Ground Ivy (Glechoma hederacea L.) Outline

Introduction

A.) Name

B.) General

 

Description (Stubbendieck et al. 1995 and Hutchings and Price 1999)

A.) Stem

B.) Leaves

C.) Flowers

D.) Seed

E.) Roots

Habitat

A.) Geographical Distribution

Ground ivy occurs throughout Europe, except in Iceland and Turkey. It is also found in Western and Northern Asia to Japan (Hulten 1971).

Ground ivy has been introduced in America where its range extends southwards from Canada to Georgia, Tennessee, Kansas, and Colorado (Gill 1979).

Ground ivy has been introduced in New Zealand, where it is found in many southern localities in the North Island and in many localities throughout the south island (Webb et al. 1988).

B.) Climate

Glechoma hederacea is a species of temperate latitudes (Ellenberg et al. 1991).

Glechoma hederacea is primarily a species of the colline-submontane zones of temperate Europe, with an oceanic to suboceanic distribution (Landolt 1977 and Hutchings and Price 1999).

The altitudinal range of Glechoma hederacea has been given from sea level in England to about 1600 meters in the Alps (Grime et al. 1988 and Hulten 1971).

C.) Substratum

Glechoma hederacea occurs mainly on damp, heavy, fertile, and calcareous soils. It is most frequent in the pH range from slightly acidic to slightly alkaline (pH 5.5 to 7.5), but can also be found on soils with a pH as low as 4.0 (Fitter 1978 and Hutchings and Price 1999).

Ground ivy does not tolerate strongly acidic soils and it is intolerant of salinity (Landolt 1977).

Glechoma hederacea is typically found in fertile habitats where potential dominant species are restricted in vigor by both shade and disturbance (Grime et al. 1988).

Glechoma hederacea becomes abundant only when there is a good phosphate and nitrate supply and a moderately rich supply of cations including Ca+2 in the soil substrate (Kreutzer and Seibert 1984 and Sinker et al. 1985).

D.) Communities

Glechoma hederacea is found in shaded, moist soils of lawns, gardens, wooded hill sides, waste places, pastures, and along streams (Stubbendieck et al. 1995).

Glechoma hederacea is typically a plant of shaded habitats including woodlands and hedgerows. It is frequent on shaded roadsides, in clear-felled and coppiced woodlands, scrub and derelict woodland, and in orchards (Hutchings and Price 1999).

Glechoma hederacea has a limited capacity to persist under tall herbs or beneath continuos tree canopy. It is a species that is most characteristic of lightly or patchily shaded habitats (Grime et al. 1988).

Competition

A.) Interspecific Interactions

The proportion of biomass allocated to stolons increased from 20% in clones without competition to 36-38% in clones subjected to short and tall grass competition (Hutchings and Price 1993).

Glechoma hederacea is a patch or carpet forming herb. Occasionally it forms extensive monoclonal stands of considerable size, but more often infiltrates, rather than dominates vegetation. It explores habitat by rapid stolon extension and establishes short lived ramets that exploit local resources (Hutchings and Price 1999).

Studies of the effects of leachates of Glechoma hederacea on Raphanus sativus and Bromus tectorum suggest strong allelopathic effects on these species. Leachates from decaying air dried leaves of Glechoma hederacea decreased seed germination, especially in Bromus tectorum (Rice 1986).

When competing with tall grass, Glechoma hederacea increased the length of its vertical petioles bearing leaves. Thus, G. hederacea escaped from competition for light in a vertical direction and responded aggressively to local above ground competition (Price and Hutchings 1996).

 B.) Intraspecific Interactions

Glechoma hederacea has a striking capacity to produce a higher yield under heterogeneous growing conditions than under homogeneous conditions providing the same quantity of resource (Birch and Hutchings 1994).

Glechoma hederacea plants growing in isolation were richly branched with shorter internodes, and thus produced more densely packed ramets (Price and Hutchings 1996).

Glechoma hederacea forms architecturally simple and short-lived ramets which allocate the majority of captured resources to stolon growth and formation of new ramets. Continuos movement through the habitat, and establishment of new ramets in new sites, provides G. hederacea with an effective method for rapidly exploiting resource rich patches in spatially and temporally variable environments, and for escaping from competition, thereby increasing the probability of persistence (Steufer and Hutchings 1994).

C.) Plasticity

Glechoma hederacea petioles are highly variable in length from 10 mm in grassland to over 200 mm in woodland with dense canopy cover and dense ground vegetation (Hutchings and Price 1999).

Clones growing entirely in full sunlight generally produce smaller leaves than clones growing in shade (Sinker et al. 1985 and Grime et al. 1988).

Internodes of the plagiotropic stolons of Glechoma hederacea lengthen under shade and other adverse conditions, including low nutrient supply. This lengthening is accompanied by significant decreases in weight per unit length and stolon cross-sectional area, mainly achieved by reducing the volume of the parenchymatous tissue. The vascular system is considerably less plastic, thus conserving its capacity for resource transport (Price and Hutchings 1992).

Many morphological characteristics are highly plastic, both within stolons and between different stolons of the same clone which experience different growing conditions (Slade and Hutchings 1987).

Phenology

Germination of Glechoma hederacea is epigeal. Emergence of some seedlings can be observed within 3 days. The radicle emerges first, followed within 1 or 2 days by the cotyledons. The first true leaves begin to expand after approximately one week (Grime et al. 1981 and Hutchings and Price 1999).

Glechoma hederacea overwinters either as small, two leafed ramets, or as 8-10 leafed rosettes produced in the autumn by rooted ramets. From April onwards, the rosettes develop two-leafed nodes at regular intervals along ascending stems. Flowers emerge in the leaf axils between April and July. After flowering the ascending stem continues its vegetative growth (Hutchings and Price 1999).

Individual ramets of Glechoma hederacea behave as annual structures, living for at most one growing season, although the clone itself survives indefinitely, through the production of overwintering daughter ramets. Ramets are normally replaced, in undisturbed natural populations, by vegetative growth of the clone rather than by seeding establishment (Slade and Hutchings 1989).

