Libythea celtis ( Laicharting [1782] )

Kawahara, Akito Y., 2013, <strong> Systematic revision and review of the extant and fossil snout butterflies (Lepidoptera: Nymphalidae: Libytheinae) </ strong>, Zootaxa 3631 (1), pp. 1-74 : 10-13

Libythea celtis ( Laicharting [1782])

( Figs. 2 , 14, 26, 39, 53, 69–72)

Papilio celtis Laicharting [1782] : 1, pl. 8, Figs. 1–3 .

Papilio cercis Fuessly [1782] .

Libythea celtis Fabricius 1807: 284 .

Hecaërge celtis Ochsenheimer 1816: 32 .

Hypatus feverus Hübner 1822: 3 .

Libythaeus [sic] celtis Boitard 1828: 299 [unjustified emendation].

Nymphalis phalerati celtis Esper 1829: 168 , pl. 87, Figs. 2, 3 .

Libythea celtis ab. obscura Millière 1879: 15 [nomen nudum].

Libythea (Libythea) celtis f. obscura: Millière 1879: 15, 16, pl. 6, Fig. 10 [new synonymy].

Libythea celtis ab. ochracea Fruhstorfer 1898: 170 [nomen nudum].

Libythea (Libythea) celtis: Pagenstecher 1901: 2 .

Libythea (Libythea) celtis ab. ochracea: Pagenstecher 1911: 5 [nomen nudum].

Libythea celtis ab. conjuncta: Schawerda 1924 [nomen nudum].

Libythea celtis ab. denudata Dannehl 1925: 64 [nomen nudum].

Libythea celtis ab. separata Dannehl 1925: 64 [nomen nudum].

Libythea celtis livida Saggara 1926: 135 [new synonymy].

Libythea celtis ab. pallida Dannehl 1927: 6 [nomen nudum].

Libythea celtis ab. pygmaea Dannehl 1927: 6 [nomen nudum].

Libythea celtis celtis: Gaede 1932: 237 .

Libythea (Libythea) celtis pygmaea: Verity 1950: 5 .

Libythea (Libythea) celtis denudata: Verity 1950: 5 [new synonymy].

Libythea (Libythea) celtis separata: Verity 1950: 5 [new synonymy].

Libythea (Libythea) celtis f. subochracea Verity 1950: 5 [new synonymy].

Libythea (Libythea) celtis f. violacea Verity 1950: 5, pl. 38, Figs. 3, 5 [new synonymy].

Libythea (Libythea) celtis f. albonervulata Verity 1950: 5, pl. 38, Fig. 5 [new synonymy].

Libythea (Libythea) celtis f. obscurenervulata Verity 1950: 5, pl. 38, Fig. 6 [new synonymy].

Libythea (Libythea) celtis f. pygmaea: Verity 1950: 5, pl. 38, Fig. 7 [new synonymy].

Libythea (Libythea) celtis f. rubescens Verity 1950: 6 [new synonymy].

Libythea (Libythea) celtis f. pallida: Verity 1950: 6 [new synonymy].

Libythea (Libythea) celtis f. latefulva Verity 1950: 6, pl. 38, Fig. 4 [new synonymy].

Libythea celthis [sic] Kostrowicki 1969: 352.

Libythea celtis f. conjuncta: Gómez-Bustillo and Fernández-Rubio 1974: 166.

Libythea celtis f. denudata: Gómez-Bustillo and Fernández-Rubio 1974: 166.

Libythea celtis f. separata: Gómez-Bustillo and Fernández-Rubio 1974: 166.

Libythia [sic] celtis Roberts 2001: 121 .

Libythea celtis platooni Korb 2005: 18 [new synonymy].

Diagnosis. Libythea celtis is sister species with L. lepita by sharing a gradually incrassate antenna, but the former can be distinguished from the latter by having: 1) a hindwing anterior margin that is distinctly lobed near the fusion of the costa and Sc + R 1, and 2) the proximal margin of the forewing dorsal orange mark between M 3 and CuA 2 is straight in L. celtis , but projects proximally along CuA 1, forming a curved or angled margin in L. lepita (Kawahara 2009) .

