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 sco 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. In both species the position of the larvae prior to hatching is with the head ve ntrally folded onto thorax and abdomen, and the oviruptor fixed on anterior margin of vertex, between mandibles. In Lerl//{{ sofiae the strongly chitinized and brown oviruptor has an aspect of a fu siform plate, more chitini zed at margins and wi th two strong, lriangled and di vergent denticles (Figs 14, IS), remaining fix ed by membranes to anterior margin of the biggesl part of the shell . The abandoned shell is translucent vivid wh ite, with mi cropyle brownish. In Nemo ptera bipennis oviruptor is very similar, but wider as Withycombe, 1925 and Fi gs 22,23 show. Knowledge about the eggs of Nemopteri nae is very scarce, spherical in Nemoptera, more ovoid but wilh very similar chorionic morphology in Len//{{ (Fi g. I) and very elongated oval and with different chorion surface in other genera as Knersvlaktia and Paimipelllw. The presence of sponge-like micropyle and aeropyles on chorion is well known in Crocinae but not V. J. Monserral : Larval Stages of European Nemopterinae 103
Figs7- 13 Just born larva of Lerl/I(I sofl(le. 7. general aspect of head. dorsal ; 8. setae as teeth of inner mandibular margin : 9. lip of maxil ar palp: 10. lip of antenna: II . tip of dorsal head dolichaster: J 2, dorsal dolichaslcrs of terminal abdomen segments: J 3. dorsal dolichasters of anlerior abdominal segments. Scale in 11m 104 DISch. enL Z., 43 (1996) 1 in Nemopterinae and the Qv iruptor is known in Nemopterinae but not in Croc inae and for hatchin g, the chorion is ruptured along a compl ete paraequatorial cut in two parts, 1/3 length for micropylar pole and 2/3 for the rest, di fferent as is known in most Crocinae (Withycombe, 1925 ; Tjeder, 1967; Picker, 1984 ; Mansell , 1977, 1980, 198 1a, 198 1b ; Monserrat, 1985). In Lertha sofiae the just born larva has a general aspect as Fi g. 16: Head hairy (Fig. 7), quadrangul ar, li ght brow n, darker in lateral and posteri or margin s. in a dorsal tri angul ar shape mark and in the ventral hind half (Fig. 16). Epicraneal structu re V-shaped (Fig. 16) and lateral suture on external margi n extending under estemata to hind margi n of labi um. Doli chasters dark brown, infundibiliform (Fig. II ), setae dark brow n, longer setae terminal li ghter, both frequent and placed as show Figs 7, 16. Eyes dark , each one compri sin g seve n stemmata, one ventral and six dorsal (fi ve in a dorsal arch and one centra l). Antennae pale brown, darker externall y, two seg mented, basal segment barrel shape with a thin and short ventral seta, di stal seg ment clavi form with an apical short and thi ck seta (Fig. 10), a structure ex ist in the in side, similar to an in vag in ated seta, and is not ex tern ally visible. M andibles strong, curved (Fi g. 17), brown reddi sh with apex orange, macrotri chi e as teeth on the internal margin (Fi g. 8) in three rows, one dorsal with 3, one medial with 2 (the bi ggest) and one ventral with 3- 4 (the small est), apex of mandible with two sensori al pits (one dorsa l subeJliptic, bi ggest and one ve ntral fu siform) and intern all y with light rows of little denticles, basal part of mandible with arched rows of microtrichi e (Fig. 17). Max illae curved and li ghtl y denti cled at internal apex (Fig. 18), cardo and stipe each one bearing a dolichaster. Labium reduced to a single medial pl ate. palps four segmented, last one fu siform, sensorial haired tip (Fig. 9). In Ne moptera bipennis the just born larva was "desc ri bed and fi gured" by Navas, 19 19 and Withycombe, 1925, but it is preferabl e to carry out a more detail ed description. It has a general aspect as Fi g. 24, very similar to th ose of L. sojiae, but more elongated. Head with rugous integument, with dolichasters of two obviously different sizes, these and setae dark brow n, both frequent and pl aced as shows Fi g. 24. Eyes dark , stemm ata as L. sofiae. Antenn ae as L. sofiae. Mandibles strongly curved (Fig. 25), brown with apex redi sh, macrotrichi a as teeth on the intern al margin as L. soflae but more in number (8 - 10), dorsal and ex tern al face with setae as teeth (Fi g. 25). Maxill ae and labium as in Lertl", sofiae (Fi g. 26). . In Lertha sojiae thorax and abdomen are pale brow n, dorsa ll y with two rows of iso lated reddish brown marks at each side of middline (Fig. 16), ve ntra ll y pale uniform . Small lateral tuberc les bearing 6- 7 long setae in thorax segments, 3 in the abdomen, last but one 2 and conical last one with a panache of shorter setae (Fi g. 16). Doli chaster dark brow n, also in fundibili form , dorsa l ones biggest and placed from the lateral tubercles to middline in a doubl e arched row of 8- 10 the anterior one and 12- 14 the posterior one (dorsall y) and of 10- 18 the anteri or one and 10- 12 the posteri or one (ventra ll y), more abundant on thorax and more uni formly di sposed on last seg ments and in all ventral face. Legs are brown, darker, shorter and stronger the prothoracic, bearing strong and dark di gger setae as spines, on tibi ae and tarsus (one dorsal in eac h segment, and ventrally in three rows, one medi al with 4 on tibi ae and two lateral with 3 on tibiae and 3 on tarsus) (Fi g. 19). Meso- and metathorac ic legs also with spines, but thinner, longer and always on ventral face, 2 at the apex of tarsus and in four rows on tibi ae , two ex tern al of 3-4 and two internal of 3-2 (Fig. 20). Tarsi one-seg mented, terminatin g in paired cl aws bearing two small notches 0 11 the inner side (Fig. 2 1). At the end of th is instar, head capsule and cephalic and thorac ic appendages are darker than just born larvae. In Nemoplera bipellnis the thorax and abdomen is pale cream brow n, dorsall y with two dark brow n spots on pronotum (Fi g. 24) and two rows of isolated, almost im perceptible li ght V. J. Monserrat : L... trval Stages for European Nemoptcrinac 105 16