Reproduction

A.) Pollination

Gynodioecious species are often insect pollinated, and restrictions in pollen production and dispersal could therefore play a part in determining the fecundity of the females, since they must outcross (Lewis 1941 and Lloyd 1974).

The flowers of Glechoma hederacea are nectariferous. They are insect pollinated, usually by bees (Garrard and Streeter 1983).

The estimated mean pollen dispersal distance was 5.9 meters when calculated on fruit set and 5.3 meters when calculated on seed set (Widen and Widen 1990).

B.) Seed Production

In Glechoma hederacea considerable quantities of resources are allocated to flowering and seed production, as shown by the larger size and longer half-lives of flowering ramets, and the long half-lives of ramets that produce seeds (Slade and Hutchings 1989).

Each flower produces up to four seeds. Flowers are borne in false whorls of 2-5. Thus, there is the potential to produce from eight to twenty seeds per node, but output per node is often well below potential (Widen and Widen 1990).

Stagg found small numbers of seeds of Glechoma hederacea in soil from exarable sites on chalk at several collection dates through the year, with a peak density in August (Stagg 1996 and Hutchings and Price 1999).

Petrov and Palkina found viable seeds of Glechoma hederacea buried in soil beneath a slightly over 40 year old spruce plantation. As Glechoma hederacea would be unlikely to set seed in dense shade, and as the seeds were buried, they had probably been in the soil for 40 years (Petrov and Palkina 1983 and Hutchings and Price 1999).

C.) Asexual Reproduction

If females divert their resources to clonal growth instead of seed production, they may gain some advantage over the hermaphrodites and thus escape from extinction (Widen and Widen 1990).

Vegetative spread is rapid, and monocultures covering several square meters are occasionally produced (Hutchings and Price 1999).

D.) Dispersal

The calyx bends down at seed maturation and the seeds are passively dispersed (barochory), although they may further be dispersed by ants (Bouman and Meeuse 1992).

On contact with water, nutlets produce moderate quantities of mucilage (myxocarpy) that fixes them to the substrate (Ryding 1992).

Control

A.) Cultural control

Research indicates that borax can be used to selectively control ground ivy in a cool season turfgrass with only minor and temporary injury to the grass. A late spring application of 610 g/100 m2 (20 oz/1000 ft2) liquid Twenty Mule Team Borax provided acceptable and consistent ground ivy control. (Hatterman et al.1996).

Control ground ivy in the spring by raising the runners with a stiff raking and by mowing close to the ground (Schultz 1989).

The best way to control ground ivy in flower and vegetable gardens is by hand pulling and hoeing. The key to effective control of ground ivy in gardens is persistence. Repeatedly pull and hoe the ground ivy until it has been eliminated (Jauron 1997).

Preliminary greenhouse experiments showed that ground ivy was extremely sensitive to borax applications compared with cool-season grasses and other broadleaf weeds that are commonly found in landscapes (Hatterman et al. 1996).

Once the ground ivy has been effectively controlled and a healthy lawn reestablished, the home gardener needs to use good mowing, fertilization, watering, and cultivation practices to maintain a dense, healthy, competitive stand of turfgrass which should help discourage future invasions of this aggressive weed (Jauron 1997).

B.) Chemical control

Super Trimec (2, 4-D + Dicamba + dichlorprop) at 560 ml/100m2 controlled ground ivy consistently (>85%) during 1991 and 1992 seasons (Hatterman et al. 1996).

The most effective control comes with the use of combination herbicides that contain the product dicamba. The best time to achieve good control is in the fall, from mid-September to early November. The next best time to control ground ivy is when it is just beginning to flower. This occurs towards the end of April. Two herbicide applications, spaced 10 to 14 days apart are usually necessary to achieve good control (Rindels 1994).

The most effective broadleaf herbicide products on ground ivy are those that contain dicamba. Trimec and Weed-B-Gone Lawn Weed Killer are two widely sold products that contain dicamba (Jauron 1997).

Usage

Ground ivy is used by modern herbalists as a remedy for kidney disease, abscesses, tumors, and eye problems (Grieve 1976 and Potterton 1985).

Before the cultivation of hops in the XVI century, ground ivy was the chief source of bitter for flavoring beer (Hyde 1976 and Garrard and Streeter 1983).

Glechoma hederacea ssp. Variegata has been sold as a horticultural plant for use in hanging baskets (Hessayon 1987).

Significance of Ground ivy as a weed problem

Ground ivy is an aggressive, low-growing, perennial invader of lawns, vegetable gardens, and flower beds (Rindels 1994).

The species is likely to remain widespread and abundant because of its success in exploiting many of the productive and disturbed habitat types that are created by modern land use (Grime et al. 1988 and Hodgson 1989).

Ground Ivy Bibliography

 

Table of Contents

 

Biology and Habitat

Antipov, N.I. 1980. The water regime of leaf galls in different species of plants. Soviet Plant Physiology. 27:115-118.

Baiderin, V. 1981. Experimental modeling of effects of winter recreation activity on ecology. Ekologiya (Moscow). 0(3):13-21.

Baltzer, H., Braun, P., and Koehler, W. 1998. Cellular automata models for vegetation dynamics. Ecological Modeling. 107:113-125.

Beerling, D.J. and Kelly, C.K. 1997. Stomatal density responses of temperate woodland plants over the past seven decades of CO2 increase: a companion of Salisbury (1927) with contemporary data. American Journal of Botany. 84:1572-1583.

Bentham, G. 1858. Handbook of British flora, a description of the flowering plants and ferns indigenous to or naturalized in the British Isles. London, Lovell Reese. pg. 418.

Bones, B.G. 1972. Fine structural observations on necrotic root apices in Glechoma hederacea L. Protoplasma. 74:41-72.

Caputa, J. 1982. Grassland weeds. Revue-Suisse-d’ Agriculture. 14:122-127.

Chen, C. 1981. The Eurasian genus Glechoma and its relationship with allied genera. Acta Botanica Yunnanica. 1(1):81-89.

Crampton, A.B., Stutter, O., Kirby, K.J., and Welch, R.C. 1998. Changes in the composition of Monks Wood National Nature Reserve 1964-1996. Arboricultural Journal. 22(3):229-245.

