55 On the Possible Myrmecophily of Nemopterinae Larvae (Neuroptera, Nemopteridae) by V.J. Monserrat & M.D. Martinez' ABSTRACf Field and laboratory studies of different species of Iberian ants with the eggs and larvae of Nemoptem bipennis and Lertha so.fiae (Neur­ optera: Nemopterldae: Nemopterlnae). suggest a previously unknown myrrnecophllous life-style In the larvae of this subfamily. A wide range of potentially edaphic prey was offered to newly hatched larvae. but growth occurred only when ant larvae were used as food. Furthermore. ant workers harvest Its eggs. as though they were seeds. and Introduce them Into their nest. Ant tolerance oflarvae inside the nest Is proved and discussed. and the acquisition of the colony odor or some other appeasement mechanisms against ants Is suggested. SImilarities In the ecological requirements of the Nemopterlnae Ima­ goes and harvester ants hav,e ,been observed and a coevolutionary process among plants (as seed producers). nemopterlnes (as plant pollinator Imago and ant-predator larva) and ants (as seed harvesters and predator lodgers) Is suggested. KEY WORDS: Myrmecophlly. larval-predation. Neuroptera. Nemopterlnae. coevolution. lNTRODUCfION Among the many myrrnecophllous arthropods deSCribed. which are symbiotic. commensals. or predators of ants. the order Neuroptera Is scarcely represented (Holldobler & Wllson 1990). Only larvae of the tribe Belonopteryglni (Chrysopldae) have been recorded as llvlng In colonies of ants. Other families such as Berothidae. Dllarldae. or Mantispldae are associated with other social Insects. as obligate larval predators oftermUe nymphs or of wasp and bee larvae (Parker & Stange 1965. Johnson & Hagen 1981. Brushweln 1987. Minter 1992). One of the most Interesting of the neuropterous famllies Is the Nemopterldae. which has specialized nectar and pollen eating Ima­ goes. It Is divided Into two subfamilies with very different morphology and habitats. The biology of the Croclnae subfamlly. with almost 50 species. Is well known Including larval stages. taxonomy. distribution

IDepartamento de Biologia Animal I. Facultad de Blo)ogia. Uruversidad Complulense E-28040 Madrid. SPAIN 56 Sociobiology Vol. 26, No.1 , 1995 and phylogeny. This subfamily has been revised recently (Holzel 1975, Mansell 1986). In contrast, the Nemoptertnae contains almost 100 species, includtng some of the biggest and most spectacular of all Neuroptera (Fig. 1). The imagoes are usually abundant, have short emergence periods, have crepuscular and diurnal flying habits, and usually have pronounced endemisms and are restricted to a single habitat. Their taxonomy and systematics need revision, and there is surprisingly a very little data on the biology of most species (1Jeder 1967, Mansell 1973), and to date, there is very little information on the morphology and biology of preimagtnal stages in this subfamily. Eggs, first or last larval stage or pupae are only known on the Mediterranean Nemoptera bipennis (Dufour 1857, Navas 1919, Withycombe 1925, Monserrat 1985), N.coa (Troger 1993), a few references of N.sinuata (Popov 1963, 1973). There are also a few references about the Austra­ lian Chasmoptera hutti (Mathews 1947) and on the South Mrican Knersvlaktia nigroptera (Picker 1984), Palmipenna aeoleoptera (Picker 1987) and Derhynchia vansoni (Mansell 1973). Nemoptertnae larvae have always been found in the ground, living freely in loose sand, sometimes at a certain depth (l5-25cm) beneath the surface. Attempts to rear them have always falled, only Troger (1993) persuaded last instars to feed on wasp, butterfly and fly larvae and pupae, while some other poSSible prey, such as small arthropods, Tlpulids, Noctulds, or Tenebrionlds have been suggested. Pupation has been observed twice, also in the ground, 5-1 Ocm beneath the surface in the fleld or O.5-lcm beneath the surface in laboratory conditions (Mathews 1947, Mansell 1973, Troger 1993). The supposed predatory behavior of larvae in this subfamily and an edaphic life-style has long been suspected (Wheeler 1929). These have been conflrmed by some of the recent findings, but many other aspects of their biology and behavior remains unknown, such as relationships with other animals, life-cycle time, the number ofins tars and molts, and sources of food in the wild. In this paper, some new data are presented on the imagoes behavior, morphology of eggs, larval instars (reared in laboratory) and cocoon of the Iberian Nemoptera bipennis (Illiger) and Lertha so.fiae Monserrat. These provide new information on the larval biology and behavior of this subfamily, and demonstrate evidences of possible myrmecophily in the Nemoptertnae larvae.

