Spilomelinae

Spilomelinae is a very species-rich subfamily of the lepidopteran family Crambidae, the crambid snout moths. With 4,135 described species in 344 genera worldwide, it is the most speciose group among pyraloids.

Description

Imagines – the adult life stage – vary considerably in size: the forewing span ranges from 11.5 mm e.g. in '' Metasia'' to 50 mm in the robust-bodied '' Eporidia''. In resting position, the moths exhibit a characteristic triangular shape, with the wings usually folded over the abdomen, the forewings covering the hindwings. Some Spilomelinae diverge from this common resting pattern, like '' Maruca'' with widely spread wings, and '' Atomopteryx'' and '' Lineodes'' with narrow wings folded along the body. All Spilomelinae moths have well developed compound eyes, antennae and mouthparts, although in the genera '' Niphopyralis'' and '' Siga'' the proboscis is lost.

Synapomorphic characters of the subfamily comprise minute or obsolete maxillary palpi, ventrally projecting fornix tympani, and the female genitalia's ductus bursae with a weak sclerotization or a granulose texture. The moths are furthermore characterized by an often bilobed praecinctorium, pointed spinula, and the absence of chaetosemata and of a retinacular hook. A gnathos or pseudognathos can be present or absent and is therefore of little diagnostic value, except for several genera of Agroterini, where the gnathos has a well-developed medial process.

Food plants

Food plant use is diverse within Spilomelinae, ranging from ferns over gymnosperms to a wide spectrum of angiosperms. The larvae of '' Niphopyralis'' live in nests of weaver ants, where they feed on ant larvae. Steniini caterpillars are detritivores.

Many Spilomelinae tribes have a narrow food spectrum, with the larvae feeding on plants of only one or a few plant families, e.g. Lineodini on Solanaceae, Hydririni primarily on Sapindaceae and Convolvulaceae, and Trichaeini on Rubiaceae.

Economic impact

A number of Spilomelinae are considered "pest species", with their larvae feeding on a variety of economically important crops. Notable representatives are the genera of '' Leucinodes'' and '' Neoleucinodes'' with larvae feeding on Solanaceae, ''Cnaphalocrocis'' and '' Marasmia'' damaging Poaceae like '' Oryza'', '' Sorghum'' and '' Zea'', the legume pod borers of the genus '' Maruca'' on Fabaceae and Amaranthaceae, and ''Spoladea'', who feeds on a variety of different agriculturally important plant families.

The box tree moth, '' Cydalima perspectalis'', whose larvae feed on box trees, a prominent ornamental plant in many parks and gardens, has been accidentally introduced to Europe in the mid-2000s and to North America in 2018.

Systematics

Until the late 1990s, Spilomelinae were included in the subfamily Pyraustinae as tribe Spilomelini.

In the past, Spilomelinae were believed to be polyphyletic. However, a recent phylogenetic study by Mally et al. (2019), based on molecular genetic and morphological data, found the subfamily to be monophyletic. The study's authors furthermore proposed 13 tribes within Spilomelinae:
* Agroterini Acloque, 1897
* Asciodini Mally, Hayden, Neinhuis, Jordal & Nuss, 2019
* Herpetogrammatini Mally, Hayden, Neinhuis, Jordal & Nuss, 2019
* Hydririni Minet, 1982
*Hymeniini Swinhoe, 1900
* Lineodini Amsel, 1956
* Margaroniini Swinhoe & Cotes, 1889
* Nomophilini Kuznetzov & Stekolnikov, 1979
* Spilomelini Guenée, 1854
* Steniini Guenée, 1854
*Trichaeini Mally, Hayden, Neinhuis, Jordal & Nuss, 2019
* Udeini Mally, Hayden, Neinhuis, Jordal & Nuss, 2019
* Wurthiini Roepke, 1916

"Non-euspilomeline" tribes

Hydririni, Lineodini, Udeini and Wurthiini share several plesiomorphic characters with the sister group of Spilomelinae, the Pyraustinae. These plesiomorphies are: absence of longitudinal strips on the male abdominal segment 8; male genitalia with a straight to concave valva costa and a phallus with an evenly sclerotized apodeme; female genitalia with a lanceolate “ediacaroid” signum in the corpus bursae and in several taxa with an appendix bursae attached to the corpus bursae.

