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Nymphalidae, Satyrinae) Contributions to Zoology, 73 (4) 293-303 (2004) SPB Academic Publishing bv, The Hague the The shape of endemics: Notes on male and female genitalia in genus Maniola (Schrank, 1801), (Lepidoptera, Nymphalidae, Satyrinae) Andrea Grill Rob de Vos Jan van Arkel , , Institute for Biodiversity and Ecosystem Dynamics / Zoological Museum Amsterdam, University of Amsterdam, P.O. Box 94766, NL-1090 GT Amsterdam, The Netherlands Keywords:: Sardinian endemic, genital morphology, Maniola nurag Abstract tion in genital morphology of the Meadow Brown has been extensively discussed (Thomson, 1973, for Butterflies of the genus Maniola are known their large 1976; Goulson, 1993). In recent decades, two new at the inter- as well as intraspecific morphological variation, Maniola species have been described (Thomson, level. Given the overlap in wing-patterns, habitat selection, and 1987, 1990). First, the island endemic, Maniola chia geographic distribution of various Maniola species, genitalia Thomson, 1987, whose distribution is restricted to morphology is sometimes the only possibility to tell specimen Maniola hali- this describe characters the Greek island of Chios. Second, apart. In paper we diagnostic to distinguish Included Bo- different Maniola species by means oftheir genitalia. carnassus Thomson, 1990, which flies on the is also the first detailed and illustration ofthe genitalia description drum peninsula (Turkey) and the Aegean island of of the Sardinian endemic Maniola nurag. Further, apparatus Nissiros. Maniola endemic nurag(Ghiliani, 1852) is we describe two Sardinian individuals with intermediate char- and to Sardinia, a third endemic has been described Maniola and acteristics between nurag and Maniola jurtina, discuss the from the island of Cyprus, Maniola cypricola propose that they are hybrids. Further, we shortly justification ofthe species status for the island endemics Maniola (Graves, 1928) (for distribution areas of species chia Maniola and cypricola. see Fig. 1). Maniola megala (Oberthiir, 1909) oc- curs on the Greek island of Lesbos, throughout southern Turkey and in Iran. Contents Although neighbouring islands would be in flight distance for all island endemics, the ranges of the Introduction 293 island-Maniola species are well confined to the Material and methods 296 borders of the respective island. In Chios, M. chia Dissection and photography 296 is said to entirely replace M. jurtina and Maniola Results 296 telmessia (Zeller, that are Discussion and conclusions 299 1847), species commonly Acknowledgements 303 found on the neighbouring islands and the Turkish References 303 mainland, which is only a few kilometers distant from Chios. In the other M. Sardinia, on hand, nurag flies sympatrically with M. jurtina. Although, the Introduction latter species is usually concentrated on the coast, whereas the Sardinian endemic has its distributional “Made of and convex hills and val- up concave centres in the mountain areas of the island (> 500 of the first of the leys”, was one descriptions geni- m), there is a zone of overlap at intermediate alti- tal of a male Meadow Maniola structure Brown, - tudes (500 900 m), where both species fly con- meant to that this jurlina (L.), emphasize species’ temporarily at the same sites (Grill, 2003). genitalia are irregularly shaped for a Satyrrd Butterflies of the Maniola known very genus are for Since varia- (Muschamp, 1915). then, geographic their large morphological variation (Fig. 2), at in- 294 Grill al. - Genitalia the A. et in genus Maniola Fig. I. Distribution areas of the six European species of the Maniola: Maniola M. genus (A) halicarnassus, (B) telmessia, M. M. M. (C) nurag,,(D) chia, (E) cypricola, (F) M. jurtina. both local ter- as well as intraspecific level, on and continental scale (Ford, 1945; Thomson, 1973). 2. Variation in in the Maniola. the Fig. wing pattern genus All Given the in selec- overlap wing-patterns, habitat specimens are in the collection of (he Zoological Museum Amsterdam, Column shows the the tion, and geographic distributionofvarious Maniola Legend: (a) upperside, (b) underside of the butterflies: I. M. jurtina France (male), 2. M. species, genitalia morphology is sometimes the only jurtina France (female), 3, M. jurtina Sardinia (male), 4. M. tell What is possibility to specimens apart. more, jurtina Sardinia (female), 5. M. telmessia (male), 6. M. telmessia can also much within genitalia shapes vary a single 7. M. 8. M. intermediate (female), nurag (male), nurag form the 9. M. 10. M. chia 11. M. chia species (Thomson, 1973). Nevertheless, spe- (male), nurag (female), (male), 12. M. 13. M. cies status of M. chia and M. cypricola has been (female), halicarnassus (male), halicarnassus (female), 14. M. cypricola (male), and 15. M. cypricola (female). justified mainly because of differences in the form of the male genitalia; in wing-patterns they resemble M. jurtina and M. telmessia, respectively. For the third the underside endemic species in this genus,M. nurag, geni- wing markings. talia structure and shape has never been described Ergo, the