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Abstracts
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Londre, R.A. and S.A. Schnitzer (in press). The distribution of lianas and their change in abundance in temperate deciduous forests over the past 45 years. Ecology.

Abstract: Lianas (woody vines) are an important and dynamic component of many forests throughout the world and increases in CO2, mean winter temperature, and forest fragmentation may promote their growth and proliferation in temperate forests.  In this study, we used a 45-year dataset to test the hypothesis that lianas have increased in abundance and basal area in the interiors of 14 deciduous temperate forests in Wisconsin since 1959.  We also censused woody plants along a gradient from the forest edge to the interior in seven of these forests to test the hypothesis that the abundance of lianas declines significantly with increasing distance from the forest edge.  We found that lianas did not increase in abundance within the interiors of temperate forests in Wisconsin over the last 45 years.  However, relative and absolute liana abundance decreased sharply with increasing distance from forest edges.  Our findings suggest that forest fragmentation, not climate change, may be increasing the abundance of lianas in northern deciduous temperate forests, and that lianas may further increase in abundance if the severity of forest fragmentation intensifies. 

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Schnitzer, S.A., S.J. DeWalt, and J. Chave (2006). Censusing and measuring lianas: a quantitative comparison of the common methods. Biotropica, 38: xxx-xxx.

Abstract: Lianas contribute to many aspects of tropical forest diversity and dynamics, and interest in liana ecology has grown substantially in recent years.  Methods to census lianas and estimate biomass, however, differ among studies, possibly hindering attempts to compare liana communities.  At Nouragues Research Station (French Guiana), we tested the extent to which liana abundance, basal area, and estimated biomass differed depending on stem diameter measurement location, inclusion of ramets, inclusion of lianas rooted within versus passing through the plot, and plot shape.  We found that the mean per-plot abundance and basal area of lianas were significantly greater when lianas were measured low on the stem, when ramets were included, and when lianas were sampled in transects (2x50m) than square plots (10x10m).  Mean per-plot liana abundance and basal area were 21% and 58% greater, when stems were measured at the largest spot on the stem compared to 130cm from the ground, respectively.  Including liana ramets increased average per plot liana abundance, basal area, and estimated biomass by 19%, 17%, and 16%, respectively.  To facilitate cross-study comparisons, we developed conversion equations that equate liana abundance, diameter, and basal area based on the measurements taken at four different stem locations.  We tested these equations at Lambir Hills National Park, Malaysia and found that they did not differ significantly between the two sites, suggesting that the equations may be broadly applicable.  Finally, we present a new allometric equation relating diameter and biomass developed from 424 lianas from five independent datasets collected in four countries.

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Kurzel, B.P., S.A. Schnitzer, and W.P. Carson (2006). The relationship between liana diameter and location in the canopy in three Panamanian forests. Biotropica, 38: 262-266.

Abstract: In three forests that differed in annual rainfall and seasonality, the probability of a liana ≥ 2.0 cm stem diameter reaching the canopy was > 50%.  Although lianas reached the canopy at significantly smaller size-classes (1.5 cm) in the wet aseasonal forest, because lianas ≥ 2.0 cm diameter were consistently in the canopy in all three forests, we suggest that 2.0 cm is the minimum stem diameter to examine the abundance and diversity of canopy lianas or canopy competition between lianas and trees.

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Gerwing, J.J., S.A. Schnitzer, R.J. Burnham, F. Bongers, J. Chave, S.J. DeWalt, C.E.N. Ewango, R. Foster, M. Martinez-Ramos, M. Parren, N. Parthasarathy, D.R. Perez-Salicrup, F.E. Putz and D.W. Thomas (2006).  Censusing lianas in tropical forests. Biotropica, 38: 256-261.

Abstract: A recent increase in published studies of lianas has been paralleled by a proliferation of protocols for censusing lianas.  This article seeks to increase uniformity in liana inventories by providing specific recommendations for the determination of which taxa to include; the location of diameter measurement points on individual stems, the setting of minimum stem diameter cut-offs, the treatment of multiple-stemmed and rooted clonal groups, and the measurement of non-cylindrical stems.   Use of more uniform liana censusing protocols may facilitate comparison of independently collected datasets and further our understanding of global patterns in liana abundance, diversity, biomass, and dynamics.

