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  • Parmenter 1988

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    MATHIX
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    Field Sampling

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    Field Sampling

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    8 visualizações8 páginas

    Parmenter 1988

    Enviado por

    MATHIX
    Field Sampling

    Direitos autorais:

    © All Rights Reserved
    Baixe no formato PDF ou leia online no Scribd
    Sinalizar o conteúdo como inadequado
    de 8
    ODYSSEY ty of Idaho Library ILL Borrower: ORZ Lending St “NTD,CKM,IOH,MNB, VWM,.CUT,EYW,IALJNAV MC,VOD,CSA,CTB,DGW,FHM Patron: Journal Title: Environmental entomology. Volume: 17 Issue: 2 Month/Year: 1988Pages: 280-286 Article Author: PARMENTER, RR Article FACTORS LIMITING POPULATIONS OF ARID-LAND DARKLING. BEETLES (COLEOPTERA, TENEBRIONIDAE) - PREDATION BY RODENTS 7) g iversit 4396: Imprint: : [Oxford] : [Oxford University Press] Uni 196 UN ILL Number: (MOA ILLiad TN: Call #: Per SB599.E55 Location: ODYSSEY ENABLED hare Maxcos Shipping Address: Portiand State University Library, Interlibrary Loan 1876 SW Park Avenue Portland, OR 97201 Fax: 503-725-4524 Ariel: 131.252.180.131 Email: ILL@pdx.edu Se rg Factors Limiting Populations of Arid-Land Darkling Beetles (Coleoptera: Tenebrionidae): Predation by Rodents ROBERT R. PARMENTER np JAMES A. MacMAHON Department of Biology and Eeology Center, Utah State University, 'UMC.5305, Logan, Utah 84522 Environ. Entomol. 17(2): 280-286 (1988) ABSTRACT The role of rodent predation as a limiting factor in the abundance of six species of darkling beetles (Tenebrionidae) was examined in a sagebrush-steppe ecosystem (Kem- ‘merer, Wyo.), Rodent predators were identified in feeding experiments using eaptive rodents and darkling beetles. Grasshopper mice (Onychomye) and deer mice (Peromyscus) success: fully captured and consumed Eleodes extricatus Say. Ground squirrels (Spermophilus) and pocket mice (Perognatkus) initially fed upon B. extricatus but developed an aversion to them after several exposures. Subsequent experiments demonstrated that the chemical aerosol produced by E. extricatus as an antipredator defense was effective in providing escape time during deer mouse attacks but not during grasshopper mouse attacks. A replicated field experiment, employing 10 rodent exclosures (15 by 15 m) and 10 control plots, showed that the removal of rodent predators resulted in population increases in five ofthe slx species of darkling beetles (Coniontis ovalis, E. extricatus, B. pimelioides, E. rileyt, and Helops dif fells), The total population of beeties exhibited « 63% increase in abundance following rodent removal. Based on 1986 field density measurements of rodents (8/he) and beetles (4.951/na), an average daily consumption of fewer than two beetles per rodent was calculated to be sulicient to produce the observed differences in darkling beetle populations on the site, Therefore, it is concluded that rodent predation functioned a8 a significant limiting factor on darkling beetle populations in the sagebrush-steppe study area, KEY WORDS DARKLING BEETLES (Tenebrionidae) are among the more common and conspicuous macro-inverte- brates of arid and semiarid ecosystems throughout the world, and: they are particularly abundant in North American deserts. Field population studies on darkling beetles have yielded density estimates ranging from a few thousand (Thomas 1979, Wise 1981a) to tens of thousands of beetles per hectare (Rickard & Haverfield 1965, Hinds & Rickard 1978). Seasonal and annual variations in darkling beetle population sizes have been linked to several abiotic variables, most notably fluctuations in pre- cipitation and temperature (Thomas 1979, Allsopp 1980, Kenagy & Stevenson 1982, Slobodchikott 1983, Sheldon & Rogers 1984, Cooper 1985, Whicker & Tracy 1987) The limiting function of abiotic variables on bee~ tle populations and activity patterns is well docu: mented, but the role of biotic interactions (e.g, competition, parasitism, and predation) in limiting, population sizes is far less clear. Wise (1981b) con- cluded