Ellenberg, H., Weber, H.E., Dull, R., Wirth, V., and Paulissen, D. 1991. Zeigerwerte von Pflanzen in Mitteleuropa. Scripta Geobotanica. 18:1-248.

Fernandes, R. 1971. Note on Glechoma hederacea L. Bot-J-Linn-Soc. v. 64 p. 72.

Fitter, A.H. 1978. An Atlas of Wild Flowers of Britain and Northern Europe. Collins, London, UK.

Fogg, J.M., Jr. 1945. Weeds of Lawn and Garden. University of Pennsylvania Press, Philadelphia. p. 215.

Gamor, F.D., Komendar, V.I., Abramova, L.M., and Mirkin, B.M. 1983. Dependence of distribution of trans-Carpathian weeds on altitude. Ukrains’kii-Botanichnii-Zhurnal. 40:54-57.

Gill, L.S. 1979. Cyto-taxonomic studies of the tribe Nepeteae (Labiatae) in Canada. Genetica. 50:111-118.

Grime, J.P., Hodgson, J.G., and Hunt, R. 1988. Comparative Plant Ecology: a Functional Approach to Common British Species. Unwin Hyman, London, UK.

Hermy, M. and Stieperacaere, H. 1981. An indirect gradient analysis of the ecological relationships between ancient and recent riverine woodlands to the south of Bruges (Flanders, Belgium). Vegetatio. 44(1):43-49.

Hodgson, J.G. 1989. What is happening to the British flora? An investigation of commonness and rarity. Plants Today. 2:26-32.

Hulten, E. 1971. The Circumpolar Plants. II. Dicotyledons. Almquist and Wiksell, Stockholm, Sweden.

Hutchings, M.J. 1999. Clonal plants as cooperative systems: Benefits in heterogeneous environments. Plant Species Biology. 14:1-10.

Hutchings, M.J. and Bradbury, I.K. 1986. Ecological perspectives on clonal perennial plants. BioScience. 36:178-182.

Hutchings, M.J. and Price, E.A.C. 1993. Does physiology integration enable clonal herbs to integrate the effects of environmental heterogeneity? Plant Species Biology. 8:95-105.

Hutchings, M.J. and Price, E.A.C. 1999. Glechoma hederacea L. Journal of Ecology. 82(2):347-364.

Hutchings, M.J. and Wijesinghe, D.K. 1997. Patchy Habitats, division of labour and growth dividends in clonal plants. Trends in Ecology and Evolution. 12:390-394.

Jeong, J.H., Kim, Y.J. and Hang, Y.P. 1983. Effect of different light intensities on the growth of several indoor ornamental plants. Research-Reports-Office of Rural Development, S. Korea, Horticulture. 25:131-136.

Klimes, L. 1997. Variation in autumnal growth of hermaphroditic clones of Glechoma hederacea originating from two geographical regions and two habitats. Preslia. 69:175-183.

Korven, N.A. 1976. Creeping Charlie an attractive weed Glechoma hederacea. Pub-Manit-Dep-Agric. p. 559.

Kreutzer, K. and Seibert, P.P. 1984. Differences in supply of phosphorus and other nutrient elements in the ash/elm lowlands of southern Bavaria. Forstwissenschaftliches Centralblatt. 103:139-149.

Landolt, E. 1977. Okologische Zeigerwerte zur Schweizer Flora. Veroffentlichungen des Geobotanischen Instituts der Eidgenossische Technische Hochschule, Stiftung Rubel, Zurich. 64:1-208.

Lau, R.R. and Young, D.R. 1988. Influence of physiological integration on survivorship and water relations in a clonal herb. Ecology. 69:215-219.

Lawrence, B.M. 1992. Chemical components of Labiatae oils and their exploitation. Advances in Labiatae Science. p. 399-436. Royal Botanical Gardens, Kew, UK.

LeTacon, F., Timbal, J. and Valdenaire, J.M. 1982. Effect of form of mineral nitrogen on the amount of herbaceous plant species of forest. Acta-Oecologica. 3:307-318.

Mitich, L.W. 1994. Ground Ivy. Weed Technology: Journal of the Weed Science Society of America. 8(2):413-415.

Pearson, J. and Soares, A. 1998. Physiological responses of plant leaves to atmospheric ammonia and ammonium. Atmospheric Environment (United Kingdom). 32(3):533-538.

Price, E.A.C. 1991. Investigations into the ecology and intraclonal physiology of Glechoma hederacea. DPhil Thesis. University of Sussex, Brighton, U.K.

Rohfritsch, O. 1971. The culture in vitro of young galls caused by Aulax glechomae L. on Glechoma hederacea L. Marcellia. 37:151-161.

Rohfritsch, O. 1974. Ultrastructure of the nutritive layer cells of the gall caused by Aulax glechomae L. on Glechoma hederacea L. Protoplasma. 81:205-230.

Rohfritsch, O. 1978. Three dimensional study of cell organelles in the nutritive tissue of a gall (Liposthebes glechomae L. on Glechoma hederacea L). Protoplasma. 95:297-307.

Rohfritsch, O. 1981. Different cell organelles observed on thick slices of tissue from Liposthebes glechomae on Glechoma hederacea. Bulletion of Society of Botany France. 128:23-25.

Siebke, K. and Weis, E. 1995. Assimilation images of leaves of Glechoma hederacea: analysis of non-synchronous stomata related oscillations. Planta. 196(1):155-165.

Siebke, K. and Weis, E. 1995. Imaging of chlorophyll-a-flourescence in leaves: topography of photosynthetic oscillations in leaves of Glechoma hederacea. Photosynthesis Research. 45:225-237.

Sinker, C.A., Packham, J.R., Treuman, I.C., Oswald, P.H., Perring, F.H., and Prestwood, W.V. 1985. Ecological Flora of the Shropshire Region. Shropshire Trust for Nature Conservation, Shrewsbury, UK.

Slade, A.J. 1986. The population biology and foraging behavior of the clonal perennial herb Glechoma hederacea. Ph.D. thesis, University of Sussex.