17
18
0,4 ~21
0,05
Fi gs 14-21 First instar larva of UrI/ill sofiae. 14, oviruptor. dorsal: 15, dillO , latcral: 16. general aspect, dorsal; 17. mandible. dorsal; 18. maxillae and labium. ve ntral ; 19. fo re leg. ve ntral: 20. hind leg. ventml; 2 1. claw. latera l. Scale in mm. 106 DISch. ent. Z., 43 ( 1996) 1
25
26
27 A 0,05
Figs 22 - 27 First instar 1•.lr va of Nelllol'tem bipellllis. 22. oviruplOr. dorsal: 23. ditto. lateral : 24. gcncml aspec t. dorsal: 25. mandible. dorsal : 26. maxillae and labium. ve ntral : 27. claw. lateral. Scale in 111111 . brown marks at each side of the middlinc, ve ntrall y pale uniform. A row of small lateral tubercles bearin g I long seta in mesonotul11 and m ctanotul11 , and 2, folded on the abdomen. in the abdominal segments, a more external row of tu bercl es bearin g I seta on pronolul11 , 2 in mesonolUm and mctanotum, and 2-3 in the abdominal segments, last segments conical with a V. J. Monserrat: Larval Stages of European Nemoptcrinae 107 panache of shorter setae more abundant ve ntrally and as dolichasters dorsall y (Fig. 24). Doli chaster on the th ora x and abdomen are dark brown, also infundibiliform. dorsal and caudal ones biggest and placed from the lateral tu bercles to middline in a double arched row of 10-24 setae on thorax and 20-24 on abdomen, more uniformly di sposed on last segments, in ventral face setae are dispossed on a median stripe in th orax and in two rows 24-44 setae on abdomen, one on each side longer as a seta. Legs as L. sojiae wi th 2- 4 more spines on tibiae. Claws wi thout small notches on the inner side (Fig. 27). At the end of this in star, head capsule and cephalic and thoracic appendages are also darker than just born larvae, and some longitudinal brown stripes are designed on dorsum (Fig. 34). Second instar similar to First, but body less globular and more elongated. Dolichasters and setae proportionall y shorter, speciall y in head and last abdominal segments, in similar positi on but more abundant and more ring disposed. Head capsule darker, its dark lateral und posteri or margi ns more extended to epicraneal sutu re. Antennae proportionall y smaller and last labial palp segment seems apically divided. Structures "mandible-shaped" on hind ve mralmarg in of seventh segment already formed. Claws of forelegs stronger than mid-h indleg. Spiracles more conspi cuous. Sutures as in First in stal'. In Lerl/w .mjiae tergi tes and slernitcs of prothorax more sclerotized and patents and longitudinal mark s of dorsum darker and more continuous, specially in two last segments. Mandible with many macrotrichie in dorsal and internal face. In Nellloplera bipellllis there are more setHe as denticles on inner mandible margin and more setae as spines on its lateral and dorsal face. General colour as shall be describe on third in star. with longitudinal rows of dark spots 0 11 dorsal face and two longitudinal rows of dark spots on ventral two first and seventh abdominal segments. Last in star in Lerlha .WJjiae (Figs 39-41) with head square, fl at dorsall y and very convexe ve ntrally, wi th V-shaped epicraneal suture more visibl e than lateral one along it s external margin and hind margin of labium, brown coloured wit h a reversed T-shaped dorsal shadow darker, latcral and hind margi ns dark brown. Trichosors dark brown, dorsal ones stronger and lanceolated. thin, longcr and fri zzled l<.lIera ll y and begin ning to appear as dolichasteron anterior and posterior regions, ve ntrally lric hosors limited to a wide central stri pe, as dolichaster the anteri or oncs and longer the posterior ones, microtrichie speciall y frequent on hind area of head. Eyes dark, haired. with very con vexes stemmata placed as in just born' larvae. Antennae (Fig. 30) pale brown, two segmented. basal segment short, subcyli nd ri cal. distal segment clavate with an api cal coni cal seta, behind whi ch there is a tegumentary sensori al structure. Mandibles (Fi g. 28) curved and strongly sclerotized at the tip. bearing small denticles in its intern al margin, setae predominantly as teeth on inner margin. lanceolated and hollow in the middle on the dorsal area and as usual on the external face, microtrichia speciall y frequent on basal inner area. Maxill ae (Fig. 29) reduced, curved and with sensoral microtrichia and sho l1 setae on mid apical, carda with one dolichaster and stipe with 3-4 dolichasters and a long seta , mi crotri chie al so on sti pe and basal palp, which is fo ur segmented, second and third segment with two setae. last one with apical hyaline papill ae, sensorial pal pi maculae conspicuous, as a drop distally and fu siform basall y, labium reduced to a single medial transverse plate. Thorax and abdomen pale cream brown. dorsall y with a wide cent ra l stripe pink-orange. more obvious on thorax, flanking out a longitudinal stripe of grey-browni sh stains, another irregul ar and more external maculated stri pe and a third striped one on abdominal spiracles (Fig. 39). Ventrall y two longitudinal rows of grey-browni sh spots on both sides of midline (Fig. 40). Dolichaster also infundibiliform, distally as a 9 points star, wide ring located, dark brown and short , longer in last segments. Di stinct sclerites onl y present in anterior proesternulTI , rudimental in prono tum, and pleural sclerites strong in legs inserti ons. Legs unequals as First instar, but di ger setae 108 DISch. ent. Z., 43 ( 1996) I 31
29 32
0 , 8 30 • 33
(o. 0 ... ~ " ...... ' . '.,.
