Revta brasil. Bot. 10:117-123 (1987)

Food web relationships involving Anadiplosis sp. galls (Diptera: Cecidomyiidae) onMachaerium aculeatum (Legum~nosae)

O. WILSON FERNANDES'. ROGÉRIO P. MARTINSl and E. TAMEIRÃO NETOl

ABSTRACT- (Food web relationships involving Anadiplosis sp. gal/s (Diptera: Cecidomyiidae) on Machaerium aculeatum

(Leguminosae». The characterization and occurrence of Anadiplosis sp. , new species, (Diptera: Cecidomyiidae) leaf galls onMachaerium aculeatum (Leguminosae) were studied. The food web centered upon the leaf galls was composed of six speciesof hymenopterans (two species of Platygasteridae, two species of Eurytomidae,one species of Tanaostjgmatidae, and onespecies of Vespidae), three species of Berytidae (Hemiptera), one species of Geometridae (Lepidoptera), and three species ofSalticidae (Aranae). The existence of herbivores utilizing galls and causing death to Anadiplosis sp. indicated asymmetricalcompetition, a "new" aspect to be explored in gall studies. The intensity of parasitism on Anadiplosis sp. larva was 64,0070(SD:t 1.4). Galls possess hairs with sticky secretions which trapped and killed insects of several orders, including parasitoidsattacking the gall maker larvat...

RESUMO -(Relações tróficas envolvendo as galhas de Anadiplosis sp. (Diptera: Cecidomyiidae) ern Machaerium aculea-tum (Leguminosae». Foram estudadas a caracterização e ocorrência das galhas foliares de Anadiplosis sp., nova espécie(Diptera: Cecidomyiidae) em Machaerium aculeatum (Leguminosae). Seis espécies de himenópteros associados (duas espé-cies d~ Platygasteridae, duas espécies de Eurytomidae, uma espécie de Tanaostigmatidae e uma espécie de Vespidae), três es-pécies de Berytidae (Hemiptera), uma espécie de Geometridae (Lepidóptera) e três espécies de aranhas Salticidae, compuserama teia alimentar baseada nas galhas foliares. A existência de herbivoros utilizando as galhas e causando a morte do cecidóge-no sugere competição assimétrica, um "novo" aspecto a ser explorado em estudos envolvendo galhas. A taxa de parasitismoem larvas de Anadiplosis sp. foi de 64,0070 (D. P. :t 1,4). Pêlos com secreção pegajosa, presentes nas paredes externas dasgalhas, foram responsáveis por aprisionar e matar insetos de diversas ordens, dentre eles, parasitóides do cecidógeno.

Key words -Anadiplosis, herbivory, insect galls, Machaerium aculeatum.

Introduction

The interest in studies on insect galls, besidestheir importance in ethnobotany (Berlin &Prance 1978, Fernandes & Martins 1985) andbiological control of weeds (Berube 1978, Hart-nett & Abrahamson 1979, Peschken 1979,Peschken et al. 1982), can provide importantcontributions to theoretical and evolutionaryecology. The knowledge of galling insects hascontributed to understanding patterns of com-munity structure, in aspects of three trophic le-vel interactions (host plant -herbivore -enemies(Price et al. 1980», and in applied ecology(agriculture, management and conservation offorest areas). Studies on insect galls are rare forthe neotropical region, notwithstanding the ap-parent abundance and diversity of these orga-nisms (Fernandes & Martins 1985).

Machaerium aculeatum Raddi (Legumino-sae), "jacarandá de espinho", is a tree speciesof widespread occurrence in the Brazilian "cer-rados" (Hoehne 1941) whose leaf galls havebeen described morphologically by Fernandeset al. (1982, 1987). These galls are caused by anew species of Anadiplosis sp. (Diptera: Ceci-domyiidae, nearA. venustaTavares, R.J. Gag-né, personal communication). The descriptionof this species is currently in preparation. Tava-res (1916) described the genus Anadiplosis andthe species A.pulchra and A. venusta. Anadi-plosis pulchra were obtained from sphericalleaf ga"s on an unidentified species of Mimo.S'a(Leguminosae), while A. venusta was also ob-tained from sphericalleaf ga"s on an unidenti-fied species of Machaerium. Besides these, twoother species, A. caetensis and A. procera, weredescribed by Tavares (1920): the former in ga"sof an unidentified legume and the latter in ga"sof another unidentified species of Mimosa (Le-guminosae).