Type. Neotype (present designation): AUSTRIA: ‘[red label] : Neotype / [printed white label]: ♂, det. A. Y. Kawahara / [original hand written white label]: 1895, Z7, Jün-Tirol’ ( BMNH). This specimen is designated as the type to clarify the taxonomic status of this species. A specimen from the type locality that resembles the holotype of Laicharting ([1782] Figs. 1–3 ) was chosen. The depository of the original name-bearing type is unknown .

Other Specimens Examined. (158♂, 99♀): AFGHANISTAN: Arwas, Transcasp , 1♀, VI ( BMNH) . AUSTRIA: 1♂, B. Neumögen ( USNM) . Steiermark, Ritzenberg : 2♀ ( AMNH) . Tirol: 1♂ ( AMNH) ; 1♀, 1890, Frey ( BMNH) ; 8♂, 1♀, 1901 ( BMNH) ; 2♀, Bates ( BMNH) ; 1♀, J. Doll ( BMNH) ; South Tirol: 2♂ ( CUIC) ; 2♂, VI-1895 ( AMNH) ; 1♂, 17-VII-1909, M. Ibarra ( AMNH) ; 1♂, 1♀, 12-15-VII-1927, M. Ibarra ( AMNH) . Eggenthal , 3♂, VII-1906, Hampson ( BMNH) . Hérault, Montpellier , 1♂, 1890, Frey ( BMNH) . Provence Alpes- Cot, Grimaud , 1♀, 5-IV-1921, A. F. Pinhey ( BMNH) . Pyrénées-Orientalis , Amélie-les-Bains-Palalda, 1♀, 1908, Wism ( BMNH) . Vernet-les-Bains , 1♂, VII-1909, R. Oberthür ( BMNH) . Roque-brune , 6♂, ♀, 24-III-1903 ( BMNH) ; 2♀, VII-1904 ( BMNH) . Sainte Maxime , 4♂, 4♀, 10-15-VI-1921, A. F. Pinhey ( BMNH) ; 1♂, 11-V- 1921, A. F. Pinhey ( BMNH) ; 1♂, 1♀, 7-13-VI-1925, C. Va [n]s ( BMNH) . Var, Hyères , 2♂, 1♀, 30-31-V-1923, A. Simmons ( BMNH) . [Austria], Klausen , 5♂ 2♀, 1904, Fruhstorfer ( BMNH) ; BULGARIA: Bulgaria , Varna: 2♂, 26-VII-1955, B. Weber ( MGCL) . 1♂, 1♀, VI-1962, Vanek ( MGCL) . FRANCE: 1♀, 14-VI-1921, P. Haig- Thomas ( AMNH) . Montpellier : 1♀, VI-1952, K. de Puységur ( CMNH) . Environs de Montpellier : 1♂, ( MGCL) . Sainte Maxime : 1♀, 10-VI-1921, A. F. Pinhey ( BMNH) ; 3♀, VI-1925, P. Haig-Thomas ( AMNH) ; 1♂, 4-VI- 1926, P. Haig-Thomas ( AMNH) . St. Georges : 1♀, VI-1935 ( TFC) . GERMANY: 3♂, 2♀ ( AMNH) ; 2♂, H. Edwards ( AMNH) . GREECE: 1♂, 16-VI-1869 ( BMNH) ; 3♂, 1901 ( BMNH) ; [Macedonia], 1♂, 10-VII-1896 ( BMNH) ; Acarnan , 1♂, 8-VI-1869 ( BMNH) ; 1♀, 8-VI-1862 ( BMNH) ; Olympus , 2500 ft [762 m]; 10♂, 4♀, VII-VIII-1935, Romei ( BMNH) ; Struma Valley, Kopriva , 2♀, 7-VI-1917, D. Blanchard ( BMNH) . HUNGARY: Pilis Szent László : 1♂, 15-IV-1962, Gaskó ( CAS) . IRAN: Shush : 1♂, VI-1899 ( MGCL) ; Karind Gorge , 6000 ft [1829 m], 1♀, 16-IV-1918, H. D. Peile ( BMNH) . IRAQ: Dahuk, Amadia , 3000-9000 ft, [914-2743 m], 1♀, 12- VI-30-VII-1936 ( BMNH) . ITALY: 1♂ ( AMNH) ; 1♂ ( CMNH) . Sardinia: 1♂, 1♀, A. H. Kraus ( AMNH) . Libiveiz : 1♂, 1953, Freyer ( USNM) . Bozen : 1♂, VI-1903 ( MGCL) ; 1♂, 21-VI-1904, F. E. Lowe ( MGCL) ; Tuscany, 2♂, 1914-1915 ( BMNH) ; South Tyrol, Abswang : 1♀, VI-1898, Hayne ( MGCL) . KYRGYZSTAN: Sary-Chelek Nature Reserve, Ail. Arkit , 1300 m: 2♂, 20-VII-2000, V. V. Dubatolov ( AYK) . PAKISTAN: Chitral , 1♀, 1903, Leslie ( BMNH) ; 7000, [2134 m], 1♂, V-1901 ( BMNH) ; Drosh , 6000-10,000, [1829-3048 m], 1♂, 23-VI-1903, A. R. C. Saunders ( BMNH) ; Utzen Nallah , 9000 ft [2743 m], 1♂, 9-VIII-190[?], G. A. Leslie and W. H. Evans ( BMNH) . REPUBLIC OF CYPRUS: Platres , 5000-6000 ft [1524-1829 m], 1♂, IX-1921, K. J. Hayward ( BMNH) . ROMANIA: Herculesbad , 2♀, 21-22-VI-1909, W. G. S. ( BMNH) . RUSSIA: Kirgizia, Kirgizaskii Ridge, Ala-Archa , Ak-Sai , 2300 m: 1♂, 17-21-VII-1979, J. A. Vaněk ( TFC) . Khojakent , 3000 m: 1♀, 7-VII-1919 ( AMNH) . Ugam Valley , 8000 m: 1♂, 8-VII-1919 ( AMNH) . SPAIN: Barcelona, Sallent : 1♂, 2- VII-1989, E. Casas ( USNM) . Barcelona, Montserrat, nr. Car terminus: 1♂, 3000 ft [1181 m], 14-VII-1901, W. Holland ( MGCL) ; 1♀ 3000–4000 ft [1181–1575 m], 15-VII-1901, E.B. Poulton ( MGCL) . Catalonia, Oliana : 1♀, III-1946, M. Ibarra ( AMNH) ; 2♂, III-1949, M. Ibarra ( AMNH) ; 1♂, VI-1949, M. Ibarra ( AMNH) . Catalonia, Sierra de Montseny, El Brull , 2600 ft [793 m]: 1♂, 13-VII-1948 ( CAS) ; 1♂, 13-VII-1958 ( LACM) . Gerona, Olot : 1♂, 23-VII-1988, I. Teixodor ( USNM) . l’Ametlla : 1♂, III-1929, M. Ibarra ( AMNH) ; 2♂, 4♀, IV- 1929, M. Ibarra ( AMNH) ; 2♀, IV-1935, M. Ibarra ( AMNH) . La Garriga : 11♂, 9♀, VI-1934, M. Ibarra ( AMNH) ; 6♂, 15♀, VI-1935, M. Ibarra ( AMNH) ; 1♀, 4-IV-1936, M. Ibarra ( AMNH) ; 5♂, 3♀, VI-1936, M.