that, during the years of his study, inter- specific competition between adult Eleodes spp. in New Mexico was not a significant factor in regu- lating population sizes. Wakeland (1926), working in Idaho, believed that dipterous and hymenop- terous parasitoids were "probably one of the factors that make E, extricatus of minor importance” as aan agricultural pest. Insecta, predation, rodents, beetles Predator-prey interactions have not been di reetly addressed in population studies of free-roam- ing darkling beetles (but see Wise (1985), despite extensive research on their antipredator defense system (e.g, Bisner & Meinwald 1966, Tschinke! 1975a,b). Most tenebrionids possess anal scent glands capable of discharging a noxious aerosol when at- tacked by a potential predator (Tschinkel 1975a,b). Laboratory tests of predator success on tenebri nids, using a variety of predators, have indicated thatthe effectiveness of the defense is variable. Ants, praying mantids, carabid beetles, spiders, sol pugids, toads, lizards, jays, armadillos, and some mice can be repelled, at least temporarily (Eisner & Meinwald 1966, Conner et al. 1985), although grasshopper mice and skunks are not deterred (e.8 Egoscue 1960, Slobodchikoff 1978). In field sit tions, tenebrionids ean be found in the diets of hundreds of predators (e.g., more than 175 species of Nearctic birds [Mca tee 1932], and a wide variety of arthropods, amphibians, reptiles, and mammals {Allsopp 1980), demonstrating that predator pro- tection is not absolute. Consequently, one can ask, to what extent does predation limit the abundances ‘of darkling beetles in the field, and which predators might be responsible? In this study, we addressed the mull hypothesis that predation of free-roaming darkling beetles by rodents has no measurable effect on beetle abun- ‘0046-25/88/0280-0286802.00/0 © 1088 Entomological Society of America April 1988 PARMENT. dances. We believed th important predators bec tivorous habits, their ¢ overlap those of darklir dance on our study plot included a series of la identified species of ro sted the effectiveneso inst each rodent st potential predator~pre lations of six species 0 and seven species of 11 ecosystem of southwest: periment involved the 1 dation pressure by excl of plots for two years, ¢ of the resident darklin dent predation functic darkling beetle populat to observe increases i rodent predation was 1 Materials Study Site. The stué sagebrush-steppe rang ‘oming, 8 km SW of Ki 110°37'0"W; elevation erages 226 mm/yr (40 snow. and is highly va peratures range from July (see Parmenter & 1 The vegetation on thi 1983). The dominant (Artemisia tridentai (Chrysothamnus visct aceasional individuals shia tridentata (Pursh; lanata (Pursh) J. T. (Amelanchier utahens brush (Tetradymia + grasses on the site inclt ‘mus tectorum L.), nee ta Trin, & Rupr.), Ind menoides (R. & 8.) Ri bluegrass (Poa spp.) The six species of t inthe study were Gomi extricatus Say, E. nig pimelioides Mannerl Helops diffcilis Hort ‘ovalis, all of these spe ical sprays (W. R. Ts cation)! Rodent species on t orous northern grass. leucogaster arcticeps } mouse (Peromyscus Coues), Great Basin parvus clarus Gold (Spermophilusarmati ly herbivorous specie ling Beetles odents of six species sus) success- iphilus) and ‘aversion to alcal aerosol ‘escape time licated field showed that Ix species of Helops dif- ve following ‘and beetles ssealeulated tions on the ant Limiting ss have not been di- 1 studies of free-roam- Wise [1985), despite antipredator defense wald 1966, Tschinkel sess anal scent glands nus aerosol when at- + (Tschinkel 1975a,b). success on tenebrio- lators, have indicated 2 defense is variable id beetles, spiders, sol armadillos, and some 4 temporarily (Eisner et al. 1985), although are not deterred (e-¢, 1978), In field situa: ‘ound in the diets of more than 175 species 32), and a wide variety epliles, and mammals ng that predator pro- quently, one ean ask, rlimit the abundances {and which predators sd the mull hypothesis 1g darkling beetles by fect on beetle abu April 1988, de PARMENTER & MACMAHON: RODENT “dances, We believed that rodents were potentially {mportant predators