I I BU LLETIN DE L'INSTITUT ROYAL DES SCIENCES NATU RELLES DE BELG IQUE ENTOMOLOGIE. 75: 71 -78. 2005 BULLETIN VAN HET KON INKL IJ K BE LGISCH INSTITUUT VOOR ATUU RW ETENSCHA PPEN ENTOMOLOG IE, 75: 71 -78, 2005

The antennae of Neopseustis meyricki: 1norphology and phylogenetic i1nplications, with special reference to the sensilla basiconica and styloconica (: Neopseustidae)

by Michel J. FAUCHEUX

Abstract grade above the level of «Dacnonypha» (comprised of the families , and Lophocor­ The antenna! flagellum of both sexes of Neopseustis meyricki Hering, onidae) and thus in cladistic te1ms representing the possible 1925 possesses seven types of sensilla: multi parous sensi lla trichodea, teroneura). multiparous large swollen sensilla basiconica, multiparous thin sensilla sister-group of Neolepidoptera ( + He basiconica, multiparous sensilla coeloconica, two types of aporous The family of Neopseustidae reveals a particular sensilla sty loconica, and aporous sensilla chaetica. The sexual di­ innovation: the formation of a double-tube proboscis morphism concerns only the number of sensilla for each of these types. (KRISTENSEN & NI ELSEN, 1981 ). However, the a nte1mal The two types ofsensilla basiconica are probably a specific structure of Neopseustidae. The two types of sensilla styloconi ca are an archaic sens01y organs have been little studied. In a recent review type, the long-haired sensilla, and an evolved type, the typical cone­ article on sensi II a and proprioceptors of Lepidoptera, this shaped sensilla. The great length of the sensilla styloconica, their great fam ily was not even mentioned (HALLBERG et al., 2003). number, their particular locali zation, and the coexistence of two both ty pes are specific characteristics of N. meyricki. Nevertheless, a preliminmy study of the neopseustid Apoplania valdiviana DAVIS & NIELSEN, 1984 revealed Key words: Lepidoptera, Neopseustidae, Neopseustis, antenna, sen­ the presence of 7 antenna! sensillum types, including a nicum, phylogeny. silla, basiconicum, styloco particul ar type of sensi ll a basiconica, the «wafer-li ke sensil1um bas iconicum» (FAUCH EU X, 1999). Since then, a more exhaustive study of the species has raised the Resume problem of interpreting «the goffered sensillum basico­ nicum with a double or triple base» which is an unique Le tlagell e a ntennaire des deux sexes de Neopseuslis meyricki Hering, A 1925 possede sept types de sensilles: d es sensilles tricl10.ides multi­ type among the Lepidoptera (FAUCHEUX, 2005a). study pores, des sensilles basiconiques mult ipores larges et entlees, des is presently being undertaken to ascertain the existence of sensilles basiconiques multipores minces, des sensilles coeloconiques a similar type of sensilla basiconica in the other species of multipores, deux types de sensilles styloconiques sans pore, des sen­ the same fami ly. We offer here the results obtained fro m silles chetiformes sans pore. Le dimorphisme sexuel concerne unique­ men! le nombre de sensilles de c hacun des types. Les deux types de Neopseustis meyricki HERING , 1925, a species considered sensilles basiconiques sont probablement une structure specifique des as being less evolved than A. valdiviana (DAV IS, 1975). Neopseustidae. Les deux types de sensilles styloconiques correspon­ dent a un type archa'ique, les sensilles a long poil , et un type evolue, les sensilles typiques a cone. La grande longueur des sensilles stylo­ coniques, leur nombre eleve, leur localisati on p articuliere et Ia Material and methods coexistence de deux types sont d es caracteristiques particul ieres de N. meyricki. The microlep idoptera come from Dr Shen-Horn Yen's Mots-clefs: Lepidopteres, Neopseustidae, Neopseuslis, antenne, sen­ collection. They were captured in Taiwan, I-Lan Co., si lles, basiconique, styloconique, phylogenese. Yuan-Shan, Fu-Shan Botanic Garden, altitude 800 m, on 18-TV -1996 for the male and 29-V -1995 for the fe­ male. For the study with scanning electron microscopy Introduction (S.E.M.), the antennae were descaled, dehydrated in ab­ solute ethanol, mmmted on specimen holders and coated In Lepidoptera, fo ur primmy clades are recognized: the with a t hin layer of gold and palladium in a JFC II 00 fam ilies Micropteri gidae, Agathiphagidae, Heterobathm ii­ sputter coater. Preparati ons were examined in a JEOL dae, and the high-rank taxon . The Glossata com­ JSM 6400 SEM at different magnifications. The termi­ prise six major clades: the famil ies Eri ocraniidae, no logy of SCH NE IDER (1964) and ZACHARUK (1985) is Acanthopteroctetidae, Lophocoronidae, Neopseustidae, used in naming the types of sensilla. The mean number and the high-rank taxa Exopori a a nd Heteroneu.ra of each sensillar ty p,e was calcul ated with S.E.M. fro m the (KRISTENSEN, 1997). lndeed, since ] 978 (DAVIS, 1978), counts on every 5[ 1 fiagellomere, and expressed as mean the Neopseustidae have been regarded as representing a +/- SO. '' 72 Michel J. FAUCHEUX