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Schnitzer, S.A. (2005).  A mechanistic explanation for the global patterns of liana abundance and distribution. American Naturalist, 166: 262-276.

Abstract: One of the main goals in ecology is determining the mechanisms that control the abundance and distribution of organisms.  Using data from 69 tropical forests worldwide, I demonstrate that liana (woody vine) abundance is correlated negatively with mean annual precipitation and positively with seasonality, a pattern precisely the opposite of most other plant types.  I propose a general mechanistic hypothesis integrating both ecological and ecophysiological approaches to explain this pattern.  Specifically, the deep root and efficient vascular systems of lianas enable them to suffer less water stress during seasonal droughts while many competitors are dormant, giving lianas a competitive advantage during the dry season.  Testing this hypothesis in central Panama, I found that lianas grew approximately seven-times more in height than did trees during the dry season, but only twice as much during the wet season.  Over time, this dry season advantage may allow lianas to increase in abundance in seasonal forests.  In aseasonal wet forests, however, lianas gain no such advantage because competing plants are rarely limited by water.  I extend this theory to account for the local, within-forest increase in liana abundance in response to disturbance, as well as the conspicuous decrease in liana abundance at high latitudes.

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Andrade, J.L., F.C. Meinzer, G. Goldstein, and S.A. Schnitzer (2005).  Water uptake and transport in lianas and co-occurring trees of a seasonally dry tropical forest. Trees: Structure & Function, 19: 282-289.

Abstract:  Water uptake and transport were studied in eight liana species in a seasonally dry tropical forest on Barro Colorado Island, Panama. Stable hydrogen isotope composition (dD) of xylem and soil water, soil volumetric water content (qv), and basal sap flow were measured during the 1997 and 1998 dry seasons. Sap flow of several neighboring trees was measured to assess differences between lianas and trees in magnitudes and patterns of daily sap flow. Little seasonal change in qv was observed at 90 to 120 cm depth in both years. Mean soil water dD during the dry season was -19‰ at 0 to 30 cm, -34‰ at 30 to 60 cm, and -50‰ at 90 to 120 cm. Average values of xylem dD among the liana species ranged from  –28‰ to –44‰ during the middle of the dry season, suggesting that water uptake was restricted to intermediate soil layers (30 to 60 cm). By the end of the dry season, all species exhibited more negative xylem dD values (–41‰ to –62‰), suggesting that they shifted to deeper water sources. Maximum sap flux density in co-occurring lianas and trees were comparable at similar stem diameter (DBH). Furthermore, lianas and trees conformed to the same linear relationship between daily sap flow and DBH. Our observations that lianas tap shallow sources of soil water at the beggining of the dry season and that sap flow is similar in lianas and trees of equivalent stem diameter do not support the common assumptions that lianas rely primarily on deep soil water and that they have higher rates of sap flow than co-occurring trees of similar stem size.

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Schnitzer, S.A., M. Kuzee, and F. Bongers (2005).  Disentangling above- and below-ground competition between lianas and trees in a tropical forest. Journal of Ecology, 93: 1115-1125.

Summary: 
1. Light is thought to be the most limiting resource in tropical forests, and thus above-ground competition is commonly accepted as the mechanism that structures these communities.  In many tropical forests, trees compete not only with other trees, but also with lianas, which compete aggressively for below-ground resources and thus may limit tree growth and regeneration.

2. Using a replicated experiment, we tested the relative strengths of above- and below-ground competition from lianas on tree saplings in a disturbed forest in Côte d'Ivoire with a heterogeneous canopy and relatively high light penetration.  We planted seedlings of three tree species and subjected them to below-ground competition with lianas (BGC), above- and below-ground competition with lianas (ABGC), or a liana-free control treatment.  After two years, we harvested the saplings and compared the amount of above-ground biomass and its relative allocation among the three experimental treatments and different tree species. 

3. Lianas competed intensely with saplings in this tropical forest, substantially limiting sapling growth.  Saplings grown in the ABGC and BGC treatments had only 18.5% and 16.8% of the above-ground dry biomass of those grown in the liana-free control treatment.