0 , 2 Fi gs 28- 30 Third inslar larvae of Lerfha sojiae. 28. mandible. dorsal (some setae magni fi ed); 29, maxille and labial pa lp, ve ntral (some mi crotrichie mag nifi ed): 30, antenn a. dorsal. Figs 3 1- 33 Third in star larvae of Nemoptera /J ipeflllis. 3 1. mandible. dorsal (some setae-magnifi ed); 32, maxile and labial palp, vent ral; 33, antenna. dorsal Scale in mm. V. J. Monserrat : Larval Stages of European Nemopterinae 109 34 •
Figs 34-38 Aspects on the biology of Nemoplera bipel/llis larvae. 34. egg. just born and eight mon ths age larva: 35. last second instar covered with sand particles. eating an ant larva: 36. dillo. particular; 37. ex uvia from second to third instar (thoracic ecdisis suture delineated) ; 38. Physogastric grav id female of Pyemoles lIelllricoslls. eCloparasit of cultured Nemopterinae larvae proportionally shorter and stronger, and claws abraded and appicall y truncated, foreleg with di gger setae stronger, in four rows, two external of four and two intern al of five and six, tarsus with two rows of three, digger setae of hindlegs similar to first instar, but inner rows with six selae and tarsal setae litlle. Adistinct dark and chitinized "mandible"-shaped structure is evident on ventral hind margin of 7th abdom inal segment (Fig. 4 1). 110 Dtsch.enI.Z .. 43 ( 1996) 1
42
Figs 39-41 Oeveloped third instar larva of Lerr/w sofiae. 39. dorsal aspect; 40. ventral aspect; 41. last abdominal segments. showing chitinized ·'mandi blc"·shaped structure on hind marg in of 7th abdominal segment, ventral. Figs 42-44 Medium developed third instar larva of Nemoprera bipe1lllis. 42. dorsal aspect: 43, ventral aspect: 44. last abdomin al segments. showing dark marks and chitinized "mandiblc"·shaped structure on 7th abdominal segment, ve ntral. V. J. Monscrrat: Larval Stages of European Nemoprcrin ac III
In Nell/op/era hipellllis last instar (Figs 42- 44) with square head, also flat dorsall y but less convex ventrall y, with V-shaped epicraneal suture as visible as lateral one, dark brown with similar mark s darker than prior instars. Trichosors dark brown, dorsal ones stronger and as spines, other setae thin, longer and fri zzled laterall y, ventrall y tric horors limited to a wide central stri pe, longer on hind half. Eyes dark, haired, with very can vexes stemmata, placed in two rows of three unities on dorsal half and one isolated on ventral half. Anten nae (Fig. 33) pale brown, two segmented, basal segment short , subcylindrical, distal seg ment seems subdivided, but without arti cul ation between the two di fferenciated parts, one subcylindrical, basal and other subovoid , distal, also with an apical coni c seta, and a tegu ment ary sensori al structu re. Mandibles (Fig. 3 1) curved, as in L. sofiae but bearing much more small denti cles as teeth in it s internal margin, and microtric hi a on dorsal and exter nal area. Maxillae and labium similar to L. soJiae (Fig. 32). Thorax and abdomen pale brown, dorsall y with centra l stri pe fl anking out th ree longitudinal stripes of dark brow stain s, the mid-one more irregul ar and the extern al darker, joining on the eighth segment (Fig. 42), on lateral margin , there is a fourth fragmented and striped row in abdominal spi racles line. Vent ra ll y two longitudinal rows of browni sh spots on both sides of midline, less marked on thorax and bi gger on seventh segment (Fig. 43), near to the "mandibles"-shaped structure whi ch is evident on ve ntral hind margin of seventh abdominal segment (Fig. 44). Dorsa-lateral tubercles almost imperceptible. losin g their setae, lateral tubercles are convex and bearin g their setae. Doli chasters infundibilimorms, distall y as 10- 11 po ints star, limited to trasversal areae, dark brown and short, longer in last segments, in the last visible one as spines on di stal half. Ventrally dolichasters in a dense and cuadrangul ar area, more dispersed on thoracic segments, also as spines on distal half of eighth segment. Distinct sclerites onl y perceptible in anteri or proesternum and in pronotum. Legs unequal as in fi rst in star, but digger setae as prior, but proporti onall y shorter and stronger, and claws abraded and so apicall y truncated. As in Nemopterinae egg morphology, the knowledge on their larvae is very poor and sometime too superfi cial, so comparati ve studies are not possible in detail , Trager (1993) made a li ght comparali on among different larvae then known. The genu s Lerllw seems very similar to Nemoplera, but there are some di fferences in extern al colouration and chealotaxy (Figs 16- 33,39-44), position of stemmata, dentic ul ati on of mandible (Fig. 28, 3 1) and morphology of antenn ae (Figs 30, 33) seem good characters to di fferenciate them. Respecting to other known Nemoptera species, larva of Nemoplera bipemlis seems darker and some small differences in the maxill ar chaetotaxy and antennae segmentati on seem be detected from description of N. coa larva (Treger, 1993).