Results base of the sensilla is sometimes deformed by S.E.M. (Figs. 9, I 0). The sensilla basiconica B I The are relatively small, with the following wing are the most numerous of the antenna) sensilla ( 40 expanse: male, 20 mm; female, 20-22 mm. The antennae to 41% ofthe total), with a very slight advantage for are long, equaling the length of forewing. The scape is the male (Table). large and swollen; the pedicel is four times shorter and 3 - The multiparous thin sensilla basiconica (type Il) are only half the width of the scape. easily identifiable thanks to their straight aspect and The flagellomeres (male, 65; female, 63) are strongly differ from the sensilla basiconica B I by their cupuliform, slightly depressed and asymmetrical (Fig. I). circular base, which is never deformed (Fig. I 0). All flagellomeres possess nmTow lamellar scales ar­ Like the latter, they offer a goffered aspect and ranged in longitudinal rows of 2-5 contiguous raised possess numerous pores in groups (Fig. 12). The sockets, and noninnervated microtrichia evenly scattered wall, 0.15 11m in width, is sometimes thinner than over the enlarged part of each segment (Fig. 3). in type I and represents I 118 of the sensillum dia­ Seven types ofsensilla were discovered on the antenna) meter (Fig. 13). What is more, they possess a spe­ flagellum of both sexes of N. meyricki. No sexual di­ cificity which has been observed neither in sensilla morphism concerning the types of sensilla was observed. 8 I , nor in any other lepidopteran antenna) sensilla The sensilla are located all around the flagellomeres but basiconica. At their base, they either narrow are twice as numerous on the ventral face. The sensillum (Fig. 16) or reveal a slender bulge followed by a types are the following: multiparous sensilla trichodea, thickening of the hair, the whole resembling a multiparous large swollen sensilla basiconica, multipar­ cupola (Figs. 14, I5). This detail will be discussed ous thin sensilla basiconica, multiparous sensilla coelo­ below. The sensilla B2 are for the most part located conica, aporous sensilla chaetica, aporous long-haired at the distal edge offlagellomeres, but they represent sensilla styloconica, and aporous cone-shaped sensilla a more 6% of the total of sensilla. styloconica. 4 - The multiparous sensilla coeloconica are small fluted cones; they number I or 2 per flagellomere - The multiparous sensilla trichodea are spread evenly (Fig. 17). Though always visible on the antenna, over both faces of the flagellomeres (Fig. 2) from they only number 3% of the antenna) sensilla. the proximal edge to the distal edge (Fig. 4). They 5 - The aporous long-haired sensilla styloconica are the can be identified by their oblique striae at the base longest and the most surprising of all the antenna! (Fig. 5) which become annular on the remainder of sensilla of N. meyricki (Fig. l ). The stylus alone is the hair (Fig. 3). The wall pores are difficult to almost as long as another sensillum. The hair is distinguish with scanning electron microscope longitudinally ridged like that of a sensillum chae­ (Fig. 5). The sensilla trichodea constitute 32% of ticum (Fig. I I). They are arranged obliquely on the all the sensilla and are more numerous in the male antenna) surface and always at the distal edge of the (Table). flagellomeres. According to estimation based on 2 - The multiparous large swollen sensilla basiconica eight successive flagellomeres, their number is as (type I) are disseminated among t he sensilla tricho­ fol lows: I, 4, I, 4, 2, 2, 4, 3. When they attain the dea (Fig. 2). They are evenly curved from their base, maximum number of 4, they are either equidistant swollen and flattened in their proximal part but from one another and arranged in the middle of the thereafter retain a regular diameter up to their tip four faces, dorsal, ventral and two lateral (Fig. 1), or (Fig. 6). Their wall offers an irregular network of grouped two by two in the middle of the dorsal and ridges with in which are located the pores thus ventral faces (Fig. 18). concentrated (Figs. 7, 8). The thickness of the wall 6 - The aporous cone-shaped sensilla styloconica are amounts to 0.13 11m and represents I II 0 of the almost as long as the previous type (Fig. 19). The diameter of the sensillum (Fig. 8). The enlarged extraordinarily long stylus is absolutely smooth and

Table - Length, basal width, average numbers and percentages ofsensilla on a male and female antenna! flagellum of Neopseustis meyricki (mean+/- S .D.)

