ZEITSCHRIFT FÜR ENTOMOLOGIE. Thomas J. Witt

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ßntomojauna ZEITSCHRIFT FÜR ENTOMOLOGIE Band 1, Heft 18 ISSN O25O-4413 Linz, 15.Dezember 198O Description of a new species and new subspecies of the genus Anaea Hübner, 1819, from the Yucatän peninsula, with notes on other species (Lepidoptera, Nymphalidae, Charaxinae) Thomas J. Witt Abstract In this paper Anaea (Memphis) maya sp.n. and Anaea (Memphis) glycerium yucatanum ssp.n., both taken by Mr E.C.Welling in the Yucatän peninsula,mexico, are described. The unknown female of Anaea (Memphis) fumata HALL, 1935, is described and the identity of the questionable female of Anaea (Memphis) aureola (BATES, 1866) is clarified. Zusammenfassung In der vorliegenden Arbeit werden eine neue Art, Anaea (Memphis) maya sp.n. und eine neue Subspezies, Anaea (Memphis) glycerium yucatanum ssp.n.,die beide von Herrn E. C. W e l l i n g auf der Halbinsel Yucatan, Mexico, erbeutet worden sind, beschrieben. Das unbekannte Weibchen von Anaea (Memphis) fumata HALL,1935,wird beschrieben, und- die Identität des- fraglichen Weibchens von Anaea 366

(Memphis) aureola (BATES, 1866) wird geklärt. 1. Anaea (Memphis) maya, sp.n. In 1978 I received a series of nine specimens belonging to the halice-qroup of the genus Anaea HÜBNER, 1819 (sensu Comstock, 1961) from Mr E.C. W e 1 1 i n g, Merida, Yucatän. Three of them proved to belong to Anaea martinezi DE LA MAZA & DIAZ.FRANCES,1978, occuring north of Oaxaca, Mexico. Six of them together with two specimens from the collection of the Allyn Museum, Sarasota, Florida, and one specimen from the collection of the British Museum (Nat.Hist.), London, already mentioned by Witt (1972), proved to represent a new species occuring in the surroundings of Piste in the Yucatän peninsula. One male was taken in Chiltepec, Oaxaca, a locality from where Anaea martinezi DE LA MAZA & DIAZ FRANCES was taken. Holotype d: Mexico, Yucatän, Municipio-Tinüm, Piste, 3O. 8.1957, leg. E. C. W e 1 1 i n g, coll. Witt, Munich (genitalia slide no. 1575, T. J. W i t t, plate 7, fig. 1). Paratypes: 1 d Mexico, Yucatän, Municipio Tinüm, Piste, 29.8.1957, leg. E. C. W e 1 1 i n g, coll. W e 1-1 i n g, Merida (genitalia slide no. 1554, T. J. W i t t) ; 1 d Mexico, Yucatän, Municipio Tinüm, Piste, 15.8. 1960, leg. E. C. W e 1 1 i n g, coll. W e 1 1 i n g, Merida (genitalia slide no. 1576, T. J. W i t t); 1 d Mexico, Yucatän, Municipio Tinüm, Piste, 6.9. 1960, leg. E. C. W e 1 1 i n g, coll. W e 1 1 i n g, Merida (genitalia slide no. 1555, T. J. W i t t),- 1 d Mexico, Yucatän, Piste, 25.10.1959, leg. E. C. W e 1 1 i n g, coll. Allyn Museum, Sarasota (genitalia slide no. 2580, L. D. M i 1 1 e r),- 1 d Mexico, Chichan Itza, 2.12.1955, leg.e.c. W e 1-1 i n g, coll. British Museum (Nat. Hist.), London (genitalia slide no. 1966-687, T. G. H o w a r t h); 1 d Mexico, Yucatän, Municipio Tinüm, Piste, 24.10. I960, leg. E.C. W e 1 1 i n g, coll. Witt, Munich (genitalia slide no. 1552, T. J. W i t t); 367

