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Insecta, Neuroptera) DISch. cnl. Z .. 43 (1996) I, 99-121 Larval Stages of European Nemopterinae, with Systematic Considerations on the Family Nemopteridae (Insecta, Neuroptera) V. J. Monserrat Departamento de Biologia Animal I, Universidad Complutense, Madrid, Spain Abstract Li]fVa l development of the European Nemoplera bipelllli,\' and Lerl/Ill sofiae have been completed in the laboratory. the morphology of their egg and larval instars arc described. New dma on their larval behaviour and biology. as embrionary development lime. larval feeding prefe re nces. way and number of moultings. strategies in their development lime and pupation are given. Some new data on their distribution and bi ology as nying ac ti vity. food. periodic ity and number of layed eggs and longevity are noted. Based on obvioll s differences in larval morphology and biology of Nemopterin ae. in regard 10 Croc in ae and other nearest Neuroptera fami li es. some aspects of their systemati c position and relation­ ships are commented, and both sub fa mil ies should be considered with fa mily rank. Key words: Ncuroptcra. Ncmopteridae. Nemoplera, Lertlw, cgg - larval morphology. larval behaviour. Iberian Penin sula. Systematics. Introduction The family Nemopteridae is one of the most cont roversial, interesting and specialized fami ly among the Ne uroptera, it is actuall y di vided into two subfamilies with ve ry different morpho­ logy and habitats. The Crocinae (with two European genera JO.l'alldrev" Navas, 1906 and PlelVcroce Withycombe, 1923), which include around 50 species of small size, is distributed in arid and desert zones on the southern borders of the West Palaearctic and West Orient al Region, and dry areas from Neotropical, Afrotropical and Australian Regions. With crepus­ cular-nocturnal fl ying activity and troglobious habits, its imaginal and preimaginal biology and morphology is acceptably well known (Tjeder, 1967; Mansell , 1976, 1977, 1980, 1981a, 198 1 b, 1983a, 1983 b ; Monserrat, 1983a, 1983 b) and its taxonomy, biogeography and phylo­ geny have recentl y been revised ( Holzel, 1975; Mansell , 1986). Whereas Nemopterinae (with two European genera Nemoptera Latreille, 1802 and Ler/lw Navas, 19 10), amounting almost 1()() species, and including some of the bi ggest and most spectacular within the Neuroptera, having a similar distribution than Crocinae, Dcpanamcnto de Biologfa Animal 1. Faculdad de Biolog fa . Uni versidi.td Complutense. 28040 Madrid, Spai n ,. 100 DISch. enl. Z., 43 (1996) 1 their imagoes are frequent ly abundant, have bri ef seasonal emergence periods, crepusc ul ar or diurnal fl ying habits, and usually with extremely pronounced endemi sms restricted to a single habitat. Its taxonomy and systematics still need a general revision, there is very lillie informati on on its biology and, curiously, the biology and morphology of its preimaginal stages, in the majority of species, is very poor or remains unknown (Tjeder, 1967; Mansell , 1973). In thi s respec t, and in stead of erroneous, doubtful or too vague and outdated references on pre imagin al stages in this subfamil y (Brulle, 1832 ; Rosenhauer, 1856; Kiesenweller, 1857; Schaum, 1857; Hagen, 1866; Blair, 192 1; Navas, 1924a, 1926), some data on the egg is only known about the West M editerranean Nemopfera bipellllis: Dufour, 1857; Navas, 1919; Withycombe, 1925 and Monserrat, 1985, about the South African Kllersvlaktia lIiglVplera: Pi cker, 1984 and PalmipellllG aeo/eop/era : Picker, 1987 and some scarce references about the East Mediterranea n Nemopfera sill/wIG: Popov, 1963, 1973. First or last larval stage have onl y been described for the Mediterranea n Nemoptera bipennis: Na­ vas, 19 19 and Withycombe, 1925, N. coa: Trtiger, 1993, and Nemoptera sil/llaw: Fri edri ch, 1953; Popov, 1963, 1973, about the Australi an C/Ulsmoptera hlllli: Mathews, 1947, about the Southafri ca n DerhYllchia vall solli : Man se ll , 1973, and onl y so me data is known about pupae of DerltYllchia vansolli: Man se ll, 1973 and Nemoplera coa: Troger, 1993. Also Monse rrat & Martinez forward some informat ion on preimaginal stages fro m Iberian Nemoptera bipellllis and Lertha sofiae, more fully developed and exposed in the prese nt contributi on. No more in formation ex ists, and so me of thi s scarce data has been repeatedl y re iterated (Navas, I 924b, 1924c; Saz, 1925 ; Tjeder, 1967; Ki s et aI. , 1970; New, 1989 ; Trtiger, 1993). The supposed preda tor behaviour of larvae in thi s subfamily is evident, due to their aspect, and an edaphic lifestyle has long been suspected (Wheeler, 1929). Both have been confirmed with some of these recent findings, however man y other aspects of it s biology and behaviour as lifestyle in relation with other animals, li fe-cycle time, number of instars and moltings, natural nourishment , parasites, have been deduced or remained almost unknown. Larvae