Pollination experiments in the Mimulus cardinalis--M. lewisii complex

Experimental sets of Mimulus cardinalis and M. lewisii plants were (1) exposed to pollinators and (2) shielded from pollinators at study sties in Red Butte Canyon and Big Cottonwood Canyon, Wasatch Mountains, Utah. The exposed plants produced 1,535 seedlings and the shielded plants only 1. Clearly, seed production is dependent upon cross-pollination. A few syrphid flies were observed visiting the flowers but no hummingbirds or bumble bees, although the latter two have been reported as the main pollinators of M. cardinalis and M. lewisii , respectively. No interspecific hybrids were produced even though the species are fully interfertile, indicating that pollinators are faithful to their species or that different parts of their bodies pick up and carry pollen to the two different species.     

On the relative importance of floral color, shape, and nectar rewards in attracting pollinators to Mimulus

Pollinator preferences were observed for the six species of section Erythranthe of the genus Mimulus using greenhouse-grown plants placed in a meadow in the Red Butte Canyon Natural Area, Salt Lake County, Utah. The principal pollinators were hummingbirds and bumble bees. Hummingbirds preferred the species with the most reflexed tubular flowers regardless of color, whereas bumblebees preferred pink. lavender, or yellow flowers to red flowers regardless of shape. Results for the six species were confirmed by observations of F 2 hybrid recombinant plants selected such that flower color could be held constant and flower shape varied and vice versa.     

Pollinator preferences for yellow, orange, and red flowers of Mimulus verbenaceus and M. Cardinalis

Red. orange, and yellow morphs of Mimulus verbenaceus and M. cardinalis were field tested for pollinator preferences. The species are closely similar except that M. verbenaceus flowers have partially reflexed corolla lobes, whereas M. cardinalis flowers have fully reflexed corolla lobes. On the basis of over 6000 bumblebee and hummingbird visits, highly significant ( p Mimulus verbenaceus and M. cardinalis .     

Feeding behavior and performance of a rabbitbrush leaf-beetle ( Trirhabda lewisii ) feeding on Chrysothamnus nauseosus regrwoth after fire

Fire often positively affects the growth and nutrient content of plants regrowing after a burn. These changes have been associated with preferential feeding by herbivores in burned areas. In this study in southeastern Wyoming, Chrysothamnus nauseosus Pursh (rubber rabbitbrush) regrowing after a fire produced new shoots with a distinct growth form. Shoots were longer than those on unburned control sites and had longer leaves with longer internodes between leaves. We conducted feeding trials to detect whether C. nauseosus shoots regrowing after fire were nutritionally superior to shoots from unburned plants for the specialist leaf beetle, Trirhabda lewisii Crotch (Coleoptera: Chrysomelidae). We also measured C:N ratios and nitrogen and water contents of leaves from burned and unburned plants. Trirhabda lewisii adults preferred shoots from burned plants when given a choice. The beetles ate similar amounts of burned and unburned plants when fed only a single type. Females that were fed either burned or unburned plants did not differ in number of eggs laid. Chemical analyses revealed no significant differences in nitrogen or water content of leaves from burned versus unburned plants. Carbon-to-nitrogen ratio of burned plants was marginally lower compared with unburned plants. In contrast to previous studies, which suggest that herbivore attraction to burned areas leads to enhanced performance, our study shows that performance is not necessarily enhanced after fire.     

Revised checklist of the vascular plants of Bryce Canyon National Park, Utah

Prior to 1960, when the senior author wrote a dissertation on the plant ecology of the Paunsaugunt Plateau forests of Bryce Canyon National Park, relatively little plant taxonomic work had been done in the area. A checklist was prepared in 1971 that included 218 species of higher plants. During the field seasons 1978, 1979, and 1980, additional plants were collected during a second plant community study of the forests. The junior author spent the summer of 1980 at the park collecting plants in additional plant communities and organizing the herbarium collection. This checklist includes the additional species collected and updates the nomenclature of the vascular plants presently known to occur within the boundaries of Bryce Canyon National Park.     

