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Oryzopsis Hymenoides) Seeds in Mixed and Monospecific Seed Patches

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Oryzopsis Hymenoides) Seeds in Mixed and Monospecific Seed Patches Western North American Naturalist Volume 65 Number 3 Article 3 7-28-2005 Comparing the effects of granivorous rodents on persistence of Indian ricegrass (Oryzopsis hymenoides) seeds in mixed and monospecific seed patches Joseph A. Veech Colorado State University, Fort Collins and University of Northern Colorado, Greeley Stephen H. Jenkins University of Nevada, Reno and University of Northern Colorado, Greeley Follow this and additional works at: https://scholarsarchive.byu.edu/wnan Recommended Citation Veech, Joseph A. and Jenkins, Stephen H. (2005) "Comparing the effects of granivorous rodents on persistence of Indian ricegrass (Oryzopsis hymenoides) seeds in mixed and monospecific seed patches," Western North American Naturalist: Vol. 65 : No. 3 , Article 3. Available at: https://scholarsarchive.byu.edu/wnan/vol65/iss3/3 This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Western North American Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Western North American Naturalist 65(3), © 2005, pp. 321–328 COMPARING THE EFFECTS OF GRANIVOROUS RODENTS ON PERSISTENCE OF INDIAN RICEGRASS (ORYZOPSIS HYMENOIDES) SEEDS IN MIXED AND MONOSPECIFIC SEED PATCHES Joseph A. Veech1,3 and Stephen H. Jenkins2 ABSTRACT.—In desert environments seeds are often heterogeneously distributed in small patches that vary in number of seed species and in seed density. Because seed harvest by rodents is often density dependent (a larger proportion of seeds is removed from high-density seed patches than from low-density patches), the proportion of residual or post-har- vest seeds should be greater in low-density patches. In addition, seed preference can affect harvest. We tested whether the residual proportion of a highly preferred seed (Indian ricegrass, Oryzopsis hymenoides) was less when in a seed patch with a 2nd species (mixed-species patch) than when in a monospecific seed patch. We predicted that the increased overall seed density due to the presence of 2 species in a patch would result in a lower residual proportion of ricegrass seeds in the mixed-species seed patches than in the monospecific patches. As predicted, the residual proportions of Indian ricegrass seeds were less each time ricegrass was paired with one of 6 other species in mixed-species patches. Similarly, the residual proportion of each of those 6 species was less when paired with ricegrass than when in a mono- specific patch. We speculate on the potential implications of these results for the population dynamics of plant species and the physical structure of plant communities. Key words: heteromyid rodent, Indian ricegrass, Oryzopsis hymenoides, density-dependent foraging, seed-tray experiment. Indian ricegrass (Oryzopsis hymenoides) is component in sustaining populations of grani- a perennial bunchgrass found throughout west- vorous rodents and, in some years, the caching ern North America. It is often associated with of seeds by rodents may serve as a source of loose substrates such as the sand typical of sand recruitment for ricegrass populations (McAdoo dunes and other arid habitats (Jones 1990). It et al. 1983, Young et al. 1983, Longland et al. shares this habitat affinity with heteromyid 2001). rodents (family Heteromyidae), such as kanga- Regardless of whether rodents have an over- roo rats (Dipodomys) and pocket mice (Chae- all positive effect on Indian ricegrass popula- todipus and Perognathus), also widespread and tions by caching seeds that subsequently ger- locally abundant in arid parts of western North minate, or a negative effect due to seed con- America. Indian ricegrass usually produces sumption, the 1st step in this interaction is the seeds during the summer months. Heteromyid harvest of seeds from the immediate area around rodents are capable of harvesting a large portion adult plants. This area may be no larger than a of the seed crop of some plant species (Soholt dinner plate as the seeds are heavy and not 1973, Nelson and Chew 1977, McAdoo et al. likely to be dispersed by wind. Furthermore, 1983, Longland et al. 2001), which they either the spatial distribution of seeds in desert eco- immediately consume or cache for later use systems is often heterogeneous (Reichman 1976, (McAdoo et al. 1983, Longland et al. 2001, Veech Nelson and Chew 1977, Price and Reichman 2001). Many rodent species prefer seeds of 1987, Henderson et al. 1988, Price and Joyner Indian ricegrass over seeds of other plant 1997, Anderson and MacMahon 2001); relatively species (McAdoo et al. 1983, Kelrick et al. 1986, dense concentrations of seeds of different Henderson 1990, Jenkins and Ascanio 1993, species occur in depressions in the soil surface Longland and Bateman 1998, Veech 2001). or against objects (e.g., small rocks, shrubs) Thus, Indian ricegrass may be an important that block the wind. Thus, the seed resource 1Department of Fishery and Wildlife Biology, Colorado State University, Fort Collins, CO 80523. 2Ecology, Evolution, and Conservation Biology Program, Department of Biology, University of Nevada–Reno, Reno, NV 89557. 3Present address: Department of Biological Sciences, University of Northern Colorado, Greeley, CO 80639. 