2. Biology and behaviour of larvae
The bi ology and behaviour of Nemopterinae preimaginal stages are almost unknown. So the data here included based on the laboratory culture of two Iberi an species seem to be interesting. Eggs, whi ch are not adhesive, were laid singly during ni ght in number by female: 0- 13, x= 6 in L. sofiae, at the same time, and 0-51, x= 25 in N. bipellllis, laid during several days, as is shown in Tabl e I, Monserrat (1985) records till 85 eggs in N. bipellllis. Data in other species are scarce, Popov (1963) recorded 6 non-adhesive eggs for N. sil/uata and Pi cker (1984) 112 Dtsch. enl. Z .. 43 ( 1996) I
Tabl e I Number of eggs (N) laid day by day from female captured to its death (+) and development period (OPO) in days
L. Jojiae Capl.day 2nd 3rd 4 5 N DPD
spec. J 0 I (+) I (+) spec. 2 0 J3 0 (+) 13 20- 22 spec. 3 0 0 0 (+) 0 spcc. 4 0 7 0 0 (+) 7 (+) spec. 5 0 II 0 0 (+) II 22 -24
N. bipellllis Capl. 2nd 3rd 4th 5th 6th 7th 8t h N DPD day
spec. 0 12 12 12 8 0 (+) 44 18-20 spec. 2 0 9 17 9 (+) 35 18- 21 spec. 3 0 10 3 1 10 (+) 5 1 18- 2 1 spec. 4 0 2 6 10 I (+) 19 18- 2 1 spec. 5 0 2 7 3 (+) 12 18- 20 spec. 6 2 3 17 7 3 (+) 32 18- 2 1 spec . 7 0 6 0 (+) 6 18- 20 spec. 8 0 7 8 5 (+) 20 18-2 1 spec. 9 0 0 0 (+) 0 spec. JO 0 0 8 (+) 8 19-21 spec. I I 0 13 J3 12 I I (+) 49 18-20 spec. 12 2 10 20 12 (+) 44 18- 20 spec. 13 0 0 I (+) 19 spec. 14 0 3 3 14 2 3 0 (+) 25 18- 19 spec. 15 0 14 3 6 6 I (+) 30 18- 19 spec. 16 0 0 0 18 5 0 0 (+) 23 18- 19 spec. 17 0 6 6 12 3 5 (+) 32 18- 19 spec. 18 0 0 18 (+) 18 18- 2 1
recorded 13 ad hesive eggs for Kn ersv/aklia lligIVptera, also ad hesive eggs are recorded for Palmipellll{l aeoloptera. by Picker ( 1987), but num ber was not indicated. Incubation period from 20- 24 days in L. sofiae and 18-2 I days in N. hipellllis, as shows Table I, not all eggs were ferti le. The onl y known data on their embrioni c period time, was recorded, without much precision, by Navas ( 19 19) for N. hipellllis, also Popov (1963) record 20 incubation days at 30 °C for N. silll/ala. Just born larva remains so me minutes resting and later is ac ti ve. The larvae dig into the sand wit h a mechanical and rhythmic Illolion, ope ning the way, head first, with closed mandibles as a shovel, but speciall y burrowing with fore legs and pushing with the othe r legs. Usually reac hing the bottom of the culture-box and remain quiet so burried during all its development. Noepigean behaviour in L. sofiae larvae has bee n observed, but sometimes larvae ofN. bipel/nis were found on sand surrace and exceptiona ll y a larva was found attached to the box blind whic h suggest a certain climbing abil ity. Cannibalism was never obse rved in L. soflae larvae, but it was seen in N. bipelillis larvae submitted to food scarcity. Thigmotaxis is evident and a certain positIve phototaXIs nave l>Cen l:IiCl:kCli up. T ile 1i.1I Vi:iC 1I 1Mull>c1l IClIlUill IIlvlivlllc:,:" illipo r - V. J. Monserrat: Larval Stages of European Nemopterin ae 11 3 turbed and pass ive, di ed like, for hours in any posture. Dataofthe di ggin g behaviour, are si milar to those noted in other kn own species (Mathews, 1947; Popov, 1963; Mansell , 1973), and some of the here related data on N. iJipeliliis are very similar to those recorded by Troger ( 1993) for N. coa. The most of preys used as possibl e food were rejected, but ant larvae were particularly predated , and only larvae fed with ants larvae grown regularl y, faster and showed a more appreciated activity, concluding its deve lop ment (Monse rrat & M artinez, 1995). Motion stimulous of prey was necessary to predation and motions of ant larvae used as food, stimul ates pos iti ve ly the mov ing of cultured larvae toward it, dead or moti onless preys were hardl y detected and predated. Larva usuall y bites the ant larva once onl y and sucks its fluids (not its gut fluids) for feeding, some predigestive, paralizing or toxic secreti ons must be applied, because the prey stops mov ing in a few seconds. During the suction, larvae are motionless with head and mandibles up turned and antennae and palps down-back turned, usuall y moves head during sucti on, creeping the prey. Type of suggested or used preys in Nemopterinae larvae have already been commented and some of the here related data agree wi th those recorded by Troger ( 1993) for N. coo. Just before of the ecdysis time, some larvae have been obse rved with the abdominal tip fixed to the botton of the culture-box with own sec reti ons. For the ecdysis, the tegument is broken in head throughout the epicranea l and lateral sutures and on the dorsal middline of thorax. As first tested res ults, two moults and three in stars have always been observed (Table 2), first moult very premature (two month s after born), second one almost one year later, both in summer (warmest time). Faster growin g during first instar and after each moult (Tabl e 2), less growing at the end of its last larval period when they do not grow, but ove rwinter, and during the autumn-winter of both years (coldest time), when food