4. Sapling biomass did not differ significantly among the ABGC and BGC treatments, suggesting that below-ground competition was the driving force behind liana versus tree competition in this forest.  Above-ground competition with lianas, however, did affect the allocation of biomass in saplings, resulting in shorter, thicker stems and a poorly developed crown. 

5. Collectively, our findings suggest that below-ground competition with lianas plays a substantial role in limiting the growth of saplings in disturbed and secondary tropical forests, while above-ground effects may have been due to a combination of both above-ground competition and mechanical stress from lianas. 

6. Disentangling above- and below-ground competition between lianas and trees is critical for a comprehensive understanding of the dynamics of naturally regenerating tropical forests, as well as formulating successful management plans for sustainable timber harvest.

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Dickie, I.A., Schnitzer, S.A., P.B. Reich, S.E. Hobbie (2005). Spatially disjunct effects of co-occurring competition and facilitation. Ecology Letters.
 Abstract: Little is known of the co-occurrence and implications of competitive and facilitative interactions within sites. Here we show spatially disjunct competition and facilitation at forest edges, with beneficial influence of trees on seedling growth via increased ectomycorrhizal infection apparent from 12 to 20 m while closer to trees seedling growth is negatively correlated with canopy closure. As a result, seedling growth is maximized at intermediate distances. Facilitative interactions were non-linear: being within 15.7 m of a tree facilitated seedling mycorrhizal infection; while competitive effects were presumably related to canopy closure, which was related to distance and generally scales with density. These patterns result in a positive correlation of tree density and seedling growth at low densities of trees, and negative correlation due to competition at higher densities. A spatial model suggests plant communities are a mosaic of positive and negative interactions, which may contribute to population homeostasis and plant diversity.
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Schnitzer, S.A., M.P.E. Parren, and F. Bongers (2004).  Recruitment of lianas into logging gaps and the effects of pre-harvest liana cutting in a lowland forest in Cameroon. Forest Ecology and Management, 190: 87 - 98.

Abstract: The abundance of lianas (woody vines) and the detrimental impact that they have on tropical rain forest trees is widely recognized.  Lianas are particularly abundant in disturbed areas of the forest, such as logging gaps, and pre-harvest liana cutting has been widely recommended throughout the tropics to reduce the impact of lianas during and following tree harvest.  The effectiveness of forest-wide liana cutting, however, is currently unresolved, particularly for reducing liana abundance in logging gaps.  Furthermore, our understanding of the dynamics and rate of liana colonization in gaps is limited.  We tested: 1) the speed at which lianas recruit into logging gaps and their dynamics afterwards; and 2) whether pre-harvest liana cutting actually reduces the abundance of lianas in post-harvest logging gaps.  To test hypothesis 1, we compared liana recruitment in new, one and six-year-old logging gaps.  For hypothesis 2, we compared liana abundance and tree infestation by lianas in one-year-old logging gaps in which all lianas had been cut nine months prior to tree felling, versus one-year old logging gaps in which lianas were not cut.  Lianas recruited heavily into logging gaps within one year, mostly by means of stem sprouts, and many of these new stems were apparently able to persist for longer than six years.  Lianas were significantly more abundant in the root/bole zone of gaps than in the canopy zone, mostly due to the vigorous regeneration of stem sprouts.  Canopy openness was highest in gaps one year after logging, possibly due to the smothering effect of the lianas on developing trees.  Although liana abundance increased significantly over the six-year gap chronosequence, direct liana infestation of trees remained the same.  Pre-logging liana cutting, however, significantly reduced the number of lianas and also the number of liana-infested trees in logging gaps.  Consequently, liana cutting appears to be an effective method to reduce the abundance of lianas and thus minimize their detrimental effects on regenerating trees in logging gaps.

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Mascaro, J., S.A. Schnitzer, and W.P. Carson (2004).  Liana diversity, abundance, and mortality in a tropical wet forest in Costa Rica.  Forest Ecology and Management, 190: 3 - 14.