Breeding biology and flower visitors of the rare White River penstemon, Penstemon scariosus var. albifluvis (Scrophulariaceae)

We studied the breeding system and flower visitors of White River penstemon, a rare endemic from the Uintah Basin of eastern Utah and western Colorado. Bagging treatments and hand-pollination treatments showed that Penstemon scariosus var. albifluvis has a mixed mating system: while some seeds and fruits are produced through selfpollination (both autogamy and geitonogamy), significantly more are produced when flowers are cross-pollinated. The primary flower visitors, and likely pollinators, were several species of native twig- and ground-nesting bees in the families Apidae, Halictidae, and Megachilidae. We found no differences in fruit or seed production between open-pollinated controls and hand-outcrossed flowers, suggesting that pollinator visits were sufficient to maximize female reproductive success. Management plans to conserve White River penstemon must recognize that full reproductive success of this rare plant taxon relies on a suite of pollinating bees, and that the species richness and abundances of bee visitors should be maintained.     

Reproduction and demography of Townsendia aprica (Asteraceae), a rare endemic of the Southern Utah Plateau

Townsendia aprica (Asteraceae: Astereae), a rare pulvinate perennial of the Southern Utah Plateau, was listed as threatened under the U.S. Endangered Species Act in 1985. Here we report on the reproductive biology and pollination of this little-known species and provide an estimate, for a single site-year, of size-specific reproductive effort. Last Chance townsendia appears a short-lived perennial that begins reproducing in its 2nd year (1.5-2.0 cm diameter). Maximum reproductive effort is attained with the 2.5-4.0 cm diameter size class: 38% of these plants produced 84% of the flower heads. Few plants survived past the 4-cm size class. The species is primarily self-incompatible: neither autogamous nor geitonogamous breeding system treatments produced a significant number of achenes. Unlike some populations of some congeners, the Ivie Creek population was not apomictic. Outcrossing is the primary means of reproduction and native solitary bees are the most important pollinators. Paramount are several species in the genus Osmia , and the ground-nesting species Synhalonia fulvitarsis , which nests among the T. aprica plants. Synhalonia fulvitarsis also visits a contemporaneous blooming phlox ( P. austromontana ), which may facilitate pollination of the rare townsendia. The Townsendia-Phlox-Synhalonia interaction may represent another example of why we must consider communities rather than individual species in our conservation efforts.     

Aquatic and semiaquatic Heteroptera of the Grand Canyon (Insecta: Hemiptera)

The aquatic and semiaquatic Hemiptera from 12 localities in Marble Canyon and Grand Canyon are reported, along with those from 1 locality in the Escalante Canyon. Fourteen species are recorded and compared to the water-bug fauna of the southwestern United States and western Mexico. Ochterus rotundus n. sp. is described from the Grand Canyon and the mountains of western Mexico.     

Geobotany of the Niobrara chalk barrens in Colorado: a study of edaphic endemism

The chalk barrens of the Niobrara Formation in the Arkansas River valley of Colorado exhibit a high degree of plant endemism and rarity. We examined their geochemistry and structure as well as the reproductive ecology of their signature species Mirabilis rotundifolius to assess factors influencing endemism. While no single component consistently identified the barren soils, the natural locations that support Mirablilis can be identified using elemental profiles from a cluster analysis of soil extracts. Because seeds germinate and plants proliferate in transplant gardens with non-calcic soils, bedrock and soil chemistry do not appear to be determinant components of endemism. Mirabilis rotundifoilus utilizes diverse pollinators but is capable of autogamy when insects are excluded; seed predation by harvester ants may be a limiting factor for population spread outside the barrens habitat. Mirabilis rotundifolius is a disturbance-tolerant species, part of a barrens cohort that shares traits of suffrutescence, woody caudices, or multibranched rhizomes penetrating the chalk strata. Its presence may be enhanced when disturbance inhibits the presence of other species that compete for limited water resources. We conclude that endemism is dictated not by unusual soil chemistry requirements but rather by structural components of the barrens. In a water-limited environment, shallow soils and thin chalk strata coupled with a functionally adaptive growth habit allow M. rotundifolius to survive when other species are excluded by disturbance or the physiochemical aspects of the barrens.     

