Flight Muscle Developtnent and Dispersal in the Life Cycle of Carabid Beetles: Patterns and Processes
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BULL ETIN DE L'INSTITUT ROYA L DES SCIE 1CES NATURELLES DE BE LGIQUE ENTOMOLOG IE. 70: 13-31 . 2000 BULL ETI N VA I-l ET KON INKLIJ K BELGISCH I STITUUT VOOR NATUU RWETENSCHAPPEN E TOMOLOGIE. 70:, 13-31,2000 Flight muscle developtnent and dispersal in the life cycle of carabid beetles: patterns and processes by Konjev DESENDER Abstract Les deviations significatives observees pour une proportion plus im portante de coleopteres avec des muscles de vols fonctionnels chez des femelles non matures en comparaison avec celles matures indiquent un Seasonal patterns of fli ght muscle development are documented fo r 27 tel syndrome chez Ia plupart des especes de Carabidae etudiees. Ceci carabid species, collected from di fferent habitats. The ph enology or s'applique surtout aux especes qui emergent aI a fin du printemps, juste timing of their life cycle is compared to t he p resence o f a functional avant leur periode de reproduction en ete-automne. Le pourcentage de fli ght apparatus during complete year cycles. The majo ri ty of the femelles matures avec des muscles alaires fonctionnels differe entre investigated ground beetles have never b een stud ied b efore in this espi:ces et peut etre utilise comme une mesure d' un "'oogenesis-fli ght respect. In most species there is a di stinct seasonal pattern of fli ght syndrome" ma ins deterministe. L'analyse globale de tous les resultats muscle functionality. Some species, with similar life cycle timing a nd/ suggere que les caracteristiques suivantes interviennent comme fac or similar habitat requirements, show the same pattern of fli ght muscle teurs expliquants les profil s o bserves au ni veau des muscles de vol: development. Inter- and intraspecific v ariability in maximal propor choix de !'habitat, evolution du cycle de vie, ainsi que d'autres eie" tions o f beetl es with functional flight muscles indicates fli ght muscle ments de Ia vie de l' insecte. Le developpement des muscles devol des dimorphism. The reproducti ve state of fe males (obtained through dis Carabidae semble di:s lors faire partie d' un ensemble de caracteres section of the ovaries) is used to test the generality of an "oogenesis coadaptes. fli ght syndrome" (trade-off between dispersal and reproduction), the null hypothesis being that functional fl ight m uscul ature and ripe ova Mots-clefs: Carabidae, pouvoir de dispersion, cycle de vie, developpe ri es occur independently. Observed signi ficant d eviations towards a ment des muscles alaires, oogenesis-flight syndrome, reproduction, hi gher proportion of beetles with fi.mctional fl ight muscles in unripe as dimorphisme alaire, adaptations, caracti:res coadaptes compared to ripe females indicate such a syndrome in most of the species studied. It is especially true for species emerging during late spring before their summer-autumn reproduction. The proportion of ripe females with functional fli ght muscles differs between species and can be used as a measure for a less deterministic version of an Introduction "oogenesis-flight" syndrome. A complex of factors must play a role in the expression of fli ght muscle development. Integrati on of all results s uggests habitat choice, evolu tion of life cycle timing and other [n many insect groups, species are known to show poly life hi story traits as ultimate factors responsible for the observed morphisms affecting their fl ight ability. The most obvious patterns of flight muscle fun ctionali ty. Flight muscle development in examples are variations in wing and fli ght muscle devel ground beetl es therefore seems to be part of a suite of coadapted traits. opment (HARR ISON, 1980). Wing polymorphisms are un Key wonls: Carabidae, dispersal p ower, life cycle, fli ght muscle der genetic and environmental control (HARRISON, 1980; development, oogenesis-flight syndrome, reproduction, wing dimor ROFF, 1986). phism, adaptations, sui te of coadapted t ra its Numerous observations indicate that in many insect species migration is limited t o t he post-teneral, pre-re productive period. These observations have led to define Resume the " oogenesis-flight syndrome" as a characteristic of insect migration (JOH NSON, 1969; for Coleoptera, see f. e. Les pro til s saisonniers du developpement des muscles alai res sont Ll NDERS et a\., 1995, MUDA et a\. , 198 1, RANKIN et a\. , reportes pour 27 especes de Carabidae, recoltees dans des habitats differents. Durant des cycles annuels complets, Ia phenologie ou Ia 1994, TADA et al. , 1991) . The inherent assumption is that periode du cycle v ital est comparee pour chaque espece i1 Ia presence mi gration (through fli ght d ispersal by means of fu nc d ' un appa reil devol fonctionnel. La majorite des especes de Carabidae tional fl ight muscles and developed hind wi ngs) and concernees n 'a jamais ete etudiee SOLI S eel angle avant ce