three main questions we address in this in and illustrated detail as yet. paper are: In this the of Are there characters in the paper, genital apparatus M. nurag 1) diagnostic genitalia is described and illustrated in detail for the first of the different Maniola species? time. We further describe two Sardinian individu- 2) What is the position of the Sardinian intermedi- whose individuals in the Maniolal als, genitalia seem to be intermediates be- ate genus M. and Is for M. chia and M. tween nurag M. jurtina. The genitalia mor- 3) species status justified phology of these Sardinian specimens is compared cypricola? the and of the in to shape structure genital organs all M. other Maniola species, except megala, as this species can be unequivocally distinguished from its congenerics by its appreciably larger size, and Contributions 73 - to Zoology, (4) 2004 295 296 A. Grill et al. - Genitalia in the genus Maniola Material and methods Results of In the main distinctive between In May 2002 we collected a series males and males, characters females of M. 3 and M. Maniola are the of the the nurag (5 males, females) species shape valvae, 3 from Sardinia and the and size of the Julien jurtina (3 males, females) (Italy), gnathos, strength organ in July 2002 M. jurtina (3 males, 3 females) from (Table 1, Fig. 3). There is also some variation in Amsterdam (The Netherlands), and in September the aedeagus, but it is difficult to use this as a char- of the same year M. chia (1 male, 3 females) from acter, as it has a rather soft structure and changes Chios These its the from where (Greece). specimens were compared shape according to angle you with specimens of M. telmessia (2 males, 2 females) look at it. The male genital apparatus also varies in and M. halicarnassus (2 males, 2 females) collected size between species. Among the individuals we in by H. van Oorschot in Turkey, present the Zoo- studied, M.jurtina has the largest and M. telmessia the smallest found differ- logical Museum Amsterdam. Two of M. nurag the genitalia. In females, we be attributed in of we dissected could not unequivocally ences the shape the ovipositor lobes and the to M. to could of the ductus bursae In all nurag; according wing-pattern they length (Fig. 4). species studied in this be a light, small M. jurtina as well as a dark large paper, except M. jurtina, the surface of the bursa contains two that consist M. nurag. Small sample sizes are sufficient, as this copulatrix signa of study aims at a qualitative and not quantitative spine-like sclerotised structures (Fig. 5). description of characters. The butterflies were con- served dry or frozen until preparation. Butterflies were identified using characteristics in their wing- Maniolajurtina (Figs. 3A, 4A) patterns following Hesselbarth et al. (1995), van Oorschot & van den Brink (1992) and Tolman & Male: Gnathos markedly swollen at the base, then Lewington (1997). All individuals studied are in quickly narrowing. Valvae bigger than all other Amster- the collection of the Zoological Museum, species except M. halicarnassus, in shape most dam. similar M. but with characteristic to nurag, a curve towards the distal distal and dorsal process; pro- cess round; ventral edge different from the other Dissection similar Julien and photography Maniola species, most to M. nurag. thick and organ always clearly visible, very strong, of Prior to dissection the abdomen the specimen can be twice the size as in congenerics. Female: was separated from the rest of the animal and soaked Length of ductus bursae comparable to in Potassium hydroxide (KOH 10%) for approxi- the other species; notably in none of the dissected mately 15 hours. To stabilize the samples for pho- females we found signa, although they were clearly in visible all female individuals the tography, they were positioned laterally a small in of other spe- drop ofethanol (30%), flattened between two glass cies. The absence of these marks might be a good M. and lids. They were photographed under the microsdppe distinctive characteristic between jurtina in (magnification x 25). In order to photograph the the other species the genus Maniola. form of the signa in the bursa copulatrix, which distinctive characteristics may show important between species, the female genitalia were dyed Maniola telmessia (Figs. 3B, 4B) with chlorazolblack. The dye was fixed in 95% For in ethanol. handling, the genitalia were kept Male: Genital apparatus clearly smaller than in all 30% ethanol, as in stronger concentrations of etha- other Maniola. Gnathos similar to M. chia, slightly nol the chitine hardens and breaks easily. For long swollen at the base, vesica round. Valvae similar term conservation the genitalia will be transferred in shape to M. chia and M. halicarnassus; distal to a or slide. similar to M. chia and M. glycerol-tube Euparol process pointed, nurag, dorsal process almost pointed. Ventral edge simi- Contributions to Zoology, 73 (4) - 2004 telmessia considerably pointed flatflat jurtinajurtina to othersothers inin cypricola gradually inin telmessia all continues thanthan in base, in curves M.
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