Extrafloral nectaries mediate the arboreal beetle community (Coleoptera) in a Neotropical rainforest

ABSTRACT

Nectaries are structures that secrete a sugary solution and can occur on vegetative and/or reproductive parts of plants. The significance of floral nectaries to reward vertebrate and arthropod pollinators is well supported. The role of extrafloral nectaries (EFNs) is more ambiguous, though research has been skewed to the ant-plant mutualism. Many other insects feed at EFNs, but these interactions are vastly understudied. This study addresses the hypothesis that EFNs may influence the occurrence and structure of Neotropical canopy beetle communities. Seven canopy trees (four families) exhibiting EFNs and their associated beetles were studied over a one-year period in southern Venezuela. In total, 6818 adult beetles identified to 868 species were recorded on 25 investigated canopy tree species (#47 individuals). Of the 868 beetle species, 150 species (517 individuals; 17.3% species) from 20 families were observed drinking from foliar EFNs on seven EFN-bearing tree species. Dietary dependence on EF nectar varied, with 95 beetle species utilising this nectar within a broader diet and 55 species found feeding exclusively on EFNs. This study demonstrates unequivocally that EF nectar is a frequently utilised food resource of many beetle adults and beetles have been a significantly underestimated visitor group. A more detailed study was conducted on six individual canopy trees of two species of Chrysobalanaceae, Licania hebantha Mart. ex Hook. f. and Moquilea subarachnophylla (Cuatrec.) Sothers and Prance. In total, 115 individual adults of 64 beetle species were collected on nectar secreted on newly-sprouted leaves. These beetle assemblages were dominated by species utilising EF nectar and were associated with distinct phenological phases of the host trees. Altogether, the beetle survey found support for the hypothesis that EFNs influence the occurrence and structure of beetle communities. These beetle-EFN relationships have implications for spatial arrangement, community assembly and evolution of both host plants and beetles. Like ant-EFN mutualism, EFN-bearing trees and beetles may also form mutualism. It is possible that the plants offer easier access to a nutritious resource that may deflect herbivory of vegetative parts.     

Food niche of Exomalopsis (Exomalopsis) fulvofasciata Smith (Hymenoptera: Apidae) in Brazilian savannah: the importance of oil-producing plant species as pollen sources

Exomalopsis are ground-nesting species, and their food-niche breadth is little known due the difficulty in locating the nests and finding efficient bait plants to attract these bees. Some species of Exomalopsis were recorded as tomato, hot pepper and eggplant pollinators. Information about the food niche could be useful to increase Exomalopsis populations, providing consistent and comparable data for the enrichment of natural and crop areas with adequate plant sources. This study aimed to determine the food niche and the role of pollen size in the diet of E. fulvofasciata. We analysed pollen loads of 28 individuals of E. fulvofasciata collected from bait plants, in two natural areas of the Brazilian savannah. Only five pollen types belonging to the families Malpighiaceae, Solanaceae, Leguminosae, Myrtaceae and Lythraceae were important for this species. This result indicates that E. fulvofasciata is probably a polylectic species. However, we noticed that the Byrsonima used as bait plants contribute significantly for its larval provision, indicating that small pollen grains were more frequently collected.     

Pollinator sharing by three sympatric milkvetches, including the endangered species Astragalus montii

Insects visiting flowers of the endangered Heliotrope milkvetch, Astragalus montii , were compared with those visiting two common sympatric congeners, A. kentrophyta and A. miser , on three sites on the Wasatch Plateau of central Utah for 2 yr. We recorded 27+ species of bees, most of which were uncommon, visiting the three species. All three species were primarily visited by native bees of the genera Osmia (15 species) and/or Bombus (4 species). Most Osmia species visited the three species of Astragalus indiscriminantly; bumblebees preferred A. miser and avoided A. montii . Our hypothesis that A. montii flowers would receive fewer total bee visits and be visited by fewer bee species than their common congeners was rejected: A. montii was intermediate to the two common species in its attractiveness to bees. Also rejected was our hypothesis that the greater similarity between A. montii and A. kentrophyta in flower size, flower morphology, and microhabitat would be associated with greater similarity of flower visitors than either had with A. miser . The data suggest that, rather than competing with each other for pollinators, the three species of Astragalus facilitate each other's visitation rates.     

Notes on significant collections and additions to the flora of Glen Canyon National Recreation Area, Utah and Arizona, between 1992 and 2004

Symphyotrichum expansum (Puepp ex Spreng.) Nesom is reported new to Utah from the Escalante River drainage. A major range extension is reported for Aralia racemosa L. in the Escalante drainage, and additional populations are reported of the rare species Imperata brevifolia Vasey in Utah, including the 1st record for the Grand Staircase--Escalante National Monument. Heterotheca grandiflora Nutt. is reported new to north central Arizona. New locations and notes on an additional 22 rare species in Glen Canyon National Recreation Area are listed.     