Figs. 1-5. 1, Morphology of a Nemopterinae imago,Lerlha sofiae; 2, Micropilar area,L.scfiae egg; 3, Newly hatched and recuperate eight months age larvae ofNemoptera bipennis; 4, Medium third instar larva of N.bipennis. eating an ant larva; 5, Enlargement of a portion of RgA. Monserrat & Martinez · Myrmecophilous Nemopterinae 57 58 Sociobiology Vol. 26, No. 1, 1995

MATERIAL & METHODS Eggs of Nemoptera bipennis were obtained from gravid females collected from different localities in Spaln during the last eight years and eggs of Lertha so.fiae from specimens collected in Spain (Almeria, Balanegra,20.VII.1991). Each female was Isolated in a plastic box of lOx 5 x 5cm, In order for her to oviposit. Some moisture, a few grains of commercial pollen and some fixed flowers were provided in each box. No records were made of temperature during the first years of unsuccessful attempts to rear larvae in the laboratory, but when an adequate feeding method was known, the monthly range of the temperatures of both years, under which the larvae were reared was noted in °c as follow: I: 14-21. II: 10-23, 1Il: 14-25, IV: 16-23, V: 19-32, VI: 15-37, VII:21-41, VIII:22-38, IX: 17-30, X: 14-22, Xl: 17-23, XlI:7-22. The photoperiod was taken to be a natural month, but In dark partial shade, not full light, and a relative humidity of 40-600A>. After birth, the larvae were Individually Isolated into plastic culture boxes of2 x 4.5 x 4.5cm, halffllled with sand. Newly hatched larvae of these species had an obvious predatory appearance (Fig. 3) and their soil dredging behavior confirmed an edaphlc life-style. Consequently soil fauna should constitute the usual diet. Nemopterins usually occur at high densities, so the larval food cannot be unusual or too temporal. However, because of their small size, different soil organisms (prefer­ ably non-predacious), as well as non-edaphlc soft ones such as aphids, and flies, beetle larvae or butterfly larvae, were buried in the sand, as possible prey for the newly hatched larvae. Food provision was mixed and renewed every 15 days. A wide range of prey was offered separately to different larVae, each kept isolated in an individual culture box. Prey included Nematoda (Rhabditlda: Rhabditldae) and the eggs, nymphs, larvae, pupae, co­ coon or imagoes of different Arthropoda (Acari: Oribatida; Thysanura: Leplsmatidae; Diplura: Japygidae; Collembola: Poduroidea; Dermaptera: Forflculidae; Embioptera: Emblldae; Isoptera: Kalotermitldae; Psocoptera: Psocomorpha; Homoptera: Aphididae; Coleoptera: Bruchldae, Tenebrionldae, Curculionldae; Diptera: Drosophilldae; lepi­ doptera: Tineidae, Pyralidae; Hymenoptera: Formicinae and Myrmicinae) (Table 1). Larvae moved and fed freely within the sand, and to study molting, during all stages of development, the sand was carefully sieved every 15 days, to look for possible exuvia. Laboratory experiments of possible interactions between ants and Table 1. List of the different prey offered to newly hatched Nemopterinae larvae, and the time (-) until their death (+) or pupation (PP). ;:: MONTHS 0 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 0 ..~ PREY '"~ ..!!!. NEMATODA (+J ;:: SOIL MITES (+J ..a. SPRING TAILS (+J ..~ SILVERFISH (+J N DIPLURA (+J ;:: EARWIG EGGS (+J '< EMBIOPTERA (+J 3 TERMITES - (+J 8 PSOCIDS (+J ";1: APH IDS 0- (+J ~ BEETLE LARVAE - (+J '"z FLY LARVAE (+J .. FLY IMAGOES (+J B MOTH LARVAE (+J ~ ANT LARVAE (PPJ ~ . ~ ANT PUPAE (+J .. ANT COCOON (+J ANT IMAGOES (+J