The aim of this study was to describe thefood web centered on Anadiplosis sp. ga"s onleaves of M. aculeatum and the intensity ofparasitism on the ga" maker by hymenopteranparasi.toids.

I. Northem Arizona University, Department of BiologicalSciences, P.O. Box 564G, Flagstaff, Arizona 86011,U.S.A.

2. Departamento de Biologia Geral, Instituto de Ciências.Biológicas, Universidade Federal de Minas Gerais, Cai-xa Postal 2486. 30000 Belo Horizonte. MO. Brasil.

o. w .Fernandes. R.P. Martins & E. Tameirão Neto118

Material and methods

Galls were observed from August 1980 to December1982. The great majority of observations occurred betweenAugust and November 1981, when ga11s were most abun-dant.

In the field. nine host plants bearing galls were labelled .with flagging tape and samples of 50 to 400 galls were col-lected from them weekly for the evaluation of attack fre-quency by gall maker and parasitoids. and for biometriccharacterization of the galls. We collected 3.500 galls be-tween August to November. 1981. Due to the low abundan-ce of galls. samples were not taken frequently in 1980 and1982.

Galls were measured. to the nearest millimete!:. alongtheir longest axis using calipers. Adults and larvae werefixed in 70C7/o ethanol and mounted on slides. Insect speci-mens were kept in the authors. collection. and at the UnitedStates Department of Agriculture (USDA). Samples of theplants were deposited in the Herbarium of the Departamen-to de Botânica of the Universidade Federal de Minas Ge-rais.

Results and Discussion

Galls were mainly located along the leafrachis, and only very rarely occurred on newstems or leaflets. The number of galls per leafwas extremely variable (figure 1). When numer-ous, galls were aggregated and displayedsmaller dimensions without the spherical shapewhich i§ a characteristic of single occurence.

200~

1501

lets were abscised and rachises becameapparently dried and hypertrophied. Galldevelopment begins the following year beforevegetative development of the host plant (figure2). Later in the season, we oberved new gallsbeing developed on new stems and leaflets.These galls were morphologically similar to theones developed by the diapausing larvae.

Anadip/osis galls were found on hostplants between the months of August and No-vember, coinciding with the period of vegetati-ve development of M. acu/eatum. Due to theannual periodicity of growth and leaf initiationin M. acu/eatum and the fact that galls are in-duced in undifferentiated tissues, Anadip/osisgalls are phenologically restricted and synchro-nized to the growing season of the host plant.Insect-host plant phenological synchronizationmay be an important factor in regulating insectand plant populations. The Anadip/osis life his-tory is completely synchronized with the lifehistory of its host plant. Brewer & Skuravy( 1980) studied the phenological synchronizationof a gall former and its host plant and showedthat plants and plant organs avoided massiveattack by the gall former by growing at diffe-rent rates at any given time. Larval diapause oc-curs from November to August in leaf rachiseswhich are apparently dry and hypertrophied.We suggest that maintenance cif diapause, be-sides occurring in cold and dry conditions, allowssynchronization of Anadip/osis adults with ve-getative growth of the host plant.

We suggest that the leaflets are selectivelyabscised because galled leaves shed their leafletsmuch earlier than do ungalled leaves. The appa-rently dry and hypertrophied shapes of the leafrachises may be related to the protectionagainst enemies, but more work is called for toelucidate the mechanisms and processes invol-ved.

100

~>~IAI-1

~

m:IAIG)2~z

50

o -9

~17

~2S

-33

-41 4' ST as 7!

The oviposition carried out in young leavesis probably related to the ability of these stilldifferentiating structures to react in a specificmanner to the influences of the galls maker, aswell as to nutrient availability in these growingorgans (Mani 1964, Rohfritsh & Shorthouse1982). According to Mani (1964), this ability toreact specifically seems to vary with the age andthe stage of development to the affected struc-ture. The ocasional development of galls onnew branches and leaflets may be due to ovipo-sitional mistakes, or to competition for oviposi-tion mistakes, or to competition for ovipositio-

NUMBE" OF GALLS

Figure I. Oistribution of Anadip/osis galIs among infectedIeaves.

