P P P p-p P ------P - - P P Advance5 In Neuropterology Proceedings of the Thlrd International Sympos~umon Neuropterology Berg en Dal. Kruger Ndtlonal Pdrk, R S A , 1988 Mamell, M W 81 Aspock, H (M\) Pretoria, R S A 1990 Pp 131-149 - - - pp p - - - - P - - --
An illustrated review of egg morphology in the families of Neuroptera (Insecta: Neuropteroidea)
Johann GEPP Institut fiir Umweltwissenschaften und Naturschutz. Graz, Austria
ABSTRACT
Thc eggs of 112 species representing 15 families of Neuroptera are listed, and a review summarizing data in 85 references is provided. The external n~orphologyof the eggs is depicted in 12 photographs and 12 drawings, whilst 20 scanning electron micrographs provide details of the micropylar and surface structures. Key words: Eggs, morphology, Neuroptera, Ascalaphidae, Berothidae, Chrysopidae, Coniopterygidae, Dilari- dae. Hernerobiidae, Ithonidae, Mantispidae, Myrmeleontidae, Nernopteridae, Nymphidae, Osmylidae, Poly- stoechotidae, Psychopsidae, Sisyridae.
INTRODUCTION
Morphological details of the eggs of Neuroptera have been discussed by several authors, particularly Withycombe (1925), Killington (1936, 1937), Balduf (1939), Popov (1 973), Hinton (1981) and New (1986). Whilst the eggs of a few families of Neuroptera, especial- ly Chrysopidae, Hemerobiidae and Ascalaphidae, are well known, only a few detailed descriptions of the egg-stage of most other families are available. The eggs of the families Brucheiseridae, Rapisniatidae and Neurorthidae are still unknown.
This contribution presents an illustrated review of present knowledge of the eggs in the families of Neuroptera. Gepp (1984a) recently provided a literature review of thc known larval stages of Neuropteroidea.
GENERAL MORPHOLOGY OF NEUROPTERAN EGGS
The eggs of most Neuroptera are oviform and one and one half to two times longer than wide. Spherical eggs are only known in the family Nemopteridae, and almost spherical eggs are found occasionally in the families Myrmeleontidae (Figs 41, 42), Ascalaphidae (Figs 14, 18) and Psychopsidae (but see also Fig. 16). Elongated eggs (two to four times longer than wide) are found in the Coniopterygidae (Fig. 2), Osmylidae (Fig. 1l), Dilari- dae (Fig. 15) and sometimes in Myrmeleontidae. Eggs with pointed micropylar regions are known in Drep.parzepteryx phuluetzoides Linnaeus (Fig. 19), in the genus Italochrysa Principi (Fig. 30), Psychopsis elegans Guerin (Fig. 16) and in some Coniopterygidae, J. Gepp
e.g. Senridalis czleyrodiformis Stephcns (Gcpp & Stiirzer 1986). The lengths of the eggs vary horn less than 0,4 mm (Sisyridae, Mantispidae) to 5,5 mm and more in Myr- meleontidae. Due to adhesion to the substrate, the underside of the egg may be flat in most species of Coniopterygidae, Osmylidae and Hemerobiidae. As far as is known, all Nymphidae, Mantispidae and, with few exceptions each, Chrysopidae and Berothidae lay their eggs on secreted stalks. Eggs of Myrmeleontidae and Ascalaphidae have two micropyles and the eggs of the remaining families one each (Fig. 19), which are more or less charac- teristically shaped (Figs 25, 26). The structure of the chorion seems to be species-specific in the varioi~sgroups, but extensive and careful comparative studies are still needed. The eggs of most central European species can be distinguished by their specific chorions, with the aid of scanning electron micrographs (Gepp, unpublished data), and the species examined are listed by Gcpp (1986). Papilla-shapcd (Chrysopidae, Figs 35-38; Myrmcleon- tidae, Fig. 43; Dilaridae, Fig 44) or sponge-like elevations (Hemerobiidae, Fig. 24) as well as net-like patterns (Coniopterygidae, Figs 1-4; Dilaridae, Fig. 15 and Nemopteri- dae, Fig. 17) have been observed. The surface of the eggs may be covered in substrate particles adhering to the egg as is the case in some Myrmeleontidae (Fig, 41), Ithonidae and Psychopsidae, or by hairs from the female as in some Sisyridae (Fig. 12). Egg-breakers invariably occur in the families Coniopterygidae (Figs 1,2) and Hemerobiidae (Figs 21,22) but have never been observed in others, e.g. Myrmcleontidae and Ascalaphidae.
