Host-range and cold temperature tolerance of Floracarus perrepae Knihinicki & Boczek (Acari: Eriophyidae), a potential biological-control agent of Lygodium microphyllum (Pteridophyta: Lygodiaceae)

Abstract The leaf galling eriophyid mite Floracarus perrepae is a widespread and damaging herbivore of Old World climbing fern, Lygodium microphyllum , across its native range in tropical and subtropical Asia and Australia; and was therefore selected as a candidate biological-control agent for the invasive fern in Florida, USA. The host testing of F. perrepae focused on Lygodium species from North America and the Neotropics, along with threatened or endangered ferns from Florida. Sporeling ferns were used for the initial no-choice screening and F. perrepae showed normal development on the Florida genotype of L. microphyllum with 33.0 ± 4.6 mites per marginal leaf roll gall, modest development on the North American native Lygodium palmatum with 29.0 ± 9.3 mites per roll, and minor development on six other fern species (4.0–0.3). Leaf rolls were induced only on Lygodium species and full rolls were common only on L. microphyllum . These same six non-target plant species were tested again as more mature plants in both no-choice and choice tests and F. perrepae developed only on L. microphyllum . Lethal minimum temperature and cold stress tests were also conducted on F. perrepae . These tests revealed that it would not likely establish north of USDA hardiness zone 8a (−9.5 to −12.2°C), which means it would not overlap with L. palmatum from temperate North America. Our studies conclude that F. perrepae is specific to L. microphyllum , poses little or no risk to native or cultivated ferns in North and South America, and therefore should be considered for release in Florida.     

Cytotaxonomic investigations on some species of the genus Galium (Rubiaceae) from the Balkans

The results of a cytological and morphological investigation on the following species of the genus Galium L. collected in the Balkans are described and discussed: Sect. Platygalium Koch: G. rotundifolium L. and G. boreale L.; Sect. Aparinoides (Jord.) Gren.: G. palustre L.; Sect. Leiogalium Ledeb.: G. heldreichii Hal., G. lovcense Ur., G. album Mill. with the ssp. album, pycnotrichum (H.Br.) Krendl and prusense (C. Koch) Ehrend. et Krendl, G. lucidum All., G. corrudifolium Vill., G. scabrifolium (Boiss.) Hausskn., G. procurrens Ehrend., G. schultesii Vest, and G. bulgaricum Velen.; Sect. Kolgyda Dumort.: G. aparine L., G. intricatum Margot et Reut., G. parisiense L., G. divaricatum Pourr. ex Lam. and G. tenuissimum Bieb.     

Host specificity ofAceria (Eriophyes) malherbe, [Acari: Eriophyidae], a biological control agent for the weed,Convolvulus arvensis [Convolvulaceae]

The host specificity of the gall mite,Aceria (Eriophyes) malherbe (Nalepa), from Greece was studied under quarantine conditions at Albany, California USA. Of the species, ecotypes, or strains tested, onlyConvolvulus andCalystegia spp. supported gall formation and mite reproduction. Although 2 of the native, North AmericanCalystegia species that served as laboratory hosts are threatened or endangered species,A. malherbe is considered safe for release in the USA as a biological control agent of the weed,Convolvulus arvensis (L.).      

Biology and host specificity of Spurgia capitigena (Bremi) (Dipt., Cecidomyiidae), for the biological control of Euphorbia esula L. in North America

Abstract: Spurgia capitigena (Bremi) was evaluated as a potential biological control agent of leafy spurge, Euphorbia esula L. (species complex) in North America. To ascertain the host specificity of this gall midge, tests were conducted in the field at Cavaillon, France, and at laboratories located in Rome, Italy and Bozeman, Montana, USA, in 1994 and 1995. Twenty-nine plant species, including eight native North American euphorbias and biotypes of E . esula , were evaluated. In the laboratory, the gall-midge was able to induce galls on seven species of spurges (all belonging to the subgenus Esula) and the three accessions of leafy spurge. In field tests, S . capitigena infested only E . esula . The biology and host specificity of S . capitigena from France appears to be similar to that observed for Spurgia esulae Gagné from Italy (= Bayeria capitigena Bremi). In France, this gall-midge was commonly observed from early May until October in habitats of very heavy and moist soils located along water channels, roadsides and in fruit orchards. Galls were induced on meristematic tissues, and thereby prevented flowering.     