Sensilla Length Basal width Numbers % Numbers % (~tm) (~tm) (male) (female)

Trichodea 26.7 +/- 4.5 1. 8 +/- 0.3 1062 32.3 949 32. 1 Basiconica I 28.3 +/- 3.9 2.0 +/- 0.4 1358 41.3 1176 39.8 .. Basiconica II 30.5 +/- 5.8 2. 7 +/- 0.2 195 5.9 183 6.2 Coeloconica 7.5 +/- 1.4 1.7 +/- 0.3 96 3.0 87 2.9 Long-haired styloconica 59.3 +/- 5.2 2.7 +/-0.2 168 5.1 159 5.4 Cone-shaped styloconica 37.5 +/- 2 .7 1.4 +/- 0.2 17 0.5 15 0.5 Aporous chaetica 28.6 +/- 4 .3 1.8 +/- 0.4 391 11.9 383 13. 1 Total of sensilla 3287 2952 Antenna! sensilla of Neopseustis 73

SUimounted by a small sensory cone (Fig. 20). They Among the ubiquitous sensilla identified on the antetma are present on the antennae of both sexes but absent of Lepidoptera, the absent sensilla are the multiparous from many flagellomeres; their frequence is of I for sensilla auricillica and the uniparous sensilla chaetica. 4 flagellomeres. These sensilla are always situated Although the lepidopteran sensilla auricillica possess vety next to a long-haired sensillum styloconicum. diverse fonns (FAUCHEUX, 2004a), it has not been possible 7 - The aporous sensilla chaetica are always straight, of to identify them, either inN. meyricki, or in A. valdiviana. the same length as the sensilla h·ichodea (Table) and They exist in families less evolved than the Neopseustidae possess a basal alveola (Fig. 21 ). Their apical end is as in the Heterobathmiidae (FAUCHEUX, 2004c), the pointed and without any pore. The wall is marked by Eriocraniidae (DAVIS, 1978) and the Lophocoronidae 8 longitudinal ridges; between them are arranged thin (FAUCHEUX, in press). Since their appearance in the transversal striae similar to those of a scale (Figs. 22, Heterobathmiidae, the Neopseustidae are the first family 23). The sensilla are located towards the distal edge of deprived of this sensillum type considered as ubiquitous each flagellomere; they are often grouped together (FAUCHEUX, 1999; HALLB ERG et a/. , 2003). The sensilla and oriented away from the antenna (Fig. 21 ). Their auricillica are unevenly represented among the families of number reaches 12% of the antenna! sensi II a. Lepidoptera. In the Exoporia () which follow the Neopseustidae, the distribution of the sensilla is like­ wise variable: they are present in Aenetus virescens and Discussion Wiseana umbraculata (FLOWER & HELSON, 1976) but absent in Afrotheora thennodes (NI ELSEN & SCOB LE , The sensilla common to two species, A. valdiviana et 1986) and Triodia sylvina (personal observations).Thus, N. meyricki are: the sensilla trichodea, the sensilla basi­ among the higher Lepidoptera, the Rhopalocera may have conica I and II, the sensilla coeloconica, the long-haired them, like Pieris rapae Linne (FAUCHEUX, 1996). sensilla styloconica, and the aporous sensilla chaetica. It may however be thought that the large swollen Those present in only one species are the cone-shaped sensilla basiconica type l of N. meyricki are close to sensilla styloconica, which were observed only in the sensilla auricillica. Indeed, they closely resemble N. meyricki and the cupuliform organs, present in the sensilla auricillica of the tineid Monopis crocicapi­ A. valdiviana. The cupuliform organs are rarely been tella Clemens. We have shown that in the keratophagous observed (FAUCHEUX, 1999); they are in fact difficult to , these sensilla could possess the shape of an ear identify and probably more frequent than is believed. or that of a long, flattened hair (FAUC HEUX, 1987). The aporous sensilla chaetica are not an ubiquitous The uniparous sensilla chaetica are a ubiquitous type type on the antennae of Lepidoptera. They have been present in families less evolved than the Neopseustidae. described with certainty only in a few families: in the Nevertheless, the Lophocoronidae which directly precede lower Lepidoptera such as the Agathiphagidae the Neopseustidae are also deprived of such sensilla. (F AUCHEVX, 1990), as well as the such as the They assure a taste function in the antetma, used by (NIEUKERKE N & Dar, 1987), the Ypono­ certain moths in the choice of site for egg-laying. The meutidae (CUPERUS, 1983), the and the antennae of N. meyricki do not consequently perform this (FAUCHEUX, 1978), and even in the Rhopalo­ function. cera such as the (FAUCHEUX , 1996). Their pre­ sence in the Neopseustidae provides their antennae with a Phylogenetic considerations particular tactile sensitivity which no doubt shows in their behaviour for most Lepidoptera are deprived of this We have not observed inN. meyricki the double or triple­ sensillum type. In patiicular, the sensilla chaetica of based sensilla basiconica present in A. valdiviana but this N.meyricki are on an average one and a half times larger latter type and the sensilla basiconica of type 1 and type 2 than those of A. valdiviana (FAUCH EUX , 2005a) and four of N. meyricki have points in common because the three times larger than those of Synempora andesae (unpub­ types possess grouped pores. This pore-pattern is unique lished observations). among the antenna! sensilla of Lepidoptera for in all other The sensilla trichodea are pheromone receptors (KAIS­ families, the sensilla basiconica belong to two types, LING et a/., 1978; (STEINBRECHT, 1987)). The sensiJla with, either with longitudinally lined pores or with basiconica respond to host plant odours in all other moths scattered pores (FAUCHEUX, 1999; HALLB ERG et a/. (SCHNEIDER & STEINBRECHT, 1968; VAN DER PERS, 1981). 2003). If this pore-pattern is confirmed in the other genera The sensilla coeloconica are olfactive and respond to of Neopseustidae, the grouped-pores sensilla basiconica short-chain organic acids (POPHOF, 1997). The cone­ may be considered as being an autapomorphy of the shaped sensilla styloconica are thermo-hygroreceptors Neopseustidae. Furthermore, it is reasonable to believe (ALTN ER et a/. , 1983 ; STEIN BREC HT & MOLL ER, 1991 ). that the single-based sensilla basiconica of N. meyricki Nothing is k11own about the function of the hair-shaped and the double or triple-based sensilla basiconica of sensilla styloconica. We believe they are tactile and A. valdiviana are similar sensilla which evolved differ­ represent a transient form between the aporous sensillum ently. The morphology of the type 2 sensilla basiconica of chaeticum and the cone-shaped sensillum styloconicum N. meyricki furnish no particular information. On the (F AUC HEUX, in press). conh·ary, certain sensilla basiconica of type I have a ,,

