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On the Fore Legs of Seed Bugs (Heteroptera: Lygaeidae): Aggression and Allometric Scaling in Scolopostethus Affinis Schilling

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21 December 2000 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY 73( I), 2000, pp. 6-10 On the Fore Legs of Seed Bugs (Heteroptera: Lygaeidae): Aggression and Allometric Scaling in Scolopostethus affinis Schilling RAFAELLUCAS RODRIGUEZ S. Department of Entomology, Haworth Hall, University of Kansas, Lawrence, Kansas 66045, USA ABSTRACT: Males and females of the seed bug Scolopostethus affinis used their fore legs with modified femora in intra-sexual aggression. Males were smaller than females, but had larger fore femora. Allometric scaling of the fore femora on body size was similar in both sexes. This argues against hypotheses that the often spiny and enlarged fore legs of seed bugs serve to move seeds or to aid in movement in confined spaces. Instead, they may function primarily in aggression. The fore legs of many seed bugs (Lygaeidae), especially among the Rhy- parochrominae, are strikingly modified with enlarged and spiny femora (Schuh and Slater, 1995) so that they seem raptorial (Borror et al., 1992). This is puzzling for bugs which are primarily herbivorous (Sweet, 1979; Schuh and Slater, 1995), and two main hypotheses have been proposed as explanations: that the fore legs are used to manipulate seeds along irregular substrates (Sweet, 1964a), or that they aid in movement within enclosed spaces (Schuh and Slater, 1995). However, another pos- sibility is that the modified legs of these bugs are used in aggressive contests, as for example in coreid bugs with their hind legs (Mitchell, 1980; Miyatake, 1993, 1997; Eberhard, 1998). There are some descriptions of fights involving the fore legs in seed bugs. Males of Neacoryphus bicrucis grapple to defend patches of their host plant (McLain, 1984). Sweet (1964a, b) reported fights in Neopamera bilobata, (see also Rodriguez S., 1998), Pseudopachybrachius basalis, Sisamnes contractus, Ligyrocoris diffusus, Scoloposthetus diffidens, S. atlanticus and S. thomsoni. Although Sweet (1964a) in- terpreted this behavior as seed defense against conspecifics, with occasional occur- rences in sexual contexts, he viewed the main function of the modified fore legs of seed bugs as moving seeds. Here I describe a case in which the fore legs of both sexes are used in aggressive interactions, in Scolopostethus affinis Schilling (Rhyparochrominae: Drymini), and I present data on the allometric scaling of the fore femora on body size. These data support the idea that the modified fore legs of seed bugs may function primarily in aggression. Materials and Methods I collected S. u.ffinis on a patch of strawberries during June and July 1998, on semi- rural Pfuhl, Neu-Ulm, Bayern, Germany. They were on the ground, on strawberries that had fallen to the ground, and on the plants themselves, especially the roots. I kept bugs in individual Petri dishes with a wet napkin and halved sunflower seeds. I observed interactions under a dissecting microscope. I placed bugs in Petri dishes containing dried bits of strawberries, in the following combinations: two males, two females, and two males with 1-2 females. I measured the width of the pronotum and the right fore femur of 50 males and 36 females with an optical grid attatched to a dissecting microscope. VOLUME 73. ISSUE I Fig I Sketch oi it male (lclt), female (nnddlet and last-in-ix nympli Ir~ght)of S ~ilfini'i,showin'; their ton, temora. Note also the mrved fore tihide which h~dbumps along the ventral side (not shown) Results BM~AVIOR:Interactions weie not very common in the Petn dishes. 1 observed one ~gressivesi~ince and one fight between ni,iiihs out 0'"I7 instaà 3 in whibii I put mates and tt'lnales ltyctliei. Theri were two iggressn7\ stances in he five ci.i c in which T put only males together. Most male encounters consisted only of brief an- tennation followed by separation. There were aggressive stances all three times that T put only females together. In aggressive intcra~[ions the hiigs assum 'i an ''act wive ~r,c" such de- scribed by Sweet (1964a. b), in which they stood on the mid 'ind hind legs, raised the head and tholax, extended the fore legs horizontally, and vibrated the antennae The fight between males consisted of thr<*ebrief en ounters ::*i 1 .iccurre4 in the piesence of dead fcrn.de. whii.h 'fray hale lh-en incidi- siial. On\ lii fie assiimr ' the aggressive stance and damped his tore legs over the other indie, which had all irp on the substrate and his side facing tile aggressor. The encounter lasted less than i U and ended in separation of the males. In fhc qecond cnm:xinterthe -ame agrt--wve male trapped the othei inale's ainrnria between his fore !