PF); Barranco de Moya, 80 m, 3 11, 3
00, 8-20-VIII-2020, leg. P. Falck (PF); Barranco de Azuaje, 270 m, 1 1, 7 00, 8-20-VIII-2020, leg. P.
Falck, DNA sample Lepid Phyl 0757PF/CILEP756-20 (PF); Barranco de Guayadeque, 700 m, 5 00,
21-VIII-4-IX-2020, leg. P. Falck (PF).
Description: Male adult. Wingspan 23.5-29 mm. Labial palp upturned, segment 2 with black and
creamy white scale-tuft, segment 3 as long as segment 2, slender, black mottled with creamy white and
with creamy white tip. Antenna black, cilia in male having length of antenna diameter, in female very
short. Head and neck blackish grey, neck with yellowish tipped scales towards thorax; tegula and
thorax blackish grey. Forewing ground colour dark grey-brown; basal patch and claviform stigmata
darker, orbicular and reniform stigmata brownish distinctly bordered with black; antemedian fascia
black; postmedian fascia jagged, black, irregularly bordered distally with white; fringe dark grey.
Hindwing of the same concolorous with forewing, paler at base; postmedian fascia indistinct, discal
spot clearly recognizable; fringe dark grey.
Female adult: Wing pattern as in male, but with black ground colour, heavily mottled with white
between subbasal- and antemedian fasciae; postmedian fascia distinct, bordered with white.
Male genitalia (Figs 30, 30a): Uncus long and spatulate. Tegumen sub-triangular. Valva short, very
broad at the base, slightly narrowing distally; apical process at the end of costa small and triangular,
ventral edge apically pointed. Ampulla long, fairly robust and evenly curved. Juxta trapezoid, anteriorly
acute. Phallus relatively short and broad, vesica with large group of small cornuti.
Female genitalia: (Fig. 36): Ostium membranous, narrow and rounded. Ductus bursae relatively
long and broad, sclerotized, transition to ostium short, constricted and membranous. Corpus bursae
membranous, pear-shaped, posterior apex pronounced, rounded and slightly sclerotized.
Molecular diagnosis (Fig. 39): We obtained full length DNA barcodes (658 bp) from three
P. FALCK & O. KARSHOLT
150 SHILAP Revta. lepid., 50 (197) marzo 2022
specimens and DNA barcode fragments of 607 bp and 528 bp from two specimens all from the island
of Gran Canaria. All the barcodes fall within Barcode Index Number (BIN) BOLD: AEE9804. The
maximum intraspecific distance is 0.17% (mean 0.05%, n=5). The minimum p-distance to nearest
neighbour N. canaria is 3.03%, with the Barcode Index Number (BIN) BOLD: AEE9801. The result
supports the status of N. vilfredi sp. n. as a separate species.
Diagnosis: N. vilfredi is closely related to N. canaria (Figs 13, 14). It can often be distinguished
by the light tipped apex of labial palps, and females are normally more contrasting and with strong
white admixture in basal part of the forewing, but a safe determination requires dissection of the
genitalia. It can be distinguished from N. simonyi and N. pinkeri (Figs 15, 16) by the longer segment 3
of the labial palps (Figs 17, 18). In the male genitalia N. vilfredi differs from N. canaria (Figs 31, 31a)
by the short and broad valva without projection of the ventral edge, and the cornuti are larger. From N.
simonyi, N. pinkeri, N. hierroana and N. maderensis it differs by the lack of the large and robust
cornutus. In the female genitalia it differs from all other members of the genus by the long and broad
ductus bursae and by the anterior rounded, sclerotized apex of corpus bursae.
Biology: Early stages unknown. The specimens were attracted to light during July to late
September at altitudes ranging from 80-1400 m.
Distribution: Only known from the central and northern part of the island of Gran Canaria.
Etymology: The species name is dedicated to the first author`s youngest grandson Vilfred.
Remarks: In the island of Gran Canaria, N. pinkeri is widespread in mountainous regions, but in
the northern part of the island the species is found in ravines near sea level in a much smaller and a
greyer form, with a wingspan of 19-21 mm (usually 24-30mm), very similar to the difference between
N. simonyi simonyi and N. simonyi debilis
In the present study we also compared the DNA barcodes of the two forms of N. pinkeri, and N.
simonyi simonyi with of N. simonyi debilis.
The result of the molecular analysis of N. pinkeri is that all barcodes (n=6) fall within the same
Barcode Index Number (BIN) BOLD: AEE9803, with maximum intraspecific variation of 0.8%, this
supports the presumption of only one highly variable species.
The result of the molecular analysis of N. simonyi simonyi (n=3) from Tenerife and N. simonyi
debilis (n=3) from Lanzarote is that all barcodes (n=6) fall within the same Barcode Index Number
(BIN) BOLD: AEE9802. The maximum intraspecific variation in N. simonyi simonyi is 0.14%, and the
maximum intraspecific variation in N. simonyi debilis is 0.17%. Diverging (minimum p-distance) by
0.82% between the two populations. This supports the assumption by BEHOUNEK & SPEIDEL
(2013: 163) that the status of N. debilis as a separate species is unlikely.