1 ö* Mexico, Yucatän, Municipio Tinüm, Piste, 27.11. I960, leg. E. C. W e 1 1 i n g, coll. W e 1 1 i n g, Merida (genitalia slide no. 1553, T. J. W i t t),- 1 ö* Mexico, Yucatän, Municipio Tinüm, Piste, 30.9. 1978, leg. E. C. W e 1 1 i n g, coll. W e 1 1 i n g, Merida; 1 ö* Mexico, Oaxaca, Chiltepec, 15.8.1964, leg. E. C. W e 1 1 i n g, coll. Allyn Museum, Sarasota (genitalia slide no. 2581, L. D. M i 1 1 e r). Male: ground-colour of upperside reddish brown, without violet sheen, brighter than all other species of the halice-group. Colour-pattern of forewings as in fumata HALL, but outer margin at right angles to inner margin. In fumata the outer margin forms an obtuse angle with the inner margin. Outer margin paler than in fumata and martinezi. Band connecting costal spot with outside margin absent, unlike rest of species-group. Shape of forewing more compact than in martinezi; the apex is not extended. There is a small white-centred black spot in the middle of the hindwing located in the area between m^ and cuj or between m2 and m^, too. Underside of forewing uniform in colour-pattern; hindwing with darker band separating the row of black spots along outer margin from the inner area. There are usually spots (one in each cell) along outer margin of the hindwing, as on the upperside. There are two males collected in 24.10.1960 and 27.11. 1960 that differ from the typical form as described above: the band connecting the costal spot with the outer margin in typical specimens is represented by dark spots, variously extended in each cell but not linked together. The underside has more white spots than typical specimens. Measurements. Holotype: wingspan 45 mm, forewing 23mm. Paratypes: wingspan 42-46 mm, forewing 23-25 mm. The author (1972) examined the material of Anaea fumata HALL in the British Museum (Nat. Hist.), London. Besides recording A. fumata as new for Mexico (Piste) and Nicaragua, I noted a male from Chichan Itza, Yucatän, differing from fumata in the absence of a band on the forewing. The genitalia of this male differed in the shape 368

of the saccus, valvae and uncus from Comstock's (1961) fig.163 of fumata. In Order to confirm these differences all available males have been dissected showing that there is a considerable individual Variation and that it may not be possible to separate maya sp.n. and fumata HALL on male genitalia. Three males of A. martinezi DE LA MAZA & DIAZ FRANCES have been dissected: 1 ö* Mexico, Oaxaca, Chiltepec, 1O.2.1963, leg. E. C. W e 1 1 i n g, coll. W e 1 1 i n g, Merida (genitalia slide no. 1577, T. J. W i t t, plate 7, fig. 2); 1 cj Mexico, Oaxaca, Chiltepec, 8.10.1961, leg. E. C. W e 1 1 i n g, coll. Witt, Munich (genitalia slide no. 155O, T.J.Witt); 1 (5 Mexico, Municipio Comaltepec, Soloyapan el Bajo, Oaxaca, 200 m, July 1963, leg. E. C. W e 1 1 i n g, coll. W e 1 1 i n g, Merida (genitalia slide no. 1551, T. J. Witt). No differences in male genitalia between martinezi and maya could be found. The halice-group now consists of eight species: A. tehuana'hall, 1917, A. chrysophana (BATES,1866), A. fumata HALL, 1935, A. maya sp.n., A.martinezi DE LA MAZA & DIAZ FRANCES, 1978, A. moretta (DRUCE, 1877), A. moretta wiegeliana WITT, 1970, A. evelina JOHNSON & COMSTOCK, 1961, A. halice (GODARD, "1819", 1824). Comstock, 1961, considers the haiice-group as "superspecies", a group of closely related species. Witt (1972) considers the group as a very recently evolved monophyletic complex of species which still is Splitting into new species. My studies of Anaea moretta and evelina showed that there are populations that cannot be asigned with certainty to either species, a clear example of "species in statu nascendi". More extensive material and knowledge of the early stages and foodplants is required allowing insight into the Variation between different generations. As suggested by Mr W e 1 1 i n g, the new species is named after the Maya people who have helped him very much in obtaining many hundreds of thousands of butterflies during the 24 years he spent collecting in the Yu- 369