Ibarra ( AMNH); 1♂, VI-1937, M. Ibarra ( AMNH) . SWITZERLAND: Ticino, Lugano , 1♀, 11-IV-1898, H. J. Elwes ( BMNH) ; 1♂, 1914-1915, F. D. Godman ( BMNH) . SYRIA: Shar-Deresy , 1♂, 1880, native collector ( BMNH) . TURKESTAN: Ugam Valley , 3000 [ft], [914 m] , 1♂, 8-VII-1919 ( BMNH) . TURKEY: Unknown locality : 1♂, Bates ( BMNH) ; [Canakkale], Erenkeui , 1♀, 22-IV-1911, P. P. Graves ( BMNH) ; Amasia, 1♂, 1♀, Franks ( BMNH) ; 1400 ft [427 m], 1♂, 9-VI-1959, K. M. Guichard ( BMNH) ; Asia Minor (Anatolia), Kedos , 3♂, 1♀, IV-VIII-1918, N. V. L. Rybot ( BMNH) ; Gallipoli (Gelibolu) , 1♂, 1-I-10-VIII-1923, W. M. J. Martin ( BMNH) ; Istanbul, Therapia , 1♀, 18-VI-1920, J. A. Graham ( BMNH) ; Muğla, Baba Dag , 6000 ft [1829 m] , 3♂, 3♀, 29-VII-1947, M. Burr ( BMNH) ; Sandras Dag , [2200 m] , 1♂, 23-VIII-1947, M. Burr ( BMNH) . UZBEKISTAN: Medeo Tien Shan Mountains , 1♂, 28-VIII-1978 ( CMNH) . Uncertain country : 3♂, 3♀ ( AMNH) ; 4♂, 1♀ ( BMNH) ; 9♂, 3♀ ( USNM) ; 4♂, 13-14-VII ( USNM) ; 1♂, 3-V-1908 ( MGCL) ; 1♂, 1914, F. E. Lowe ( MGCL) ; 1♂, 18-VI-1916 ( MGCL) ; 1♂, 30-VI-1975 ( TFC) ; 1♀, ‘ Europe’ ( MGCL) ; ‘ Southeastern Europe :’ 1♂, F. R. Mason ( CMNH) .