XYLENINAE
Caradrina rebeli Staudinger, 1901 (Figs 19, 20)
Caradrina rebeli Staudinger, 1901. Cat. Lep. Pal.: 196
The species is widespread in all of the larger Canary Islands and is often very common from sea
level to mountain regions. It is highly variable both in adult appearance and in the genitalia. Males from
La Palma, Tenerife and Gran Canaria have greyish brown ground colour of the forewing and almost
solid dark grey hindwing with discal spot visible, whereas males from Lanzarote and Fuerteventura are
more whitish grey in ground colour of the forewing and the hindwing is dark grey becoming almost
white towards base, with discal spot and postmedian fascia visible, there are no clear differences in
adult females from different islands. It is not surprising that more subspecies and one additional species
have been described: Caradrina lanzarotensis Pinker, 1962, C. rebeli grancanariae Pinker, 1962, C.
rebeli lapalmae Pinker, 1962, C. rebeli hierrensis Pinker, 1969 and C. lanzarotensis fuerteventurensis
Pinker & Bacallado, 1975. In the revision of the genus Caradrina Ochsenheimer, 1816 the subspecies
of C. rebeli and C. lanzarotensis were synonymized (HACKER, 2004: 380-384, figs 426-428, pls. 22).
In this study we have examined the DNA barcode of specimens from La Palma, Tenerife, Gran
Canaria, Lanzarote and Fuerteventura.
NEW DATA ON NOCTUOIDEA FROM THE CANARY ISLANDS, SPAIN
SHILAP Revta. lepid., 50 (197) marzo 2022 151
Molecular results (Fig. 39): We obtained DNA barcode fragments of 592 bp from one specimen
from the island of La Palma, 587 bp and 586 bp from two specimens from the island of Tenerife, 598
bp and 622 bp from two specimens of the island of Gran Canaria, 598 bp and 688 bp from two
specimens from the island of Lanzarote and 633 bp and 632 bp from two specimens from the island of
Fuerteventura. The result of the molecular analysis is that all barcodes (n=5) from specimens from La
Palma, Tenerife and Gran Canaria fall within the same Barcode Index Number (BIN) BOLD:
AEE7063. The maximum intraspecific divergence in BIN (p-distance) is 0.52%, and it is between
populations from La Palma and Tenerife. The barcode from the specimens (n=4) from Lanzarote and
Fuerteventura fall within Barcode Index Number (BIN) BOLD: AEE7064. The maximum
intraspecific divergence in BIN (p-distance) is 0%. Diverging (minimum p-distance) by 1.87%
between the two populations from La Palma, Tenerife, Gran Canaria and Lanzarote, Fuerteventura.
The results support the assumption by HACKER (2004: 383) that at least the subspecies C. rebeli
grancanariae and C. rebeli lapalmae are synonyms of C. rebeli rebeli and C. lanzarotensis
fuerteventurensis is a synonym of C. rebeli lanzarotensis. Based on adult appearance and DNA results
it is concluded that the species occurs in two subspecies: C. rebeli rebeli Staudinger, 1901 and C.
rebeli lanzarotensis Pinker, 1962.
Cosmia affinis (Linnaeus, 1767)
Noctua affinis Linnaeus, 1767. Syst. Nat., ed., 12: 848
The species is mentioned by VIVES MORENO (2014) as Cosmia affinis canaria (Pinker, 1974)
by mistake, but it refers to Cardepia affinis antinea Rungs, 1972 (= deserticola canaria Pinker, 1974).
Cosmia affinis (Linnaeus, 1767) should be removed from the list of Lepidoptera found in the
Canary Islands.
Polymixis aurora (Turati, 1924) (Fig. 21)
Pseudopolia aurora Turati, 1924. Atti Soc. ital. Sci. nat., 63: 83
Material examined: SPAIN, FUERTEVENTURA, Corralejo, 10 m, 13 11, 7-27-XI-2017, leg. P. Falck
(PF). New to the Canary Islands.
Distribution: North Africa from Morocco to Egypt (FREINA & BEHOUNEK, 1996: 29).
Biology: Early stages unknown.
Remarks: The specimens from Fuerteventura belong to P. aurora commixta (Rungs, 1943). It is
probably a migrant from Africa.
Nonagria typhae (Thunberg, 1784) (Fig. 22)
Noctua typhae Thunberg, 1784. Diss. Ent. sistens Insecta Suecica, (1): 3
Material examined: SPAIN, GRAN CANARIA, Barranco de Azuaje, 270 m, 1 1, 1 0, 8-20-VIII-
2020, leg. P. Falck, genitalia slide 3486PF (PF). New to the Canary Islands.
Distribution: Europe, Turkey, Russia, Middle East, Iraq, Iran and Central Asia.
Biology: The larvae feeds on Typha sp. and Schoenoplectus sp. (ZILLI et al., 2005: 86).
Remarks: N. typha is probably a resident of the Island of Gran Canaria.