Checklist of the mosses of Grand Teton National Park and Teton County, Wyoming

Symphyotrichum expansum (Puepp ex Spreng.) Nesom is reported new to Utah from the Escalante River drainage. A major range extension is reported for Aralia racemosa L. in the Escalante drainage, and additional populations are reported of the rare species Imperata brevifolia Vasey in Utah, including the 1st record for the Grand Staircase--Escalante National Monument. Heterotheca grandiflora Nutt. is reported new to north central Arizona. New locations and notes on an additional 22 rare species in Glen Canyon National Recreation Area are listed.     

Afrotropical mistletoe butterflies: Larval food plant relationships of Mylothris Hübner (Lepidoptera: Pieridae)

A detailed list of larval food plants of Mylothris based on review of published records in the literature is presented. Food plants have been reported for some 23 species (40%), although in only eight species (14%) of butterflies have plants been identified to species level. The food plants represent two unrelated orders, Santalales and Polygonales, although nearly all records (93%) are from four closely related families in the former group. Within the Santalales, the family Loranthaceae, a diverse group of aerial stem‐parasitic shrubs (mistletoes), is most frequently exploited. Only two species of Mylothris, M. bernice (Hewitson) and M. rubricosta (Mabille), which are closely related and possibly comprise a single species, are associated with Polygonaceae (Polygonales). Comparison of the morphology of the early stages of M. bernice, and other evidence, clearly indicates that M. bernice/rubricosta belong to Mylothris and do not represent a separate genus or sister group. Comments are made on these diverse food plant relationships within the context of a modern phylogeny of the Santalales and the evolution of parasitic plant feeding in butterflies in general. It is suggested that Mylothris had its major period of evolution and adaptive radiation on the Loranthaceae, probably sometime after the differentiation of most of the families or lineages in the Santalales (rather than on an earlier ancestral branch at or near the root of the Santalales evolutionary tree), and that this association has facilitated host switching multiple times. While more field data are clearly needed to ascertain the full extent of host specificity and range of mistletoe species exploited by Mylothris, this is unlikely to significantly change the patterns of food plant utilization seen at the higher taxonomic (ordinal and familial) levels. Mylothris represents one of only a few diverse groups of Lepidoptera, and insects in general, known to feed predominantly on African mistletoes.     

Chironomidae (Diptera) of the Colorado River, Grand Canyon, Arizona, USA, I: systematics and ecology

We describe the chironomid midge fauna of the Colorado River between Glen Canyon Dam and Lake Mead, Arizona. This depauperate fauna, consisting of 38 species, is dominated by euryecious Nearctic or Holarctic orthocladine taxa. In addition, a small Neotropical faunal component is represented by Polypedilum obelos Sublette & Sasa and Rheotanytarsus hamatus Sublette & Sasa. The following new synonyms are given: Protenthes riparius Malloch 1915 with Tanypus bellus Loew 1866 [= Procladius (Psilotanypus) bellus (Loew)]; Cricotopus olivetus Boesel 1983 with Cricotopus (Circotopus) annulator (Goetghebuer) 1927; Cricotopus edurus Sublette & Sublette 1971 with Orthocladius infuscatus Malloch 1915 [= Cricotopus (Cricotopus) infuscatus (Malloch)]. The following new species are described: Cricotopus (Cricotopus) blinni Sublette, Cricotopus (Cricotopus) hermanni Sublette, Metriocnemus stevensi Sublette, and Cladotanytarsus marki Sublette. We discuss the distribution and ecology of each chironomid species in this large, regulated, aridlands river.     