ifl 60 Sociobiology Vol. 26, NO. I, 1995

eggs or lruvae were made using Chauvin type ant-culture cyllndrtcal boxes (l2cm In diameter x 5cm high), In which ant colonies were maintained and fed with a Bhatkar diet. Field expertments with nemoptertne eggs - lruvae and ants were carded out on different locations within the Murcia and Madrtd provinces, where 7 newly hatched lruvae were released on small plots of sand, Isolated by an open metallic cylinder 50cm long and 30cm In diameter, vertically engraved Into the sand. For hruvestlng expert­ ments, 10 Nemoptertnae eggs were placed near different ant-nests (In the field) and 5 eggs on the foraging surface of Chauvin ant culture boxes (In the laboratory). The ants used In these expertments are noted In Table 2. and ant lruvae used as food were taken In the field are noted In Table 3. RESULTS The eggs obtained were not adhesive and were Isolated Into small vials to observe their development time. The number of eggs laid was 0-13. x = 6 by L.so.fiae and 0-51, jf = 25 by N.bipennis. Electron microscope studies were made of several (Fig. 2) . A) Laboratory Experiments After 20-24 egg development days, the hatched lruvae rest for some minutes, then burrow head first, vertically Into the sand, to a depth of 5-lOcm, where they remain without further burrowing. When dis­ turbed, the lruva lay motionless, as If dead, for hours In any posture,

Table 2: Ant species used, and time spent by workers, to harvest and to introduce in their nests (+) len (in field) or five (in Chauvin boxes) Nemoplerinae eggs layed near by their nests. (""": granivorous, .": omnivorous, *: flouidophagous ant species).

DAYS ANT SPECIES 2 3 4 5 6 7

Messor barbarus"· + Messor stllJCto,·" + Lasius niger·· + + Lasius flavus·· + + Tetramoriumhispanicum ... + + Aphaenogasterseni!is·· + + lridomyrmex humilis·· + + Pheidole pallidula"" + + Tapinoma nigerimun u + + Camponotus Belhiops" + + + CamponotuslateraJis'" + + + Camponotuspilicomis·· + + + + + Plagiolepispygmaea" Monserrat & Martinez - Mynnecophilous Nemopterinae 61

Table 3. Larvae-Ant Species used as food of Nemopterinae larvae.

Tetramorium hispanicum Emery T.semilaeve semi/seve Andre T.semilaeve depressum Forel Messor structor (Latreille) M.barbarus (Linnaeus) M.minor maurus Santschi MonomOrium sa/omonis (Linnaeus) Fonnies pyrenaea Bondroit Lasius alienus (Foerster) Aphaenogaster senilis Mayr and then briefly burrowed actively. Most larvae died In a few days (Table 1). as they were unable to feed on most of the prey Items offered. Some other larvae (fed with aphids, beetle and butterfly larvae and formlclne cocoons) lived for some months, but with either no appre- Table 4. Growth (Iinearbody length) ofdifferentspecimensofLerths sofiaBfromfreshlyeclosed larvae to their death (+) or pupation (P) fed with ant, moth, or beetle soft larvae. Arrows which correspond roughly to the symbols of the food indicate ecdysis; the direction of the arrow is irrelevant. E-egg, P-pupa, and I-imago. mm 17 .- -.- .- --r- - -- - ,- r -,.· -- - - ._- - 16 .- _.. - --- - I- 1- - --.- - -- I- ~ - , p I ~ . : ' 15 . - . -- -- r -, • • • - 14 --- - r- - i - 13 -, 12 ._- --- 11 -- ANTS 10 - -.- 1-- •® MOTHS 9 -- -- . -- - - ~ - ~ . - --- 0 f- B .-. - - -t ~ - ._- - BEETLE ~ ,- . -- •_. .- - - . 7 M t I- I-- - 1- - --