Evaluation of Cyphocleonus trisulcatus (Coleoptera: Curculionidae) as a potential biological control agent for Leucanthemum vulgare in North America

Leucanthemum vulgare, a perennial forb native to Eurasia, has become invasive in several other parts of the world. Since there is a lack of methods suitable for sustainable management of this plant across invaded landscapes, a biological control project has been initiated in North America and Australia. We evaluated the potential of the root-feeding weevil Cyphocleonus trisulcatus as a biological control agent for L. vulgare in North America by investigating its impact on L. vulgare and studying its larval host range under no-choice, multiple-choice cage and open-field conditions. Our impact experiment revealed that larval feeding by C. trisulcatus can severely damage and even kill plants. No-choice tests conducted with 41 non-target species showed that C. trisulcatus can develop on five closely related species. In a multiple-choice cage test established with three of these species, the medicinal plant Matricaria chamomilla and the native North American M. occidentalis were also attacked, but to a lower degree than L. vulgare. An open-field test conducted with four varieties of the ornamental Shasta daisy (Leucanthemum × superbum and L. × maximum) revealed similar attack on Shasta daisy as on L. vulgare. Due to the popularity of Shasta daisies, we suggest that C. trisulcatus, despite its potential to suppress L. vulgare populations, is not a suitable biological control agent for North America. However, C. trisulcatus may still be a potential biological control agent for L. vulgare in regions where Shasta daisy is less popular, such as Australia. Further host-range tests with species native to Australia as well as with additional species important for the horticultural industry in Australia are currently under way.     

Evaluation of host range and larval feeding impact of Chrysolina aurichalcea asclepiadis (Villa): considerations for biological control of Vincetoxicum in North America

A biological control program has been initiated against European swallow-worts Vincetoxicum nigrum (L.) Moench. and V. rossicum (Kleopow) Barbar., which are invasive in North America. A population of the leaf beetle Chrysolina aurichalcea asclepiadis (Villa) originating from the western Alps has been under evaluation as a part of this program. The preliminary host range of C. a. asclepiadis was determined among 37 potential host plants. In addition, a prerelease impact study was conducted to determine the effect of larval feeding on the performance of V. nigrum. Under no-choice conditions beetle larvae completed development on nine plant species within the genera Artemisia and Tanacetum (Asteraceae) and Asclepias and Vincetoxicum (Apocynaceae). The host range of adults is broader than larvae (13 plant species within five genera received sustained feeding). Three of the six nontarget species supporting larval development are native to North America, however in separate oviposition tests, female beetles failed to produce eggs when confined to these hosts. In multiple-choice tests, neither larvae nor adults preferred Vincetoxicum spp. to nontarget species. Larval damage by C. a. asclepiadis at densities at and above five larvae per plant substantially reduced growth, biomass, and delayed reproduction of V. nigrum. However, this population of C. a. asclepiadis is polyphagous and unsuitable for biological control of Vincetoxicum because of potential risk of attack to Asclepias tuberosa L. and native North American Asteraceae, particularly Artemisia.     

Biology and host specificity of Rhinusa pilosa,a recommended biological control agent of Linaria vulgaris

Linaria vulgaris Mill. (Plantaginaceae), common or yellow toadflax, is a Eurasian short-lived perennial forb invasive throughout temperate North America. Rhinusa pilosa (Gyllenhal) (Coleoptera, Curculionidae) is a univoltine shoot-galling weevil found exclusively on L. vulgaris in Europe. Under no-choice test conditions, 13 non-native Linaria species exposed to R. pilosa were accepted for oviposition and most were found to be suitable, to varying degrees, for gall and larval development. Adult feeding and survival was minimal on native North American species in the plant tribe Antirrhineae which includes the target plant. In no-choice tests with 63 native North American species and 24 other non-target species outside Linaria, oviposition was limited to four native North American species. Only three larvae developed to the adult stage on Sairocarpus virga (A. Gray) D.A. Sutton, with no negative impact on plant growth. Risks to native flora from the release of R. pilosa are therefore expected to be minimal. The Technical Advisory Group for the Biological Control of Weeds (TAG—BCW) has recommended release of R. pilosa in September 2013.     

