. Much confusion exists in the English-language literature on plant invasions concerning the terms ‘naturalized’ and ‘invasive’ and their associated concepts. Several authors have used these terms in proposing schemes for conceptualizing the sequence of events from introduction to invasion, but often imprecisely, erroneously or in contradictory ways. This greatly complicates the formulation of robust generalizations in invasion ecology.

Based on an extensive and critical survey of the literature we defined a minimum set of key terms related to a graphic scheme which conceptualizes the naturalization/invasion process. Introduction means that the plant (or its propagule) has been transported by humans across a major geographical barrier. Naturalization starts when abiotic and biotic barriers to survival are surmounted and when various barriers to regular reproduction are overcome. Invasion further requires that introduced plants produce reproductive offspring in areas distant from sites of introduction (approximate scales: > 100 m over  6 m/3 years for taxa spreading by roots, rhizomes, stolons or creeping stems). Taxa that can cope with the abiotic environment and biota in the general area may invade disturbed, seminatural communities. Invasion of successionally mature, undisturbed communities usually requires that the alien taxon overcomes a different category of barriers.

We propose that the term ‘invasive’ should be used without any inference to environmental or economic impact. Terms like ‘pests’ and ‘weeds’ are suitable labels for the 50–80% of invaders that have harmful effects. About 10% of invasive plants that change the character, condition, form, or nature of ecosystems over substantial areas may be termed ‘transformers’.

/pdf/naturalization-and-invasion-of-alien-plants-concepts-and-43idu8swf7.pdf

Naturalization and invasion of alien plants: concepts and definitions

Summary
1 The invasion of habitats by non-native plant and animal species is a global phenomenon with potentially grave consequences for ecological, economic, and social systems. Unfortunately, to date, the study of invasions has been primarily anecdotal and resistant to generalization.

2 Here, we use insights from experiments and from long-term monitoring studies of vegetation to propose a new theory in which fluctuation in resource availability is identified as the key factor controlling invasibility, the susceptibility of an environment to invasion by non-resident species. The theory is mechanistic and quantitative in nature leading to a variety of testable predictions.

3 We conclude that the elusive nature of the invasion process arises from the fact that it depends upon conditions of resource enrichment or release that have a variety of causes but which occur only intermittently and, to result in invasion, must coincide with availability of invading propagules.

https://besjournals.onlinelibrary.wiley.com/doi/epdf/10.1046/j.1365-2745.2000.00473.x

Fluctuating resources in plant communities: a general theory of invasibility

What determines the number and size of the seeds produced by a plant? How often should it reproduce them? How often should a plant produce them? Why and how are seeds dispersed, and what are the implications for the diversity and composition of vegetation? These are just some of the questions tackled in this wide-ranging review of the role of seeds in the ecology of plants. The authors bring together information on the ecological aspects of seed biology, starting with a consideration of reproductive strategies in seed plants and progressing through the life cycle, covering seed maturation, dispersal, storage in the soil, dormancy, germination, seedling establishment, and regeneration in the field. The text encompasses a wide range of concepts of general relevance to plant ecology, reflecting the central role that the study of seed ecology has played in elucidating many fundamental aspects of plant community function.

The ecology of seeds

Summary 1. Chronosequences and associated space-for-time substitutions are an important and often necessary tool for studying temporal dynamics of plant communities and soil development across multiple time-scales. However, they are often used inappropriately, leading to false conclusions about ecological patterns and processes, which has prompted recent strong criticism of the approach. Here, we evaluate when chronosequences may or may not be appropriate for studying community and ecosystem development. 2. Chronosequences are appropriate to study plant succession at decadal to millennial time-scales when there is evidence that sites of different ages are following the same trajectory. They can also be reliably used to study aspects of soil development that occur between temporally linked sites over time-scales of centuries to millennia, sometimes independently of their application to shorter-term plant and soil biological communities. 3. Some characteristics of changing plant and soil biological communities (e.g. species richness, plant cover, vegetation structure, soil organic matter accumulation) are more likely to be related in a predictable and temporally linear manner than are other characteristics (e.g. species composition and abundance) and are therefore more reliably studied using a chronosequence approach. 4. Chronosequences are most appropriate for studying communities that are following convergent successional trajectories and have low biodiversity, rapid species turnover and low frequency and severity of disturbance. Chronosequences are least suitable for studying successional trajectories that are divergent, species-rich, highly disturbed or arrested in time because then there are often major difficulties in determining temporal linkages between stages. 5. Synthesis. We conclude that, when successional trajectories exceed the life span of investigators and the experimental and observational studies that they perform, temporal change can be successfully explored through the judicious use of chronosequences.

