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Oecologia © Springer-Ver!Ag 199Q Oecologiu (\990) 82:408-413 Oecologia © Springer-Ver!ag 199Q Host-associated differences in fitness \vithin and between populations of а seed beetle (Bruchidae): effects of plant variabllity David Н. Siemens and Clarence Dan Johnson Department of Biological Sciences, Northern Arizona University, Flagstaff, AZ 86011-5640, USA Received December 19, 1988/ Received in revised form September 25,1989/ Accepted November J, 1989 Summary. Experiments were co11ducted with the sexually Кеу words: Fitness in bruchid beetles - Specialization reproduci11g seed beetle Stator limbatus a11d its lюsts i11 to specific hosts - Seed beetles - Bruchidae - Plant vari­ 11orth-ce11tral Arizona to determine if it was sнbstrнc­ aЬility tured i11to нnits, each specialized for higher fitness 011 а specific host species. Unlike many studies, we incorpo­ rated scale, I.e., conducting experiments betVv·een and within beetle popнlations 011 seeds from withiп апd be­ Several authors have focused on the genetic substructur­ tween plant species. Of particular interest ;vas whether iпg of species that feed оп mot·e thaп one species of intraspecific plant variability preYented beetle specializa­ host plant (e.g., Futuyma апd Peterson 1985). For infer­ tion vvithin beetle populatioпs. Resнlts sнggest that S. ences about the evolution of specialization to specitic limbatus is specialized to certaiп hosts. 011 the palo verde hosts it is important to bave corresponding measures Cercidium f/oridum, beetles origiпally reared from this of fitпess for these species. lп this study we tested for host had significantly higher emergence compared to host-associated differences in fitness within and between beetles transferred from other hosts. We did not test populations of the sexually-reproducing seed beetle Sta­ directly for а genetic basis for this. Alternative hypothe­ tor limbatus (Horn). For sexually reproduciпg insects, ses of variation in symblotic microorganisms in larval most of the evidence fог host-associated geпetic differ­ gнts and maternal effects were assessed. Essentially по ences in fitness сошеs fгom populations s~parated geo­ bacteria, yeast or protozoa were found, and maternal graphically оп locally abundant hosts (Hsiao 1978; effects as expressed Ьу varyiпg egg weights were not Rausher 1982; Scriber 1983; Т:J.bashnik 1983). There are detected; however, other шicroorgaпisшs might have по coпviпciпg examples of geпetic host-associated differ­ ences in fitпess for sexually reproducing iпsects within Ьееп present and maternal effects through iпdнciЬle eп­ zymes was possiЬle. Caution, theп, is needed in any ge­ local populatioпs, even though theorists сопсеrпеd with speciatioп nettc шterpretatioпs of our resнlts. The differences on sympatric have predicted it (Maynard Smith С. floridum were detected from tests between апd withiп 1966; Bush 1975; Endler 1977; White 1978; Udovic beetle populations. Evidence for specializatioп was поt 1980; Felsenstein 1981; Lande 1982; Kondrashov 1983; detected on the other hosts, Ce,·ciclium microphyilum and Rice 1984). Acacia greggii. On the other hosts. beetles performed Evolution iп local host utilization might Ье better well regardless of their source. Significant differences understood Ьу iпcorporati11g fiпe scale eпvironmental were detected amoпg individual plants of С. f/OJ'idшn factors into empirical studies. Coпspecific plaпt variaЬii} 1ty IS such а factor and has been shown to Ье significant as to the suitaЬility of their seeds Гоr developmeпt оГ S. limbatus. No such differences were detected among in preveпting insect species from adapting to а plaпtJ species (Pimeпtel апd Bellotti 1976; Vап der Plank 1984; the other host plaпts. These patterns of conspecific plant Whitham апd Slobodchikoff 1981 ), but which has not variaЬility are opposite of what is expected if plant vari­ been iпcorporated into studies of sympatric speciatioп. aЬility preveпts specialization of beetles to particular Неге we were concerned with conspecific variaЬility of species of hosts. Thus, the data suggest seed variaЬility plaпts in relation to the aЬility of S. limbaшs to become amoпg plants does поt preveпt specialization to host specialized with host species. We also co11sidered the pos­ species in this systeш. We discuss how the patterпs of host use in this study relate to the hypothesis of sympat­ siЬility that S. /imbatus could Ье specialized on certain ric host race formation. conspecific plants varying in quality. This гesearch was stimulated Ьу the above and specif­ ically Ьу the knowledge that Stator limbatus, which has at least 40 host plaпts (Johпsoп апd Kingsolver 1976), Offprint requests to: D. Siemens has а greateг mortality rate iп seeds of the palo verde. ~~~~r· 409 ТаЬiе 1. Expcrimcnts conducted to detcrminc if diffcrcnccs in litncss cxist bctwccn populations оГ Stator !imhatus. Тlн~sс cxperimcnts "crc also desigпcd to detect variation among conspccific plaпts, а t~н:tor \Vhich might prcvent spccialization оГ bectles. Bcetles were. tг.1nsfcrred onto sceds of individual plants thcy werc not originally rcarcd from. Thcse transfers wcrc comparcd to bcctlcs that werc oriJ;Iin;.11ly rearcd from those plants (controls). Ifthe controls outperformcd (see tcxt) thc tr;:шsfcrs. wc took this as cvidcncc tOr spccialization Expt:riment Source of beet\es Seeds uscd Location Host location Host Verde А. gгeggii •.'"' P<~.radise C.jlщ-idum .11 tr;шst'crs .4.