catän peninsula. 2. Anaea (Memphis) glycerium yucatanum, ssp.n. Anaea (Memphis) glycerium (DOUBLEDAY, 1850) is a widespread species. Records listed bycomstock (1961) include Arizona, Mexico, Guatemala, Nicaragua, Costa Rica, Panama and the Panama Canal Zone. Anaea (Memphis) glycerium comstocki WITT, 1972, was described from spe-- cimens collected in Venezuela, Colombia and French Guiana. This subspecies differs from the nominate subspecies in the paler ground-colour of both sexes and the larger size. In 1978 I received two pairs of Anaea glycerium from Mr W e l l i n g who took them in the Yucatän peninsula. He wrote: "They are exceedingly rare here. We have always taken one or two every five years or so, among many tens of thousands of specimens." These specimens represent the first record for the Yucatän peninsula. They prove to be subspecifically distinct from the nominate race and ssp.comstocki WITT and are described as follows: Holotype cj: Mexico, Yucatän, Nuevo X-Can, Quirftana Roo, 3.11.1978, leg. E. C. W e 1 1 i n g, coll. W i t t, Munieh. Allotype 9: Mexico, Yucatän, Municipio Tinüm, Piste, 19. 8.1978, leg. E. C. W e 1 1 i n g, coll. Witt, Munich. Paratypes: 1 6 Mexico, Yucatän, Municipio Tinüm, Piste, 13.9.1978, leg. E. C. W e 1 1 i n g, coll. W i t t, Munich; 1 9 Mexico, Yucatän, Nuevo X-Can, Quintana Roo, 17. 9.1978, leg. E. C. Welling, coll. W i t t, Munich. Male: Ground-colour of upperside slightly darker than in nominate subspecies and ground-colour of underside darker. Measurements. Anaea glycerium males from Mexico ränge from 27 to 34 mm, males of yucatanum ssp.n. from 28 to 29 mm. Female: Colour-pattern as in nominate subspecies, but marginal band and apical area dark brown, the area between them strong ochre brown in contrast to glycerium 370

where this area is yellowish ochre and comstocki where it is paler. The inner area proximal the brown margin, which is dark-brown, is also strong ochre-brown towards the base in contrast to glycerium and comstocki. On the hindwing this strong ochre-brown colour is present at the base and is more extensive along the anal area and towards the outer margin; Single small white-centred black spots are present between veins 1113 and cui and CU2 and two spots between CU2 and pcu. An important difference is that in glycerium the outer margin of the hindwings is a strong ochre-brown while in yucatanum ssp.n. this outer margin is identical in colour to the ground colour of the wing. The pattern of the underside is more densely speckled and darker brown. Measurements. Length of forewing ranges from 32 to 33 mm. That of nominate subspecies ranges from 3O to 37 mm. 3. The female of Anaea (Memphis) fumata HALL, 1935 From Mr H. G e r s t n e r, Schweinfurt, West Germany, who has sent me important material of the genus Anaea HÜBNER for years, I received an interesting series of A. fumata from Venezuela containing the hitherto unknown female of fumata. The distribution of this species was represented by C o m s t o c k (1961) and Witt (1972). 5 66 3 99 Venezuela, Pick Avila, 16. and 19.12.1973, coll. Witt, Munich (one male genitalia slide no. 1556, T. J. W i t t). The female belongs to the group of females without white spots on the forewings, like tehuana and chrysophana. The' band connecting the costal spot with the outside margin is unbroken; but the outside margin forms an obtuse angle with the inner margin as in the male figured in C o m s t o c k (1961) and the band is a little diffuse towards the outside margin. The ground-colour is paler than in the male, but is unlike that of the yellowish female of A.chrysophana BATES as figured in Comstock (1961, plate 12, fig.4). The wing shape of the female of A.tehuana HALL is more compact,the tail of the hindwing (vein 1113) is shorter and broader in tehuana. 371

4. The female of Anaea (.Memphis) aureola (BATES, 1866) From Mr Gerstner I received 2 ö\3 1 9 of this rarely captured species, taken by one of his collectors and labelled "Mexico,Puebla, September 1979". The female of aureola was unknown to C o m s t o c k (1961). It was first described by G o d m a n & S a l v i n (1884) in Biologia Centrali-Americana. The hand-coloured plate shows a specimen with dark green ground-colour and a broad, diagonally transverse band on the forewing. Although the latin description refers to the colour of this band as white ("anticis autem fascia transversa albida ultra cellulam"), the band of the specimen figured is of a pale green colour. Röber (1916) in: Seitz, Die Gross-Schmetterlinge der Erde, figured a female with a white band on the forewing. In addition to this difference to the illustration in G o d m a n & Salvin's female, the tail (1113) is more than twice as long in the R ö b e r figure. The picture in Seitz shows the metallic sheen of the fore- and hindwings very well.in the Godman & S a l v i n illustration this sheen is represented by another tone of green. Mr R. S m i 1 e s, British Museum (Nat. Hist.), London,was so kind as to send me coloured diapositives of the two females deposited in the British Museum's collection. I was surprised to be able to identify one of them to be the Single female mentioned and figured in Biologia Centrali-Americana: 1 9 Polochic Valley, F. D. G. & 0. S.; R. Polochic, Vera Paz, H a g u e; sp. figured; B. C. A. Lep. Rhop., Anaea aurelia, Bates. G o d m a n - S a l v i n Coll. 1917-2; The transverse band proved to be rather white and is little speckled with metallic light blue scales and differs from the hand-coloured drawing by the colour of the band which is of a pale green colour. This difference only can be interpreted as the result of wrong hand-colouring. The tails (1113) are broken. 1 9 Vulcan Chiriqui; J o i c e y Bequest Brit. Mus. 1934-12O; This second female has very long tails. The white band 372