REVIEW OF THE FAMILIES Coniopterygidae Species with described eggs: Coniopteryx exigua Withycombe ( = pusana Withycombe): Narayanan 1942 Coniopteryx parthenia (Navas & Marcet): Killington 1936 Coniopteryx tinefirmis Curtis: Killington 1936; Figs 1, 3 Coniopteryx califorrzica Meinander ( = hageni Banks): Quale 19 12 Conwentzia pineticola Enderlein: Collyer 1951 Conwentzia psociformis (Curtis): Figs 2, 4-6 Conwentziu sinica Yang: Yang 1974 Parusemidalis fuscipennis (Reuter): Withycombe 1923a Semiclnlis albata Enderlein: Yang 195 1 Semidulis aleyrodiformis (Stephens): Killington 1936; Gepp & Stiirzer 1986 Semidalis vicinu (Hagcn) : Muma 1967 The Coniopterygidae have mainly elongated eggs (Figs 1 & 2) which, in most cases, arc whitish or reddish initially but may also be green, grey or darkly-coloured. The chorion may manifest various patterns (Figs 3 & 4), whilst the micropyle (Fig. 5) and fine struc- tures (Fig. 6) provide species-specific characters with the aid of a scanning electron micro- scope. The bottom is flat or adapted to the surface. Empty cgg shells are white.
Ithonidae Species with described eggs: Ithone&.scu Newman: Tillyard 1922; Withycombe 1925; Riek 1970 Oliarces clarc1 Banks: (Faulkner, unpublished data)
The eggs of I. &sca (Fig. 7) arc deposited in sand or loose on the surface (sizc: 1,7 X 0,9 mm), and the number of eggs laid by a single female varies from about 200 to almost Egg morphology in Neuroprera
Fig- I -t~. SEL1 (1 t tg;c 131 tl~c l;~rni[yC'on~uptcrycrtl,ic.: I . C;c.r~cr;ll view II~egg-hhcll of Cor~rtrytt.r?:rtitltifi>rrr~i, CurIir (Icnpth 0.5 1 nlnl): 2. Gcncriil view uf an cpy-shelt ofCorrn~er~r:ta psnrifnr- IIII.~(Cunis) (Icnglh (1.5? rnlub; 3, Hnneyrcrrlth pattern (>Cur! cgg of C. rirrr,ifi>rrr~is:4. Honcycrrmh pallcm of an cgg of C. />.corililrn~i.r-:5. MlcropyEc ol' an egg or C. p.~,sric~fnrx~ris;h. Chr>rionic niicro-pattern of a lioneycornh of an cgf nf C, p.cr>rifi~rt~ris. Figs 7-10. Oviform egg-types of the families (frorn left to right) Ithonidae, Osmylidae, Polystoechotidae and Sisyridae: 7. Eggs of lrhont~,fisc.aNewman (length 1,7 mm), the surracr is quite smooth, creamy-white, without sculpturing but covered with sand and soil particles after laying ((redrawn from Tillyard 1922); 8. Egg of S~c,nosmylu.s fenuis (Walkcr) (length 0.85 rn~n),pale ycllow when fresh, darkening to pale-brown then dark-brown heforc hatching (redrawn from New 1974); 9. Egg of Poly- .storchorrspuncmtus Fabricius (length 0.93 mrn), the surface is ~moolhand free of surface structul-es (hut ~ninutelygranular), chalky-white, later green (rcdrawn from Hungcrford 193 I); 10. Egg of Clirncrc.in cir.eo1uri.c (tfagen) (length 0.34 mm), is glistening, whitish and wnii-tranrparent when frehhly laid (drawn from a photo by Brown 1952). 300. The anterior end of the egg bears a raised disc-like micropylar knob. The surface is quite smooth, without visible sculpturing, but coated with sand. Faulkner (pers. comm.) knows the eggs of another undescribed North American species, which are laid close together in sloping rows; the micropyles are white, the egg shells colourless and translucent. Osmylidae Species with described eggs: 0.srrlylu.s fulvicephulus (Scopoli): Withycombe 1925; David 1936; Killington 1936; Ward 1965 St~rlosmylustenuis (Walker): New 1974, Fig. 8 The eggs of 0. &lvicephalu~ are relatively large and are laid on the upper surfaces of leaves (Fig. 11). The eggs are attached to the substrate by their flat side and are laid in close contact, side by side, to form a straight or slightly curved row. They are rarely laid singly.Withycombe (1925), Killington (1936, 1937) and New (1974, 1986) provide fur- ther information on this family. Polystoechotidae Species with described eggs: Polptoechntes punclatus Fabricius: Welch 19 14; Withycombe 1925; Hungerford 193 1.