availability in nature is probably smaller. One specimens got two moults at first year (Table 2), suggesting the occasional possibili ty of reaching a shorter development and to mi x genome from two different generations, as Troge r ( 1993) suggest for N. coo and some of the here related data agree with those recorded for last in stars of thi s species. Growing mensurement s and time between moult s during the larvae ' development was shown by Monserrat & Martinez ( 1995) for L. sofiae, and those for N. bipellllis are shown in Table 2. At the end of the development the larvae start the pupation, making a soft spheri cal silk cocoon ( 12 mm in diameter for L. sofiae and 13 mm in diameter for N. bipennis) wi th accreted sand particles similar to Myrmeleontidae, but much less silk paded internall y, spe nding 7-10 days to co mplete its constructi on. The cocoon of N. coa and D. va l1solli see m smaller (Mansell , 1973 ; Troger, 1993). Time as pupae could not be noted cause a virulent infestati on of "hay itch" mites PyemoteJ vellfrieoSIlS (Newport , 1850): Acari , Pyemotidae (Fig. 38), a mite usually recorded as ectoparasite of development stages of Lepidoptera, Coleoptera and Hymenoptera (Owen Evanset aI. , 1961; Hu ghes, 1976, etc.), mites of this genu s were used as biolog ica l c0 11lrol of different in sect pest, al so of fire ants (Solel/opsis iI/ view Buren), and larvae and pupae of ants are used to breed this parasite (Gerson & Smiley, 1990; Holl dobler & Wilson, 1990), the infestation into culture-boxes could come th rough some field ant-larvae used as food. Different famili es of mites as Pygmepiloridae, Tetrllllyehidae, Erythraeidae, Thrombidiidae etc. have been recorded as parasites on Neuroptera (Hartze ll, 191 8; Smith, 1922; Killington, 1936; Principi , 1956; Parfin & Gurney, 1956; Asp6ck et aI. , 1980; Pantaleoni, 1983; Hagley & Miles, 1987), but previous records do not exist on Nemopterinae mite parasites, nor thi s mite family in Neuroptera. Infested larvae and pupae were immobilized and its development was interrupted, however imagoes fl y in nature
8 DIsch. cnt. Z .. 4.\(1996) I 114 Disch. ent. Z.. 43 ( 1996) I
Tab le 2 Time-growing graph (li near body length ) of three specimens of Nemoplera bipellllis fro m just born larvae, to their pu pation (P), showing in one specimen a cel1ain posibiliLy of overl ap two generations. (Arrows show the ecdysis from an instar to other. E: egg, P: pu pe, I: imago), mm 17 ...... :: ...... - P I 16 • .. - I 15 .. 14 13 .- I• 12 ::.. 11 Ie ---- - u· THIRD INSTAR 10 I . ' • Si!COND IN STA.. 9 .- ! • I- • ''',aT INSTAR 8 - • • 7 ! '" ! 6 • 5 4 "' 3 • 2 E 1 o VII VIII IX X XI XII I II ttl IV V VI VII VIII IX X XI XII I II tt l IV V VI FIRST YEAR SECOND YEAR
30- 40 days laler (Table 2), and some dala can be applied from those recorded by Troger (1993) for N. coa.
3. Biology and behaviour of imagoes
The bi ology of imagoes in Nemoplerinae is much more unknown than in Crocinae. Us uall y restri cted to a single habitat, with pronounced endemisms, the most of species have been scarcell y collected, usuall y prefer arid, sandy or stony countries, with main ly low vegetation, usually crepusc ul ar or nocturnal in their habits, other wings pi gmented genera as Nemoplera, Palmipemw or Knersvlaktia fl ying in bright sunshine. Usua ll y feed pollen and have poor fli ght ability (Tjeder, 1967; Picke r, 1984, 1987). Other in fo rmation on the bi ology, habitat pre ferences, seasonal and altitude di stributi on for European genera, was noted by Popov ( 1970) and AspOck et al. ( 1980). Im agoes of L. sofiae onl y have been associated to windy and dry subdeserti c coast plains and hi ll s, from 10-300 m hight, onl y during 10- 15 days a year always in Ju ly, brief seasonal emergence periods and narrow habitat preference in Nemopterinae have usuall y been recorded V. J. Monserrat: Larval Stages of European Nemoptcrinae 115
(Tjeder, 1967; Picker, 1984). Im agoes of N. hipellllis are locall y frequent and usually fl ying at the end of spring, sooner or later according with latitude and altitude, are assoc iated to open , sunny and not wet plains. hills and mountains, from coast to almost 2000 m, in habitats with abundant low or annual vegetation , from bi g wood clearings to coastal dunes and always in habits under mediterranean influence. Imagoes of L sojiae are active fl ying onl y for one hour or less,just between sunset to almost ni ght dark, in the short wind pau se between see breeze (during day) and contin ental breeze (during ni ght), no specimen have been seen fl ying at day light and none have been coll ected at light-trap, even in the same place and date when, some minutes ago, fl yi ng specimens have been observed. The positive phototropic 10 light in the evening has been recorded in other genera (Tjeder, 1967). Whereas, imagoes of N. hipellllis tl y during dayli ght and are specially acti ves to hiter hours. Win g coloured Nemopterinae ha ve this diurnal activity (Tjeder, 1967; Popov, 1970; Asptic k et aI. , 1980; Picker, 1984, 1987). The almost