Molecular ecology of some Cecidophyopsis mites (Acari: Eriophyidae) on Ribes species and evidence for their natural cross colonisation of blackcurrant (R. nigrum)

Ribosomal DNA from Cecidophyopsis mites from different Ribes species was amplified using the polymerase chain reaction and the products digested using restriction enzymes. After separating the DNA fragments on gels, it was possible to identify specimens of mites obtained from field samples by comparing the profiles of their DNA banding patterns with those of known Cecidophyopsis species. Using this analysis, a non-gall forming mite found infesting blackcurrant buds in New Zealand was identified as the gooseberry mite (C. grossulariae). On wild red currant (Ribes spicatum) from Finland showing two sizes of galled buds, the red currant gall mite (C. selachodon) was identified in the larger galls located at the tips of branches and a distinct mite in the smaller galls located on the lower parts of the branches. A mite with a DNA banding profile indistinguishable from this latter mite from R. spicatum was also identified in galled buds of blackcurrant genotypes growing in Finland, including those containing the blackcurrant gall mite (C. n'ftw)-resistance genes P or Ce. The DNA banding profile of this mite resembled most closely that of C. ribis , but was distinct from it. The occurrence of C. grossulariae and this distinct Cecidophyopsis mite on blackcurrant has implications for the genetic control of Cecidophyopsis mites and possibly for the spread of the reversion disease agent in this crop.     

Antifeedant activity of an anthraquinone aldehyde in Galium aparine L. against Spodoptera litura F

The insect antifeedant anthraquinone aldehyde nordamnacanthal (1,3-dihydroxy-anthraquinone-2-al) was identified in Galium aparine L., and isolated from the root powder of akane (Rubia akane), a member of the Rubiaceae. Structure-activity relationship (SAR) studies using a series of anthraquinone analogues suggested that the aldehyde group on the anthraquinone was more important than the quinone moiety for antifeedant activity against the common cutworm (Spodoptera litura). High levels of nordamnacanthal were found in the seed leaf stage and in callus tissue induced from seedlings of G. aparine, but its concentration decreased with plant development. Since these compounds are natural pigments for dying textiles, we also evaluated the antifeedant activity against the carpet beetle (Attagenus japonicus ), a textile pest was also evaluated. While nordamnacanthal had strong antifeedant activity against the common cutworm, it did not show any antifeedant activity against the carpet beetle. The most effective antifeedant against the carpet beetle was the major constituent in the extract of R. trictorum, lucidin-3-O-primeveroside, a food pigment.     

Post-fire management targeting invasive annual grasses may have inadvertently released the exotic perennial forb Chondrilla juncea and suppressed its biocontrol agent

Top-down and bottom-up factors affecting invasive populations are rarely considered simultaneously, yet their interactive responses to disturbances and management interventions can be essential to understanding invasion patterns. We evaluated post-fire responses of the exotic perennial forb Chondrilla juncea (rush skeletonweed) and its biocontrol agents to landscape factors and a post-fire combined herbicide (imazapic) and bacteria (Pseudomonas fluorescens strain MB906) treatment that targeted invasive annual grasses in a sagebrush steppe ecosystem. Biocontrol agents released against C. juncea in previous decades included Cystiphora schmidti (gall midge), Aceria chondrillae (gall mite), and Puccinia chondrillina (rust fungus). C. juncea abundance was greater in sprayed than unsprayed plots, and where soils were coarser, slopes faced southwest, solar heat loads and topographic water accumulation were greater, and cover of deep-rooted native perennials was lower. Mite infestation was greater in unsprayed plots, midge infestation was greater at higher elevations on steeper slopes, and midges were more abundant while rust was less abundant on gravelly soils. Biocontrol infestation levels varied considerably between years and could not be predicted in 2019 from 2018 infestation levels. Multiple biocontrol species were often present at the same plots but were rarely present on the same C. juncea individuals. These results suggest that spatial patterns of invasion by C. juncea are related to deep-soil water availability, warmer conditions, and alleviation of competition. Treatments designed to reduce invasive annual grasses may inadvertently release C. juncea by both reducing plant competition for soil resources and affecting biocontrol agent (mite) abundance.