Slade, A.J. and Hutchings, M.J. 1987. The effects of nutrient availability on foraging in the clonal herb Glechoma hederacea. Journal of Ecology. 75:95-112.

Slade, A.J. and Hutchings, M.J. 1987. The effects of light intensity on foraging in the clonal herb Glechoma hederacea. Journal of Ecology. 75:639-650.

Steufer, J.F., During, H.J., and Kroon, H.D. 1994. High benefits of clonal integration in two stoloniferous species, in response to heterogeneous light environments. Journal of Ecology. 82:511-518.

Stubbendieck, J., Friisoe, G.Y., and Bolick, M.R. 1995. Weeds of Nebraska and The Great Plains. Nebraska Department of Agriculture. Lincoln, Nebraska. Pg. 322-323.

Timonin, A.K. and Cherkashin, A.A. 1996. The role of cell size in determination of morphogenesis of the stomata in leaves of several lamioid Labiatae. Izvestiya-Akademii-Nauk-Seriya-Biologicheskaya-Moscow. 0(6):687-697.

Webb, C.J., Sykes, W.R. and Garnock-Jones, P.J. 1988. Flora of New Zealand, Vol. IV. Naturalised Pteridophytes, Gymnosperms, Dicotyledons. DSIR, Christchurch, New Zealand.

Wijesinghe, D.K. and Hutchings, M.J. 1996. Consequences of patchy disribution of light for the growth of the clonal herb Glechoma hederacea. Oikos. 77(1):137-145.

Wijesinghe, D.K. and Hutchings, M.J. 1999. The effects of environmental heterogeneity on the performance of Glechoma hederacea: the interactions between patch contrast and patch scale. Journal of Ecology. 87:860-872.

Zennie, T.M. and Ogzenalla, C.D. 1977. Absorbic acid and vitamin A content of edible wild plants of Ohio and Kentucky. Economic Botany. 31:76-79.

Control

A.) Cultural

Eaton, F.M. 1994. Deficiency, toxicity, and accumulation of boron in plants. J. Agr. Res. 69:237-277.

Gadoury, H. and Watson, A.K. 1987. Biological control of lawn weeds. Cahier-des-Journes-Horticoles-Ornamentals (French). 3:9-15.

Gupta, U.C., Jame, Y.W., Campbell, C.A., Leyshon, A.J., and Sicholaichuk, W. 1985. Boron toxicity and deficiency: A review. Can. J. Soil Science. 65:381-409.

Hatterman - Valenti, H. and Christians, N. 1992. Ground Ivy control with borax. Ornamental and Turf Newsletter. 1(2):5.

Hatterman - Valenti, H., Owen, M.D.K., and Christians, N.E. 1996. Ground Ivy (Glechoma hederacea L.) control in a Kentucky bluegrass turfgrass with borax. Journal of Environmental Horticulture. 14(2):101-104.

Jauron, R. 1997. Borax on Ground Ivy: Boon or Bane? Horticulture and Home Pest News. Iowa State University, Ames, IA. p. 132-133.

Jennings, V.M. and Fawcett, R.S. 1977. Weed control: ground ivy (Glechoma hederacea L.). Iowa State Universtiy of Science and Technology, Ames. Cooperative Extension Service. April 1977 p. 2.

Schultz, W. 1989. The chemical free lawn. Rodale Prss, Emmaus, Pennslyvania. p. 122-125.

B.) Chemical

Eue, L., Metzger, K., and Faust, W. 1973. A new total herbicide based on thiadiazole. Compte-Rendu-de-la-7e. Conference du Columa. p. 14-21.

Feldwick, G.A. 1987. The use of the growth retardant mefluidide for maintaining acceptable grass height and for improving species diversity in a sward. British Crop Protection Council. 36:119-126.

Gange, A.C., Brown, V.K., and Farmer, L.M. 1992. Effects of pesticides on germination of weed seeds: Implications for manipulative expirements. Journal of Applied Ecology. 29:303-310.

King, J.E. 1992. Control of broadleaf weeds in cool season turfgrass with confront herbicide. Down to Earth. 47:10-13.

Meyer, J. 1972. Evaluation of a combination of methazole and simazine for weed control in orchards and vineyards. Proceedings of the 6th International Velsicol Symposium, Brighton, UK. p. 13.

Mijatovic, K. and Veljkovic, B. 1984. Study of some properties of the weed association under the conditions of the use of herbicides. Zastita-bilja (Yugoslavia). 35(170):379-387.

Olsen, D.D. and Hall, A.B. 1988. Postemergence broadleaf weed control in turf with non-phenoxy herbicide combinations. Proceedings, 42nd annual meeting of the Northeastern Weed Science Society, p. 185-186.

Rick, S.K., Gasperini, F.A., and Rainger, G.C. 1986. Broadleaf weed control in fine turf with DPX L5300 and DPX M6316. Proceedings, North Central Weed Control Conference. 41:25-26.

Rindels, S. 1994. Ground Ivy. Horticulture and Home Pest News. Iowa State University, Ames, IA. p. 48.

Ruger, H. 1991. Experiments on reducing herbicide use in fruit growing. Obstbau-Bonn. 16:346-350.

Turgeon, A.J. 1994. Susceptibility of broadleaf weeds to turf herbicides. Turf Weeds and Their Control. p. 235. Crop Science Society of America, Inc. Madison, WI.

Growth and Development

Alpert, P. and Stuefer, J.F. 1997. Division of labour in clonal plants. The Ecology and Evolution of Clonal Plants, p. 137-154. Backhuys Publishers, Leiden, the Netherlands.

Balzter, H., Braun, P.W., and Kohler, W. 1998. Cellular automata models for vegetation dynamics. Ecological Modeling. 107:2-3, 113-125.

Birch, C.P.D. and Hutchings, M.J. 1992. Analysis of ramet developement in the stoloniferous herb Glechoma hederacea using a plastochron index. Oikos. 63:387-394.

Bones, B.G. 1961. Inequality in the development of the axillary members in Glechoma hederacea L. Ann-Bot. 25:391-406.