Abstract: Lianas can have a large impact on the diversity, structure, and dynamics of tropical forests, yet they remain essentially unknown even in some of the most intensely studied tropical forests, such as La Selva Biological Station in Costa Rica.  We quantified the diversity, abundance, and mortality of lianas in primary and selectively logged forest at La Selva for over three years, from January 1999 until July 2002.  We measured, identified, permanently marked, and mapped all lianas ≥ 1.3 m in length and 2 mm in diameter, whether climbing or free-standing, in nine, 24 x 36 m (864 m2) plots.  There were no significant differences in density, diversity, or mortality between primary forest and areas that were selectively logged approximately 50 years prior to our study.  We found a mean density of 1493 lianas ha-1 and a mean species richness of 23 species per 864 m2 plot.  Annual mortality was 9.4% over all size classes, but was highest for the smallest individuals (< 2 cm in diameter).  Annual mortality for larger individuals (≥ 5 cm) was much lower over the 3.5 year period (3.2% per year) and the five most abundant species suffered no mortality in this size class.  In contrast to many lowland neotropical forests, where Bignoniaceae and Fabaceae are reported to be the dominant liana families, at La Selva we found that Sapindaceae was the most speciose family and Dilleniaceae the most abundant.  Moutabea aculeata (Polygalaceae) was the most abundant species, constituting approximately 17% of the individuals and having the lowest mortality of all 60 species.  The ten most abundant species at La Selva accounted for more than 60% of all individuals.  Compared to other lowland sites in the neotropics, including other wet forests, the abundance and diversity of lianas at La Selva are very low.

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Holmgren, M. and S.A. Schnitzer (2004).  Science on the rise in developing countries. PLoS Biology, 2: 10-13.

Abstract: The disparity between scientific output in the developed and undeveloped regions is dramatic, with developing countries contributing just 12% of the publications worldwide.  Here we demonstrate that this grim picture may be improving.  We studied the dynamics of scientific publication since 1990 using the Americas as a case study.  Our analyses revealed that the relative increase in the number of publications has been far greater in Latin America than in North America, especially when corrected for the investment in research and development.  Although the number of publications from the developing world still lags behind those of the developed world, our findings suggest that this disparity is shrinking.

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Schnitzer, S.A., M.P.E. Parren, and F. Bongers (2004).  Recruitment of lianas into logging gaps and the effects of pre-harvest liana cutting.  Forest Ecology and Management, 190: 87-98.

Abstract:  The abundance of lianas (woody vines) and the detrimental impact that they have on tropical rain forest trees is widely recognized.  Lianas are particularly abundant in disturbed areas of the forest, such as logging gaps, and pre-harvest liana cutting has been widely recommended throughout the tropics to reduce the impact of lianas during and following tree harvest.  The effectiveness of forest-wide liana cutting, however, is currently unresolved, particularly for reducing liana abundance in logging gaps.  Furthermore, our understanding of the dynamics and rate of liana colonization in gaps is limited.  We tested: 1) the speed at which lianas recruit into logging gaps and their dynamics afterwards; and 2) whether pre-harvest liana cutting actually reduces the abundance of lianas in post-harvest logging gaps.  To test hypothesis 1, we compared liana recruitment in new, one and six-year-old logging gaps.  For hypothesis 2, we compared liana abundance and tree infestation by lianas in one-year-old logging gaps in which all lianas had been cut nine months prior to tree felling, versus one-year old logging gaps in which lianas were not cut.  Lianas recruited heavily into logging gaps within one year, mostly by means of stem sprouts, and many of these new stems were apparently able to persist for longer than six years.  Lianas were significantly more abundant in the root/bole zone of gaps than in the canopy zone, mostly due to the vigorous regeneration of stem sprouts.  Canopy openness was highest in gaps one year after logging, possibly due to the smothering effect of the lianas on developing trees.  Although liana abundance increased significantly over the six-year gap chronosequence, direct liana infestation of trees remained the same.  Pre-logging liana cutting, however, significantly reduced the number of lianas and also the number of liana-infested trees in logging gaps.  Consequently, liana cutting appears to be an effective method to reduce the abundance of lianas and thus minimize their detrimental effects on regenerating trees in logging gaps.

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Mascaro, J., S.A. Schnitzer, and W.P. Carson (2004).  Liana diversity, abundance, and mortality in a tropical wet forest in Costa Rica.  Forest Ecology and Management, 190: 3-15.