because of their known insec- tivorous habits, their diel activity patterns (that Overlap those of darkling beetles), and their abun- ance on our study plots. Our experimental design {ncluded a series of laboratory experiments that identified species of rodent predators, and evalu- ated the effectiveness of the beetles’ defense tactics against cach rodent species. We then examined potential predator-prey interactions among, pop- Flations of six species of large-bodied tenebrionids find seven species of rodents in the shrub-steppe ecosystem of southwestern Wyoming. The field ex- periment involved the manipulation of rodent pr Ration pressure by excluding rodents from a series Of plots for two years, and measuring the response Gf the resident darkling beetle populations. If ro- dent predation functions as a limiting factor on darkling beetle populations, then we would expect to observe increases in beetle populations when rodent predation was removed. Materials and Methods Study Site. ‘The study plots were located in the sagebrush-steppe rangelands of southwestern Wy- ming, 8 km SW of Kemmerer, Wyo. (41°43'0°N, 110°37°0°W; elevation 2,100 m). Precipitation av- cerages 226 mm/yr (40-yr mean), arrives mostly as Snow, and is highly variable. Mean monthly tem- peratures range from ~8°C in January to 17°C in Jaly (ee Parmenter & MacMahon 1984 for details). “The vegetation on thessite was shrub-steppe (West 1983). ‘The dominant shrubs were big sagebrush (Artemisia tridentata Nutt.) and rabbitbrush (Chrysothamnus viscidiflorus (Fook.) Nutt), with ‘occasional individuals of antelope bitterbrush (Pur- Shia tridentata (Pursh) DC.), winterfat (Ceratoides anata (Pursh) J. T. Howell), Utah serviceberry (Amelanchier utahensis Koehne), and gray horse- brush (Tetradymia canescens DC). Common grasses on the site included cheatgrass brome (Bro- ‘mus tectorum L.), needle-and-thread (Stfpa coma- ta‘Trin. & Rupr.) Indian rice-grass (Oryzopsis hy. menoides (R. & S.) Ricker), and several species of bluegrass (Poa spp.) "The six species of tenebrionid beetles included inthe study were Coniontis ovals LeConte, Eleodes extricatus Say, E. nigrinus perlongus Blaisdell, E. pimeltoides Mannetheim, E. riley! Casey, and elope dificilis Horn, With the exception of C. ‘vals, all of these species possess defensive chem- ieal sprays (W. R. ‘Tschinkel, personal communt- cation) Rodent species on the site included the insectiv- ‘orous northern grasshopper mouse (Onychomys leucogaster arcticeps Rhoads), the omnivorous deer mouse (Peromyscus maniculatus nebrascensis Coues), Great Basin pocket mouse (Perognathus parous’clarus Goldman), Uinta ground squirrel (Spermophilus armatus Kennicott),and threechief- ly herbivorous species, the least chipmunk (Eu- PREDATION ON DARKLING BEETLES 281 tamias minimus consobrinus J. A. Allen), the long- tailed vole (Microtus longicaudus longicaudus (Merriam)), and the sage vole (Lagurus curtatus levidensis (Goldman)). Predation Tests with Captive Rodents. Feeding trial experiments were performed to address three specific questions concerning rodent predation on darkling beetles: Which rodents eat darkling bee- tles? Can darkling beetles escape predation if re- fugia are nearby? How effective is the beetle’s chemical defense against rodents? Rodents and E. extricatus were live-trapped from ‘areas near the field study and maintained outdoors in shaded cages under natural light cycles and tem- peratures. Captive rodents were given water, bird- Seed mix, and fresh lettuce ad libitum. Beetles re- ceived fresh lettuce and rolled oats. ‘OF the six species of tenebrionids examined in this study, only E. extricatus was collected in suf- ficient numbers to run all the feeding trials. How- ever, we believe that this species isa suitable rep- resentative of the resident tenebrionid assemblage, ‘because like almost al of the other darkling beetles on the study site, E. extricatus possesses defensive secretions E. extricatus is also approximately the Same size as most of the other tenebrionids in the study area (only E. nigrinus is substantially larger). Finally, E. extricatus displays a predator-

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