is raore extensively speckled with metallic blue scales, the underside is darker. The transverse band of my specimen is white, there are metallic blue scales and there is no doubt that these three females belong to the same species. Like other Anaea-species, the underside is highly variable. The extension of the metallic blue scales in the white band and of the metallic sheen on the fore- and hindwings is variable, too. According to the collector, my males and the female were taken in the same locality in September and I am satisfied that the sexes are properly matched. Because of the confusion resulting from the differing illustrations in Godman & Salvin and R ö- ber inrseitz, Comstock (1961) was unable to clarify the identity of the female. As the work of Comstock was done during and shortly after the Second World War, he had little occasion to have a look into the British Museum's collections. Acknowledgements The author is grateful to Mr E. C.Welling, Merida, for loans and gifts of material, to Mr H. G e r - s t n e r, Schweinfurt,for delivering the fumata-series, to Dr L. D. Miller, Allyn Museum, Sarasota, for sending photographs of the questionable specimens, to Mr R. S m i 1 e s, British Museum (Nat. Hist.), London, for sending diapositives of the females of Anaea aureola, to Mr E. Lehmann, Zoologische StaatsSammlung, Munich, for dissecting specimens, to Mr B. Aus s e m, Großhelfendorf, for drawing the genitalia, to Mrs M. M ü 1 - ler, Zoologische Staatssammlung, Munich, for her photographic assistance, to Dr W. F o r s t e r, Zoologische Staatssammlung, Munich, for lending me the paper of de la Maza and Diaz Frances and to Mr A. W a t s o n, British Museum (Nat. Hist.), London, for reading the manuscript. 373

References C o m s t o c k, W. P. - 1961. Butterflies of the American Tropics. The genus Anaea, Lepidoptera, Nymphalidae. - New York, American Mus.Nat.Hist.: V-XIII+ 214 pp. ill. Godman, F. D. & 0. Salvin - 1884. Biologia Centrali-Americana, Insecta, Lepidoptera-Rhopalocera, London, Vol.I. de la Maza, E. J. & A. Diaz Frances - 1978. Dos nuevas especies del genero Anaea Hübner de Mexico (Nymphalidae: Charaxinae). - Rev.Soc.Mex. Lep., Vol.IV, No.1:29-38. R ö b e r, J. - 1916. in: Seitz, Die Gross-Schmetterlinge der Erde. - Stuttgart, Vol.5, p.591. W i t t, T. J. - 1972. Beiträge zur Kenntnis der Gattung Anaea Hübner, 1819 (Lep.,Nymphalidae). -Mitt.Münchner Ent.Ges., Vol.62:163-183. author's address: Thomas J. W i t t Tengstr. 33 D-8OOO Munich 40 West-Germany 374