Abstract:  Lianas can have a large impact on the diversity, structure, and dynamics of tropical forests, yet they remain essentially unknown even in some of the most intensely studied tropical forests, such as La Selva Biological Station in Costa Rica.  We quantified the diversity, abundance, and mortality of lianas in primary and selectively logged forest at La Selva for over three years, from January 1999 until July 2002.  We measured, identified, permanently marked, and mapped all lianas * 1.3 m in length and 2 mm in diameter, whether climbing or free-standing, in nine, 24 x 36 m (864 m2) plots.  There were no significant differences in density, diversity, or mortality between primary forest and areas that were selectively logged approximately 50 years prior to our study.  We found a mean density of 1493 lianas ha-1 and a mean species richness of 23 species per 864 m2 plot.  Annual mortality was 9.4% over all size classes, but was highest for the smallest individuals (< 2 cm in diameter).  Annual mortality for larger individuals (* 5 cm) was much lower over the 3.5 year period (3.2% per year) and the five most abundant species suffered no mortality in this size class.  In contrast to many lowland neotropical forests, where Bignoniaceae and Fabaceae are reported to be the dominant liana families, at La Selva we found that Sapindaceae was the most speciose family and Dilleniaceae the most abundant.  Moutabea aculeata (Polygalaceae) was the most abundant species, constituting approximately 17% of the individuals and having the lowest mortality of all 60 species.  The ten most abundant species at La Selva accounted for more than 60% of all individuals.  Compared to other lowland sites in the neotropics, including other wet forests, the abundance and diversity of lianas at La Selva are very low.

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Schnitzer, S.A. and F. Bongers (2002).  The ecology of lianas and their role in forests. Trends in Ecology and Evolution, 17: 223-230.

Abstract:  Recent studies have demonstrated the increasingly important role of lianas (woody vines) in forest regeneration, species diversity and ecosystem-level processes, particularly in the tropics. Mechanisms responsible for the maintenance of liana species diversity could yield new insights into the maintenance of overall species diversity. Lianas contribute to forest regeneration and competition, not only by competing directly with trees, but also by differentially affecting tree species and thus changing how trees compete among themselves. In addition, they contribute considerably to ecosystem-level processes, such as whole-forest transpiration and carbon sequestration. As the rate of tropical forest disturbance increases, they are likely to increase in relative abundance throughout the tropics and the importance of lianas to many aspects of forest dynamics will grow.

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Schnitzer, S.A., P.B. Reich, B. Bergner, and W.P. Carson (2002).  Herbivore and pathogen damage on grassland and woodland plants: A test of the Herbivore Uncertainty Principle. Ecology Letters, 5: 531-539.

Abstract:  Researchers can alter the behaviour and ecology of their study organisms by conducting such seemingly benign activities as non-destructive measurements and observations. In plant communities, researcher visitation and measurement of plants may increase herbivore damage in some plant species while decreasing it in others. Simply measuring plants could change their competitive ability by altering the amount of herbivore damage that they suffer. Currently, however, there is only limited empirical evidence to support this ‘herbivore uncertainty principle’ (HUP). We tested the HUP by quantifying the amount of herbivore and pathogen damage in 13 plant species (> 1400 individuals) at four different visitation intensities at Cedar Creek Natural History Area, Minnesota, USA. Altogether, we found very little evidence to support the HUP at any intensity of visitation. Researcher visitation did not alter overall plant herbivore damage or survival and we did not detect a significant visitation effect in any of the 13 species. Pathogen damage also did not significantly vary among visitation treatments, although there was some evidence that high visitation caused slightly higher pathogen damage. Based on our results, we question whether this phenomenon should be considered a ‘principle’ of plant ecology.

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Schnitzer, S.A. and W.P. Carson (2001).  Treefall gaps and the maintenance of species diversity in a tropical forest. Ecology, 82: 913 - 919.