Due to these plesiomorphies, these four Spilomelinae tribes are referred to as "non-euspilomeline" tribes as opposed to the monophyletic "euspilomeline" group that represents a more derived group of Spilomelinae. The "non-euspilomeline" tribes form a paraphylum as they do not comprise the "euspilomelines".

"Euspilomeline" tribes

The tribes Agroterini, Margaroniini, Spilomelini, Herpetogrammatini, Hymeniini, Asciodini, Trichaeini, Steniini and Nomophilini form the monophylum of "euspilomelines" (Greek ''eu-'' for "good" or "true"), all sharing a common ancestor. The synapomorphies of euspilomelines are: male abdominal tergite 8 with an emarginate anterior edge; male genitalia with partly sclerotized hair pencils on the anterior edge of vinculum-tegumen connection, with a convex valva costa, and the phallus without a coecum and the sclerotization of the phallus apodeme reduced to a ventral, longitudinally sclerotized strip along the manica; female genitalia with a longitudinal membranous strip in the antrum, and the lack of a strongly sclerotised colliculum between antrum and ductus seminalis.

Genera currently unplaced in any Spilomelinae tribe

Based on the morphological synapomorphies and characteristics of these tribes, Mally et al. (2019) assigned numerous Spilomelinae genera to these proposed tribes, so that about two thirds of the 339 Spilomelinae genera are placed in these tribes, leaving 132 genera currently unplaced:

*'' Aboetheta'' Turner, 1914
*'' Acicys'' Turner, 1911
*''Aediodina'' Strand, 1919
*'' Agrammia'' Guenée, 1854
*'' Almonia'' Walker, 1866
*'' Ametrea'' Munroe, 1964
*'' Archernis'' Meyrick, 1886 (= ''Chrysommatodes'' Warren, 1896, ''Metoportha'' Meyrick, 1894, ''Protonoceras'' Warren, 1890)
*'' Arxama'' Walker, 1866
*'' Atelocentra'' Meyrick, 1884
*'' Auchmophoba'' Turner, 1913
*'' Bacotoma'' Moore, 1885 (= ''Epactoctena'' Meyrick, 1937, ''Platamonia'' Lederer, 1863, ''Platamonina'' J. C. Shaffer & Munroe, 2007)
*'' Cangetta'' Moore, 1886
*'' Carthade'' Snellen, 1899
*'' Chabula'' Moore, 1886
*'' Chromodes'' Guenée, 1854
*'' Cissachroa'' Turner, 1937
*'' Coelorhyncidia'' Hampson, 1896
*'' Coptobasis'' Lederer, 1863
*'' Coremata'' Amsel, 1956
*'' Cotachena'' Moore, 1885
*'' Criophthona'' Meyrick, 1884
*'' Camptomastix'' Warren, 1892 (= ''Camptomastyx'' Hampson, 1896)
*'' Cangetta'' Moore, 1886 (= ''Blechrophanes'' Turner, 1937)
*'' Carthade'' Snellen, 1899 (= ''Chartade'' Neave, 1939)
*''Cavifrons'' Zeller, 1872
*'' Ceratarcha'' Swinhoe, 1894
*'' Chabula'' Moore, 1886
*'' Chromodes'' Guenée, 1854
*'' Cissachroa'' Turner, 1937
*'' Coelorhyncidia'' Hampson, 1896
*'' Coptobasis'' Lederer, 1863
*'' Coremata'' Amsel, 1956 (= ''Culcita'' Amsel, 1957)
*'' Cotachena'' Moore, 1885 (= ''Mesothyris'' Warren, 1892, ''Syndicastis'' Meyrick, 1889)
*'' Criophthona'' Meyrick, 1884 (= ''Conoprora'' Turner, 1913)
*'' Daulia'' Walker, 1859 (= ''Girtexta'' Swinhoe, 1890)
*'' Deuterarcha'' Meyrick, 1884
*'' Deuterophysa'' Warren, 1889
*'' Dichocrocis'' Lederer, 1863 (= ''Zebrodes'' Warren, 1896)
*'' Discothyris'' Warren, 1895
*'' Dracaenura'' Meyrick, 1886
*'' Duzulla'' Amsel, 1952
*'' Ebuleodes'' Warren, 1896
*'' Ectadiosoma'' Turner, 1937
*'' Elbursia'' Amsel, 1950
*'' Eudaimonisma'' T. P. Lucas, 1902
*'' Eulepte'' Hübner, 1825 (= ''Acrospila'' Lederer, 1863)
*'' Eurybela'' Turner, 1908
*'' Eustenia'' Snellen, 1899
*'' Furcivena'' Hampson, 1896
*'' Gadessa'' Moore, 1885
*'' Gethosyne'' Warren, 1896
*'' Glauconoe'' Warren, 1892
*'' Goniorhynchus'' Hampson, 1896
*'' Hemopsis'' Kirti & Rose, 1987
*'' Heterudea'' Dognin, 1905
*'' Hyalea'' Guenée, 1854
*'' Indogrammodes'' Kirti & Rose, 1989
*'' Ischnurges'' Lederer, 1863 (= ''Nesolocha'' Meyrick, 1886)
*'' Legrandellus'' J. C. Shaffer & Munroe, 2007
*'' Lepidoneura'' Hampson, 1896
*'' Leucinodella'' Strand, 1918
*'' Leucochromodes'' Amsel, 1956
*'' Leucophotis'' Butler, 1886
*'' Lipararchis'' Meyrick, 1934
*'' Luma'' Walker, 1863 (= Pelena Moore, 1886, Pelina Hampson, 1897, ''Petena'' Neave, 1940)
*'' Mabra'' Moore, 1885 (type species: ''Asopia eryxalis'' Walker, 1859, by original designation)
*'' Macaretaera'' Meyrick, 1886 (= ''Trichoptychodes'' Swinhoe, 1894, ''Trigonophylla'' Turner, 1937)
*'' Macrobela'' Turner, 1939
*'' Malaciotis'' Meyrick, 1934
*'' Malickyella'' Mey & Speidel, 2010
*'' Meekiaria'' Munroe, 1974
*'' Merodictya'' Warren, 1896
*'' Mesocondyla'' Lederer, 1863
*'' Metallarcha'' Meyrick, 1884 (= ''Panopsia'' Turner, 1913)
*'' Metoeca'' Warren, 1896
*'' Metraeopsis'' Dognin, 1905
*'' Microgeshna'' J. C. Shaffer & Munroe, 2007
*'' Microphysetica'' Hampson, 1917 (= ''Falx'' Amsel, 1956, ''Falcimorpha'' Amsel, 1957)
*''Mimudea'' Warren, 1892
*'' Mukia'' Amsel, 1954
*'' Myriostephes'' Meyrick, 1884
*'' Myrmidonistis'' Meyrick, 1887