HADENINAE
Mniotype loslobensis (Fischer, Saldaitis & Ivinskis, 2007) (Fig. 23)
Eremobastis loslobensis Fischer, Saldaitis & Ivinskis, 2007. Atalanta, 38(3/4): 377, figs 1-2, pl. 21, fig. 1
Material examined: SPAIN, LANZAROTE, Mojón Blanco, Orzolá, 20 m, 2 11, 5-XI-2018, leg. and
coll. B. Skule and C. Hviid, same data but 10 11, 4 00, 21-X-10-XI-2019, leg. P. Falck, genitalia slide
3287PF, 3394PF, 3478PF, DNA samples Lepid Phyl 0596PF/CILEP595-20, 0596PF/CILEP595-20
(PF).
The species was described on the basis of three males from the island of Los Lobos 2 km NE of
Fuerteventura (FISCHER et al., 2007). M. loslobensis is very similar to M. dimorpha (Rungs, 1948)
P. FALCK & O. KARSHOLT
152 SHILAP Revta. lepid., 50 (197) marzo 2022
from Morocco both in the appearance of the adults and in the male genitalia. It can be distinguished by
the lack of the prominent basal streak in the forewing, and in the male genitalia by the lack of several
small triangular spines in the phallus diverticulum.
Female genitalia (Fig. 38): Ostium narrow, funnel-shaped, membranous, laterally sclerotized.
Ductus bursae relatively long, heavily spinulated and with longitudinal folds. Corpus bursae
membranous and oval.
Remarks: We obtained DNA barcode fragments of 633bp and 635pb from two specimens, the
barcodes fall within Barcode Index Number (BIN) BOLD: AEE1222. The p-distance to nearest
neighbour Mniotype fulva (Rothschild, 1914) is 4.33%, with the Barcode Index Number (BIN) BOLD:
AEC7959. It was not possible to compare the DNA barcode with Mniotype dimorpha as no barcode
seems to exist so far.
The genus Eremobastis Pérez-López & Morente-Benítez, 1996 was synonymised with Mniotype
Franclemont, 1941 by FIBIGER & HACKER (2007: 230).
Hecatera sancta (Staudinger, 1859)
Hadena sancta Staudinger, 1859. Stettin. ent. Ztg., 20(7-9): 213
This species is mentioned by VIVES MORENO (2014) as Hecatera sancta canaria (Pinker, 1974)
but by mistake, it actually refers to Clytie sancta canaria Pinker, 1974 (= Clytie illunaris (Hübner,
[1816])).
Hecatera sancta (Staudinger, 1859) should be removed from the list of Lepidoptera found in the
Canary Islands.
Mythimna languida (Walker, 1858) (Fig. 24)
Hadena languida Walker, 1858. List. Spec. Lepid. Insects Colln Br. Mus., 15: 1728
Material examined: SPAIN, GRAN CANARIA, Los Tilos de Moya, 500 m, 3 11, 11-24-VI-2018,
leg. P. Falck, same data but 1 0, 17-30-IX-2018; Carreteria, 455 m, 1 1, 2 00, 8-20-VIII-2020, leg. P.
Falck; Barranco de Azuaje, 270 m, 5 11, 2 00, 8-20-VIII-2020, leg. P. Falck; San Filipe, 25 m, 1 1,
24-X-13-XI-2020, leg. P. Falck; Tenerife, Las Americas, 40 m, 1 0, 21-V-3-VI-2019, leg. P. Falck (all
PF). New to the Canary Islands.
Distribution: Europe, Africa, Cape Verde, and along the southern border zone of the Palaearctic
and the Oriental Regions eastwards to Nepal (HACKER et al., 2002: 188).
Biology: The larva and pupa are figured in LEPIFORUM (2008-2021).
Remarks. M. languida is a resident of the Canary Islands.
Leucania zeae (Duponchel, 1828) (Fig. 25)
Noctua zeae Duponchel, 1828. Hist. nat. Lépid. Fr., 7(1): 363, pl. 122, fig. 4
Material examined: SPAIN, FUERTEVENTURA, Caldereta, 120 m, 1 0, 27-II-19-III-2018, leg. P.
Falck; Vega de Rio Palmas, 245 m, 3 01, 6-26-I-2020 leg. P. Falck; Gran Canaria, Pie de la Cuesta,
500 m, 1 0, 17-30-IX-2018, leg. P. Falck (all PF). New to the Canary Islands.
Distribution: Europe (surroundings of the Mediterranean Sea) and from North Africa through the
south-western part of the Palaearctic region to Mongolia and Northwest China (HACKER et al., 2002:
201).
Biology: The larva feeds on Phragmites but is also regarded as a pest of Zea mays L. (HACKER
et al., 2002: 201).
Remarks: L. zeae is a resident of the Canary Islands.
Agrotis corralejoi Falck & Karsholt, sp. n. (Fig. 26)
Holotype 1: SPAIN, FUERTEVENTURA, Corralejo, 10 m, 7-27-XI-2017, leg. P. Falck, genitalia slide
3479PF, DNA sample Lepid Phyl 0763PF/CILEP762-21 (ZMUC).