Grime, J.P., Mason, G., Curtis, A.V., Rodman, J.,Band, S.R., Mowforth, M.A.G., Neal, A.M., and Shaw, S. 1981. A comparitive study of germination characteristics in a local flora. Journal of Ecology. 69:1017-1059.

Holdsworth, G. 1999. The effects of defoliation and rooting on the clonal growth of Glechoma hederacea. DPhil Thesis. University of Sussex, Brighton, UK.

Hutchings, M.J. and Price, E.A.C. 1999. Glechoma hederacea L. Journal of Ecology. 87(2):347-364.

Jeune, B. 1984. Position and direction of mitosis in the organogenetic zone of young leaves of Fraxinus excelsior, Glechoma hederacea, and Lycopus europaeus. Canadian Journal of Botany. 62:2861-2864.

Marcinkowska, N., Szlacheta, W., and Prabucki, A. 1998. Planting density of herbaceous perrenials. Folia Universitatis Agriculturae-Stetinensis. 70:65-71.

Marshall, C. 1990. Source-sink relations of interconnected ramets. In: van Groenendael J, de Kroon, H. (eds.) Clonal growth in plants: regulation and function. SPB Academic, the Hague, p. 23-41.

Marshall, C., Vuorisalo, T., and Hutchings, M.J. 1996. Sectoriality and physiological organization in herbaceous plants: an overview. Plant sectoriality: causes and consequences. Vegetatio. 127:9-16.

McPhee, C.S., Bonner, S.P., and Aarssen, L.W. 1997. The role of apical dominance in the interpretation of adaptive architecture in prostrate plant species. Ecoscience. 4(4):490-500.

Pelling, V.J. 1994. Factors influencing the vegetative growth of clonal perennial species in the family Lamiaceae. DPhil Thesis. University of Sussex, Brighton, UK.

Pitelka, L.F. and Ashmun, J.W. 1985. Physiology and integration of ramets in clonal plants. Population Biology and Evolution of Clonal Organisms. Yale University Press, New Haven, p. 399-437.

Price, E.A.C. and Hutchings, M.J. 1992. Studies of growth in the clonal herb Glechoma hederacea. II. The effects of selective defoliation. Journal of Ecology. 80(1):39-47.

Price, E.A.C., Marshall, C., and Hutchings, M.J. 1992. Studies of growth in the clonal herb Glechoma hederacea. I. Patterns of physiological integration. Journal of Ecology. 80(1):25-38.

Price, E.A.C., Marshall, C., and Hutchings, M.J. 1996. Causes and consequences of sectoriality in the clonal herb Glechoma hederacea. Vegetatio. 127:41-54.

Wijesinghe, D.K. and Hutchings, M.J. 1997. The effects of spatial scale of environmental heterogeneity on the growth of a clonal plant: an experimental study with Glechoma hederacea. Journal of Ecology. 85(1):17-28.

Zukowski, W., Latowski, K., Ninova, D., Popora, M., Cesmedziev, I., Uzmanov, B. and Klimko, M. 1984. Tendencies in anatomical variation of genus Glechoma as determined by population analysis. Fragmenta Floristica Geobotanica (Cracow). 28:341-362.

Interspecific Interactions

Ash, J.E. and Barkham, J.P. 1976. Changes and variability in the field of a coppiced woodland in Norfolk, England. Journal of Ecology. 64:697-712.

Cantele, A. and Zanin, G. 1983. Results of six years of weed control in a vineyard in Venezia. Compte-rendu-de-la-12e-conference-du-Columa-Tome-II. p. 281-291.

Crampton, A.B. and Stutter, O., Kirby, K.J., and Welch, R.C. 1998. Changes in the composition of Monks Wood National Reserve (Cambridgeshire, UK.) 1964-1996. Arboculture Journal. 22:229-245.

Feldwick, G.A. 1987. The use of the growth retardant mefluidide for maintaining acceptable grass height and for improving species diversity in a sward. British Crop Protection Council. 36:119-126.

Gao, G.Y., Boggs, J.F., Bennett, P.J., Martin, J.C., Chatfield, J.A., Rose, M.A., Rimelspach, J.W., Zondag, R.H., Strect, J.R., and Pomd, W.E. 1999. Weed problems in Ohio turf, landscapes, and nurseries. Ornamental Plants. (Special-Circular Ohio Agricultural Research and Development Center.) 165:55-58.

Hutchings, M.J. and Price, E.A.C. 1993. Does physiological integration enable clonal herbs to integrate the effects of environmental heterogeneity? Plant Species Biology. 8:95-105.

Leeflang, L., Marshall, C., and Price, E.A.C. 1999. Are stoloniferous plants able to avoid neighbors in response to low R:FR ratios in reflected light? Special Issue. Clonal Plants and Environmental Heterogeneity - space, time, and scale. Plant Ecology. 141:1-2, 59-65.

Mika, V., Swital, F., and Nerusil, P. 1998. Prediction of botanical composition of grasslands using spectroscopy in near infrared region (NIRS). Rostlinna-Vyroba. 44:245-249.

Mundel, G. 1993. Extensively used lowland-fen grassland in the Havell and ische Luch, Brandenburg. Archiv-fur-Naturschutz-und-Landschaftsforschung. 32:161-169.

Novak, J. 1998. Changes in grassland after additional sowing of grass mixture with dominance of valuable herbs. Rostlinna Vyroba UZPI (Czech Republic). 44(3):123-131.

Price, E.A.C. and Hutchings, M.J. 1996. The effects of competition on growth and form in Glechoma hederacea. Oikos. 75(2):279-290.

Pyle, L.L. 1995. Effects of disturbance on herbaceous exotic plant species on the flood plain of the Potomac River. American-Midland Naturalist. 134:244-253.

Rice, E.L., Putnam, A.R., and Tang, C.S. 1986. Allelopathic growth stimulation. The science of allelopathy. Jon Wiley and Sons Inc. New York, U.S.A. p. 23-42.

Slade, A.J. and Hutchings, M.J. 1989. Within and between population variation in ramet behavior in the gynodioecious clonal herb, Glechoma hederacea (Labiatae). Canadian Journal of Botany. 67:633-639.