Plate 1 Fig.l: Anaea maya sp.n. d, Holotype Mexico, Yucatän, Mpio. Tinüm, Piste, 30.8.1957, leg. W e 1 1 i n g, coll. Witt, Munich (gen. slide no. 1575, T. J. W i t t) Fig.2: Anaea maya sp.n. d, Paratype Mexico, Yucatän, Mpio. Tinüm, Piste, 29.8.1957, Leg. W e 1 1 i n g, coll. W e 1 1 i n g, Merida (gen. slide no. 1554, T. J. W i t t) Fig.3: Anaea maya sp.n. d, Paratype Mexico, Yucatän, Mpio. Tinüm, Piste, 6.9.1960, leg. W e 1 1 i n g, coll. W e 1 1 i n g, Merida (gen. slide no. 1555, T. J. W i t t) Fig.4: Anaea maya sp.n. d, Paratype Mexico, Yucatän, Mpio. Tinüm, Piste, 24.10.1960, Munich (gen. leg. W e 1 1 i n g, coll. Witt, slide no. 1552, T. J. W i t t) Fig.5: Anaea martinezi DE LA MAZA & DIAZ FRANCES, 1978 d Mexico, Oaxaca, Chiltepec, 8.10.1961, leg. W e 1-1 i n g, coll. W i t t, Munich (gen. slide no. 155O, T. J. W i t t) Fig.6: Anaea martinezi DE LA MAZA & DIAZ FRANCES, 1978 d Mexico, Oaxaca, Chiltepec, 10.2.1963, leg. W e 1-1 i n g, col?.. W e 1 1 i n g, Merida (gen. slide no. 1577) Fig.7: Anaea fumata HALL, 1935 d Venezuela, Pick Avila, 19.12.1973, coll. Witt, Munich Fig.8: Anaea fumata HALL, 1935 9 Venezuela, Pick Avila, 16.12.1973, coll. Witt, Munich Plate 2 underside of plate 1 Mu- Plate 3 Fig.l: Anaea glycerium glycerium (DOUBLEDAY, 1850) d Mexico, Morelos, October 1968, coll. Witt, nich 375

Fig.2: Anaea glycerium glycerium (DOUBLEDAY, 185O) 9 Mexico, Puerto ELigio, Mpio. Oaxaca, Comaltepec, 7DOm,7.9.1964, leg. W e 1 1 i n g, coll. W i t t, Munich Fig.3: Anaea glycerium yucatanum ssp.n. d, Holotype Mexico, Yucatän, Nuevo X-Can, Quintana Roo, 3.11. 1978, leg. W e 1 1 i n g, coll. Witt, Munich Fig.4: Anaea glycerium yucatanum ssp.n. 9, Allotype Mexico, Yucatän, Mpio. Tinüm, Piste, 19.8.1978, leg. W e l l i n g, coll.witt, Munich Fig.5: Anaea glycerium comstocki WITT, 1972 d, Holotype Colombie, Pereira (Cauca), coll. Witt, Munich Fig.6: Anaea glycerium comstocki WITT, 1972 9, Allotype Colombie, Pereira, Cauca, coll. Witt, Munich Plate 4 underside of plate 3 Plate 5 Fig.l: Anaea aureola (BATES, 1866) d Mexico, Puebla, Sept. 1979, coll. Witt, Fig.2: Anaea aureola (BATES, 1866) 9 Mexico, Puebla, Sept. 1979, coll. Witt, Munich Munich Plate 6 Plate 7 underside of plate 5 Fig.l: Anaea maya sp.n. d, Holotype Mexico, Yucatän, Mpio. Tinüm, Piste, 30.8.1957, leg. Welling, coll. Witt, Munich (gen. slide no. 1575, T. J. W i t t) Fig.2: Anaea martinezi DE LA MAZA & DIAZ FRANCES, 19 78 d Mexico, Oaxaca, Chiltepec, 10.2.1963, leg. Welling, coll. Welling, no. 1577, T. J. W i t t) Fig. 3: Anaea fumata HALL,. 1935 ö* Matagalpa, Nicaragua, coll. L e M o u 1 Witt, Munich (gen.. s.lide no.138, T.J. 376 Merida (gen. slide coll. i t t)

Plate 1 WL 377

Plate 2 4 ß 378

Plate 3 379

Plate 4 380

Plate 5 381

Plate 6 382

Plate 7 1) Gen Präparat 1575 2) Gen Präparat 1577 3) Gen Präparat 138 383

Druck: im Eigenverlag Eigentümer, Herausgeber, Verleger und für den Inhalt verantwortlich: Maximilian Schwarz, Konsulent für Wissenschaft der O.ö. Landesregierung, Eibenweg 6, A-4052 Ansfelden Redaktion: Erich Di1ler, DenkenhofStraße 6a, D-8OOO München 6O Max Kühbandner, Marsstraße 8, D-8O11 Aschheim Wolfgang Schacht, Scherrerstraße 8, D-8081 Schöngeising Thomas Witt, Tengstraße 33, D-8OOO München 4O