The eggs of P. punctutus are laid singly, they are stalkless, ovoid (O,9 X 0,s mm), chalky- white with a fincly granular surface (Welch 1914) and each with a button-like micropyle (Fig. 9). Sisyridae Species with described eggs: Climacici arcoluris (Hagen): Brown 1952 Slcjm fi.sca (Fabricius): K~llington1936 Sr~xrcrterrnintili.\ Curtis: Withycombc 1923a, 1925 Egg morphology in Neuroptera
Sisyridae lay very small eggs in small clusters in depressions, especially along the veins of leaves overhanging water. The egg of S. fi4.sc~ru is elongate-oval (length 0,35 mm) and bears one ~rlicropylarknob; the female occasionally deposits some hairs on the egg (Figs 10 & 12). Berothidae Species with described eggs: Lorriutnyia .fluvic-(1rni.s(Walker): Smith 1923 Lornurtiyia lalipctznis Carpenter: Toschi 1964; Tauber & Tauber 1968 S~~crrnophor~lludi.sseminatu Tillyard: Tillyard 19 16 S1)erinophorella rnuci~htissimaTillyard: T~llyard19 16 The eggs of S. disscminutu are deposited on stalks: several batches of such eggs were found, ranging fio~n30 to 50 eggs each, which closely resembled the fruiting bodies of small clulnps of moss. The stalk is hollow throughout except at the base and tip. and is so delicate that it sometimes bends under the weight of the egg (Fig. 27) (Tillyard 1916). The eggs of L,. lutipentzis are also stalked and are laid in small clumps comprising fewer stalks than eggs (Fig. 28) Toschi 1964. Tauber & Tauber (1968) report that the stalks are apparently very flexible and do not support the eggs as do those of the eggs Chrisopidae, but serve to suspend the eggs from a substrate. Smith (1923) described an urntalked egg of LomamyiaJlavicornis; the egg was infertile and the only one which was laid by a female in captivity. Gurney (1947) is certainly correct in assuming that the unstdked condition may have been abnormal. Mantispidae Species with described eggs: Climacicllu brzrntieu (Say): Balduf 1939 Clitt~nciellutncrgtzu (Miyake): Kuroko 196 1 Mnntispu intcrrzrptu Say: Hungerford 1936 Mantispa jrlpotzica McLachlan: Kuroko 1961 Muntispi~s~qi Ranks: Smith 1934 (in Balduf 1939) Mnntispu .styriacu (Poda): Brauer 1852; Schrenimer 1959 Mantispu viridis Walker: Davidson 1969 Mantispa vittura Guerin: McKeown & Mincharn 1948 The author has seen eggs of Inany species (all belonging to the subfamily Mantispinae) from different parts of the world; they are all short-stalked (Fig. 29) and very small, elon- gated ovoid and all appear very similar. The stalks of M. suyi are 0,66 mm long and the eggs 0,39 mm long (Balduf 1939). In some species of Mantispidae the number of eggs laid is enormous, for example more than 8000 in M. irztcrruptn (Hungerford 1936) and even more than 35 000 in M. uhleri Banks (Redborg & MacLeod 1984). Mcrtztispu stjriir- t.a deposits several hundred eggs within three square centimetres within a few hours (Fig. 13), but a female repeats this several times, so the total number of eggs laid by this spe- cies may exceed 6000 (Schremmer 1959). The stalks are apparently (as in Berothidae) much weaker than in Chrysopidae, and eggs are laid in such a rnanncr that thcy hang their stalks. Brauer (1887). Lewis (191 l), Bristowc (1932), Balduf (1939) and Poivre (1976) provide further information on the eggs of Mantispidae. Dilaridae Species with described eggs: Dilur turcicu.~Hagen: Popov 1973; (Gepp & Popov, in preparation) Na1lachiu.s umc~r-itunms(McLachlan): Gurney 1947; MacLeod & Spiegler 1961 J. Gepp
Fig% I I- 13. Eggc rrf the fnrnilies Osmylidac, Sisyridae, Manrispidac and Ascalaphidac: I I. Egg clusters or OJIII~L~.\jrh~irepl~nlti.~ (Scopoli) on fhc uppcr surface oC a lcaf (egp lcngth 1 .R mm): I?. Egg clusters rrfS~s,vrlrjrr<(crr~(Fabricius) (egg lcnprh 0.35 mm): 13. Clutch of Mnlrrir~srtriarrr IPrda) egg\ U 11h shon sr:tlka (egg length O,4 nlnlE: 14. Ilpp oTIibc/korif~.~nrurunnritr.~ IL~nnacu~) la~d nn a prass %ten](egg length I .X mlnl (photos I. Gcpp).