Takahashi, Y., Saitoh, S., Otani, I., Uozumi, S., Hagino, K., and Igarashi, R. 1995. Studies on allelopathic interactions among some grassland species, 6: Screening of allelopathic activities from native grassland species by using the lettuce (Lactuca Sativa) seed bioassay with the plant shoot extracts. Journal of Japanese Society of Grassland Science. 41:232-239.

Intraspecific Interactions

Birch, C.P.D., Hutchings, M.J. 1994. Exploitation of patchily distributed soil resources by the clonal herb Glechoma hederacea. Journal of Ecology. 82(3):653-664.

Birch, C.P.D., Hutchings, M.J., Marshall, C., and Price, E.A.C. 1999. Clonal segmentation: the development of physiological independence within stolons of Glechoma hederacea L. Plant Ecology. 141:21-31.

Huber, H., Lukacs, S., Watson, M.A., Marshall, C., and Price, E.A.C. 1999. Spatial stucture of stoloniferous herbs: an interplay between structural blue print, ontogeny and phenotypic plasticity. Plant Ecology. 141:107-115.

Hutchings, M.J. and Wijesinghe, D.K. 1997. Patchy habitats, division of labor and growth dividends in clonal plants. Trends in Ecology and Evolution. 12:390-394.

Klimes, L. 1997. Variation in autumnal growth of hermaphroditic clones of Glechoma hederacea originating from two geographical regions and two habitats. Preslia. 69:175-183.

McPhee, C.S., Bonner, S.P., and Aarssen, L.W. 1997. The role of apical dominance in the interpretation of adaptive architecture in prostrate plant species. Ecoscience. 4:490-500.

Price, E.A.C. 1991. Investigations into the ecology and intraclonal physiology of Glechoma hederacea. DPhil Thesis. University of Sussex, Brighton, UK.

Price, E.A.C. and Hutchings, M.J. 1996. The effects of competition on growth and form in Glechoma hederacea. Oikos. 75(2):279-290.

Slade, A.J. and Hutchings, M.J. 1987. An analysis of the costs and benefits of physiological integration between ramets in the clonal perennial herb Glechoma hederacea. Oecologia. 73:425-431.

Slade, A.J. and Hutchings, M.J. 1989. Within and between population variation in ramet behavior in the gynodioecious clonal herb, Glechoma hederacea. Canadian Journal of Botany. 67:633-639.

Steufer, J.F. and Hutchings, M.J. 1994. Environmental heterogeniety and clonal growth: a study of the capacity for reciprocal translocation in Glechoma hederacea L. Oecologia. 100(3):302-308.

Wijesinghe, R.K. and Hutchings, M.J. 1996. Consequences of patchy distribution of light for the growth of the clonal herb Glechoma hederacea. Canadian Journal of Botany. 67:633-639.

Plasticity

Huber, H., Lukacs, S., Watson, M.A., Marshall, C., and Price, E.A.C. 1999. Spatial structure of stoloniferous herbs: an interplay between structural blue print, ontegeny and phenotypic plasticity. Plant Ecology. 141:107-115.

Hutchings, M.J. 1995. Morphological plasticity in clonal plants: the foraging concept reconsidered. Journal of Ecology. 83:143-152.

Hutchings, M.J. and de Kroon, H. 1994. Foraging in plants: the role of morphological plasticity in resource acquisition. Advances in Ecological Research. 25:159-238.

Hutchings, M.J. and Slade, A.J. 1988. Morphological plasticity, foraging and integration in clonal perennial herbs. Symposium of British Ecological Society. 28:83-109.

Hutchings, M.J. and Turkingston, R. 1995. Plasticity of branching patterns in the clonal herbs Trifolium repens L. and Glechoma hederacea L. Experimental and molecular approaches to plant biosystematics. p. 173-186. Missouri Botanical Garden, St. Louis, MO.

Price, E.A.C. and Hutchings, M.J. 1992. The causes and development effects of integration and independence between different parts of Glechoma hederacea clones. Oikos. 63:376-386.

Slade, A.J. and Hutchings, M.J. 1987. Clonal integration and plasticity in foraging behavior in Glechoma hederacea. Journal of Ecology. 75:1023-1036.

Silvertown, J. 1998. Plant phenotypic plasticity and non-cognitive behavior. Trends in Ecology and Evolution. 13:255-256.

Reproduction

Baker, H.G. 1948. Corolla size in gynodioecious and gynomonoecious species of flowering plants. Proceedings of the Leeds Philosophical and Literary Society. 5:136-139.

Birch, C.P.D. and Hutchings, M.J. 1992. Stolon growth and branching in Glechoma hederacea L.: an application of a plastochron index. New Phytologist. 122(3):545-551.

Birch, C.P.D. and Hutchings, M.J. 1999. Clonal segmentation: the development of physiological independence within stolons of Glechoma hederacea L. Plant Ecology.

Caraco, T. and Kelly, C.K. 1991. On the adaptive value of physiological integration in clonal plants. Ecology. 72:81-93.

Charlesworth, B. and Charlesworth, D. 1978. A model for the evolution of dioecy and gynodioecy. American Natural. 112:975-997.

Garrard, I. and Streeter, D.T. 1983. The Wildflowers of the British Isles. MacMillan, London, UK.

Handel, S.N. 1985. The intrusion of clonal growth patterns on plant breeding systems. American Natural. 125:367-384.

Hutchings, M.J. and Price, E.A.C. 1999. Glechoma hederacea L. Journal of Ecology. 87(2):347-364.

Lewis, D. 1941. Male-sterility in natural populations of hermaphrodite plants: The equilibrium between females and hermaphrodites to be expected with different types of inheritance. New Phytology. 40:56-63.

Lewis, D. and Crawe, L.K. 1956. The genetics and evolution of gynodioecy. Evolution (Lawrence, Kansas). 10:115-125.

Lloyd, D.G. 1974. Theoretical sex ratios of gynodioecious angiosperms. Heredity. 32:11-34.

