Description of the reduced mouth parts of Coleophora micronotella Toll ( Lepidoptera , Coleophoridae ) , with a new synonym

The reduced mouth parts of Coleophora micronotella Toll (Coleophoridae) are described and illustrated. The proboscis is lacking, but rudimentary maxillary and labial palpi are present. The mouth parts of C. micronotella are compared with those of other species of Coleophora, including some others with reduced mouth parts, and their possible adaptive value is discussed. Ramidomia Falkovitsh, 2005 and Ramidiomia Falkovitsh, 2005, a misspelling, are synonymized with Coleophora Hübner, 1822. Résumé. La morphologie des pièces buccales très réduites de Coleophora micronotella Toll (Coleophoridae) est décrite et illustrée. Cette espèce présente des palpes maxillaires et labiaux rudimentaires, mais aucun proboscis. Les pièces buccales de C. micronotella sont comparées avec celles d’autres espèces de Coleophora, incluant quelques-unes ayant des pièces buccales réduites, et la possible valeur adaptative de la réduction des pièces buccales est discutée. Ramidomia Falkovitsh, 2005 et Ramidiomia Falkovitsh, 2005, à l’orthographe fautive, sont mis en synonymie avec Coleophora Hübner, 1822.


Introduction
In Glossatan Lepidoptera, which comprise over 99.9% of the species of the order, the mouth parts typically consist of a labrum bearing a pair of lateral pilifers anteriorly, a haustellum (proboscis) formed by the modified galeae of the maxillae and a pair of maxillary palpi medially, and a pair of labial palpi posteriorly.Mandibles are fully formed only in the non-glossatan Micropterigidae and Heterobathmiidae.In some scattered groups of species of various families and in some larger groups, such as a good proportion of Lasiocampoidea and Bombycoidea, however, a reduction of the mouth parts can be observed.This phenomenon has not been dealt with comprehensively in the literature and we offer some observations in an attempt to explain it in one small species.
As shown in detail by Câpuçe (1971), the members of Coleophoridae, although generally quite small, usually have well-developed mouth parts, especially in the form of prominent, recurved, tri-segmented labial palpi and a functional haustellum.Some species with reduced mouth parts are nevertheless known, such as C. minipalpella Baldizzone, 1998, described from Andalusia, Spain and known from only four specimens.Here we describe and illustrate the ves- tigial mouth parts of Coleophora micronotella Toll, 1956, for which there are more specimens available for study.
In the original description, Toll (1956) mentioned the colour of the head and antennae, and indicated that the palps are missing in all of the nine specimens of C. micronotella that he had at hand, but said nothing about the lack of proboscis.Falkovitsh (2005) erected the monotypic genus Ramidomia, here synonymized with Coleophora Hübner, 1822 (along with Ramidiomia Falkovitsh, 2005, a misspelling) to accommodate C micronotella.Along with Baldizzone et al. ( 2006) and others, supported by the phylogenetic analysis of Bauer et al. (2012), we find it more coherent to use one large genus Coleophora ( 1 340+ described species) than a classi- fication of Coleophora subdivided in numerous small genera that have been erected without comprehensive phylogenetic inference, as exemplified by Falkovitsh (2005).The latter author noted that C. micronotella has short palpi, with the third segment shorter than the second, but did not mention the lack of proboscis.Subsequent records of the species did not add any more morphological information (Baldizzone 1994;Baldizzone et al. 2006).

Methods
The head of one specimen was mounted on slide in Euparal after maceration in about 15% KOH at 60°C in an oven for an hour, cleaning in 30% ethanol, staining in Orange G and chlorazol black, and dehydrating in baths of 95% ethanol and isopropanol for one hour each.
The photographs of the scaled head as well as that of the slide-mounted whole head were taken with a Leica MZ APO dissecting scope and AutoMontage, the close-ups of the mouthparts on slide with a Zeiss microscope and AutoMontage, and the scanning electron micrographs with a Zeiss DSM 940-A scanning electron microscope in the Muséum d'histoire natur- elle de Genève (MHNG).Description of head.Frontal aspect with appressed scales, with short, apparent scaled palpi, without proboscis.Ventral edge of fronto-clypeus slightly sinuate.Labrum not apparent.Maxillary palpi unsegmented, very short, with few scale sockets and setae, with apical sensillum styloconicum.Labial palpi slightly longer than maxillary palpi, reaching apices of subgenal apophyses, with two segments visible, apical segment about half as long as basal segment and lacking vom Rath's organ.With sac-like rounded structure laterally from labial palpi.Notes.The halophilous biotope of La Caletta, where three of the specimens of C. micronotel- la studied were collected, is about one kilometer away from the Mediterranean, on the sides of a small canal where tamarix {Tamarix sp., Tamaricaceae) grows.The specimens were collected with a black light in a salt marsh where Salicornia (Amaranthaceae) and other halophilous herbaceous plants were growing.Hundreds of specimens came to light, but could not come to rest on the sheet because of the presence of thousands of staphylinid beetles and flies.Falkovitsh (2005) recorded that the larva of C. micronotella feeds as a borer, without making a case, in 'green branches (assimilating shoots)' of Halostachys caspica CA.Mey. and Halocnemum strobilaceum (Pallas) Bieb.(Amaranthaceae).