The unstalked eggs of Dihr fnr-cic~~sHagen are laid singly or in small groups in bark crcviccs (Fig. 15 1. Although they are sometimes substantially deformed by insertion into small spaces, it apparently does not affect their viability. The eggs are 3,5 times as long as broad, conspicuously elongated for a neuropteran, but have one typically knobbad micro- pyle and a chorionic pattern resembling that of some Coniopterygidae. with a fine struc- hrre like Chrysopidac (see Figs 35 and 44). The eggs of Ni~ll~chiusamerimnirs(McLachlan) are laid closeEy packed toecther and ate pale prey ish-while with a very thin unmarked chorion (MacLeod & Spicgler 1961).
Psychopsidae
Species with described eggs: Psydtup.~i.~~rk~gons (Gudrin) ( = rie~vnoni) Froggatt: Gallard 19 14; Tillyard 191 9 P.~~c/~np.sisrr~il~lirn Newman: Froggatt 1902; Gallard 1923 Eggs of P. e1qgan.s are laid singly. or in groups of two or three. on the bark of EEIMI~~IUS trees (Fig. 16). The oviform eggs (1,O x 0.45 mm) have a perceptible micropyle. The eggs of Cnhralis gloriostrs Nariis are almost sphcrical and covered with a greenish secre- rim, presumably consisting of plant tissue collected in the genital Iracr ofthe female. The ovoid egp of S. nrciilrn and S. mursti~lli,laid in captivity usually have a sand covering (Minter, Mansell. Gepp. unpubIished data) and Tjeder (1960). 0l5e..;$:-
F ,-
,$ '4 S,, .-' L: :g;.;-
l:#*, - h,!?:.,-;,."J'< 11.. !+(. tj*; -; l?7*..'.. , l '7; ",:, . ?,l>-. : g b.;:::;-,'i 3 *;:/ l 3 Figs 15.1%. Eg~sof thc rarnilrcr I!cCt 10 right) Dilaridac. Psychopsidac. Ncmtipleridae ;and A\c;sl;~pludac I5 Egg ol Ililor rrrrr?rrrs Hascn with a dcl~c;~~choncycomh pairrrn tlunglh 0.8, mm): !h. E$? or P.rv
FI?\ 1') A 20. 14. I'\%rl c?? clurtc~\ ui I)rc~prrrfcprcr>I plrrrlrrc.rrt)~rk.r Il.~nn.~cur).20 I?yp 114 Ht.rrfrrrdritr.5prni Sicphsn5 on ,i xpnic~ncrtllc (Ilcnpth 0.H nm).
Spccies with dcscrihed eggs: Drcpa~icrrt-Uhirloc~rlr~ (Ncwnlan): Ncw 1975 Drcpa~~epr~grphlil(~etroidc.c (Linnacus): Mnrrnn 1910 Hemrmhirr.~krrrnti1intr.s Linnacus: Smith 1923: Killington 4 437 Hememhius rnargitrr~rlrsStephens: Figs 22, 24, 26 H~tnerohittsnirid~dus Fahricius: Wit hycombe 1923a Hertrerol>iuspc~c~fir-tls Banks: Neuenschwander 1975 Hrm~rr~/)iusp~r~legntis Stephens: Killington 1936, 1937 Hrmerol)ius pini Stephens: Killington 1937 Herrrrrnhit~ssfigmo Stcphens: Withycombe 1922 Megu/nm~rshintu (Linnaew): Killington 1937 Mirrotntrs angiilnrus (Stephens): Miermont eL Canard 1975 J. Gepp
Microniu.~[)(IKCI~L(S (bnnaeus): Ki 11 inglon 1936 Microrrius posticus (Walker): Smith 1922 Micronii~stnstnaniar (Walker): Husscin 1984 Micrornrns variegatz4s (Fabricius): Duelli 1986 Nesornic~rotnuswgus Perkins: Terry 1908 Psectra tlipt~rcr(Burmeister): Killington 1936 Syrnphc.rohiw arnicrilus (Filch): Smith 1923 Syn7pherohiu.s arnicus Navas: (= jirllux Navris): Rivnay 1943 Synlpherohiu.~elegnns (Stephens): Withycombe 1923a Syrriph~rohiusfrrscescetzs (Wallengren): Withycombe 1923a Synzpherobius pyyrnaeus (Ranlb-ar): Withycombe 1923a We.smaelius c,oncinnu.s (Stephens): Withycombe 1923a; Figs 21. 