travail. Chez Ia plupart des especes, il y a un cycle saisonnier de Ia foncti onnali te des reproduction are alternating physiological states. This muscles alaires. Certaines es peces qui ont un cycle de vie e t/ou des view was strengthened by the fact that migration in many exigences environnementales similaires demontrent le meme profil cases is associated with or induced by conditions promo saisonnier au ni veau des muscles devol. Si l'on considi:re Ia periode du cycle de vie otl le developpement des muscles de vol est le plus ting a dult diapause (i.o. w. delaying reproduction), for important, on constate une vari abilite inter- et in traspecifique, ce qui example a s horter photoperiod, lower t emperatures or suggere un dimorphisme de muscles alai res. L 'eta! reproducti f d es poor f ood conditions (RANKIN et a\. , 1986). Migrating fe melles (obtenu apres dissection des ovaires) est utilise pour t ester insects moreover often are physiologically comparabl e Ia presence d' un " oogenesis-fl ight syndrom.e" (balance entre d isper sion et reproduction), !' hypothese nulle etant qu· une muscul ature alaire to cli apausing insects in their possession of hypertrophic fonctionnell c et Ia maturi te des ovaires apparaissent independamment. fat bodi es and immature ovaries. Studies on the physio- 14 Konjev DESENDER '' logy and endocrinology of flight muscle degeneration and production) and is related to habitat change (TIETZE, regeneration, in a few insect species, have suggested a 1963 ; VAN Hui ZEN, 1977). Flying females were always control by juvenile hormone (FAIRBAIRN & Y ADLOWSKI, unripe. According to DEN BOER eta!. ( 1980) some species I 997; ZERA et al., 1997). A strong negative correlation (Amara.familiaris, Anisodactylus binotatus, Calathus ro has commonly been observed between flight muscle mass tundicollis and Nebria brevicollis) deviate more or less and ovarian mass of insects (ROFF, I 986), suggesting that from an "oogenesis-flight" syndrome. In Nebria brevi the construction and maintenance of the flight apparatus col/is developing flight muscles were found in immature competes with egg production for a limited internal nu beetles (i.e. in spring) more than in reproducing beetles trient pool (ZERA & DENNO, 1997). The metabolic rates of (in autumn), while at least some flight activity was ob flying insects can be 20-100 times that of resting animals served during autumn too (N ELEM ANS, 1987). MATALIN and are among the highest known (RANKIN & BURCHSTED, ( 1994, 1997) and ZHANG et a!. ( 1997) studied Harpalus 1991 ). Nevertheless, several authors ( cf. GATEHOUS E & rujipes and came to the conclusion that flight was more ZHANG, 1995) have chall enged the assumption that the prevalent in (but not limited to) young beetles before oogenesis-flight syndrome is a general phenomenon. reproduction. MEIJER ( 1974) suggested variation in flight Ground beetles are well-known for their varying de behaviour of the small Bembidion varium (always posses gree of hind wing development. Some species are con sing functional flight muscles): in spring relatively more stantly winged, others show a wing polymorphism or unripe females were recorded flying as compared to wing dimorphism and some species always possess re reproducing ones. duced wings. Carabid beetles consequently exhibit a large Clearly, there is a considerable lack of empirical data amount of variation in their respective dispersal power. on most carabid beetles in this respect. To this end, we Wing development is largely under direct genetic control have dissected large numbers of ground beetles collected in the few ground beetle species studied so far (cf. Au during complete year cycles in a multitude of habitats and KEMA, 1986; DESENDER, 1989a; LINDROTH, 1946). About belonging to 27 different species (20 constantly macro three-quarters or as much as 280 carabid species from pterous, seven wing dimorphic or polymorphic). These Belgium are known to be constantly macropterous or full species can be roughly divided into spring breeders (adult winged. Nevertheless wing development and, as a con hibernators) on the one hand and summer-autumn bree sequence, fli ght capacity are extremely variable between ders (larval or larval/adult hibernators) on the other hand. winged species too, as has been shown on the basis of a In this paper, we will first look at seasonal patterns of biometric approach (DEN BOER et al., 1980; DESENDE R et fli ght muscle development by comparing the phenology a!., 1986; DESENDER, 1989b ). Many winged ground beet or timing of the life cycle in each species to the relative les moreover not necessarily possess functional flight occurrence of a functional flight apparatus. Secondly, the muscles. Below a certain value of relative wing size, reproductive state of females (obtained through dissec functional flight muscles apparently are only rarely ob tion of the ovaries) will be used to test the generality of served (D EN BOER et a!., 1980; DESENDER, 1989b ).