~. ~С'.+, <j)( .:ontro1s С.·",.......,<;,.(:'~ Paradisc C..floridum " i\ Paradisc C.jloridum Ь) tr:ш~Ccrs C..f.-:;> Д c.t _ Paradise С. jloridtm1 \ \-'erde А. greggii conrrols А ·5. "> 1\~ . Verde А. greggii ) "'}-'~ Yerdc А. greggii J с) tr;.шs!Crs А ·5- -'9 с. N\ Verde А. greggii j Paradise С. micropltyllum controls с,......, ~с_>'>). Paradise С. microphy!lum Paradisc С. micrupllyf!um Cercitlium jloгidum, than it does in many of its other ТаЬiе 2. Experimcnts conducted to determine if differences in fit­ h'"" (Johnson 1981). Janzen (1980) hypothesized that пess exist within populations of S. limbatus. The experiments are thoro "·ere sympatric host strains of S. limbatus. If there similar to those bet\\·een populations (ТаЬ\е 1) except that seeds arc indeed specialized strains of S. limbatus, the beetles апd beetles \vere from one site reared from their original hosts should perform better Experiment Source Seeds used on these hosts. In particular, we hypothesized this for of beetles Sraror limbatus on Cercidiumjlд1·iclum. Thus our primary questions were: (1) Do some а) Roosevelt Lake groups of S. limbatus have higher fitness in the seeds transt'ers А. greggii C.jloгitlum ot' some of their hosts in an experimental en\·ironment? controls С. jloridum С. flмidum 12) variaЬility and !s there intraspecific in the protective Ь) Paradise Valley mochanisms of the seeds of the hosts to protect them transfers С. mici'Opiiyllum С. jlщ·idum !'r(·!:. the depredations of S. !imbaшs? controls C.jloridum C.jloridum с) Paradise Yalley transfers C.jloridum .\\а terials, methods, and study sites с microplty!lum controls С. miaop/Iyllum с microp/Iyllum ln gcncral, our experiments involved rearing Sraror !imbatus from sceds collected in the wild and then ц11owing them to oviposit ТаЬiе 3. Experiments conducted to determine if fitncss diПerences and develop in seeds from which they \Vere reared and in seeds exist between and within populations of Stшor limbatlls occurring from \""hich they wcrc not reared. Each experiment \\·as two-factori­ on Cercidium jloridum. Experiment а tests differences bet\•;een a! in design and was analyzed Ьу t\VO-\.,.·ay ANOV А. The factors groups of S. limbatus from geographically separate stands оГ С "'~rc: 1) Groups of S. limbatus designared Ьу seed source and jloridum (Paradise Valley and Roosevelt Lake). This experiment ~~ ~..:eds of conspecific plants. Thus \\ie were <~.Ые to determine \Vas aJso designed to test differences between the seeds t'rom the di:":~~:::nccs between beetlc groups, which invo\Yed symp<ttric and t\VO stands. In experiment Ь beetles were transferred among indiYid­ a!k·poltric components, and diП'erences in seeds о[ conspecific host ual p!ants at one location (plants no more than 400 m apart). From plants. Various experiments were conducted iп \Vhich beetles were each plant (А through Н) beetles were reared and put back on trans(erred from host to host (ТаЫеs 1-3). seeds from that plant (controls). Beetles from every plant \\-·ere Cercidiumjloridum (Ыuе palo-verde) and С. microphyl!um (\it­ also transferred onto all the other plants (transfcrs). Capitalletters t!c-Jeaf palo verde) are small trees and Acacia grcggii (catclaw aca­ symbolizc individual plants ciaJ is а large shrub whose seeds are fed uроп Ьу 5. limhatus in north-central Arizoпa. Their distributions arc mostly overlapping Experiment Source of beetles Sceds used and they inhabit washes in desert plain~ and mountains (Carter 1974: Benson and Darrow 1944). They fruit at about the samc а) Between populations tJmc so their seeds arc availaЬ\e to prcdators at about the samc ttme. transfers Paradise RooseYelt Lake ':" ~eds of Cercidium jloridum, С. micropil\-!lum und А. greggii Rooseve\t Paradise ··"с:~: -.:oi!t:ctcd from Paradi~e Va1lcy, MaricoPa County, Arizona, contro1s Paradise Paradisc .н:..З Roosevelt Lakc, Gila County, Arizona. Thcre arc no species Rooseve\t RooseYelt ui Cercidium there, but Acacia greggii \vas a.\so collected in the \"erde Valley, Yavapai County, of north-centra! Arizona. Each site Ь) Within population v.as approximately 500m2. The relative Гrequencics of С. jloridum, Plants А-Н Plants А-Н С. micropltyllum, and А. greggii at Paradisc Va!ky were 279, 94, <1nd 1 respectively; and at Roosevc!t Lake 1. 2. and 5 respectively. То obtain specimens for experiments, seeds wcre put into papcr at а temperature of 27.7 degrces (±0.57; mcan±standard error) "<lcks in the field then transferred into jars in the Jaboratory (John­ and tl re\ative humidity о!' 69.5 (± 1.7%). Experiments \.,.·erc con~ ~on 19S1). Thus, all experiments in the labor<ttory \\·ere conducted ducted in petri dishes. cach with about 50 sccds and about 15 1 ':ng ~eeds Гrom nature.
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