Paratypes: SPAIN, FUERTEVENTURA, Corralejo, 10 m, 3 11, 12-23-XI-1997, leg. N. M. Hall
NEW DATA ON NOCTUOIDEA FROM THE CANARY ISLANDS, SPAIN
SHILAP Revta. lepid., 50 (197) marzo 2022 153
(NHMUK), same data but 25 11, 29-XI-11-XII-2000, leg. N. M. Hall (NHMUK), same data but 23
11, 7-27-XI-2017, leg. P. Falck, genitalia slide 3477PF, 3480PF, 3481PF, DNA samples Lepid
Phyl0764PF/CILEP763-21, 0765PF/CILEP764-21 (PF, MNCN).
Description: Wingspan 26.5-34 mm. Labial palp porrect, segment 2 whitish mottled with grey and
with large, bushy scale tuft, segment 3 short, white. Antenna creamy white, strongly bipectinate. Head,
neck, tegula and thorax light grey mottled with white and darker grey. Forewing ground colour pale
grey mottled with white, especially along dorsum and costa, and with reddish brown between
postmedian fascia and subterminal fascia and around the stigmae; orbicular-, reniform- and claviform
stigma quite distinct creamy white, often more or less bordered with black, discal spot diffuse grey;
postmedian fascia jagged, sometimes indistinct, subterminal fascia distinct, jagged, distally white.
Fringe light grey. Hindwing whitish often with a darker, diffuse subterminal fascia: discal spot small
and indistinct. Fringe grey.
Female unknown
Male genitalia (Figs 34, 34a, 34b): Uncus long slightly spatulate. Valva parallel sided, ventral- and
dorsal edge almost straight, apex rounded; harpe with a broad base, tapering distally, apex almost
pointed, length of sacculus and harpe about 2/3 the length of valva. Saccus rounded. Phallus straight;
vesical with a smaller basal swelling, tapering towards apex.
Female genitalia unknown.
Molecular diagnosis: We obtained full lenght DNA barcodes (658 bp) from one specimen and
DNA barcode fragments of 632 bp and 621 bp from two specimens. The barcodes fall within Barcode
Index Number (BIN) BOLD: AEI1694
The maximum intraspecific distance is 0.32% (mean xx5%, n=3). The p-distance to nearest
neighbour Agrotis venerabilis Walker, 1857 is 3.55%, with the Barcode Index Number (BIN) BOLD:
ABZ1938. The minimum p-distance to A. aistleitneri is 7.94%, with the Barcode Index Number (BIN)
BOLD: AEW1944 and the minimum p-distance to A. boeticus is 4.26%, with the Barcode Index
Number (BIN) BOLD: AEF1653.
Diagnosis: Externally, A. corralejoi resembles A. boeticus (Boisduval, 1837) (Fig. 27) and A.
aistleitneri Behounek & Speidel, 2009 (Fig. 28). It is distinguished by the pale grey ground colour, the
distinct subterminal fascia and the white area distally; furthermore, it differs from A. aistleitneri by the
much more strongly pectinate antenna. In the male genitalia A. corralejoi is distinguished from A.
boeticus (Figs 35, 35a, 35b) by the slender and pointed harpe and by the lack of the small plate with
cornuti in the phallus, and from A. aistleitneri by the longer and broader harpe and by the straight
ventral and dorsal margin of the valva. The genitalia of A. aistleitneri are figured by BEHOUNEK &
SEIDEL (2009: 127).
Biology: Early stages unknown. The adult is recorded from October to the beginning of February
at light (HACKER et al., 1996: 207).
Distribution: A. corralejoi is only known from the island of Fuerteventura (Jandia, Costa Calma
and Corralejo) (HACKER et al., 1996: 207; LEPIFORUM, 2021).
Etymology: The species is named after the town of Corralejo, situated at the northern tip of the
island of Fuerteventura. The name was originally suggested by Michael Fibiger and Martin Honey.
Remarks: FIBIGER & SKULE (2004-2021) listed Agrotis corralejoi Fibiger & Honey, 2004 (a
misspelling of A. corralejoi) as a valid species. In 2009 A. aistleitneri Behounek & Speidel, 2009 was
described from the Cape Verde Islands and A. corralejoi was at that time erroneously regarded as
conspecific with A. aistleitneri (LEPIFORUM, 2008-2021), so the taxon was never described or
published in a scientific publication. A. corralejoi has also been confused with A. boeticus, due to
misidentification (LEPIFORUM, 2008-2021).
Numerous males of A. corralejoi were attracted to light in the sand-dunes south of Corralejo, but
no females were found, in spite of searching for it in the surroundings. We assume that the female is
brachypterous, and it may hide in the loose sand, as it is known also from other species of Noctuidae
occurring in deserts. Females of A. aistleitneri and A. boeticus (Boisduval, 1837) are fully winged.
P. FALCK & O. KARSHOLT
154 SHILAP Revta. lepid., 50 (197) marzo 2022
A. boeticus (Boisduval, 1837) and A. aistleitneri Behounek & Speidel, 2009 should be removed
from the list of Lepidoptera found in the Canary Islands.