     

Interaction between the non-myrmecochorous herb Galium aparine and the ant Formica polyctena

Non-myrmecochorous plant species are able to colonize Formica polyctena mounds. Galium aparine has been chosen as a model to study this kind of ant-plant association. In order to understand benefits to ants and plants, we have studied removal rates of fruits by ant workers, and compared soil seed material, seedling pattern and development of plants of the nest versus forest.Ant workers remove fruits at very low rates; removal time for half of the sample of 10 seeds was on average 5 h. Removed fruits were usually lost within 20–30 cm distance of the ant route. The soil samples from the nests contained a higher number of fruits, than those from randomly chosen sites in the forest. Seeds start to germinate in late August; on the F. polyctena mounds significantly earlier than at surrounding places. The seedling density was higher within experimental quadrats located on mound sites especially in late September–early October. The highest density of seedlings was on the slope of the mound. Plants from mounds are significantly longer, have a higher number of nodes during all stages of development and a higher mortality rate than that at surrounding places.The interactions between G. aparine and F. polyctena ants are asymmetrical. In spite of higher mortality rate of seedlings on mounds, ants provide some benefits to the plant. Firstly, workers are additional dispersal agents for G. aparine fruits. Furthermore, the soil peculiarities of ant mounds are beneficial for seed germination and seedling growth. G. aparine does not provide ants with energetically valuable food like plants with elaiosome-bearing seeds. Moreover, the high density of seedlings and plants on the mounds shades the nest.     

Invasive Galega officinalis (Goat's rue) plants in Canada form a symbiotic association with strains of Neorhizobium galegae sv. officinalis originating from the Old World

The toxic legume plant, Galega officinalis, is native to the Eastern Mediterranean and Black Sea regions. This legume is considered to be a noxious weed, and its establishment in Canada may have resulted from ornamental planting and/or field trials. In its native range, a highly specific nitrogen‐fixing symbiosis with the bacterium, Neorhizobium galegae symbiovar (sv.) officinalis, is required for normal growth. In North America, nothing is known about the bacterial symbionts of G. officinalis. Our purpose was to determine the species and symbiovar identity of symbiotic bacteria associated with invasive plants of G. officinalis at five sites in the province of Ontario, Canada.  Sequence analysis of four housekeeping (16S rRNA, atpD, glnII, and recA) and two symbiosis (nodC and nifH) genes showed that all 50 bacterial isolates from root nodules of G. officinalis at the five Canadian sites were identical to strains of N. galegae sv. officinalis originating either from Europe or the Caucasus. Plant tests indicated that soils collected from four Canadian sites without a history of agriculture or presence of G. officinalis were deficient in symbiotic bacteria capable of eliciting nodules on this plant. Collectively our data support the hypothesis of anthropogenic co‐introduction of G. officinalis and its specific symbiotic bacterium into Canada from the Old World. Factors that may limit the spread of G. officinalis in new environments are discussed.     

Biology and host specificity of the chromolaena stem gall fly, Cecidochares connexa (Macquart) (Diptera: Tephritidae)

Abstract The stem-galling tephritid fly Cecidochares connexa (Macquart) was investigated as a potential biocontrol agent against the weed Chromolaena odorata (L) King & Robinson in Indonesia. Adults were tested in choice and no-choice tests, on 55 non-target plant species in 17 families. No oviposition was recorded on 53 of the species, while oviposition but no larval development was recorded on two. Field releases commenced in 1995 and establishment was immediate. The gall fly is now widely established in most Indonesian provinces, where gall parasitism and predation is generally low.     

Does microhabitat structure affect foliar mite assemblages?

1. Habitat structure is an important factor influencing population dynamics and trophic organisation of terrestrial invertebrates. The phylloplane zone on vascular plant leaves is topographically complex, containing a multitude of microhabitats such as leaf hairs, lesions, and structural refugia such as domatia, which may modify interactions between resident invertebrate communities, colonisers, and subsequent trophic relationships. Leaf domatia are small indentations on the underside of leaves and are often inhabited by potentially beneficial mites and other arthropods. 2. This study investigated the relationship between domatia availability and foliar mite assemblages in contrasting habitats (native forest, plantation forest, and pasture) using a standard test plant (the endemic New Zealand shrub Coprosma lucida, J.R. & G. Forst.). 3. Diverse woody native vegetation types supported higher numbers of mite species than either plantation forest or pastoral grasses. The highest number of mite species occurred in the native forest (63%), plantation forest (38%), and pastoral grasses (25%). In the native vegetation type, experimental C. lucida leaves with domatia supported higher mite densities, greater colonisation success, and more diverse mite assemblages than those without domatia. Mite assemblages within the pastoral site were significantly different from the other two vegetation types. Only one fungivorous mite species, Orthotydeus californicus, occurred compared to five mite species in native and plantation forests. 4. This study indicated that foliar mite assemblages in native vegetation on experimental C. lucida shrubs are influenced by domatia availability, resident foliar mites, and local mite assemblages.     