Anadip/osis sp. began oviposition in leafbuds at the end of August, soon after the emer-gence and mating of the adults. At this time, M.acu/eatum was producing new stems and leaveswhere the females oviposit. Eggs were laid intothe plant tissues. Eggs hatched a few days laterand the first instar larva entered diapause fromNovember to early July. After oviposition, leaf-

Food web relationships in galls 119

nal sites with displacement of the less competi-tive females. Shifts in the host organ attackedand in the time of attack could eventually leadto the formation of a new population or race ofAnadip/osis isolated from the prime populationby developmental time and/or behavior. Toanswer this question, experimental studies onthe relative abundance of ovipositional sites,and studies of the behavior and fitness of thetwo "populations" of Anadip/osis should beperformed.

Anadip/osis galls are greenish in color ,with short hairs' which are distributed alI overthe spherical surface. Hairs are the first structu-res that appear at the oviposition site. During

the early developmental stages of the gall, smalldroplets of a sticky secretion (from 0.2 to0.6mm in diameter) are produced at the apicesof the hairs. This secretion ends as the gall ma-ker ends feeding activity. Once gall develop-ment is complete, a small circular area (about2mm diameter), which is lighter in color thanthe rest of the gall, appears. From this circular ,portal-area the adult gall maker later emerges.Generally, after emergence of adults, the pupa-rium remains in the emergence hole (figure 2).Subsequently, the galls dry up and the affectedleaves are abscised.

An average of 10.81 (SD :t 10.24) galls perleaf were found (range of 1 to 73 galls per leaf,

Figure 2. Life cycle of Anadiplosis, new species, (Diptera: Cecidomyiidae) on its host plant Machaerium aculeatum (Legu-minosae). (A) After mating, females oviposit in leaf buds of the host plant. (B) First larval instar enter in diapause from No-vember to August in the apparently dry and hypertrophied leaf rachis. Leaflets are selectively abscised during gall develop-ment. .(C) Anadiplosis larva initiate gall formation before vegetative growth begins in the host plant. (D) Cross-section sho-wing larva inside a gall; arrows in D show direction expansion of galls. (E) The adults emerge from a small, circular portal,area leaving their puparia in the emergence hole.

o. w .Fernandes, R.P. Martins & E. Tameirão Neto120

n = 308). Figure I shows the frequency distri~

butiôn of the number of galls per leaf. The samedistributional pattern of galls among infestedleaves was observed by Washburn & Cornell(1979) in Acraspis harta galls on Quercus pri-nus. The mean diameter of gall size was 5.lmm(SD ::!: 0.7, n = 1,522, figure 3). Each gall con-

tained a single cavity.with only one larva in it.

.00 -

and/ or consumed by the inquiline larvae whichtakes over the gall. Of 1,107 observed gal1s, 706were parasitized corresponding to a total of64.0OJo (SD :t 1.4) parasitism in 6 host plants(table 1). Platygasteridae species " A" produced

multiple larvae in each gall parasitized. Anaverage of 13.70 (SD :t 5.37, n = 201) Platy-gasteridae species ' , A' , were reared per A nadi-

plosis larva. The frequency distribution of thenumber of the Platygasteridae species " A" per

larva of the gall maker is shown in figure 4. Forthe remaining parasitoids, only one larva perhost was found.

The high percentage of parasitism (table 1)indicates that parasitism is probably importantin population regulation of the gall former. Thealmost normal distribution of the number ofPlatygasteridae sp. " A" per larva of Anadiplo-

sis (figure 4) suggests a behavioral variability inthe pattern of oviposition 'of this parasitoid.Variation may be due to positively graded eggnumber increasing with host-gall size, effects ofegg depletion over several oviposition bouts, orpolyembryony. These possibilities cannot be se-parated at present.

1/1 500

:1., 400...O8: 300"'~2 200jZ

100

o3,0 3,~ 4,0 4,5 ~p 5,5 sp 6,5 1,0

DIAMETER(mm)

Figure 3. Oistribution of leaf gall diameter of Anadiplosis.