23, 25 W~smrreliusjc1c~li.s (Banks): MacLcod 1960 Wesrnt~eliusnervosus (Fabricius): Killington 1937 Westriae1iu.s quatlrifuscitrtus (Reuter): Killington 1937 Wesmtre1iu.s .suhnebulosus (Stephens): Withycombe 1923a The eggs of Hemerobiidac are elongate-oval and bear a small terminal rnicropylar knob. Egg stalks are unknown in this family, and the eggs of most species are laid on their sides in direct contact with the substrate (Fig. 20). Specific factors that determine the position of eggs are known. Species of the genus Sytnpherohius Banks lay small conspicuous eggs in small clusters. he central ~urb~eans~&ies of ~ernerohiusLinnaeus show s~ec&- specific deposition sites (Fig. 20), but there are almost no differences in the forms of the clusters. Drepa~z'pteryx pha1uetzoidc.s (Linnaeus) lays magnificent star-like egg-groups on leaves (Fig. 19). Psectru Hagen, Micrornus Rambur and We::mnelius Kyiiger Iay mostly single eggs. Eggs of the gcnus We.srrzuelius have conspicuous micropyles. In L:! genera, the micropyles (Figs 25, 26) are white during the entire egg stage, the larval segtlientation pattern gradually becoming visible through thc patterned chorion (Figs 21-24).
Chrysopidae
Described eggs of more than 50 species of Chrysopidae are listed in the following papers: Duelli 1984; Gepp 1984a, b; Gepp (in press). Our knowledge of oviposition, egg-stalk and egg morphology in Chrysopidae is extraor- dinarily coillprehcnsive compared to other families. Anlong the central European species, five egg deposition forms can be distinguished: (a) single deposition, c .g. Ti:>cierir7ngracilis (Schneider), (b) in more or less regularly spaced rows or star-like around twigs, e.g. Nothochrystr ,fi*lvic.tys (Stephcns): Fig. 3 1. (c) in a loose group, e.g. C/zrysoprrlr~ctri-nea (Stephens): Fig. 34, (d) as a cluster with ceparated stalks. c g. Yirirto ~qun/kirmm~r!sis{pictet): Fig. 3?, (e) as a cluster with iinkeci stalks, e.g. Mullurla j/tr\ijroiz.s (Rrauer): Fig. 32. Whcn laid in clusters, each egg also has its own stalk (Fig. 40). Specics with short-stalked eggs cicposit their eggs mainly on pine-needle tips and leaf cdges. Nothochrysa ccipitata (Fabricius) distributes an oily secretion on the egg stalks. Unstalked eggs are known from Anotnak)chnsa sp., but could also be pathologically induced in other species (Duelli 1984). The number of eggs in each clutch may vary widely within the same species. The micropylar knob is somewhat flattened (Fig. 39) and saucer-shaped. The chorion has a waxlike sur- face (Figs 35-38), with its fine structure and dimensions being characteristic of the spe- cies. The eggs are generally ovoid (Figs 31-34). but are spear-like in the genus Itrtlochrysa Principi (Fig. 30). Fipf 21-26, SEM of cgp of thc family Hemcruhiidac: 'I. Egp-\hcll\ of Syr~rpltcmhirrr r>yyn~(rcrrrIRarnhtsri flcnfth 0.46 nini): ??. Epp->hell rrT Ilrott,rohi~~strtrrr,vitrtrtrrs Slcphcns (lcnplh n.R? mm}: 23. Chor~on~uStnlcturcs of vn cyg of S /ygnwrrrrr: 14. Chor~onicslruuturcn of an crp or H. mnr- vrlrrrrsrs: 25. Micropylar bnoh of an cpg c1C.5. pcmtclt*lc~r:16. Micropylar knob of an cgg al' 1-I. srnr~ir~nna. F~gi17-30. Stalked egg\ of the f:~nlilic\ (fro111 left to right) Herothidac, Mrintispidae and Chrysopidae: 27. Eggs of Spe~r-rirr~~~hor~~lleclis.\c~trli~rutr~ Tillyard (length wilhout stalk 0,75 m~n),the stalk ih exceedingly fine and creani-coloured l~kethe cgg (redrawn from Tlllyard 1016): 28. Egg clusters of Lr)iiitririyici lcznpc2irnisCarpenter (egg length 0,6 mrn, stalk 6 nim), the chorion is dull white and has a honey- cotnbed appearance (drawn from a photo in Toschi 1964); 29. Egg of Climuric~lluri~ujincl (M~yake) (length 0,37 mm),ivory in colour, sniooth with faint pearl) lustre, but later turns dark purpli\h-brown (redrawn from Kuroko 1961): 30. Egg of I~i~lnc.hrysciitnlrc,c~ (Koss~) (length 0.47 mm) pediccl partly shown (redrawn from Pr~ncipi1916).