321 322 WESTERN NORTH AMERICAN NATURALIST [Volume 65 of granivorous rodents can be viewed as existing tion dynamics of plants producing seeds that within distinct small spatial patches. Rodents are foraged by rodents. forage for seeds in and among these patches. We predicted that the residual number of The harvesting of seeds by heteromyid Indian ricegrass seeds would be greater in rodents is sometimes density dependent (Price monospecific patches than in mixed-species and Heinz 1984, Brown 1988, Bowers 1990, seed patches. We also compared the residual Mitchell and Brown 1990, Veech 2000, 2001). seed numbers of the other 6 species when In the present study we define density-depen- paired with Indian ricegrass and when in a dent foraging as the harvest (i.e., removal) of a monospecific seed patch. Again, we predicted larger proportion of seeds (of a given species) a greater number of residual seeds in mono- from patches with high overall seed densities specific seed patches because their lower total than from patches with low densities. Overall seed density induces less foraging. seed density is defined as the combined den- sity of seeds of all species. If the proportion of METHODS seeds harvested is density dependent, then Description of the proportion of seeds that are not harvested Study Site is also density dependent. We refer to this lat- ter proportion as residual seeds. We tested for We measured rodent harvesting of seeds of differences in the residual number of Indian Oryzopsis and 6 other plant species at a study ricegrass seeds in patches consisting solely of site about 80 km northeast of Reno, Nevada, ricegrass (monospecific patches) and patches USA. These species were Astragalus cicer, consisting of ricegrass and seeds of 1 of 6 other Panicum miliaceum, Sphaeralcea coccinea, Stan- species (mixed-species patches). The initial leya pinnata, an unidentified species of Lupi- number of ricegrass seeds was the same in both nus, and an unidentified species of Penstemon. types of patches; thus, the mixed-species patches All of the species were found at or near the represented a higher initial seed density than study site except for Panicum and Astragalus, the monospecific patches. although a congener of A. cicer (A. lentigeno- We were primarily interested in testing sus) was found at the study site. We used Pan- whether the presence of a 2nd seed species icum (millet) as a proxy for a seed type that is could, by increasing overall seed density, affect highly preferred by rodents; Panicum is often the residual number of Indian ricegrass seeds. used to trap rodents and to study their forag- Previously, it was demonstrated that density- ing behavior. We collected seeds of Lupinus, dependent harvest occurs when rodents forage Penstemon, Sphaeralcea, and Stanleya at the in monospecific patches with substantially dif- study site while seeds of the other 3 species ferent initial seed densities (Brown 1988, were obtained from a commercial supplier. Mitchell and Brown 1990, Veech 2000). Indeed, The heteromyid rodent community at the this type of density-dependent foraging is some- study site was diverse. Extensive trapping by what easy to demonstrate. However, density- other researchers has revealed the existence of dependent harvest may also occur under a dif- 8 species: Chaetodipus formosus, Dipodomys ferent (and perhaps more realistic) scenario deserti, D. merriami, D. microps, D. ordii, Micro- where differences in seed densities among dipodops pallidus, Perognathus longimembris, patches are due to the presence of additional and P. parvus (Breck and Jenkins 1997, Jones seed species, not merely to differences in the and Longland 1999). We live-trapped rodents density of a single seed species. That is, the on 21 June 1998 to confirm the existence of actual type of patch (monospecific seed patches rodents on our study blocks (see next section). versus patches with seeds of multiple species) We caught 38 individuals representing 5 species might influence harvest rates and residual seed in a total of 200 traps. The relative abundances densities. The residual seeds not harvested of each species in terms of percentage of total from a patch may, in some cases, act as a chan- individuals captured were D. merriami (71.1), nel of recruitment for a plant population. Thus, D. microps (10.5), M. pallidus (2.6), P. longi- we also present our study as an example of the membris (7.9), and P. parvus (7.9). Although potential need to consider density-dependent there were also granivorous ants and birds at foraging in studies and models of the popula- the study site, most of the seed harvesting that 2005] RODENTS AND RICEGRASS 323 occurred during the seed-tray experiment (see higher overall seed density also always con- below) could be attributed to rodents because tained 2 seed species.
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    Panicoideae ① Micrairoideae Eriachne Eriachneae Coelachne A Isachne Isachneae Micraira Micraireae Arundo Monachather Arundinoideae Amphipogon Elytrophorus Eragrostis Phragmites Hakonechloa Molinia Austrodanthonia-1 Joycea Rytidosperma Austrodanthonia-2 Karroochloa Tribolium Danthonioideae Schismus Merxmuellera-1 Cortaderia-1 Cortaderia-2 Notochloe Cortaderia-3 Danthonia Chaetobromus Chionochloa Pentaschistis Cortaderia-4 Merxmuellera-2 Centropodia Centropodia clade Ellisochloa Neyraudia Triraphis Triraphidieae Eragrostis-1 Pogonarthria Eragrostis-2 Eragrostis-3 Eragrostis-4 Eragrostis-5 Eragrostis-6 Eragrostis-7 Ectrosia Eragrostis-8 Eragrostideae Uniola Entoplocamia Fingerhuthia Tetrachne Cottea Enneapogon Schmidtia Spartina Calamovilfa Sporobolus-1 Thellungia Crypsis Zoysieae Sporobolus-2 Sporobolus-3 Zoysia Pappophorum Tridens PACMAD Muhlenbergia Sohnsia Bouteloua Distichlis Eragrostis-9 Allolepis Hilaria Jouvea Scleropogon Swallenia Blepharidachne Erioneuron Dasyochloa Munroa Tragus Willkommia Chloridoideae Oropetium Tripogon Melanocenchris Dactyloctenium Brachychloa Neobouteloua Odyssea Monodia Plectrachne Triodia Kengia Perotis Ctenium Trichoneura Leptothrium Bewsia Cynodonteae Gymnopogon Triplasis Gouinia Vaseyochloa Tuctoria Neostapfia Orcuttia Aeluropus Leptochloa-1 Acrachne Leptochloa-2 Coelachyrum Dinebra Astrebla Eleusine Chloris-1 Enteropogon-1 Chloris-2 Lintonia Chloris-3 Enteropogon-2 Tetrapogon Trichloris Lepturus Oxychloris Rendlia Harpochloa Microchloa Eustachys Chloris-4 Brachyachne Cynodon Aristidoideae Stipagrostis
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