tran sparent imagoes of L. sojiae are not easy to observe, specially for the spare li ght in this moment, fl y fast and irregul arly over low bush, resting on them (probably feeding, but no many flower existing around du ri ng the fl ying period) with its dark brown hind win gs hanging, but when so me danger is detected, hind wings are opened at maximum , making it s silhouette more di srupti ve, if danger increases, it lifts it s hind win gs to make the other part of the body less vulnerable to an eventual attack. Whereas, the we ll coloured ye llow-brow n wings of Nemoplera agrees with its diurnal activity, and the di srupti ve pi gmentation of its wi ngs imagoes of N. bipellllis remai n motionless on vegetati on if danger is prese nt. The use of wings as cryptic, as visual signalling function and anti-predator roles agai n robber flies was studied in Pa/mipellllll and Kllersv/aktia by Picker ( 1984, 1987) and Picker et al. (199 1, 1992), the robber fli es NeomOClherus (Celt/isIII S) j/avicomis (Ruthe, 183 1) was very frequent in L. sofiae habitat, and Dasypogoll (Se/idopogoll ) diadema (Fabriciu s. 178 1) was frequent in some N. bipennis habit s, and it was seen attacking thi s species. No pollen have been found in the gut of the onl y dissected specimen of L. sofiae, but large amount of pollen of different pl ant fami li es have been found in the gut of some dissected spec imens of N. bipellllis. belonging to Ca mpanul aceae (Campanu/a, i asiolle) and Compositae (Bellis, Cardus, Helialllhus, Taraxacum ), Caryophyll aceae, Cruciferae, Leguminosae and Gramineae, also some of Conifers and spores of molinials fungus. Monse ITat (1977) record imagoes of thi s especies feeding on flowers of Lllvcmdu/a and SarolluUllI lII S and we have seen specimens apparently Iibating on Caryophyll aceae (Silelle scahrij/ora 8ro!.), Compositae (Ambya/a il11 egrifo/ia L.), Labiatae (La vQfu/u/a stoechas pedw1Cu/ata Miller, Th ymus masti c/zina L.), Umbelliferae (Tapsia villom L.) and Campanulaceae (Jasione //lantana L.), usuall y feeding directly on fl owers and also sucking polien stuck to its fore legs. Feeding large quantities of different kind of pollen in many Southafrican genera (Tjeder, 1967), and on Compositae and associated to two kinds ofpolien, probably Mesembryanthemaceae, have been recorded in Palmipenna by Picker (1984, 1987). No mating has been obse rved , and no spermatophore, prese nt in Barbibucca and Nemopte reI/a (Tjeder, 1967) have been detected in Iberian genera. We have not observed in nature fema les laying eggs, but observati ons on female on substrate agree with the possibility of laying on the ground as Picker ( 1984, 1987) records in other species. Number oflaid eggs in laboratory have been commen ted and are indicated in Table I. Longevi ty of imagoes has not been recorded, but some data in laboratory (probably greater in nature), can also be extracted from Table I. ,. 116 Dtsch.enI.Z.. 43 ( 1996) I
4) Distribution and new faun is tics records
Two genera and five species are known in Europe, of these Lertha Jofiae is only kn own from S.E. of the Iberian Pen in sula (Monserrat, 1988) and new records now annoted, and Nemopfera bipen"is was considered as an Iberian species, with a wide distribution on it, but records in north Ibe rian areas with Eurosiberian influence would must be checked, the species have also been collected and recorded long time ago in South of France (Marseille) and in North Africa (Algerie) as N. IlI siwllica or N. eoa as well as N. hipellllis (Burmeister. 1839; Brauer, 1876; Hagen, 1886; McLachlan, 1886; Kirby, 1900; Ko lbe, 1900; Navas, 1903, 19 10, 1926; Lacroix, 19 13; Auber, 1955), recently these o ld records have not or have been d ubiously considered (Popov, 1970 ; AspOck et aI. , 1980), now the french record is confirmed, and it is also recorded in Morocco, so a more West- Mediterranean, that onl y Iberian distribution must be assigned to thi s species.
Lertha sofiae Monserrat, 1988. SPA IN: AlmerIa. El RIo Chico, 21. VII. 199 1, 9 de Y. J. Mon serrat. Rambla de Ba lanegra. 19. VII. 1988.3 00. I 9 v. J. Monserrat, 20. VII. 199 1, I I 00, 5 99 v. J. Monserra!.
Nemoptera hipellllis (Illiger, 18 12).
FRANCE: Marseille. MI . Sl. Ponl. 189 1. l o A. David (MP). MOROCCO: Timadit, V. 1925. 1 Q Escalera (MCNM ). PORTUGAL: Alfeite, Quinlado Brasileiro. VI - VII . 12 00 P. Mcndcx,;a. Campomaior, Embalsc de Caya, 3. VI. 1990. I 0, leg.? (UB). Caparica, 23. VI. 1969. I 9 P. Mend""a. Corroios. Seixal. 6. VI. 1985 12 00, 4 9 9 J. dos Santos. 10. VI. 1985, 4 00,4 9 9 A. Figueira. Riofrio, 14. VI. 192 1, I 9 Wallison (N HM ). Setubal, VI. 1904, I 0 L. Navas (MZB). Sines. I. VI. 1970, 2 99 P. Mend""a. Tras os Montes. Vill a Rea l, w.d., I Q McLachlan (N HM ). Vilanova de Mil fontcs. 3. VII. 197 1. I Q P. Mend""a. 9. VII. 197 1. I 9 P. Mend""a. SPAIN: Alicante. Aspe, VI. 1989, II 00.9 99 T. Garcia, Biar, 24.- 27. VI. 1963, I 0 F. Espanol (MZB). Elche, VI. 1989. 12 00.4 99 T. Garcia. Almeria. Cabo de Gala. 20. VI. 1990. I & v. J. Monserral. San Juan, 30. VI. 1983. 