Molecular insights into the taxonomy of Glyceria (Poaceae: Meliceae) in North America

Eighteen Glyceria species grow in the United States and Canada, with 16 being native to the region. We used data from morphology and three chloroplast DNA intergenic regions to address taxonomic questions concerning Glyceria in North America, particularly the status of G. declinata, G. occidentalis, G. fluitans, G. striata, and G. elata in western North America. The chloroplast data confirmed the presence of two European species, G. declinata and G. fluitans, in western North America. Glyceria occidentalis was exceptional among the taxa studied in having chloroplast genotypes that fell into two different clades, one of which contained G. fluitans and the other the North American species G. leptostachya. The morphological data showed G. occidentalis to be intermediate between G. fluitans and G. leptostachya with respect to their distinguishing characters. Based on these results, we hypothesize that G. occidentalis consists of hybrids between G. fluitans and G. leptostachya. Glyceria elata and G. striata, which have sometimes been treated as a single species, had different chloroplast genotypes, supporting their recognition as distinct taxa. DNA data from all three intergenic regions would be needed for unequivocal identification of the non-hybrid species examined.     

外源茉莉酸及枸杞瘿螨危害诱导的枸杞防御反应

利用植物的诱导抗性防治病虫害越来越受重视, 为寻求枸杞害虫防治的新方法, 探索外源茉莉酸对枸杞的诱导反应及机制, 本研究测定了外源茉莉酸(jasmonic acid, JA)及枸杞瘿螨Aceria pallida Keifer危害对枸杞叶蛋白酶抑制剂(protease inhibitors, PIs)、 超氧化物歧化酶（superoxide dismutase, SOD）、 苯丙氨酸解氨酶（phenylalanine ammonia-lyase, PAL）活性及多糖（polysaccharide）、 木质素（xylogen）含量的变化。结果表明： 外源JA处理或枸杞瘿螨危害均使枸杞叶PIs, SOD和PAL活性及木质素含量显著提高, 却使多糖含量显著降低。胰凝乳蛋白酶抑制剂（chymotrypsin inhibitors, CI)和胰蛋白酶抑制剂(trypsin inhibitor, TI)在喷洒JA或枸杞瘿螨危害的第5 天增幅最大, 且在JA浓度、 瘿螨危害间差异显著(P<0.05), 其中, 0.1 mmol/L JA处理使CI增加了96%, TI增加了128%, 枸杞瘿螨危害使CI增加了94%, TI增加了122%。枸杞叶多糖含量在JA处理或瘿螨危害后显著降低(P<0.05), 且与JA浓度有关, 浓度越高, 降低越明显。 JA或瘿螨危害使枸杞叶木质素增加的最大值出现在处理后的第5 天, 高浓度JA、 瘿螨危害及低浓度JA分别使其增加了70%, 41%和36%, 三者之间差异显著(P<0.05), 第10 天增加幅度减低。 3种处理同样使SOD和PAL活性显著增加(P<0.05), 第5天时增幅最大, 高浓度JA 处理使枸杞叶SOD活性由对照的70.77 U增加到128.98 U, PAL活性由10.91 U增加到20.59 U, 第10天时呈现下降趋势。这些与抗性有关物质的变化及其变化趋势, 说明外源茉莉酸及枸杞瘿螨均可诱导枸杞产生防御反应, 而且反应的程度与茉莉酸浓度及处理后的时间有关。     

Mechanical adaptations of cleavers (Galium aparine)