Discussion
The Noctuidae are among the most popular groups of nocturnal Lepidoptera. This is both because
of their relatively large size, their often-concealed way of life and the diversity of the group. With more
than 40,000 species worldwide, the Noctuoidea are the most diverse superfamily of Lepidoptera. As it
is often seen the diversity in oceanic islands is much lower and the most recent list of Canary Island
Lepidoptera (VIVES MORENO, 2014) listed only 133 species of Noctuidae. This is certainly not due
to under-sampling because the islands are, with their pleasant climate, a favourite holiday destination
and lepidopterist from several European countries have undertaken fieldwork in the islands for more
than 100 years.
In the present paper we add 15 species of Noctuoidea for the Canary Islands, three of which are
described as new to science. The latter were previously found by other lepidopterists but were
misidentified, and we regard all three to be endemic to the Canary Islands. Some of the newly recorded
species are resident in the islands, and the occurrence of others are most likely the result of migration.
It is assumed that some species arriving the Canary Islands as migrants, or introduced due to human
activities, can establish temporary populations in the islands (sometimes for longer periods of time).
It is of course of high interest to record additions to the fauna of a certain area (here the Canary
Islands), but it is also very important to keep the list of taxa occurring there updated. We have therefore
critically assessed the current list of Canary Island Lepidoptera (VIVES MORENO, 2014), and we
suggest that five species are removed from the list: 3 of them due to misidentifications and 2 of them
due to nomenclatorial confusion. The list of Noctuoidea of the Canary Islands thus includes 143
species.
Acknowledgements
We are grateful to Hermann Hacker (Bad Staffelstein, Germany) for pointing out the synonymy of
Zebeeba with Gerarctia and allowing us to publish it, and for help with determination of other
Noctuoidea from the Canary Islands, to Dr. Axel Hausman (Zoologische Staatssammlung, Munich,
Germany) for giving access to DNA barcode of Agrotis aistleitneri and for permission to use
photographs of that species, to Dr. Antonio S. Ortiz Cervantes (Departamento de Zoología y
Antropología Física, Facultad de Veterinaria, Universidad de Murcia, Murcia, Spain) for giving access
to DNA barcodes of Agrotis boeticus, to Norman M. Hall, Reading, England, for permission to include
his records of Agrotis corralejoi in the paratype series, to Carsten Hviid, Per Stadel Nielsen, Danny
Nilsson and Bjarne Skule (all Denmark) for loan and donation of specimens, and to Dina Soliman and
other staff of the Canadian Centre for DNA Barcoding for sequencing the samples and for continuous
help in management of our BOLD records, and to Thomas Simonsen, Naturhistorisk Museeum, Århus,
for his help interpretation of the DNA results. Dr. Martin Honey, The Natural History Museum,
London, U. K. kindly commented on and improved the English language of the final version of the
manuscript. We are moreover grateful to Dr. Antonio Vives, Madrid, Spain for translating the abstract
into Spanish, for editing our manuscript, and for his kind help with obtaining permission to collect
Lepidoptera in the Canary Islands into the Scientific Project of SHILAP.
BIBLIOGRAPHY
BÁEZ, M., 2010.– Orden Lepidoptera. Pp. 302-318.– In S. ARECHAVALETA, N. RODRÍGUEZ, N. ZURITA & A.
GARCÍA (eds). Lista de especies silvestres de Canarias (hongos, plantas y animales terrestres). Gobierno de
NEW DATA ON NOCTUOIDEA FROM THE CANARY ISLANDS, SPAIN
SHILAP Revta. lepid., 50 (197) marzo 2022 155
Canarias, La Laguna. 577 pp. Available from http://www.gobiernodecanarias.org/medioambiente/piac/
descargas/Biodiversidad/Listas_Especies_Silvestres.pdf.
BEHOUNEK, G. & SPEIDEL, W., 2009.– Eine neue Agrotis-Art von Cabo Verde: Agrotis (Powellinia) aistleitneri
sp. n. (Lepidoptera, Noctuidae).– Entomofauna, 30: 121-128. https://www.zobodat.at/pdf/ENT_0030_0121-
0128.pdf.
BEHOUNEK, G. & SPEIDEL, W., 2013.– Contribution to the knowledge of the genus Nyctobrya Boursin, 1957
(Lepidoptera: Noctuidae: Bryophilinae) in the Macaronesian archipelago, with description of a new species
from Gran Canaria.– Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen, 65: 157-166.
FALCK, P. & KARSHOLT, O., 2019.– New data on Praydidae, Oecophoridae, Stathmopodidae and
Cosmopterigidae from the Canary Islands (Insecta: Lepidoptera).– SHILAP Revista de lepidopterología,
47(186): 325-340.
FALCK, P., KARSHOLT, O. & SLAMKA, F., 2019.– New data on Pyraloidea from the Canary Islands
(Lepidoptera: Pyraloidea).– SHILAP Revista de lepidopterología, 47(185): 33-48.
FIBIGER, M. & HACKER, H., 2007.– Amphypyrinae, Condicinae, Eriopinae, Xyleninae (part).– Noctuidae
Europaeae, 9: 1-504.