Nernopteridae Species with dcscribcd eggs: Crow ,filipenni.s Westwood: Lefroy 19 10; Im~ns19 1 1 Lcrirrller\,asiil rlu~nibicuMansell: Mansell 1980 Lrrlwhcr~sci.siri srtuccw (Klug): Mansell 1980 Cot7c.r-act cupcrzsis Tjedcr: Mansell 198 l a Concroce parvcz Mansell: Mansell 198 lit Conc.rocr wulkeri Tjeder: Mansell 198 1a Tjedcrin hrevicor-rlis Mansell: Mansell 198 1 b Tjrllrrirl nurnrryuen.si.s Mansell: Manscll 198 1b Jo,suizrlrc~~usazi NavBs: Monscrrat 1983a; 1985 Ptc,roc.roc.e c,~zpillnris(Klug): Monserrat 1983b K~rei~vl(ikti~~tligropteru Picker: Picker 1984 iVc,rno/~tc~nrbiprtlnis (Illiger): Monserrat 1985 The ~~nstalkedeggs of Nc~nopteridaeare deposited singly. They are small, spherical or ovoid, with one micropyle. Withycorilbe (1925) reported an egg of Crow McLachlan without a micropyle. The surface of the chorion differs considerably between the various species, being smooth or with regularly spaced the structures (Fig. 17). Nymphidae Species with de\cribed egg\ N,mnphes m~rr~zeleonzclee.,Leach Gallard 1936, New 1986 O\rnjlop~Banks. New 1,986 Em morphology in Neuroptera
FI~3 1-34, V:~renu\ cgp-rleprlc~luu+ ~ncll~rrdcin Chrysopidac: 31. Two egg< ot Nnrlrr~t~l~r~x~r fitI\-j~-ep~(Srcphenl): 32. Egg ulusbcr r>d ,+t~~llt~(lnfiv~r~r~,r(Rrnucrl with linked stalks: 33 Egg clurrer or Nitwtn A~i~rrrk~irrrne~rtir(l'ictet, with ?;cp:tralc stalkr: 34. A Irmrc egg-group nf Clr~.top~rlrrC~~TIIPO (Srcphcnrl (phntoh J. Gcpp). Fip 35-38. SE%!\ oI' p.ipjtlit-4iuputl chtrrronic niicrtlpatlcrnh or cp~sof Clrrysttpidac: 35. Cltr~.+op~~111Lt-ri %fcl-a~,l~l;~n : M. UI~Y.~,IJ!~I)/J~~~ (.iIitili# (Wchm:~c!): 37. M61llffdi1 t~rtlw~li~(Cud is): 3R. hlitrt.rtr rrrrprrtll,rrrrlr ( Rcutcr). ht~l?lph~~.\tt~)lyrt~l~lro)li(l~*.\~ litys eggs (length 2 mm) in a rernarkable.ur;uallyresular, pattcrn in o horseshoe-shnpcd arranycmcnt of stal ked eggs lin kcd by brid_ecsof non-stal ked eggs. E;~chhi~tch contains about 30 cggs (Gallard 1936)
Myrmeleont idae Spccics with dcwrihed cgp: Arwtirh~tc-lisi.~I?(~~ritn Ramhur: Principi I947 Frrr~l~~r~t~plttrnherrs (Olivicr): Gcpp & Hiilzel (1989) E1rrr)i~iwtttosrrris I Fourcsoy ): Piotrowski 1969 GI~~rrol~~n~ihrrer.t)ptrr?.r Gcrstaccker: W ithycornhc 1925 Myrtac*lcorr ~-~*IP/?I,II.Y~.TM~IC h l;ln : Rong-Me:tng 1 930 IWyrttt~l~ot~,fi>l-tt~ir+~it-itl~ Linnncuh: Mcissncr 1909 Uy~itii)c+~~rt~tic~~nrir>:srrf~ (Schncidcr): Gepp & Popov (in prcpararion). Despite numerous artempts at observation, there is very little information on the numbers of eggs and oviposidon behaviour in Myrmelcontidae. Egg morphology in Neuroptera
FIRS 39 Rr 40. 39. SEM of egg of Mullfidn $'orfins (Rraucr) (egg lenglh 1.0 mm): 40. $EM of the base of thc egg sralks of Ninrrtt flww (Scopnli).