. - - . _. _. Ii> .~ - -- 6 .. ~- p- - 5 -. Ii> p: tS: . Q": !+> r-- ,- - -. - - -- . 4 .,~ ~: ~ _.- 1- - - -- I------~ . 3 . ~ - - . --_ . _- -- 2 f-- 1-- - ,- .• --t- - r- E~ -_. 1 -1-- ...... - .. --_ .. -- - ._- - -''-1------o VII I IX X Xl XII I II III IV V VI VII VIII IX X XI XII I II IV V VI VII FIRST YEAR SECOND YEAR'" 62 Sociobiology Vol. 26, No.1, 1995

ciable growth or with very Irregular growth. These died In their second instar (Table 4). Only those fed with myrmlclne pupae, and especially with ant larvae (Figs. 4,5) grew properly and underwent early ecdysis (Table 4). Those individuals reared on ant larvae throughout their development had 2 molts and 3 Instars, growth measurements and the time between molts Is given In Table 4. When full-grown (2 years later), the larvae start to pupate (Table 4), making a soft spherical silk cocoon of 12mm In diameter, covered with accreted sand particles. B) Field Experiments Involving Ants and Nemopterine Eggs and Larvae The first hint of a relationship between ants and larvae was the recovery In the field of a well-grown first Instar larva of N.bipennis In association with ants. Some newly hatched larvae of this species, which had been released In July 1982 on a small plot of sand In coastal dunes were Isolated In open metallic cylinders, vertically Inserted Into the sand. The larvae burrowed vertically Into the Isolated sand, and disappeared Immediately. Every two months, a half-liter sand sample was taken from the sand cylinder and the sample was washed and filtered, In an attempt to recover some larva. In February 1983 (eight months later), a full-grown first larva was obtained (Fig. 3), associated with many specimens of lridomyrmex humilis (Mayr), a very common ant In this and all' previous samples. No other larva was obtained In other samples. If Nemopterinae larvae are predators of ant-larvae underground. rather than predators of ant-Imagoes on the surface (like Myrmeleontidae), they must have evolved a mechanism for entering ant nests In order to feed. The possible way of reaching the ant nest was demonstrated In the field. where several groups of ten Nemopterinae eggs were laid on the ground. near nests of different ant species. Workers harvested the eggs and carried them Into their nests. with a frequency depending on the particular diet of the ant species. Similar experiments were made In the laboratory, with different ant species In Chauvin boxes. In this case. both eggs and newly hatched larvae were cropped and Introduced Into these artificial nests. Table 2 shows the variety of ant species used. In the field. we lost track of the harvested and Introduced eggs. but these are probably stored as seeds In the nest. and newly hatched larvae could move freely within the colony searching for ant larvae to feed on. But In Chauvin culture boxes, these larvae have no possibility of burying and hiding. and eggs and larvae were eventually devoured by ants. So. as a further study. some eggs or larvae were placed on the Monserrat & Martinez· Myrmecophilous Nemopterinae 63 surface of an earthenware plant pot (26cm Wgh x 26cm In diameter) half filled with sand. where a colony of Tapinoma nigeTimun Nylander had been established. Workers were seen to harvest the eggs and Introduce them Into the nest. Seven months later. the colony had died. but the nest was washed and filtered. using two sieves of 2 and 5mm mesh. In an attempt to recover some living larva. However. Just one exuvla (from first to second Instar) was obtained. DISCUSSION Eggs of the well-known Nemopterlnae species and those reported In tWs paper are lald individually. and they are not adhesive. Self- gluing eggs are recorded only In Palmipennaand Knersvlaktiaby Picker ( 1984. 1987). Nothing is known about how females lay eggs In nature. despite many hours of field observation. The way In which females of many species rest and settle on the ground. sand or rocks. wWch are clear of vegetation cover suggests an edaphlc Oviposition (Popov 1963. 1)eder 1967. Picker 1984 and personal observations). If the hypothesis of the myrmecopWly In these larvae Is correct. they have three possible ways of reaching the ant nest: 1) Directly by females laying eggs near the entrance or Into the nest (this seems Improbable because of the female risk. the remote possibility of locating the tiny ant nest hole for a not-very-preclse fiylng Insect. and the unspeclalized female terminal genitalia). 2) By the larva positively searching for an ant nest. while excavating Into the sand or ground (this Is Improbable since larvae do not walk on the surface and they dig for only a short time after hatching. Moreover. females lay low numbers of eggs. 3) Freely lald eggs are harvested by foraging ants. and so directly Introduced Into the nest. where. after hatching. they will find their food. This last hypothesis seems the most likely. due to the seedlike appearance and hardness of the egg. Moreover. the conspicuous micro pile resembles an elalosome (Fig. 2J. and can be easily mistaken for seeds by ants. Table 2 shows the time taken by different ant species to Introduce Nemopterlnae eggs Into their nests. As expected from the ant diet. the relative size of the ant and Nemopterlnae eggs. and the maturity and number of ant workers of the colony In Chauvin ant nests. the eggs were harvested faster In granivorous ants. They took more time to be harvested by omnivorous ants. and were not harvested at all by fiuld feeding or by small ant species (Table 2). The morphology of cropped eggs. resembling seeds. harvested by ants from other Insects Is similar to the Nemopterlnae. and their introduction Into the nests have been recorded frequently (Carroll & Janzen 1973. Compton 64 Sociobiology Vol. 26. No.1. 1995