Coleophora micronotella
Described from Tozeur, Tunisia, the species is also found in Italy (mainland, Sardinia, Sici- ly) and Spain.It has also been reported from Iran (Baldizzone 1994), and Algeria, Kazakhstan, and Tadjikistan (Falkovitsh 2005).However, a specimen from the Iranian series mentioned in Baldizzone (1994) was re-examined and found to represent another, probably undescribed spe- cies.Its mouth parts are more developed than in C. micronotella, with the labial palpi about as long as 3/4 the widest diameter of the compound eye, but without apparent proboscis.Also, the male valva differs slightly in the distally larger and more evidently separated sacculus from the basally narrower valvula, and the phallus is narrow, needle-like, instead of spatulate.
©Societas Europaea Lepidopterologica; download unter http://www.biodiversitylibrary.org/ und www.zobodat.atOther species of Coleophora with vestigial mouth parts Coleophora galligena Falkovitsh, 1970, described from Uzbekistan, has vestigial palpi and no proboscis.Coleophora daeva Baldizzone, 1994, C. minipalpella Baldizzone, 1998, and C   zagella Falkovitsh, 1972, respectively described from Iran, Spain, and Mongolia, have short proboscis and palpi.These four species and C micronotella can be associated with the 8* group of Toll (1953Toll ( , 1962)), based on their simple genitalia, but Toll (1956) only mentioned that C micronotella is close to C plurifoliella Chrétien, and never associated this species with any group subsequently.These species are associated with halophilous habitats and deserts and they were not treated by Bauer et al. (2012).

Discussion
The sac-like rounded structures that protrude laterally from the labial palpi in C. micronotella may represent vestigial mandibles.Rudimentary mandibles were found in all species of Co- leophora studied by Càpuçe (1971), with variation in shape and size.Càpuçe (1971) noted that the mandibles are usually sclerotized, but he found at least one species in which they were membranous (C.asthenella Constant).Càpuçe (1971) did not study C. micronotella, but for comparison it may be noted that among the 1 00 species studied he mentioned that the maxillary palpi were missing in one only (C.asthenella).Although it is often difficult to count the number of maxillary palp segments precisely because the limits between segments may be not easily discernible, Capuçe found that the number of segments varied from one to five, the one-segmented species numbering 20, the 2-segmented 3 1 , and the other cases in lower frequencies.
Càpuçe (1971) also found that C. ciconiella Zeller, a species with two maxillary palpomeres, has the apex of the maxillary palpi adorned with a sensillum styloconicum.
The labial palpi in the Lepidoptera groundplan are tri-segmented (Kristensen 2003).This is the condition found in Coleophora, but the overall length of the palpi, length of individual palpomeres, shape of terminal palpomere, and vestiture vary across species.Càpuçe (1971) did not study any species with markedly reduced labial palpi.The lack of vom Rath 's organ is unusual in Coleophoridae and presumed here to be linked with the reduction of the labial palpi.
Regarding the proboscis, Capuçe ( 1 97 1 ) recorded no species in which it is lacking, but a few in which it is slightly shorter than the labial palpi, and membranous.No species with reduced mouth parts were found from the Western Hemisphere (J.-F.Landry, pers.comm.to G. Baldiz-  zone), but several undescribed species from the Afrotropical region have reduced proboscis and palps.Bauer et al. (2012) presented the first phylogeny of Coleophoridae based on a formal analy- sis of molecular characters; one hundred and two European species were analysed, but neither C. micronotella, any of the above-mentioned species with reduced mouth parts, nor any species of Toll's 8"" group were included.Nevertheless, two of the species groups recovered, the albel- la and alticolella groups, have species feeding on Amaranthaceae, with the larvae of the albella group being internal miners in the first larval stages, until they become too large to be fully hid- den in the nutritive tissue, after which they build a tubular silk case.This host plant choice and the internal feeding habits may point to an affinity of C. micronotella with the albella group.