Abstract:  The maintenance of species diversity by treefall gaps is a long-standing paradigm in forest ecology. Gaps are presumed to provide an environment in which tree species of differing competitive abilities partition heterogeneous resources. The empirical evidence to support this paradigm, however, remains scarce, and some recent studies even suggest that gaps do not maintain the diversity of shade-tolerant species. Although there is evidence that gaps maintain the diversity of pioneer trees, most of this evidence comes from studies that did not make comparisons between gaps and intact forest sites (controls). Further, nearly all studies on the maintenance of diversity by gaps have ignored lianas, an important component of both old-world and neotropical forests. We tested the hypothesis that treefall gaps maintain shade-tolerant tree, pioneer tree, and liana species diversity in an old-growth forest on Barro Colorado Island (BCI), Panama. We compared the density and species richness of these guilds between paired gap and non-gap sites on both a per-area and a per-individual (per capita) basis. We found no difference in shade-tolerant tree density and species richness between the gap and non-gap sites. Both pioneer tree and liana density and species richness, however, were significantly higher in the gap than in the non-gap sites on both a per-area and a per-individual basis. These results suggest that gaps maintain liana species diversity and that this effect is not merely a consequence of increased density. Furthermore, our data confirm the long-held belief that gaps maintain pioneer tree species diversity. Because lianas and pioneer trees combined account for ~43% of the woody plant species on BCI, and in other forests, our results are likely to be broadly applicable and suggest that gaps play a strong role in the maintenance of woody species diversity.

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Schnitzer, S.A., J.W. Dalling and W.P. Carson (2000).  The impact of lianas on tree regeneration in tropical forest canopy gaps: Evidence for an alternative pathway of gap-phase regeneration. Journal of Ecology, 88: 655 - 666.

Summary:
1. Regeneration in forest canopy gaps is thought to lead invariably to the rapid recruitment and growth of trees and the redevelopment of the canopy. Our observations, however, suggest that an alternate successional pathway is also likely, whereby gap-phase regeneration is dominated by lianas and stalled in a low-canopy state for many years. We investigated gap-phase regeneration in an old-growth tropical forest on Barro Colorado Island (BCI) in Panama to test the following two hypotheses: (i) many gaps follow a pathway in which they remain at a low canopy height and are dominated by lianas and (ii) the paucity of trees in this pathway is a function of liana density.

2. We surveyed a total of 428 gaps of varying ages (c. 5, c. 10, and 13 years old) and identi®ed those which followed the conventional pathway of regeneration and others that remained stalled in a low-canopy state for many years and were dominated by either lianas or palms. Each of these pathways will likely have different successional trajectories that will favour the growth of a distinct suite of mature species and ultimately result in contrasting species composition.

3. The successional pathway of liana-dominated, stalled gaps is common throughout the forest. We estimate conservatively that 7.5% of the gaps that form each year will follow this pathway, probably due to the suppression of tree regeneration by lianas, and that many of these stalled gaps will persist for much longer than 13 years. Consequently, a high proportion of gaps in the forest at any given time will be stalled. Furthermore, liana tangles, which persist in the tropical forest understorey for extended periods of time, almost certainly originate in these gaps.

4. Liana abundance was positively correlated with pioneer tree abundance and diversity while negatively correlated with non-pioneer tree abundance and diversity. Thus, lianas appear to inhibit non-pioneer tree survival while indirectly enhancing that of pioneer trees.

5. Lianas are abundant in many types of tropical and temperate forests and a successional pathway involving liana-dominated, stalled gaps may therefore be frequent and widespread.

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DeWalt, S.J., S.A. Schnitzer and J.S. Denslow (2000).  Density and diversity of lianas along a seasonal tropical forest chronosequence in central Panama. Journal of Tropical Ecology, 16: 1 - 19.

Abstract:  The abundance and diversity of lianas were examined along a tropical forest chronosequence at the Barro Colorado Nature Monument, Panama. Lianas M 0.5 cm diameter were sampled along transects in two replicated stands in second-ary (20, 40, 70 and 100 y after abandonment) and old-growth (>500 y) forests. Ordination of stands based on relative abundance, but not presence-absence, showed a significant separation of stands by age. Lianas were signi®cantly more abundant and diverse (Fisher's a) in younger forests (20 and 40 y) than in older forests (70 and 100 y, and old-growth). The decline in liana abundance with stand age was offset by increased mean basal area per individual, resulting in a relatively constant total basal area and estimated biomass across stand age. The proportions of tendril climbers decreased and stem twiners increased over stand age. Decline in liana abundance and changes in liana composition may be related to changes in support and light availability. Although lianas are recognized as playing an important role in the early secondary succession of many tropical forests, these results have shown that their important contribution to total basal area and biomass can continue as the forest matures, even as the number of established lianas declines.

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Last modified: April 11, 2003