& Ware 1991. Hughes & Westoby 1992a}. These field and laboratory experiments may also demonstrate It for the Nemopterinae. The biochemistry of the chorion has still to be studied. The adhesive eggs laid by females of other Nemopterlnae genera (Picker 1984.1987) suggest a more specific oviposition In these spe­ cies. or that young larvae search directly for ant nests. or that the larvae themselves are harvested and Introduced Into the nest. The harvesting of myrmecophilous Insect larvae by ants Is well known. for example among lycaenld butterflies. We also demonstrated that foraging ants harvest Nemopterlnae larvae In the laboratory. and the dead-like behavior. of newly hatched Nemopterlnae larvae for such long periods. could be an adaptation for this. Evidently. predators and omnivorous ants could adopt Nemopterinae eggs or larvae. but granlvorous ants are probably more disposed to harvest and store them as seeds. especially as It Is known that granlvorous ants Introduce other materials. including Insects Into their nests (Carroll & Janzen 1973. Risslng & Wheeler 1976. Whitford et aL 1976. 1980. Whitford 1978. Holldobler & Wilson 1990). These eggs may be eaten by ants. but only the capitulum (as elaiosome) Is eaten by ants In some other myrmecophilous Insects. similar to some types of seeds. Other types of seeds are not always consumed. or are not eaten Immediately (Carroll & Janzen 1973. Whitford 1978. Brown et aL 1979. Holldobler & Wilson 1990. Compton & Ware 1991). Thus egg viability In the nest cannot be reduced. or a certain period of time may allow the larvae to hatch. bury themselves. and hide. The eating of eggs and larvae In our Chauvin boxes has been explained. and can probably also be Induced by the protein requirement In the nourish­ ment offered to ants. The experiment In using a sandy ant-nest In a flower pot. demonstrates that not all eggs are devoured. and that larvae can live In a nest without any food source other than ant-larvae. This fact was also demonstrated In the laboratory feeding experiments (Tables 1. 4). Therefore some defensive mechanism must be attributed to the larvae. It Is Interesting to speculate that larvae possess defence mechanisms against ants. Obviously. larvae born with the colony odor are less likely to be attacked. but other chemicals cannot been discounted. such as neurotoxins. allomones. Immobilizing. repelling or adhesive secretions recorded In other Neuroptera larvae associated with social Insects (Prlncipl 1946. Kennet 1948. Johnson & Hagen 1981. MacLeod & Redborg 1982. Brushweln 1987). The ants resulting from uneaten pupae that had been placed as food In the culture-box. were observed reorganizing sand and other pupae. and even walking on the larva. but showed no aggressive behavior against them. Other -