FIBIGER, M., RONKAY, L., STEINER, A. & ZILLI, A., 2009.– Pantheinae- Bryophilinae.– Noctuidae Europaeae,
11: 1-410.
FIBIGER, M., RONKAY, L., YELA, J. L. & ZILLI, A., 2010.– Rivulinae, Boletobiinae, Hypenodinae,
Araeopteroninae, Eublemminae, Herminiinae, Hypeninae, Phytometrinae, Euteliinae and Micronoctuidae.
Including Supplement to volumes 1-11.– Noctuidae Europaeae, 12: 1-451.
FIBIGER, M. & SKULE, B., 2004-2021.– Noctuidae.– In O. KARSHOLT & E. J. VAN NIEUKERKEN (eds):
Lepidoptera.– Fauna Europaea, version 2017.06. Available form https://fauna-eu.org [last update of
Noctuidae: version 2.4 January 2011] (accesed 21 April 2021).
FISCHER, H. & FREINA, J. J., 2014.– Nyctobrya hierroana sp. n., eine weitere Art der makaronesischen
Nyctobrya simonyi (Rogenhofer, 1889)-Artengruppe (Lepidoptera: Noctuidae: Bryophilinae).– Mitteilungen
der Münchner Entomologischen Gesellschaft, 104: 139-143.
FISCHER, H., SALDAITIS, A. & IVINSKIS, P., 2007.– Eine neue Art der Gattung Eremobastis Pérez-López
& Morente-Benítez, 1996 von der Kanareninsel Los Lobos (Lepidoptera, Noctuidae).– Atalanta, 38(3/4):
377-380.
FREINA, J. J. & BEHOUNEK, G., 1996.– Beitrag zur Noctuidenfauna der Wüstenregion Südmarokkos: Das
Artenspektrum in der gemäßigten Jahreszeit November bis April (Lepidoptera, Noctuidae).– Esperiana, 5: 11-
38, pl. A-B.
HACKER, H., 2004.– Revision of the genus Caradrina OCHSENHEIMER, 1816, with notes on other genera of the
tribus Caradrini (Lepidoptera, Noctuidae).– Esperiana, 10: 7-690.
HACKER, H., 2016.– Systematic and illustrated catalogue of the Macroheterocera and Cossoidea Leach, [1815],
Zygaenoidea Latreille, 1809, Thyridoidea Herrich-Schäffer, 1846 and Hyblaeoidea Hampson, 1903 of the
Arabian Peninsula, with a survey of their distribution (Lepid.).– Esperiana, 20(1):1-742; (2): 1-430.
HACKER, H., 2019.– Biogeography; Boletobiinae (Lepidoptera, Noctuidae, Erebidae).– In H. HACKER (ed).
Moths of Africa. Systematic and illustrated Catalogue of the Heterocera (Lepidoptera) of Africa, 1: 1-816.
HACKER, H., 2021.– Rivulinae, Hypeninae, Herminiinae and Hyponodinae (Erebidae). Evolution of diversity of
the African flora and fauna.– In H. HACKER (ed). Moths of Africa. Systematic and illustrated Catalogue of
the Heterocera (Lepidoptera) of Africa, 2: 1-720.
HACKER, H., RONKAY, L. & HREBLAY, M., 2002.– Hadeninae 1.– Noctuidae Europaeae, 4: 1-419.
HACKER, H. & SCHMITZ, W., 1996.– Fauna und Biogeographie der Noctuidae des makaronesischen Archipels
(Lepidoptera).– Esperiana, 4: 167-221, pls. L-O.
HENRIKSEN, K. L., 1921-1937.– Oversigt over Dansk Entomologis Historie.– Entomologiske Meddelelser, 15(1):
1-48 (1921), (2): 49-96 (1922), (3): 97-144 (1924), (4): 145-192 (1925), (5): 193-240 (1926), (6): 241-288
(1927), (7-10): 289-480 (1936), (11-12): 481-578 (1937).
KÖHLER, J., 2014.– Scythocentropus inquinata (Mabille, 1888) auch auf La Gomera nachgewiesen - Erste
Beobachtungen zur Biologie einer eremisch verbreiten Art (Lepidoptera, Noctuidae).– Atalanta, 45: 179-181.
LEPIFORUM, 2008-2021.– E. RENNWALD, J. RODELAND et al. (ed.): Lepiforum: Bestimmungshilfe für die in
Europa nachgewiesenen Schmetterlingsarten. Lepiforum e.V. Available from http://www.lepiforum.de/
lepiwiki.pl?Schmetterlingsfamilien_Europa (accesed 24 April 2021).
PINKER, R., 1965.– Interessante und neue Funde und erkenntnisse für die Lepidopterenfauna der Kanaren. III.–
Zeitschrift der Wiener Entomologischen Gesellschaft, 50: 153-172.
P. FALCK & O. KARSHOLT
156 SHILAP Revta. lepid., 50 (197) marzo 2022
RATNASINGHAM, S. & HEBERT, P. D. N., 2007.– BOLD: the barcode of life data systems.– Molecular Ecology
Notes, 7: 355-364.