74 Michel J. FAUCHEUX

Figs. 1-1 0 - Antenna! sensilla of Neopseustis meyricki. l. 24-27 111 descaled male flagell omeres showing long-haired sensill a ' styloconica (arrows); 2. sensill a tri chodea (T) and type I basiconi ca (8), scale (S); 3. female sensill a trichodea (T), mi crotriches (M); 4. male sensillum trichodeum; 5. obli que ridges of th e wall sensillum; 6- 10. type I sensillum basiconi cum; 6. broken sensillum; 7. detai l of pores; 8. break showing the width of th e wall ; 9. base of sensillum; 10. bases of type I (8 I) and type II (82) sensill a basiconica. Antenna! sensill a of Neopsei.1sti s 75

Figs. 11-1 8 - Antenna! sensill a of Neopseustis meyricki. 11-16. sensill a basiconi ca type II ; 11 . th e w hole sensillum (arrow) a nd base of a long-haired sensillum styloconi cum (ST); 12. detail of pores; 13 . broken b ase showing the thin wall (arrow); 14. kind of alveola at the base (arrow); 15. idem in oth er sensillum; 16. basal n arrowing (arrow); 17. sensillum coeloconi cum; 18. 42'11 fe male fl agell omere s howing th e d istal locati on of f our long-haired sensilla styloconi ca whose two are broken (arrows).