Late quaternary plant zonation and climate in southeastern Utah

Plant macrofossils from packrat middens in two southeastern Utah caves outline development of modern plant zonation from the late Wisconsin. Allen Canyon Cave (2195 m) and Fishmouth Cave (1585 m) are located along a continuous gradient of outcropping Navajo Sandstone that extends from the Abajo Mountains south to the San Juan River. By holding the site constant, changes in the floral composition for a plot of less than one hectare can be observed, even if sporadically, over tens of millennia. At Allen Canyon Cave, Engelmann spruce-alpine fir forest was replaced by the present vegetation consisting of pinyon-juniper woodland on exposed ridgetops and cliffside stands of Douglas fir, ponderosa pine, and aspen. Xerophytic woodland plants such as pinyon, plains prickly pear, and narrowleaf yucca arrived sometime in the middle Holocene between 7200 and 3400 B.P. At Fishmouth Cave, Utah juniper in Holocene middens replaced blue spruce, limber pine, Douglas fir, and dwarf and Rocky Mountain junipers in late Wisconsin samples. Disharmonious associations for the late Wisconsin occur only at the lower site with the xerophytes Mormon tea, plains prickly pear, and narrowleaf yucca growing alongside subalpine conifers. One possible explanation involves the late Wisconsin absence of ponderosa and pinyon pines from the Colorado Plateaus. Released from competition at their lower limits, subalpine conifers were able to expand into lower elevations and mix with xerophytic plants found today in understories of pinyon-juniper and ponderosa pine woodlands. Quantitative climatic estimates are derived for the late Wisconsin by applying vertical gradients for temperature and precipitation to the amount of vegetation depression. The Fishmouth Cave sequence indicates a minimum lowering of 850 m for blue spruce, limber pine, and dwarf juniper. A depression of at least 700 m for Engelmann spruce and alpine fir is suggested for the Allen Canyon locality. Use of conservatively low gradients for stations below 2080 m yields a 3–4 C cooling from present mean annual temperature and 35 to 60 percent more rainfall than today. Steeper gradients associated with more mountainous terrain suggest a 5 C lowering in temperature and up to 120 percent increase over modern precipitation.        

Bird community composition in relation to habitat and season in Betatakin Canyon, Navajo National Monument, Arizona

Bird species density, diversity, and species richness in relation to habitat and seasonal variations were studied in the Betatakin Canyon area of Navajo National Monument, Arizona. The two most prominent habitat types are a riparian forest deep in the canyon bottom and a mature pinyon-juniper woodland on the mesas and slopes above the canyon. One hundred thirty-five species of birds were encountered during the study and recorded by season and habitat. The avifauna assemblages demonstrate definite habitat selection into groups associated with the riparian and pinyon-juniper woodland communities. Diversity, density, and species richness were greatest in the riparian habitat during the spring and summer months. The differences in bird community composition were greater between seasons than between habitats. Pinyon pine and Gambel oak were highly selected as perch sites in the pinyon-juniper and riparian areas, respectively.     

Western balsam bark beetle, Dryocoetes confusus Swaine, flight periodicity in northern Utah

The flight periodicity of western balsam bark beetle ( Dryocoetes confusus Swaine) in Big Cottonwood Canyon, Utah, was studied during the summer months of 1992, 1993, and 1994. Contents of baited funnel traps were tallied by species up to 3 times weekly. Two main periods of flight activity were observed each year. The first and, generally, largest occurred in early summer soon after flight was initiated for the season. A 2nd period was observed in late summer, generally August. Timing of the 2 periods was influenced by unusually warm or cool weather in each study year. The 1st period had more males than females while the 2nd period had a majority of females. Except during periods of cool or wet weather, western balsam bark beetles were found to be active at least at minimal levels from June through September.     

A report on two large collections of the squat lobster Munidopsis platirostris (Decapoda,Anomura, Munidopsidae) from the Caribbean,with notes on their parasites,associates, and reproduction

The reproduction and parasite associates of the squat lobster Munidopsis platirostris (A. Milne Edwards and Bouvier, 1894) were investigated based on collections made in the Bahamas and Curaçao with grassmat and bundled fishing net traps used to collect sipunculan worms and other small invertebrates. Size of ovigerous M. platirostris was significantly correlated with clutch size for females from both localities but females from the Bahamas produced significantly more eggs (on average 10.1 eggs/clutch) than females from Curaçao (on average 6.6 eggs/clutch). Early embryos of M. platirostris from the Bahamas were 0.74–0.82 mm in diameter, similar to some other species of Munidopsidae as well as Chirostylidae. Two species of crustaceans, another squat lobster and a leptostracan, as well as a limpet mollusc, were collected with M. platirostris in the Bahamas, while a sipunculan was an associated species in a Curaçao collection. One specimen of M. platirostris had an unidentified cryptoniscoid epicaridean isopod, possibly representing a new genus and species. Two specimens of M. platirostris each had one rhizocephalan externa of a species belonging to Lernaeodiscus Müller, 1862 but their morphology does not match that of L. schmitti Reinhard, 1950, the only species in the genus known from squat lobsters in the western Atlantic. Additional materials and tools, such as DNA analysis, are needed to describe these potentially new parasites and we suggest that use of these traps may be an effective method to obtain additional samples.