2 I:( Q v. J. Monserrat, Sierra Carbonera, 3. VI. 1911. 30 O. I 9 J. Jacobs (N HM). Avila. Arenas de San Pedro, V. - VI. 1927.5 00. 5 99 A. Schmidt (MCNM), Navaescorial. 16. VII. 1985, I 9 F. Marin. Navalguijo. 27. VII. 1985.2 99 J.L.Nieves. Paradorde Gredos, IS . VII. 1930, 1 0,299 L.Nav;!s(MZB), 18. VII. 1930, 1 0 L. Navas (MZB). 18. VII. 1988, I O. I 9 E. Clemente. Sierra de Gredos. La Cebcdilla. 6. VII. 1985. I 0 J. L. Viejo. Gredos. 18. VII. 1930. I 0 L. Navas (MZB). Badajoz. Valle de Matamoros. 20. VI. 1985.2. Q 2 E. Clemente. Caceres, Guadalupe. V. 1927. 1 0 M. Escalera (MCNM). l arandilla de la Vera. 18. VI. 1990,2 00 C. Ornosa. Plasencia. VI. 19 19, 1 0 L. Navas (MZB). CMiz. Almorai ma, 23. V. 19 11 . I 0 leg. ?(NHM ), cadiz, w.d .. I 9 McLachlan (N HM ), Gibraltar. w.d. , 3 00, 2 99 Walker (N HM ), La Linea. 28. V. 19 11 , I 0 leg. ? (N HM). 20. V. 1993, 4 00. 2 9 9 J. L. Torres. San Roque. 27. V. 199 1. 2 99 J. Ramirez. Ciudad Real. Almaden, VI. 1950. I 0 E. Morales (MCNM). Mestanza, VI. 1950. I 0 E. Morales (MCNM). Puertollano. VI. 1950, 13 00.20 9 9 E. Morales (MCN M). Vi ll arta de San Juan. VI. 1932.3 00.3 99 E. Morales (MCNM). Cuenca. VI. 1953. I 0 L. Higgins (N HM ). Granada, Alfacar. 22. VI. 1927, I O. I 9 B. H. Cooke (N HM ), 19. VI. 1968, I O. 1 9 K. Sauler & DJ. Carter (N HM). Huescar. 8. VI. 1896 I O. I 9 w. N. (NHM), Sierra Nevada, w.d .. I 0, I 9 L. Bequest (N HM). Fuente Hervidero. L1 Zu bia, VII. 1963. 2 99 F. Fernandez Rubio (MCNM). 3. VII. 1989. 6 99 Due ll i & Studer (CPD). Penones de San Francisco. w.d. , 4 00. 3 29 M. Baena, Valle del rio Fardes, Diezma, 18. VI. 1968. 2 9 9 K. Sattler & D. J. Carter (N HM), 22. VI. 1968, 1 0 K. Sattler & D. J. Carter (N HM ). Venta del Molini llo. 6. VII. 19895 99 Duell i (CPD). Huelva. Donana, V- VI. 1957, I 9 G. Mountfort (NHM). Jaen. Linares, VIII. 1934. I 0 L. Navas (MZB). lOrida, Camarrasa. 3. VII. 1909, 2 00 L. Nav;!s (MZB). Madrid. Aranjuez. VI. 1906. I 0 Arias (MCNM). 23. VI. 1975. 2 99 V. J. Monserrat, 28. V. 1987, I 0 V. J. Monserra!. Cenic ien tos, I I. VI. 1972. l o R. Outerelo. Cercedilla. 28. VI. 1934.2 99 L. Navas (MZB). 15.- 26. VII. 1953. I 9 H. Noack (MH). EI Escori al, 29.-30. VI I. 1879, 2 99 L. Bledse (N HM), 12. V. 1919, I 0 L. Navas (N HM). 3. VII. 1934. V. J. Monserrat : Larval Stages of Europea n Nemopterinae 11 7
2 22 L. NAVAs (MZB), EI Pardo, w.d., I 2 BOLI VAR (MCNM), Fuencarral, VI. 1945, I 0 leg.? (MCNM ), Galapagar, 18. VI. 1975, I <3 V. J. Monserral, Hoyo de Manzanares, 23. VI. 1980,2 00 , 2 22 V. J. Monserrat, 9. VI. 1990, I <3, 5 22 V. J. Monse rral, 28. VI. 1992,8 <30, 6 22 V. J. Monserral, 27. VI. 1993, 16 <3 <3, 16 22 v. J. Monserral, 3. VII. 1993,4 <3 <3 .7 22 v. J. Monserral. La Cabrera. 28. VII. 1975. I 2 V. J. Monserral, u.guna de Onligola, 25. V. 1982,300, 3 22 V. J. Monserral, La Pedriza, 19. V. 1934. I 2 L. Navas (MZB), Madrid, 1908, I 2 L. Navas (N HM ), Miranores de la Sierra, 16. VI I. 1977, I 2 J. Plaza. Monlarco, VII , I 0 L. Navas (NHM), II. V. 19 15. I 0 L. Navas (MZB), 15. VI. 1924, 4 00, 16 22 L. Navas (MZB), I 2 L. Navas (NHM ), Navaccrrada, VII. 1964, I 2 F. Fernandez Rubio (MCNM), Pelayos, 18. VI. 1992. I <3, I 2 R. OUlerelo, Pinto, 9. VI. 1975.400 V. J. Monserrat , PUCl10 de Navaeerrada, w.d. , I
S. Systematic considerations
Systemati cs and phyloge netic re lat ionships among different families of Neuroptera on the basis of larva l characters have man y times been forgotten, and most of the cla ss ic general c1a ss itications are mainl y based on im agoes characters. From Withycombe ( 1925) larval morphology, as a crucial point, starts it s modern phyloge netic class itication, and larval morphology has rece ntly induced ve ry interestin g new arguments, discuss ions and hypo lhesis (Popov, 1973 ; Henry. 1978; New, 1989; Brooks & Barnard, 1990; Manse ll , 1992; A spock, 1992, 1993; Aspoek e l a I. , 1980), bUI systemalics and relationship w ithin Ihe Neuro plera, are by no means clear al all (Tauber & Adam, 1990; A spock, 1992, 1993). Without co ming into a genera l di sc ussions, and circumscribin g to Nemopteridae, its syste matic position, as a natural family. is mainl y based on the prese nce of long hind wi ngs and speciali zed mouthparts for pollen and nectar co ll ection, being considered as a monophyletic group close to M ynne leontidae , A sealaphidae and Nymphidae (Ho lzel, 1975; He nry, 1978; Mansell, 1986, 1992). Afler to know belter some mo rpho logical characlers of Nemoplerinae larvae, it is ev ident , that the "close relationships" between Croci nae and Nemopterinae see m to be no more th an artificial. The presence - abse nce of ovi ruptor, ki nd of egg-rupture and aeropyles on egg, as much as so me differencial larval characters as: two seg mented - multi segmented antenna, absence - presence of inner mandibular teeth , four - three seg mented labial palp, 0 10 7 - 610 8 ste mmala, prothorax normal o r specialized and scleroli zed , legs short and digger - long and wa lker, with fore leg tibia fu ssed to tarsus - independent to tarsus, with without