.. I I 76 Michel J. FAUCHEUX

Figs. 19-23 - Antenna! sensilla of Neopseustis meyricki. 19. 33th male flagellomere with numerous scales, two long-haired sensilla styloconica (arrows) and one cone-shaped sensillum styloconicum (star); 20. distal edge of the 53th female fl agell omere with a cone-shaped sensillum styloconi cum (star), a long-haired sensi llum styloconicum (arrow), sensilla basiconica type 1 and II (8 I, 82), sensi ll a trichodea (T); 21 . three sensilla chaetica on the latero-distal part of the 18th female flagellomere; 22. sensillum chaeticum in the middle of the distal edge of the 5th male flagellomere; 23. detail of the wall.

particular basal structure which has not so far been frequently met type in most Lepidoptera. The long-haired observed in any multiparous olfactive sensillum: a nar­ type exists only in the , Micropterix rowing followed by a widening or the presence of a bulge calthella Linne (FAUCHEUX , 1997). The cone-shaped type simulating the formation of a cupola or of a stylus, appears in the sister-group, as in Sabat inca sterops TUlner marking a discontinuity between the base and the hair. (Micropterigidae) (FAUCHEUX, 2004b), where in fact it We propose the hypothesis that, starting from this dis­ coexists with the long-haired type (FAUCHEU X, in press). continuity, the base of certain sensi ll a basiconica has In the following families, the Agathiphagidae, Hetero­ spread out before dividing into two or three bases, result­ bathmiidae, Eriocraniidae, Acanthopteroctetidae, and Lo­ ing in the particular sensillum observed in A. valdiviana, phocoronidae, the two types of sensilla styloconica are a more evolved species than N. meyricki. absent, except in the last family which possesses a unique The presence of two types of sensilla basiconica, thin type, the long-haired type. It is in the Neopseustidae that and large swollen, is common to N. meyricki and the the sensilla styloconica reappear in force. A. valdiviana family of Lophocoronidae which directly precedes possesses only the long-haired type. On the contrmy, the Neopseustidae (FAUCHEUX , in press). The same is true N. meyricki is provided with both types. The long-haired of the Hepialidae (FLOWER & HELSON, 1976). This diver­ type is the most numerous on the same antenna but the sity of sensilla basiconica which appears with the Lopho­ cone-shaped type regularly appears on the fl agellomeres. coronidae and is confirmed by the Neopseustidae In themselves, the long-haired sensill a styloconica, (N. meyricki and A. valdiviana) will continue in the hi gh­ which are of exceptional length, are enough to identify er Lepidoptera (FAUCHEUX , 1999). the species N. meyricki. They are twice as long as in The sensi lla styloconica of N. meyricki also raise an A. valdiviana. It is also the first time that such a large interesting problem. They belong to two types, one being number of sensill a styloconi ca has appeared per flagell a­ consi dered as archaic (the long-haired type) and the other mere, 4 in N meyricki. The most common number is one as evolved (the cone-shaped type). The latter is the most sensillum per flagell omere in the Lepidoptera. The Antenna! sensilla of NeopseuSt'is 77

Satumiidae, among the higher Lepidoptera, are the only the incomplete covering of the flagellomeres, due to ones known sometimes to possess six (F AUCI-IEUX , 1978). the narrowness of scales. This character is paralleled in In fami li es more evolved than the Neopseustidae, the the Lophocoronidae (NIELSEN & KRISTENSEN , 1996). long-haired type of sensilla styloconica disappears com­ the absence of sensilla auricillica, pletely and only the cone-shaped type remains. Never­ the presence of two types of sensill a styloconica, theless, the cone-shaped type may offer a few variants the presence of a particular type of sensilla basiconica such as a longer cone in the tineid Trichophaga tapetzella with groups of pores. Linne (FAUCHEUX, 1989) but the latter is different from The above results, obtained in A. valdiviana and N. the hair of a long-haired sensillum styloconicum. meyricki, need to be confirmed by a comparative study The coexistence of both types of sensilla styloconica, of Synempora andesae, a South-American neopseustid. primitive and evolved, in N. meyricki accords with the claims by DAVIS (1975): «The general morphology of the head (of Neopseustidae), as reviewed by KRIST EN SEN (1968), displays a mixture of primitive and specialized Acknowledgments features». In conclusion, in accordance to the results obtained in A. valdiviana and N. meyricki, the neopseus­ We wi sh to acknowledge the courtesy of Dr Shen-Hom Yen for tid antenna! sensillum and scale features that may even­ providing th e specimens of Neopseustis meyricki. We are indebted to Dr Yves Gruet (University of Nantes, France), for faciliting use of tually prove phylogenetically significant, perhaps as fa­ SEM, A. Barreau (Centre of SEM , Uni versity of Nantes), Dr Joseph mily autapomorphies, include the following: Baudet and Vittorio Ballardini for technical assistance.

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