According to previous authors, the numkrs of eggs of Euroleon nosfras (Faurcroy), for example, range between five and cight with no rcferencc to the deposition of more than 1 1 cggs. The known eggs of European species arc oviform. about 1.3 to 1,7 times longer than wide and are mostly brown or pale green. The chorion is finely sculptured, with a dull sheen (Fig. 43) and the surface of Gymnocnemia variegafa (Schneider) eggs resem- bles that of Chrysopidac (Gepp & Popov, in preparation). The two mil ky-whire micro- pyles occur at each cnd of the egg and are perceptible only in newly laid eggs. During oviposition, the surface of the egg is coated with a glandular secretion to facilitate the adhcsion of substratc particlcs to the cgg (Figs 4 1, 42).
Ascalaphidae
Species with described eggs: Asmloptym frrrci~er(MC Lachlan): Henry 1972 Asculorphnr itnpuvi(1n Walker: New L 97 1 ICnrd~rlecenrsnlopcrittrrs Rurmeister: New 197 1 ; Hcnry 1978 Colol?npfenr.rdi.wintilis McLach tan: New E 97 1 Epispur~fre.~nrennstrs Walker: New 1 93 1 H~licornitwsdicar Wal kcr: Ghosh 19 13 Helirot~tir~rsfestivu (McLachIan): Henry 1978 Hybris s~rbjncmrWal kcr: Sonan 1938 tihelloides coccajus (Denis 8 Schiffermiiller): Henry 1978 Lilwdloi~Eestnacr!ronft4.~ (Scopal it: Brauer 1 854; Wcstwood 1 888 Supknlrmitus rnala.vantts jar~otrer~sisWecle: Henry 1978 Uliiloris !?~acl~ayonaiijnhata Burrne~stcr: Ncw 197 1 Ulitlncles a~lrifc.rn Mc Lach Ian: Ncw 1 97 1 Utitlo(1es Iry(11infl ( = set~csBurmeistcr): McClendon 1902 Uiulode.~me,ricann (McLachlan): Hcnry 1972: 1973: 1978 The cggs of most species of Ascalaphidae are ovoid (Fig. 14). some have longish-ovoid or almosr square eggs. They are relat lvely long: e.g. Hr~licontirusjesti~srs (Mchchlan). 1,2 mm; Corduirc~rrtsalopecinus Rurmeister, 2,3 mm (Henry 1978). They have two micropyles, onc at each end and the cggs arc usually attachcd at one end to !he suhstsate. J. Gepp
Fips 11-44. 41. Epg uf Cri,ulca~r plrt~nl~crrr IOtiviur) cnvercd with rand: 42. Eggr ol phmrh~rtsInicl In a \andlcc:, Gtuation in ~ht.latmralury: 13. SEM oFpapilla-shad uhorionic micro- patlern rrf an ugg 111 G~tt~rrorrrrrrrinlmirrfrgt:t(l (Schnc~dcr): 44. SEM of uhnrionic n~lrrtl-palternclP an cgg ol Dtlnr trrr<-rrlrr Hagen.
Eggs are often deposited in two parallel rows on a straight grass stern as in Sib~lilluloides Iorrgirnrnis (Linnaeus) or laid in clouble rows in transverse groups around twigs (Balduf 39.193. Many Ascalaphidae in the New World have interest in^ oviposition habits where egg hatches bf some species arc hordcred at each end by repag;la (Fig. l U) (Henry 1972, IL177. 1973).