have always been found in the ground li vin g freely in loose sand surfaces, at times to a considerable depth ( 15-25 em beneath the surface), usually wi thout being ab le to breed them in laboratory conditions, some poss ible preys such as small arthropods and Tipulids, Noctuids or Tenebrionids have been suggested, onl y Popov (1963, 1973) during 10 months in feeding first in stars of N. silllUlfa offering it different ani mal-plant substances and Trtiger ( 1993) succeeded last instars of N. coa with wasp, butterfly and fl y larvae and pupae. Monserrat & Martinez ( 1995) conclude the myrmecophily as the possible lifestyle in larvae of thi s subfamily, breeding all in stars of N. bipellllis and L. sofiae with ant larvae as food, observ ing some similar ecological req uirements between Nemoplerinae im agoes and grain cropperants. Pupation has only been observed twice, in the ground, to a depth of 5- 10 cm beneath the surface in fi eld or 0.5- 1 cm beneath the surface in the laboratory (Mathews, 1947; Mansell , 1973; Trtiger, 1993). In thi s co ntribution, new data on imago behav iour, eggs, larvae instars (bred in the laboratory) and cocoon morphology of the Nemoptera bipennis (Illiger, 1812) and Lertlw sofiae Monserrat, 1988 are described, giving new data on the larval bi ology and behaviour of this subfamil y. On the basis of these data, the systemati c pos ition of subfamil y Nemopterinae as regards to Croc inae and other Neuroptera famili es is disc ussed, and thei r considerati on with family rank is proposed. 102 DISch. enl. Z .. 43 ( 1996) 1 Fi gs 1- 6 Egg of Lertha sojiae. I. general aspect: 2. micropylar area : 3. micropyle: 4. aspect of chorion al veoles; 5. p::ul icul ar of thei r border crest: 6. particul ar of surface Qv'lri an follicle cells impressions V. 1. Monserrat : Larval Stages of European Nemopterinac 101 Material and methods Eggs of Nemoprera bipellllis were obtained from gravid females collected at different points in Spain during the last eight years and those of LertlUl sojiae from specimens collected in Spain: Almeria, Balaneg ra 20. VIl.I 99 1. Females were indi vidually isolated into plastic boxes of 10 x 5 x 5 cm, in order to obtain eggs. Some drops of water, a few grains of commercial pollen and some fi xed fl ower were introduced into each box. The eggs laid were isolated into small vial s to observe their dcvelopment time. Several were fi xed for posterior microscopy studies. Temperature conditions of the laboratory during the first eight years of unsuccessful attempts to breed them were not noted, but when the adequate way of feeding them was known (Monserrat & Martinez, 1995), the monthl y range of the thermi c conditions of both years, under whi ch the larve were bred is noted in "C as follo w: I: 14-21, II : 10-23, III : 14-25, IV : 16-23, V: 19-32, VI: 15-37, VII : 21-41, VIII : 22 -38, IX: 17-30, X: 14- 22, XI: 17- 23, XII : 7- 22. The photoperiod was taken to be a natural month, but in dark: penumbra (shade, not full light), and RH : 40-60 %. After bil1h , larvae were indi vidually isolated into plastic culture boxes of 2 x 4.5 x 4.5 cm, half filled with sand and throughout all their development they were fed on al1l larvae burried into thc sand. Data on results obtained with other offcred preys in Monserrat & Martfnez ( 1995). The larvae move and feed freely within the sand and, in order to get information on mOilin g, every 15 days, the sand was revised to look for possible ex uvies. Field observation on imagoes have been noted throughou t our professional acti vity and fauni stical records, here included, mainly belong to the author'S collecti on, but al so ot her material from these Institutions (CPO: Collection P. Duelli , Birmensdorf, MCNM : Museo Nacional de Ciencias Naturales, Madrid, MH : Zoologisches Museum , Hamburg, MP : Mu seum Nati onal d'Histoire Naturelle, Paris, MZB: Museu de Zoologfa, Barcelona, NHM: Natural Hi story Mu seum , London, UB: Uni versidad de Ex tremadura. Badajoz), whose indicated abbreviati ons will be ll sed in the faun isti cal chapter. Results and discussions 1. Description of preimaginal stages of Iberian Nemopterinae In Lerl"a sofiae the egg is ovoid in shape (Fig. I), size 1100- 1200 I'm length , 750- 770 I'm wide, micropyle conical with a di scoidal and conspicuous apex (Figs 2, 3), surface of chorion covered with convex ovarian follicle cell impressions, irregularly hexagonal and bordered by subcylindrical crests (Fig. 4- 6). While yellowish when laid, rosaceous some days later and dark greyish priorto hatching. The egg of Nellloplera hipellllis were described by Dufour, 1857; Navas, 1919 ; Withycombe, 1925 and more fully by Monserrat, 1985, but il mu st be added that its colour is li ght green at first, brown-rosaceous with white conspicuous embryonic stripe later and dark greyish at the end, non fertil eggs are whiter or yellowish.
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