Larvae of Anadip/osis are parasitized bytwo unidentified platygasterids, and two uni-dentified eurytomids (Hymenoptera). In addi-tion, at1acks by one species of Tanaostigmati-dae have been observed. Most of the NewWorld tanaostigmatids are gall makers, withsome records of seed infesters (Gomes 1942),and a few species whose biology is unknown(LaSalle, personal communication). Gallswhich are parasitized by tanaostigmatid waspsare recognizable by their tougher consistency,larger size and different shape. It has beenshown that some inquilines modify the normalpattern of gall development (Shorthouse 1973,1980). No gall maker larva was found in gallswhich contained the tanaostigmatid wasp. Thececidomyiid larvae are presumably killed

...w .w .w NUMBER OF PLATYGASTERIDAE sp A

Figure 4. Oistribution of Platygasteridae sp. " A.. on Ana-

dip/osis larvae.

Table 1. Overall gal1 number and percentage o! parasitized Anadiplosis gal1s.

INDIVIDUALS OBSERVED*PLANT

TOTALc D E FA B

No. of larvae or gallsexamined per plant. 270 1107168 36 124 38471

No. and (percentage)

of parasitized larvae. 248 (92) 706 (64)274 (58) 115 (68) 18(50) 35 (28) 16 (42}

.Ali measurements were taken during November 1981

121Food web relationships in galls

The phytophagous hemipterans Jalysus SO-brinus, Parajalysus pal/idus, and P. spinosus(Berytidae) feed on the galls. Previously, onlyadults of P. pal/idus were described, howeverwithout any records of host plant and life cycle(T. Henry, personal communication). For J.sobrinus, only the host Nicotiana tabacum (So-lanaceae) has been described (Wheeler & Schae-fer 1982). Females probably lay their eggs onthe galls of M. aculeatum and the nymphs andadults feed on gall tissues. The berytids extendtheir proboscis and penetrate the gall with thestylets in order to obtain the fluids containedtherein. This renders the attacked galls softerthan unattacked ones, and results in the deathof the Anadiplosis larva. The death of the larvaoccurs indirectly through the feeding activity ofthese phytophages .

A geometrid (Lepidoptera: Geometridae)larva chews through the gall walls, opening thelarval chamber and causing the larva to fallfrom the gall. Larvae which fall from galls pre-sumably die from exposure and/or predation.

Gall tissues are a modified product of thehost plant which are induced by the gallmaker .Most of the gall maker life is spent inside theseniodified tissues. There, the gall maker findsrich and abundant food, protection against cli-matic factors, and in some cases protectionfrom natural enemies. However, galls are notfree from being eaten by other organisms, orpredators. The gall "predators" should be con-sidered competitors of the gall makers, as theycompete with the gall former for the gall re-source. The existence of a geometrid speciesand three berytid species utilizing the galls andcausing death to Anadiplosis indicates a possi-bility of strong asymmetrical competition(Lawton & Hassel 1981) by these herbivoresagainst the gall maker. Since the gall maker iskilled by the feeding activity of the competitors,'Ye believe 1Jlat the dominant species are the ift-~~~~hile the gall former is the subordina-te species in the system. The mechanism: bywhich the gall maker is excluded from the foodsource by the geometrid is probably exploita-tion of resources and habitat modification. Theinteraction between the berytids and the gallmaker is more difficult since the Anadiplosislarva may be killed by habitat modification, re-source exploitation or by direct aggression anddeath. unfortunately, these interactions cannotbe cla!ified without extensive additional work.Studies of gall communities may offer num-

erous examples of competition between gall ma-kers, inquilines and other associated arthro-pods. This contrasts with the view of Lawton &Strong (1981), who state that although ants andbees provide many good examples of competi-tion, competition for food between phytopha-gous species is unusual.

One species of Vespidae (Hymenoptera)preys on the pupae of Anadiplosis and Platy-gasteridae sp. ' , A' , .These wasps cut through

the gall walls to reach the pupae of Anadiplosisand they make multiple cuts in order to obtainali the pupae of the Platygasteridae sp. "A".After opening the galls and locating the pupaeof Anadiplosis, these are held with the mandi-bules and masticated before being ingested. AI-ternatively, the wasp might withdraw the pupaeof the Platygasteridae sp. "A" from their pu-paria, before mashing and ingesting them.