Monserrat & Martinez· Myrmecophilous Nemopterinae 65 strategies to minimize ant aggression seem to exist. These can be classed as passive mechanisms. such as the clYPtic body color and the presence of dolichasters on Its Integument which help to fix sand particles on the body (Figs. 4. 5) or as active mechanisms. such as the death feigning behavior and the obvious burrowing activity. These make larvae less conspicuous In the nest or allow them to hide In the sand outside nest galleries. Other aspects of the morphology and biology of Nemopterlnae larvae and Imagoes support our myrmecophily hypothesis. In the 3rd Instar the combination of a relatively soft Integument. and a large body relative to the legs. make It too weak and voluminous to be supported or to move on Its own (Fig. 4), let alone to move through almost compact soil. Such larvae would not need to move or displace themselves often If there Is abundant food nearby (as In an ant nest). or If they do move only along galleries made and opened by other soil organisms (as In an ant nest). Among the Nemopterlnae larvae found In loose desert sand. this may be not a problem. but the depth where larvae were found (Mansell 1973). suggests a . very difficult and tiresome progression for free and blind dredgers. so the myrmecophily hypothesis may also be applied here. In other related families such as Myrmeleontidae. Nymphldae. or Ascalaphidae. even In Nemopterldae (Croclnae). with psammophilous and edaphic free predator larvae. the body and legs are different. The legs are much longer In Croclnae which live on the floor among sand and dust. frequently with burled abdomens. but with their necks and heads outside. These are able to move forwards as well as backwards along the surface. The legs are much bigger and stronger ___ In other families which allows the body to burrow backwards Into the surface like Myrmeleontidae which live Just under the free sand surface. or which sometimes build sand-pits. In other species. the body Is flattened as some Nymphidae and Ascalaphldae which live In litter or under bark or stones. However none have such soft bodies and short legs as were described for the Nemopterlnae (lJeder 1967. Pieper & Willmann 1980. Mansell 1980. 1983a. 1983b. Monserrat 1983a. 1983b. Stange & Miller 1990). Having developed In the ant-nest. the last question to answer Is how the Imagoes reach the open air. Pupation occurs In the field In soil 5- 10 cm beneath the surface. Decticous and exarate pupae and an spherical sand-cover cocoon are known for all Myrmeleontoldea. and obviously In this subfamily (Mansell 1973. Trager 1993 and In the species here studied). Nemopterldae pupae cut the cocoon. move and walk. reaching the soil surface by themselves. from whence the Imagoes later fly. In the Nemopterlnae pupae the nest galleries could 66 Sociobiology Vol. 26, No.1, 1995 make It easier, and some of the defence mechanisms of larvae against ants may also apply here. Finally, It is known that Nemopterlns are usually abundant insects (Tjeder 1967), so their larval food cannot be unusual or too temporal. Ant colonies satisfY this. Ant colonies are also consistent with the extended development time and high food consumption of the larva. These require abundant and permanent food resources which granivo­ rous ants, in particular, with their large and permanent colonies can provide. These ant nests are also found in the dry, sunny and open biotopes where Nemopterinae imagoes fly and where myrmecochory is decisive for many plants (Carroll & Janzen 1973, Brown et aL 1979, Hiilldobler & Wilson 1990, Hughes & Westoby 1992b). The ant toler­ ance to larval predation in the colony can be recompensed by the imagoes pollination of plants whose seeds are later harvested by ants, in an evident and venerable coevolutionary process among plants­ ants-nemopterlnes. This may explain the highly specific and localized habitats of most Nemopterlnae. Thus despite no Nemopterlnae larvae ever having been found in nature in ant-nests, these experiments, taken as a whole with the direct or indirect arguments, suggest the myrmecophily as the Ufe­ style among the Nemopterinae. ACKNOWLEDGMENTS We wish to express our cordial thanks to all our friends who patiently, throughout these three years, helped us to search in the field for ant larvae to feed the nemopterid larvae. REFERENCES Brown, J.H .. O.J.Reichnan. & D.W.Davidson 1979. Granivory in desert ecosystems. Ann. Rev. Eco!. Syst. 10:201-227. Brushwein, J.R. 1987. Bionomics of Lomamyia hamata (Neuroptera: Berothidae). Annals entomo!. Soc. Amer. 80:671-679. Carroll, C.R.& D.H. Janzen, 1973. Ecology of foraging by ants. Ann. Rev. Eco!. Syst. 4:231-257. Compton, S.G. & A.B. Ware 1991. Ants disperse the elaiosome-beartng eggs of an African stick insect. Psyche 98:207-213. Dufour, L. 1857. Fragments d'anatomie entomologique. !. Sur l'appareil digestif et les ovaires du Nemoptera lusitanica. Ann. Sci. Nat. 4, VIlI:5-IO, pl.! . HOIldobJer, B. & E.O. Wilson 1990. The Ants. Belknap Press of Harvard University Press, Cambridge, Mass. 732 pp. HoJzel, H. 1975. Revision der Netztlugler-Unterfamilie Crocinae (Neuroptera: Nemopteridae). Entomo!. Germ.2,1: 44-97. •

Monserrat & Martinez· Myrmecophilous Nemopterinae 67

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