BACKGROUND AND AIMS: Cleavers (Galium aparine) is a fast-growing herbaceous annual with a semi-self-supporting, scrambling-ascending growth habit. Mature plants often use upright species for support. It is common in hedgerows and on waste ground. This study aims to characterize the mechanical behaviour of the stem and roots of cleavers and relate this to the arrangement of structural tissue, the net microfibrillar orientations in the cell walls, and plant growth habit. METHODS: The morphology and mechanics of mature cleavers was investigated using plants grown in pots and ones collected from the grounds at the University of Lincoln, Lincoln, UK. Tensile tests were carried out on the stem and the basal section of the first-order lateral roots. The net orientation of cellulose microfibrils in the cell walls was investigated using polarized light microscopy. KEY RESULTS: Results show that the basal regions of the stem and first-order lateral roots were highly extensible. Breaking strains of 24 +/- 7% were recorded for the stem base and 28 +/- 6% for the roots. Anatomical observations showed that the lower stem (base + 100 mm) was circular in cross-section with a solid central core of vascular tissue, whereas further up the stem the transverse section showed a typical four-angled shape with a ring-like arrangement of vascular tissue and sclerenchyma bundles in the corners. The net orientation of wall microfibrils in the secondary xylem diverges from the longitudinal by between 8 and 9 degrees . CONCLUSIONS: The basal region of the stem of cleavers is highly extensible, but the mechanism by which the stem is able to withstand such high breaking strains is unclear; reorientation of the cellulose fibrils in the stem along the axis of loading is not thought to be responsible.     

Always on the bright side: the climbing mechanism of Galium aparine

Galium aparine is a herbaceous climbing plant that attaches to host plants mainly via its leaves, which are covered by hooked trichomes. Although such hooks are found on both leaf surfaces, the leaves of G. aparine are mainly positioned upon the leaves of supporting plants and rarely beneath. In order to understand the mechanism underlying this observation, we have studied structural and mechanical properties of single leaf hooks, frictional properties of leaf surfaces, turgor pressure in different leaf tissues and bending properties of the leaves in different directions. Abaxial and adaxial leaf hooks differ significantly in orientation, distribution, structure and mechanical properties. In accordance with these differences, friction properties of leaves depend on the direction of the applied force and differ significantly between both leaf surfaces. This results in a ratchet mechanism. Abaxial leaf hooks provide strong attachment upon the leaves of adjacent plants, whereas adaxial hooks cause a gliding-off from the underside of the leaves of host plants. Thus, the leaves of G. aparine can function as attachment organs, and simultaneously orient themselves advantageously for their photosynthetic function. Further adaptations in turgor pressure or concerning an anisotropy of the flexural stiffness of the leaves have not been found.     

Making host specificity testing more efficient: Exploring the use of abridged test plant lists

Testing the specificity of candidate agents is a key component of risk analysis in weed biological control. This step is often time-consuming due to the numerous plant species that need to be tested under quarantine conditions in the invaded country of the weed species. Here, we examined whether an abridged phylogenetically based test list could be used in the weed's native range to quickly screen the host specificity of candidate agents. Ten plant species were used to test the host specificity of a promising candidate for the biological control of Sonchus oleraceus in Australia, the gall midge, Cystiphora sonchi. No-choice and choice tests were carried out in the native Mediterranean range of the midge. The results showed the midge has potential to threaten native Australian species, as those species showed high infestation levels in no-choice tests and produced significantly higher numbers of galls in choice tests. As a result of this approach, C. sonchi was rapidly discarded from the list of agents to be imported into Australian quarantines for further tests. This study demonstrates that testing a few key phylogenetically related species in the native range may save cost and effort in a weed biological control programme.     

Native cover crops suppress exotic annuals and favor native perennials in a greenhouse competition experiment

In a greenhouse experiment, we examined the effectiveness of four native cover crops for controlling four exotic, invasive species and increasing success of four western North American grassland species. Planting the annual cover crops, annual ragweed (Ambrosia artemisiifolia) and common sunflower (Helianthus annuus), reduced the biomass of the exotic species cheatgrass (Bromus tectorum), Japanese brome (Bromus japonicus), Canada thistle (Cirsium arvense), and whitetop (Cardaria draba). The annual cover crops also reduced the desired species biomass in competition with the perennial exotics, but either increased or did not affect the desired species biomass in competition with the annual exotics. Planting the perennial cover crops, Canada goldenrod (Solidago canadensis) and littleleaf pussytoes (Antennaria microphylla), rarely inhibited exotic species, but did increase the desired species biomass. Field experiments are needed to test the cover crops under more ecologically relevant conditions, but our results suggested that the annual cover crops may be effective for controlling invasive annuals and for facilitating native perennials.