*''Nacoleia'' Walker, 1859 (= ''Aplomastix'' Warren, 1890, ''Orthocona'' Warren, 1896, ''Semioceros'' Meyrick, 1884)
*'' Nankogobinda'' Rose & Kirti, 1986
*'' Nausinoe'' Hübner, 1825 (= ''Lepyrodes'' Guenée, 1854, ''Phalangiodes'' Guenée, 1854)
*'' Nausinoella'' J. C. Shaffer & Munroe, 2007
*'' Neostege'' Hampson, 1910
*'' Nevrina'' Guenée, 1854
*'' Nistra'' Walker, 1859
*'' Notesia'' Yamanaka, 1992
*'' Ommatobotys'' J. C. Shaffer & Munroe, 2007
*'' Orphnophanes'' Lederer, 1863 (= ''Syntomodora'' Meyrick, 1894)
*'' Orthospila'' Warren, 1890
*'' Osiriaca'' Walker, 1866 (= ''Myriotis'' Meyrick, 1885)
*'' Otiophora'' Turner, 1908
*'' Paramecyna'' Amsel, 1961
*'' Pectinobotys'' Munroe, 1959
*'' Pelinopsis'' Dognin, 1905
*'' Physematia'' Lederer, 1863
*'' Piletocera'' Lederer, 1863 (= ''Alutefa'' Swinhoe, 1900, ''Danaga'' Moore, 1885, ''Diplotyla'' Meyrick, 1886, ''Ellogima'' Turner, 1913, ''Erebangela'' Meyrick, 1886, ''Graphicopoda'' Butler, 1886, ''Hormatholepis'' Butler, 1886, ''Ptilaeola'' Meyrick, 1886, ''Rinecera'' Butler, 1884, ''Sematosopha'' Meyrick, 1937, ''Strepsimela'' Meyrick, 1886)
*'' Plateopsis'' Warren, 1896
*'' Platygraphis'' Dyar, 1918
*'' Pleonectoides'' Hampson, 1891
*'' Polythlipta'' Lederer, 1863
*'' Praeacrospila'' Amsel, 1956
*'' Praephostria'' Amsel, 1956
*'' Pramadea'' Moore, 1888
*'' Preneopogon'' Warren, 1896
*'' Prionopaltis'' Warren, 1892
*'' Prorodes'' Swinhoe, 1894 (= ''Idiostrophe'' Warren, 1896)
*'' Proternia'' Meyrick, 1884
*'' Protonoceras'' Warren, 1890
*'' Pycnarmon'' Lederer, 1863 (= ''Aripana'' Moore, 1886, ''Entephria'' Lederer, 1863, ''Eutrichotis'' Swinhoe, 1900, ''Pyralocymatophora'' Strand, 1918, ''Satanastra'' Meyrick, 1890)
*'' Pylartes'' Walker, 1863
*'' Pyradena'' Munroe, 1958
*'' Ravanoa'' Moore, 1885
*'' Rehimena'' Walker, 1866
*'' Rhimphaliodes'' Hampson, 1893 (= ''Rhimphaleodes'' Hampson, 1896)
*''Rhynchetria'' Klunder van Gijen, 1913
*'' Sagariphora'' Meyrick, 1894
*'' Sameodesma'' Hampson, 1918
*'' Sedenia'' Guenée, 1854
*'' Sericophylla'' Turner, 1937 (= ''Sericophora'' Turner, 1937)
*'' Stenorista'' Dognin, 1905
*'' Syllepte'' Hübner, 1823 (= ''Arthriobasis'' Warren, 1896, ''Haitufa'' Swinhoe, 1900, ''Haliotigris'' Warren, 1896, ''Neomabra'' Dognin, 1905, ''Nothosalbia'' Swinhoe, 1900, ''Polycorys'' Warren, 1896, ''Subhedylepta'' Strand, 1918, ''Sylepta'' Hübner, 1825, ''Syllepta'' Hübner, 1826, ''Troctoceras'' Dognin, 1905)
*'' Syngropia'' Hampson, 1912
*'' Syntrita'' Dognin, 1905
*'' Tabidia'' Snellen, 1880
*'' Tanaophysopsis'' Munroe, 1964
*'' Tasenia'' Snellen, 1901
*'' Torqueola'' Swinhoe, 1906
*'' Trichoceraea'' Sauber in G. Semper, 1902
*'' Trigonobela'' Turner, 1915
*'' Trithyris'' Lederer, 1863
*'' Troctoceras'' Dognin, 1905
*'' Tylostega'' Meyrick, 1894
*'' Voliba'' Walker, 1866 (= ''Gabrisa'' Walker, 1866, ''Stereocopa'' Meyrick, 1885)
*'' Xanthomelaena'' Hampson, 1896
*'' Zagiridia'' Hampson, 1897

Genera excluded from Spilomelinae

'' Aporocosmus'' Butler, 1886 was transferred to Odontiinae, ''Orthoraphis'' Hampson, 1896 to Lathrotelinae, '' Hydropionea'' Hampson, 1917 and '' Plantegumia'' Amsel, 1956 to Glaphyriinae, and '' Prooedema'' Hampson, 1896 to Pyraustinae.

See also

* List of crambid genera

References

External links

{{Taxonbar, from=Q2310651

Moth subfamilies
Taxa named by Achille Guenée