Owens, S.J. and Ubera-Jimenez, J.L. 1992. Breeding systems in Labiatae. Advances in Labiatae Science. p. 257-280. Royal Botanical Gardens, Kew, U.K.

Slade, A.J. and Hutchings, M.J. 1987. An analysis of the influence of clone size and stolon connections between ramets on the growth of Glechoma hederacea L. New Phytologist. 106:759-771.

Southwick, E.E., Loper, G.M., and Sadwick, S.E. 1982. Nectar production, composition, energetics, and pollinator attractiveness in spring flowers of western New York, USA. American Journal of Botany. 68(7):994-1002.

Widen, B., Cronberg, N., and Widen, M. 1994. Genotypic diversity, molecular markers and spatial distribution of genets in clonal plants, a literature survey. Folia-Geobotanica-et-Phytotaxonomica. 29:245-263.

Widen, B. and Widen, M. 1990. Pollen limitation and distance dependent fecundity in females of the clonal gynodioecious herb Glechoma hederacea (Lamiaceae). Oecologia. 83:191-196.

Seed

Bouman, F. and Meeuse, A.D.J. 1992. Dispersal in Labiatae. Advances in Labiatae Science (eds R.M. Harley and T. Reynolds), p. 193-202. Royal Botanical Gardens, Kew, UK.

Brenchley, W.E. 1918. Buried weed seeds. Journal of Agricultural Science. 9:1-31.

Chancellor, R.J. 1976. Weed changes over 11 years in Wrenches, an arable field. Proceedings of the 1976 British Crop Protection Conference-Weeds, p. 681-686.

Chancellor, R.J. 1985. Changes in the weed flora of an arable field cultivated for twenty years. Journal of Applied Ecology. 22:491-501.

Ferrero, A. and Maggiore, T. 1992. Dissemination of weeds by irrigation. International Symposium on Crop Protection, Gent, Belgium. May 5, 1992. 44(4):1093-1098.

Hutchings, M.J. and Price, E.A.C. 1999. Glechoma hederacea L. Journal of Ecology. 87:347-364.

Lewis, D. and Crowe, L.K. 1956. The genetics and evolution of gynodioecy. Evolution (Lawrence, Kansas). 10:115-125.

Odum, S. 1965. Germination of ancient seeds. Dansk Botanisk Arkiv. 24:1-70.

Petrov, V.V. and Palkina, T.A. 1983. The content of dormant viable seeds in the soil of a broad leaved forest and spruce plantation. Vestnik Moskovskogo Universiteta, Biologya. 38:31-35.

Roberts, H.A. 1981. Seed banks in soils. Advances in Applied Biology. 6:1-55.

Ryding, O. 1992. The distribution and evolution of myxocarpy in Lamiaceae. Advances in Labiate Science (eds R.M. Harley and T. Reynolds). p. 85-96. Royal Botanic Gardens, Kew, UK.

Slade, A.J. and Hutchings, M.J. 1989. Within and between population variation in ramet behavior in the gynodioecious clonal herb, Glechoma hederacea (Labiatae). Canadian Journal of Botany. 67:633-639.

Stagg, P.G. 1996. Aspects of the restoration of chalk grassland vegetation on ex-arable land. DPhil Thesis. University of Sussex, Brighton, U.K.

Thompson, K., Bakker, J.P., and Bekker, R.M. 1997. The soil seed banks of Northwest Europe. Cambridge University Press, Cambridge, U.K.

Widen, B. and Widen, M. 1990. Pollen limitation and distance-dependent fecundity in females of the clonal gynodioecious herb Glechoma hederacea. Oecologia. 83:191-196.

Variability

Gill, L.S. 1979. Cyto-taxonomic studies of the tribe Nepeteae (Labiatae) in Canada. Genetica. 50:111-118.

Ninova, D.I., Cheshmedzhiev, I.V., Popova, M.T., and Kuzmanov, B.A. 1984. Variability of Glechoma hirusuta Waldst et Kit and Glechoma hederacea L. in Bulgaria. 1. Anatomical study. Fitologiia (Bulgaria). 24:3-28.

Ninova, D.I., Cheshmedzhiev, I.V., Popova, M.T., and Kuzmanov, B.A. 1984. Variabiltiy of Glechoma hirusuta Waldst et Kit and Glechoma hederacea L. in Bulgaria. 2. Anatomical investigation. Fitologiia (Bulgaria). 25:13-40.

Ninova, D.I., Kuzmanov, B.A., Ruseva, R.P., and Kurteva, M.E. 1984. Variability of Glechoma hirusuta Waldst et Kit and Glechoma hederacea L. in Bulgaria. Morphological study. Fitologiia (Bulgaria). 26:3-59.

Orlenko, M.L. 1990. Use of factor analysis for the study of ecotypic variation of Glechoma hederacea. Byulleton-Glavnogo-Botanichiskogo-Sada (Russian). 153:35-40.

Rutland, J.P. 1941. The Merton Catalogue. A list of chromosome number of British plants. New Phytologist. 40:210-214.

Stahl, E. and Datta, S.N. 1972. New sesquiterpenoids of the ground ivy (Glechoma hederacea L.) Justus-Liebigs-Ann-Chem. 757:23-32.

Zukowski, W., Latowski, K., Ninova, D., Popova, M., Cesmedzijev, I., Kuzmanov, B., and Klimko, M. 1985. Tendencies in anatomical variation of genus Glechoma as determined by population analysis. Fragmenta Floristica et Geobotanica. 28(3):341-362.

Zukowski, W. And Slowinska, T. 1981. Chromosome numbers of angiosperms of northwestern Poland. Fragmenta Floristica et Geobotanica. 25(4):477-484.

Other

A.) Foraging: Definition - to make a search; to strip of provisions

Cain, M.L. 1994. Consequences of foraging in clonal plants species. Ecology. 75(4):933-944.

Cain, M.L., Dudle, D.A., and Evans, J.P. 1996. Spatial models of foraging in clonal plant species. American Journal of Botany. 83(1):76-85.

Hutchings, M.J. 1995. Morphological plasticity in clonal plants: the foraging concept reconsidered. Journal of Ecology. 83:143-152.