% mur and tbbi.i, and ;!I > !her male walked backwaids and pulled free Finally, the aggressive male trapped the other male's thorax with both his fore legs, one over and the other under ! and flipped him over This has not been previously reported: fight- ~isuallyiwml the males siniply stiiking each other with hi-izontal ir~i~\crncnl~(Sweet, Rodriguez S., 1998). Ou subscqucnt encounicrs in me ~ollowi~ig30 min, tt~.as gressive male assumed the aggressive stance and the othe~male walked &)way. All three aggressive interactions between females involved on? single apressne lcinale and ~liree"sithn~~ssive' tcniales, iihi ived in soiiience. I'l-, aggrew* c fe- JOURNAI OF THE KANSAS ENTOMOLOGICAL SOC IETY proroturn width (mm) Fig. 2. Males had sigiticitiit!y wider fore feiniwa than females (i = 3.7, P = 0.0004) ailhougli their pr bnotum widili fi.i'. -.ii~iiificantlyliroi" :.i -- 6.2. P < O.OiW , Hm the log-log sir!'. of fore femur wiJili in pronoturn v>,.l'h w, I-. not signifi-'.iiiiiy ,h,i ;rent between ~ii;)lc", :nd females (iii.;ii;,inpc = 1.4, A'-- 50, :male slope -- 1. I \ -= 35; F, 8, = d ti!. 1' > 0.75). ~v,ileassumei.! [hi- iggressive i ¥-rapped th uhmissive fei~v'lt"with her antibt) sue and app t ).iiylit. ;, at whicl i.ioini subi-ni: i \ e it nale retrentvd i i one encounto the aggressivi I in:ile stood on 1 i mid and hisiil !>-5 .ind dropred {)it top of the otht r fermie Finally she approi.u:hed the 0th female again, briefly trapped one of her an- tennae between her foic femur and tibia, climbed on tup of her and there assumed the aggressive stance! I did not ubsi'w- S affini: ii~t-ivrseeds with ihei- tore legs. 1 ide.ut they caused them on tht. itsait urn, which ;,.is kneeled antetsorl~, about 45 with the horizontal L ,e even when i r 8 ul o mait \ valk on an img~iLusurface whil~ s, and then hghtly pro them with a forceps. The irregular surface consi.stcd of sand and small branches placed on the Petn dish, to imitate the obsta- les of Sweet (19643) that should favor the use of the tore lees in rruning seeds. iowever, 1 nl'sc \cd two rn,j,i'-flmt appearell u he trying to HKIW ~i dead feirilit: .uound by ,litnbni~ on her iwij. iiid in this c:isi. .ill the males' 1- 3 were broiigbi li-iio action. MORPHOLOGY: Adult males had a significantly wider fore femur than females, al- though they were significantly srnailei (Figs. l, 2). Howe~cr,the slopes of fore femora on hndy sixdid nut differ hetween males and females (Fiy The fore tibia curved (PH- 1) and had hm,.ili bumps al(ti):r I!\ central sidi k~i7~i.hseems well suited for ag~~th~:.i~~einteraciiiitss, especially trap! ii;r the body oi 'he opponent tx- tween fore lemui and tibia. i~ymplisof 5. uiJtrus did not have the enlarged hie femora ot the adults (Fig. 1 ). Discussion My observe~~~~issuggest U~illie fore legs r! > (~jfin/,vfun~~on in ~ggression If 1 hey served iv unh to move ,ccd + )r to aid in mt merit in cuiitiih~tl.spaces, one would expect nymphs to aho have enlarged fore femora. which was not the case in VOLUME 73. ISSUE 1 9 S. uffinis. However, other seed bugs do have nymphs with enlarged fore femora (J. A. Slater, pers. comm.), although this may sometimes be incidental to development for the adult stage. Behavioral observations to evaluate the prevalence of aggression are neccesary to determine whether it explains the modified fore legs of seed bugs or not. Estimates of sexual dimorphism alone may be misleading because, as in the present case, dif- ferent methods may yield different answers and both sexes may engage in aggres- sion. However, it may be common that the sexes engage in aggression to different degrees or with different structures. For example, males of Acanthocephala declivis and Leptog1o.s.~~~australis (Heteroptera: Coreidae) have modified hind femora that are larger and scale differently on body size than in females, and use them for de- livering squeezes in male-male contests (Miyatake, 1993, 1997; Eberhard, 1998), whereas female A. declivis have hind tibiae with a modified plate that is wider and scales differently on body size than in males, and use them in aggressive displays between females (Eberhard, 1998). Another prediction of the aggression hypothesis is that it should be more common in species of seed bug with modified fore legs. For example, the coreid bug Lep- toglossus phyllopus is monomorphic in hind leg structure, and the males did not en- gage in aggression (Mitchell, 1980). However, males of the seed bug Neacoryphus bicrzlcis do not have modified fore femora, but did engage in aggression with other males (McLain, 1984). Aggression may occur in several contexts, such as intra-specific competition for food, mates or other resources, or competition when several species co-occur on a temporarily plentiful resource (J.