DISCUSSION
Mare than 60 years have elapsed since Withycombe's (1925) phylogenetic arrangement of the Neurnpteta based on immature stages. and the number nf species with described eggs ha< increased [rum 30 to mare than 150. Despite some fundamental discussion on relationships of e~gsin thc families of Neuroptera (e.g. Killington 1936, Popw 19731, no attempt has hitheno been made to provide a general review. Table 1 provides a com- parison predominant and sp~radiccharacteristics of the eggs of each family in the Neu- roptera. Egg morphology in Nc~uroptera
Oviposition habits cover a wide range. Merrlbers of most families deposit eggs on leaves or on the bark of trees. Egg dispersal in flight is not reliably documenteci. but possible in some species of Myrmeleontidae. Psychopsidae and Nemopteridae. Deposition in sand or soil is only known in the families Ithonidae, Myrmeleontidae, Nemopteridae and Psy- chopsidae. The total number of eggs laid by one fernale differs from less than 20 in Eu- rolcon nosrrcls Fourcroy (Myrmeleontidac) (Piotrowski 1969) to over 35 000 in the family Mantispidae (Redborg bi MacLeod 1984). A large number of eggs per deposition has been reported for most species of Mantispidae, Ascalaphidae and, as far as is known, Dilari- dae. The number of eggs per batch may evcn differ within the sallie genus, from singly deposited eggs to clusters of several dozen eggs clumped together eg. as in Nineta Nav6s (Chrysopidae). Only species in the family Ascalaphidae show a definite arrangeincnt in their deposition of eggs. The development of stalked eggs has presumably been indepcn- dcntly evolved in the diffcrent families, and can be traced back to the existencc of adhe- qivc secretion glands (see also Duelli 1986). The cxisteiice of two niicropylar-like knobs on each pole of the egg in the families Myrrneleontidae and Ascalaphidae has been fre- quently discussed (e.g. Hinton 1981), but it has never been conclusively proven whether
T:ible l MORPHOLOGICAL
NEUROPTERAN EGGS
a predominant sporadic
Families:
Rerothidae Mantispidae ---.------p----- Dilaridae
p----p- Psychopsidae Brucheiseridae
-- ---p-- Hemerobiidae Chrysopidae -- - .. - .-p---- Nemopteridae --.---p------.~-~- Nymphidat!
- ---p---- Myrmel.eontidae
p-- .--p ------. AscaI.aphj.dae one of them has an aeropylar function. Mansell (1980, 1981) terms as aeropyles, the globu- lar or bower-shaped structures on the chorion in some species of Crocinae, and a similar terminology may be applied to structures in a few species of Hemerobiidae (e.g. Fig. 24). It is interesting too, that there are only a few observations of egg-laying in some species- rich groups such as Myrmeleontidae, where there are only short accounts of oviposition behaviour (Piotrowski 1969), and no detailed paper on egg production. Egg-breakers are present in the families Coniopterygidae (Figs 1, 2) and Hcmcrobiidac (Figs 21, 22), but have never been observed in the families Myrmeleontidae and Ascalaphidae. They may be present or absent in different species of the same family, as in Nemopteridae. The shape of the egg-breakers could also be useful indicators of groups (Coniopterygidae, Hemerobi- idae), but this possibility has never been thoroughly investigated.
ACKNOWLEDGEMENTS
I owe special thanks to the Zentrum fiir Elektronenmikroskopie in Graz, (leader W HR Dr. H. Horn) for the production of the SEM photos. I thank Miss Franziska Feichter (Graz) and Miss Carol Veenstra (Graz) for some preparations (Figs 21-26) and Dr Alexi Popov (Sofia) for loan of material (Figs 43, 44). I am grateful to Mag. Wilhelm Draxler (Graz) for the carefully prepared drawings.
REFERENCES
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Address of author: Univ.-Doz. Dr. Johann GEPP Institut fiir Umweltwisscnschaften ~~ndNaturschutz A-8010 Gru, Heinrichctrassc 5 Austria Bibliography of the Neuropterida
Bibliography of the Neuropterida Reference number (r#): 7264
Reference Citation: Gepp, J. 1990 [1990.??.??]. An illustrated review of egg morphology in the families of Neuroptera (Insecta: Neuropteroidea). Pp. 131-149 in Mansell, M. W.; Aspöck, H. (eds.). Advances in Neuropterology. Proceedings of the Third International Symposium on Neuropterology (3-4 February 1988, Berg en Dal, Kruger National Park, South Africa). South African Department of Agricultural Development, Pretoria. 298 pp.
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Notes:
File: File produced for the Bibliography of the Neuropterida (BotN) component of the Lacewing Digital Library (LDL) Project, 2010.