Distribution. Southern Europe and northwestern Africa ( Algeria and Tunisia) to southern Russia and western India.

Biology. Primary larval host plants include: Celtis australis L., C. caucasica L., C. formosana Hayata , C. glabrata Steven ex Planch. , and C. tournefortii L. ( Celtidaceae ), For additional host plant records and a detailed discussion on biology, see Kawahara (2006a).

Remarks. The holotype is believed to have been lost, and specimens from the original type series could not be located during this study. For this reason, a neotype is designated. Numerous taxonomic studies included lepita as a subspecies of L. celtis (e.g., Pagenstecher [1901, 1902, 1911]; Shields [1984a]; Okano [1987]). Yoshimoto (1999) discussed the possibility of L. celtis and L. lepita as distinct species. In a book on the butterflies of Pakistan, Roberts (2001) treated the two species as allopatric. Since there are morphological differences between the two, and there are no intermediates found in northern India and Pakistan, the western population (Western India, Pakistan to Europe) is here recognized as L. celtis , and the eastern population (east India to Japan and Taiwan) as L. lepita . Phylogenetically, L. celtis and L. lepita are sister species, being grouped together because both share a gradually incrassate antenna (Kawahara 2009). Two characters distinguish L. celtis from L. lepita : 1) the prominent lobe along the hindwing anterior margin near the fusion of the costa and Sc + R 1, and 2) the straight proximal margin of the forewing dorsal orange mark between M 3 and CuA 2. An orange mark on the dorsal surface of forewing 1A + 2A is always present on L. celtis , but this mark can be absent in L. lepita . For additional discussion on the delimitation of these two species, see the ‘remarks’ section of L. lepita . Additional illustrations of the external morphology of L. celtis can be found in Kawahara et al. (2012).

Figures

FIGURE 1. Phylogeny of snout butterflies based on morphology (after Kawahara 2009). Numbers above and below each rectangle refer to the characters and states in Kawahara (2009). A dark rectangle indicates a unique character state change (synapomorphy), a white rectangle indicates homoplasy. Large numbers indicate branch support (Bremer Support/jackknife). Due to limited space, outgroups are not shown in this figure.

FIGURES 2−13. Lateral view of the male eighth abdominal tergum of Libythea and Libytheana with scales and hairs removed. 2. Libythea celtis; 3. Libythea collenettei; 4. Libythea geoffroyi; 5. Libythea labdaca; 6. Libythea laius; 7. Libythea lepita; 8. Libythea myrrha; 9. Libythea narina; 10. Libytheana carinenta; 11. Libytheana fulvescens; 12. Libytheana motya; 13. Libytheana terena.