Hutchings, M.J. and de Kroon, H. 1994. Foraging in plants: the role of morphological plasticity in resource acquisition. Advances in Ecological Research. 25:159-238.

Hutchings, M.J. and Slade, A.J. 1988. Morphological plasticity, foraging and integration in clonal perennial herbs. Symposium of British Ecological Society. 28:83-109.

Slade, A.J. 1986. The population biology and foraging behavior of the clonal perennial herb Glechoma hederacea. Ph.D. Thesis, University of Sussex.

Slade, A.J. and Hutchings, M.J. 1987. Clonal integration and plasticity in foraging behavior in Glechoma hederacea. Journal of Ecology. 75:1023-1036.

Slade, A.J. and Hutchings, M.J. 1987. The effects of light intensity on foraging in the clonal herb Glechoma hederacea. Journal of Ecology. 75:639-650.

Slade, A.J. and Hutchings, M.J. 1987. The effects of nutrient availability on foraging in the clonal herb Glechoma hederacea. Journal of Ecology. 75:95-112.

B.) Medicinal Uses

Bergendorff, O., Franzen, C., Jeppsson, A.B., Sterner, O., and Waldeck, B. 1995. Screening of some European medicinal plants for spasmolytic activity on isolated guinea pig trachea. International Journal of Pharmacognosy. 33:356-358.

Bergeron, J. and Jodoin, L. 1983. Effects of secondary plant compounds on body weight and the weight of certain organs in the laboratory mouse. Canadian Journal of Zoology. 60(8):1855-1866.

Franke, W. and Lawrenz, M. 1981. Contents of protein and its composition of amino acids in leaves of some medicinal and spice plants, edible as greens. Herba Hungarica. 19:71-82.

Milovanovic, M. 1995. Chemical investigation of medicinal spices and herbs. Fundamental research in food technology, Yugoslavia. p. 35-42.

Okuyama, E., Yamazaki, M., and Ishii, Y. 1984. Isolation and identification of ursolic acid-related compounds as the principles of Glechoma hederacea having an antiulcerogenic activity. Shoyakugaku Zasshi. 37:52-55.

Sagimoto, K., Sakuri, N., Shirasawa, H., Fujise,Y., Shibata, K., Shimada, K., and Sukata, J. 1992. Bovine cases of urolithiasis treated with traditional herbal medicine, p-3. Journal of Veterinary Medicinal Science. 54:579-582.

Severnet, T. 1991. Looking for new drugs: what criteria. Journal of Ethno Pharmacology. 32:83-90.

Tokuda, A., Ohigashi, H., Koshimizu, K., and Ito, Y. 1987. Inhibitory effects of ursolic and oleanolic acid on skin tumor promotion by 12-O-tetradecanoylphorbol-13-acetate. Cancer Letters. 33(3):279-286.

Zieba, J. 1973. Isolation and identification of flavonoids from Glechoma hederacea L. Polish Journal of Pharmacology and Pharmacy. 25:593-597.

Zieba, J. 1973. Isolation and identification of non-heterside triterpenoids from Glechoma hederacea L. Polish Journal of Pharmacology and Pharmacy. 25:593-597.

C.) Etc.

Boer, R. 1981. Genetic affinities between spider mite Tetranychus urticae (Acarina: Tetranychidae) populations in a non-agricultural area. Glass house plants, Glechoma hederacea in dunes. Entomologia Experimentalis Applicata (Netherlands). 28:22-28.

Fisher, C. 1995. Horse care: perilous pasture plants. Rural Heritage. 20(2):44-45.

Grieve, M. 1976. A Modern Herbal. Peregrine Books, Harmondsworth, UK.

Henry, D.Y., Gueritte-Voegelein, F., Insel, P.A.,Ferry, N., Bouguet, J., Potier, P., Sevenet, T., and Hanoune, J. 1987. Isolation and characterization of 9-hydroxy-10-trans, 12-cis-octadecadienoic acid, a novel regulator of platelet adenylate cyclase from Glechoma hederacea L. European Journal of Biochemistry. 170:389-394.

Hessayon, D.G. 1987. The Gold Plated House Plant Expert. Century, London, UK.

Hornig, R. and Buenemann, G. 1996. Ground covers and fertigation in IP apple orchards: III. Biomass yield and nutrient uptake of the ground covers and influence on soil moisture and nutrients in the soil. Gartenbauwissenschaft. 61:164-173.

Hyde, M. 1976. Hedgerow Plants. Shire Publications, Aylesbury, UK.

Iwasuki, A. 1997. Notes on Pytomyza leaf miners (Diptera: Agromyzidae) on Lamiaceae in Japan. Japanese Journal of Entomology. 65:500-501.

Komprda, T., Stohandlova, M., Foltyn, J., Pozdisek, J., and Mika, V. 1999. Content of p-coumaric and ferulic acid in forbs with potential grazing utilization. Archives of Animal Nutrition. 52:95-105.

Kuhn, H., Wiesner, R., Alder, L., and Schewe, T. 1989. Occurence of free and esterified lipoxygenase products in leaves of Glechoma hederacea L. and other Labiatae. European Journal of Biochemistry. 186:155-162.

Milovanovic, M., Picuric-Jovanovic, K., and Vrbaski, Z. 1995. Flavonoids antioxidant activities from the plant species Glechoma hederacea. Modern Trends in Food Technology, (Proceedings from the 2nd Yugoslav symposium of food technologists, Belgrade, Yugoslavia, Feb. 9-10, 1995). Pg. 219-225.

Potterton, D. 1985. Culpepper’s Colour Herbal. Guild Publishing, London, UK.

Reimherr, P., Hanke, H., Linse, H., Rost, J., Dotterweich, B., and Schobert, W. 1996. An endogenous bedding plant species, Glechoma hederacea ‘Variegata’. Gaertnerboerse, Germany. 96(3):112.

Tucker, K., Sage, R.B., Buckley, G.P. 1997. Introducing other plants into short rotation coppice willow. Biomass and Energy Crops. Aspects of Applied Biology. 49:293-299.