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    NINE Phylum ARTHROPODA SUBPHYLUM HEXAPODA Protura, springtails, Diplura, and insects ROD P. MACFARLANE, PETER A. MADDISON, IAN G. ANDREW, JOCELYN A. BERRY, PETER M. JOHNS, ROBERT J. B. HOARE, MARIE-CLAUDE LARIVIÈRE, PENELOPE GREENSLADE, ROSA C. HENDERSON, COURTenaY N. SMITHERS, RicarDO L. PALMA, JOHN B. WARD, ROBERT L. C. PILGRIM, DaVID R. TOWNS, IAN McLELLAN, DAVID A. J. TEULON, TERRY R. HITCHINGS, VICTOR F. EASTOP, NICHOLAS A. MARTIN, MURRAY J. FLETCHER, MARLON A. W. STUFKENS, PAMELA J. DALE, Daniel BURCKHARDT, THOMAS R. BUCKLEY, STEVEN A. TREWICK defining feature of the Hexapoda, as the name suggests, is six legs. Also, the body comprises a head, thorax, and abdomen. The number A of abdominal segments varies, however; there are only six in the Collembola (springtails), 9–12 in the Protura, and 10 in the Diplura, whereas in all other hexapods there are strictly 11. Insects are now regarded as comprising only those hexapods with 11 abdominal segments. Whereas crustaceans are the dominant group of arthropods in the sea, hexapods prevail on land, in numbers and biomass. Altogether, the Hexapoda constitutes the most diverse group of animals – the estimated number of described species worldwide is just over 900,000, with the beetles (order Coleoptera) comprising more than a third of these. Today, the Hexapoda is considered to contain four classes – the Insecta, and the Protura, Collembola, and Diplura. The latter three classes were formerly allied with the insect orders Archaeognatha (jumping bristletails) and Thysanura (silverfish) as the insect subclass Apterygota (‘wingless’). The Apterygota is now regarded as an artificial assemblage (Bitsch & Bitsch 2000).
  • Pseudotsuga Menziesii

    Pseudotsuga Menziesii

    SPECIAL PUBLICATION 4 SEPTEMBER 1982 INVERTEBRATES OF THE H.J. ANDREWS EXPERIMENTAL FOREST, WESTERN CASCADE MOUNTAINS, OREGON: A SURVEY OF ARTHROPODS ASSOCIATED WITH THE CANOPY OF OLD-GROWTH Pseudotsuga Menziesii D.J. Voegtlin FORUT REJEARCH LABORATORY SCHOOL OF FORESTRY OREGON STATE UNIVERSITY Since 1941, the Forest Research Laboratory--part of the School of Forestry at Oregon State University in Corvallis-- has been studying forests and why they are like they are. A staff or more than 50 scientists conducts research to provide information for wise public and private decisions on managing and using Oregons forest resources and operating its wood-using industries. Because of this research, Oregons forests now yield more in the way of wood products, water, forage, wildlife, and recreation. Wood products are harvested, processed, and used more efficiently. Employment, productivity, and profitability in industries dependent on forests also have been strengthened. And this research has helped Oregon to maintain a quality environment for its people. Much research is done in the Laboratorys facilities on the campus. But field experiments in forest genetics, young- growth management, forest hydrology, harvesting methods, and reforestation are conducted on 12,000 acres of School forests adjacent to the campus and on lands of public and private cooperating agencies throughout the Pacific Northwest. With these publications, the Forest Research Laboratory supplies the results of its research to forest land owners and managers, to manufacturers and users of forest products, to leaders of government and industry, and to the general public. The Author David J. Voegtlin is Assistant Taxonomist at the Illinois Natural History Survey, Champaign, Illinois.