RATNASINGHAM, S. & HEBERT, P. D. N., 2013.– A DNA-based registry for all animal species: The Barcode
Index Number (BIN) System.– PLOS ONE, 8(8): e66213. doi:10.1371/journal.pone.0066213.
REBEL, H. & ROGENHOFER A., 1894.– Zur Lepidopternfauna der Canaren.– Annalen des Naturhistorischen
Museums in Wien, 9: 1-96, pl. 1.
ROBINSON, G. S., 1976.– The preparation of slides of Lepidoptera genitalia with special reference to the
Microlepidoptera.– Entomologist’s Gazette, 27: 127-132.
TAMURA, K., STECHER, G., PETERSON, D., FILIPSKI, A. & KUMAR, S., 2013.– MEGA6: Molecular
Evolutionary Genetics Analysis version 6.0.– Molecular Biology and evolution, 30: 2725-2729. Available
from https://doi.org/10.1093/molbev/mst197.
VIVES MORENO, A., 2014.– Catálogo sistemático y sinonímico de los Lepidoptera de la Península Ibérica, de
Ceuta, de Melilla y de las islas Azores, Baleares, Canarias, Madeira y Salvajes (Insecta: Lepidoptera): 1184
pp. Suplemento de SHILAP Revista de lepidopterología, Improitalia, Madrid.
ZAHIRI, R., KITCHING, I. J., LAFONTAINE, D., MUTANEN, M., KAILA, L., HOLLOWAY, J. D. &
WAHLBERG, N., 2010.– A new molecular phylogeny offers hope for a stable family level classification of the
Noctuoidea (Lepidoptera).– Zoologica Scripta, 40: 158-173.
ZAHIRI, R., LAFONTAINE, D., HOLLOWAY, J. D., KITCHING, I. J., SCHMIDT, B. C., KAILA, L. &
WAHLBERG, N., 2012.– Major lineages of Nolidae (Lepidoptera, Noctuoidea) elucidated by molecular
phylogenetics.– Cladistics, 2012(1): 1-23.
ZAHIRI, R., LAFONTAINE, D., HOLLOWAY, J. D., KITCHING, I. J., SCHMIDT, B. C., KAILA, L. &
WAHLBERG, N., 2013.– Major lineages of Nolidae (Lepidoptera, Noctuoidea) elucidated by molecular
phylogenetics.– Cladistics, 2013(29): 337-359.
ZILLI, A., RONKAY, L. & FIBIGER, M., 2005.– Apameini.– Noctuidae Europaeae, 8: 1-323.
*P. F. O. K.
Aarsdalevej, 22 Zoological Museum
DK-3730 Neksø Natural History Museum of Denmark
DINAMARCA / DENMARK Universitetsparken, 15
E-mail: per.falck@live.dk DK-2100 Copenhagen
https://orcid.org/0000-0002-0030-9214 DINAMARCA / DENMARK
E-mail: okarsholt@snm.ku.dk
https://orcid.org/0000-0002-6969-2549
*Autor para la correspondencia / Corresponding author
(Recibido para publicación / Received for publication 1-VI-2021)
(Revisado y aceptado / Revised and accepted 3-VIII-2021)
(Publicado / Published 30-III-2022)
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SHILAP Revta. lepid., 50 (197) marzo 2022 157
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158 SHILAP Revta. lepid., 50 (197) marzo 2022
Figs 1-8.– 1. Zebeeba orzolae Falck & Karsholt, sp. n., 1, Lanzarote, 14.5 mm. 2. Zebeeba poliotis (Hampson,
1905), 1, Tenerife, 22 mm. 3. Epharmottomena eremophila (Rebel, 1895), 0, Fuerteventura, 23 mm. 4.
Eublemma thermobasis Hampson, 1910, 1, Gran Canaria, 11 mm. 5. Eublemma candidana (Fabricius, 1794),
0, Tenerife, 13.5 mm. 6. Araeopteron ecphaea (Hampson, 1914), 0, Gran Canaria, 11 mm. 7. Gnamptonyx
innexa (Walker, 1858), 0, Gran Canaria, 28 mm. 8. Rhabdophera acrosticta (Püngeler, 1904), 1, Fuerteventura,
30 mm.
12
34
56
78
NEW DATA ON NOCTUOIDEA FROM THE CANARY ISLANDS, SPAIN
SHILAP Revta. lepid., 50 (197) marzo 2022 159
Figs 9-16.– 9. Nycteola columbana (Turner, 1925), 1, Fuerteventura, 22 mm. 10. Amyna axis Guenée, 1852, 1,
Lanzarote, 25 mm. 11. Nyctobrya vilfredi Falck & Karsholt, sp. n., 1, Gran Canaria, 28 mm. 12. Nyctobrya
vilfredi Falck & Karsholt, sp. n., 0, Gran Canaria, 27 mm. 13. Nyctobrya canaria (Alphéraky, 1890), 1,
Tenerife, 24 mm. 14. Nyctobrya canaria (Alphéraky, 1890), 0, Tenerife, 24 mm. 15. Nyctobrya pinkeri
Behounek & Speidel, 2013, 1, Gran Canaria, 27 mm. 16. Nyctobrya pinkeri Behounek & Speidel, 2013, 0,
Gran Canaria, 27 mm.