Besides this predator, three species of Sal-ticidae (Aranae) (here named Salticidae speciesA, B, and C) have been observed which preyupon both the adults of Anadiplosis and of Ta-naostigmatidae. A food web based on Anadi-plosis galls is shown in figure 5.

The hairs of the galls display sticky secre-tions that trap insects of several orders, inclu-ding Diptera, Hymenoptera, Homoptera, Co-leoptera and Thysanoptera, that have an aver-age body size less than 2mm. The most frequent

insects trapped were dipterans, hymenopteransand homopterans. Platygasterid parasitoids ofthe gall forming larvae were frequently seentrapped and killed by the hair secretions.

The secretions present in the hairs of mostAnadiplosis galls may act in the deterrence ordestruction of parasitoids on Anadiplosis. Hairsecretions may, by trapping potentially phyto-phagous insects of the gall and/or plant, conferprotection to the gall former as well as to thehost plant. There are examples of galls secretingsubstances that can trap small insects (Bequaert1924, Cornell 1983, Darlington 1975, Mani1964). The ecological significance of these hairsecretions to plant and/ or insect protectionmay be important to studies on interactionsamong three trophic levels, as proposed by Priceet al. (1980), though little work has been do-

ne on this aspect to date (but see Washburn

1984).Galls are associated with a broad spectrum

of plant taxa, in. ali plant organs, and withmany different habitats. In addition, they pre-sent astonishing variations in morphology, ana-~

122 G. w .Fernandes. R.P. Martins & E. Tameirào Neto

SALTICIDAE SD. A

SAL TICIDAE sp. B SAL TICIDAE sp. C

""'I"-_/ANADIPLOSIS sp.

(adult)

MACHAERIUM ACULEA TUM

(host plant)

PARAJALYSUS PALLIDUS-

GEOMETRIDAE

VSUS SPINOSUSVESPIDAE

1:;1:.::;:;:;:;:.:;;;:;:

JALYSUS SOBRINUS~,

ANADIPLOSIS sp.:!i!!!i!iJii[

~

~ ? (larva) ::;;;:;;:'

':;:;:;:;;;;;;:/00

/~

EURYTOMIDAE sp. A

EURYTOMIDAE sp. B\\\PLATYGASTERIDAE sp. A PLA TYGASTERIDAE sp. B

T ANAOSTIGMA TIDAE

Fjgure 5. Food web based upon Anadiplosis sp. leaf galls on Machaerium aculeatum. Observe that the vespids. eurytomids

and platygasterjds are Anadiplosis predator and parasjtojds. respectively. while1he berjtids and geometrjd larvae are actually

feeding on gall tjssue. hence bejng here calledcompetitors of the gall forming cecidomyjjd. The tanaostigmatjd species feeds{)n thp a"ll ti"",IP 1."n1icl linp) "ncl n{)""ihlv c{)n",lmp" thp a,,1I f{)rmina in"pct I"rv" (rl""hpci linp)

123Food web relationships in galls

tomy, chemistry, seasonality and ecology,which makes the study of the gall maker/gallcomplex and their associated organisms ofgreat interest in the developmen.t of evolutio-nary and applied ecology.

Acknowledgements -The authors wish to thank J.M. Fer-rari (Botany Department -U.F.M.G.) and L. Kinoshita(Herbarium of the University of Campinas), for the deter-mination of M. aculeatum; R.J. Gagné(U.S.D.A.) for thedetermination of Anadiplosis sp.; A.A.P. Fidalgo (Fun-danción Miguel Lillo, Tucumán -Argentina) for the familydeterminations of the hymenopteran parasitoids; T. Henry(U.S.D.A.) for the determination of Berytidae; J. LaSalle(University of California -Riverside) for helpful informa-tion on the Tanaostigmatidae; J .H. Kirkbride and v. Ruddfor information on M. aculeatum; to draughtsmen J. Bit-tencourt Neto and T. Dougi, for the drawings. In addition,we wish to thank R. Bronner, T. Craig, P .W .Price. C. Sac-chi. R. Woodman. and to an annonymous reviewer for theircriticisms and assistance with the English language duringthe preparation of this manuscript. Also, we thank the De-partamento de Biologia Geral, ICB-U.F.M.G., for generalsupport, and the Ralph M. Bilby Research Center, N.A.U.for providing the facilities.

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