lateral thoracic and abdom in al tubercle s, presence - absence of chitini zed structures in hind ventral 7th abdom in al segment, as much as known differences on larval biology and behaviour, ca n no longer sustain the actua ll y considered Nemopteridae as a natural and monophyleti c taxa , many other we ll es tabli shed and defi ned fami lies in Neuroptera have less, and less important differences among their larvae than betwee n Nemopterinae and Crocinae. The two unique argued characters as the specia li zed mouthparts of nectar and poll en nutrition was considered as plesiomorphic character (Brooks & Barnard , 1990) and it is present in other 11 8 DISch. ent. Z.. 43 ( 1996) 1 fam ilies (New, 1989; Mansell , 1992), and the presence of hindwings has also evident funct ional and morphological differences (broad apical dilation - fi lamentous tip, presence - absence of Media vein , presence - absence of pterostigma and freq uent and diverse di fferenciated win g selation and structures do not ex isting in Nemopterinae are prese nt in Crocinae). The prese nce of long hindwi ngs has not an enough phylogenetic weight, by the same character woul d be unthinkab le to reunite the all normal-winged neuroptera into one "natural" family. So, both subfamil ies, (erected by Navas (19 10) with tri be category) Nemopterinae and Crocinae, must be considered with fa mil y rank, fact some time suggested, recently by Popov ( 1973). The description of fa mil y Ne mopteri dae, in Burmeister ( 1839) sense, has a mixed (Crocin ae + Ne mopterinae) concept ion, a wrong origin al nomination (as Ne matopteridae, that it is an incorrect an unjustified emendati on of original latinization of Nemoptera Latrei lle, 1802) and genus Nematopterll as type genus by monotypy. By priori ty law, the famil y name Nemopteridae of Burmeister (1839) remains, but its concept mu st be restricted to the Nemopterini (in sense of Navas ( 19 10) tribe designation), adding their larval characters now exposed. The subfamily Crocinae (in sense of Navas ( 19 10) tribe designation) must be erected wi th a fa mil y status, with the same concept, but adding their larval characters now exposed. So two different fa milies must be considered:
N e m o p t e r id ae Burmeister, 1839 = (Nematopteridae Burmeister, 1839 partim ).
Ty p e ge nu s: Nemoplera Latreille, 1802.
C r oc i d ae Navas, 19 10 = (Nematopteridae Burmeister, 1839 partim), sial. n.
Ty p e ge nu s : Croce McLachl an, 1885.
The systematic pos ition within the Neuroptera of these, now separately considered families, mu st be co mmented. Usually the mi xed charac ters of Croc inae and Nemopterin ae all told, have caused prob lems in the systemat ic position and relationship of the prev ious concept yet of the Ne mopteri dae fa mil y, as well as their two subfamilies (Holzel, 1975; Mansell , 1986, 1992). Under our po int of view, Croc idae, in its new famil y status, must remai n behind, but inside of the same Mansell 's ( 1992) cluster, wit h Myrmeleontidae, Ascalaph idae and Nymphi dae, because the apomorphic traits 10 & I I (long hindwings, speciali zed mouthparts) gived in Mansell 's ( 1992) c1 adogram, have been questioned and trait 15 (developed larval inner mandibular teeth) cannot exclude to Croc idae, becau se its presence is ev ident in genera } osandrevo, Prerocroce, Ausrrocroce, and Ca mavialltl, existing (and recorded as inn er pap il lae) in other genera as Laurhervasia, Thysanocroce, COl1 croce and Tjederia (M onserrat, 1983 a, b; Mansell , 1980, 198 1a, b, 1983 b). On the other hand, some characters of Nemopteridae, in the proposed status, as the presence of oviruptor, lateral tubercles bearing setae and the myrrnecophil y in the larvae, must be added, as other new arguments, to those exposed by Mansell (1992) when di sc uss the close the relationshi p between Nemopteridae and Chry sopidae, long suggested (Navas, 19 10). Also the character 5 (multisegmented fl agellum of larvae) of Asp6cks ( 1992) c1 adogram, would exclude Nemopteridae (i n the proposed status) from M yrmeleonti formi a where it was included, to Hemerobi iformi a where Chrysopidae is included. However, the systemati c position of the Nemopteridae (in the proposed status) and its relationship wi th other families is still not full clear, and new evidences and new larval studies would be necessary to clarify this or any other hypothesis. V. J. Monserrat: Larval Stages of European Nemopterinae ll9
Acknowledgements
I would like to express my cordial thanks to Ora. M. O. Martinez, Dr. J. Barquln, Dr. L. S. Subfas, Dr. M. Portillo, and Ora. C. G6mez for the identification of the ants, mites, robber fli es and pollen here recorded, also to refered InstilUtions and persons who let me study material of the recorded species and to all those friends thal patiently, helped me to search in the field, ant larvae to feed the nemopterids larvae. Also to Ora. L. Diaz-Aranda and Ora. M.e. Roldan for their photograph help and to L. Ghi o for his linguistic improvement.
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