910
11 12
13 14
15 16
Figs 17-22.– 17. Nyctobrya vilfredi Falck & Karsholt, sp. n., palps, Gran Canaria. 18. Nyctobrya pinkeri
Behounek & Speidel, 2013, palps, Gran Canaria. 19. Caradrina rebeli rebeli Staudinger, 1901, 1, Tenerife, 31
mm. 20. Caradrina rebeli lanzarotensis Pinker, 1962, 1, Fuerteventura, 27.5 mm. 21. Polymixis aurora (Turati,
1924), 1, Fuerteventura, 31 mm. 22. Nonagria typhae (Thunberg, 1784), 0, Gran Canaria, 46 mm.
P. FALCK & O. KARSHOLT
160 SHILAP Revta. lepid., 50 (197) marzo 2022
17 18
19 20
21 22
NEW DATA ON NOCTUOIDEA FROM THE CANARY ISLANDS, SPAIN
SHILAP Revta. lepid., 50 (197) marzo 2022 161
Figs 23-29.– 23. Mniotype loslobensis (Fischer, Salditis & Ivinskis, 2007), 0, Lanzarote, 33.5 mm. 24.
Mythimna languida (Walker, 1858), 1, Gran Canaria, 29 mm. 25. Leucania zeae (Duponchel, 1828), 1,
Fuerteventura, 35 mm. 26. Agrotis corralejoi Falck & Karsholt sp. n., 1, Fuerteventura, 30 mm. 27. Agrotis
boeticus (Boisduval, 1837), 1, Algarve, Portugal, 30 mm. 28. Agrotis aistleitneri Behounek & Speidel, 2009, 0,
Cape Verde Islands. 29. Zebeeba orzolae Falck & Karsholt, sp. n., 1, Lanzarote, GP3473PF. 29a. Phallus,
Lanzarote, GP3473PF.
23 24
25 26
27 28
29 29a
P. FALCK & O. KARSHOLT
162 SHILAP Revta. lepid., 50 (197) marzo 2022
Figs 30-33.– 30. Nyctobrya vilfredi Falck & Karsholt, sp. n., 1, Gran Canaria, GP3294PF. 30a. Phallus, Gran
Canaria, GP3462PF. 31. Nyctobrya canaria (Alphéraky, 1890), 1, Tenerife, GP3292PF. 31a. Phallus, Tenerife,
GP3292PF. 32. Mniotype loslobensis (Fischer, Salditis & Ivinskis, 2007), phallus, Lanzarote, GP3287PF. 33.
Mniotype dimorpha (Rungs, 1948), phallus, Morocco, GP3385aPF.
30 30a
31 31a
32 33
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SHILAP Revta. lepid., 50 (197) marzo 2022 163
Figs 34-35.– 34. Agrotis corralejoi Falck & Karsholt sp. n., 1, Fuerteventura, GP3481PF. 34a. Phallus,
Fuerteventura, GP3481PF. 34b. Phallus with everted vesica, GP3477PF. 35. Agrotis boeticus (Boisduval, 1837),
1, Algarve, Portugal, GP3484aPF. 35a. Phallus, Algarve, Portugal, GP3484aPF. 35b. Phallus with everted
vesical, Algarve, Portugal, GP3475aPF.
34 34a
35
34b
35a 35b
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164 SHILAP Revta. lepid., 50 (197) marzo 2022
Figs 36-38.– 36. Nyctobrya vilfredi Falck & Karsholt, sp. n., 0, Gran Canaria, GP3300PF. 37. Nyctobrya
canaria (Alphéraky, 1890), 0, Tenerife, GP3304PF. 38. Mniotype loslobensis (Fischer, Salditis & Ivinskis,
2007), 0, Lanzarote, GP3478PF.
36 37
38
Fig. 39.– Neighbour-joining tree of Nyctobrya vilfredi Falck & Karsholt, sp. n., related Nyctobrya species and
Caradrina rebeli.
NEW DATA ON NOCTUOIDEA FROM THE CANARY ISLANDS, SPAIN
SHILAP Revta. lepid., 50 (197) marzo 2022 165
... The genus Nyctobrya Boursin, 1957 (Noctuidae) is treated in three relatively recent papers: Behounek & Speidel (2013), Fischer & de Freina (2014) and Falck & Karsholt (2022) with description of three new species. Hitherto six species are known: N. simonyi (Rogenhofer, 1889), N. canaria (Alphéraky, 1889), N. maderensis (Bethune-Baker, 1891), N. pinkeri Behounek & Speidel, 2013, N. hierroana Fischer & de Freina, 2014and N. vilfredi Falck & Karsholt, 2022 Fieldwork undertaken in the Canary Island, El Hierro from the end of July until the beginning of August 2022 by the author revealed several new records of Lepidoptera. ...
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