/pdf/the-use-of-chronosequences-in-studies-of-ecological-1c5s5077we.pdf

The use of chronosequences in studies of ecological succession and soil development

Natural disturbances such as lava flows, landslides and glacial moraines, and human-damaged sites such as pavement, road edges and mine wastes often leave little or no soil or biological legacy. This 2003 book provided the first comprehensive summary of how plant, animal and microbial communities develop under the harsh conditions following such dramatic disturbances. The authors examine the basic principles that determine ecosystem development and apply the general rules to the urgent practical need for promoting the reclamation of damaged lands. Written for ecologists concerned with disturbance, landscape dynamics, restoration, life histories, invasions, modeling, soil formation and community or population dynamics, this book will also serve as an authoritative text for graduate students and a valuable reference for professionals involved in land management.

/pdf/primary-succession-and-ecosystem-rehabilitation-46xfnlnrj7.pdf

Primary Succession and Ecosystem Rehabilitation

Landslides are common in mountainous regions of the Caribbean and are triggered by heavy rains and earthquakes, and often occur in association with human disturbances (e.g., roads). Spatially heterogeneous removal of both substrate and vegetation is responsible for a variety of patterns of ecosystem development and plant successional trajectories within Caribbean landslides. Soil nutrient pools in exposed mineral soils reach levels comparable to mature forest soils within 55 yr but soil organic matter recovers more slowly. Plant colonization of landslides depends on the availability of propagules and suitable sites for germination, soil stability, and the presence of residual or newly deposited soil organic matter and associated nutrients. Once initial colonization occurs, the rate and trajectory of plant succession on landslides is strongly affected by plant/plant interactions. We present two conceptual models of landslide succession that summarize the major processes and pathways of ecosystem development and plant succession on landslides. Additional work is needed to characterize interactions between spatially heterogeneous zones, controls over soil development, impacts of key plant species, and the role of animals on Caribbean landslides.

https://www.jstor.org/stable/info/2389097

Ecosystem development and plant succession on landslides in the caribbean

In this review I present results from pattern analysis and field experiments in New Jersey, United States concerning tree invasion and establishment in old fields. Hutcheson Memorial Forest Center (HMF) is the New Jersey study site and a logical choice to anchor a review of this topic. HMF contains the longest and largest old field permanent plot study in the USA including extensive data concerning tree invasion and establishment. HMF has also been the site of considerable old field tree experimental research. In addition, I review other investigations of old field tree invasion and establishment of the eastern deciduous biome and contrast them with HMF. I identify critical questions concerning tree invasion and establishment, indicate what progress has been made towards answering these questions and propose future experimentation. I also synthesize results into a conceptual model and investigate the relative importance of seed and seedling processes in determining tree dynamics in old fields.

Tree invasion and establishment in old fields at Hutcheson Memorial Forest

Field experiments were conducted to investigate the effect of light, moisture, temperature, and litter on the regeneration ot two early-, one mid-, and two late-successional tropical tree species. High light and litter seem to be universally good cues for regeneration, increasing seed/seedling survival for all species except for Cecropia (an early-successional species) whose small seeds may not be able to penetrate the litter layer. In addition, the high temperature environment in both artificially shaded and nonshaded area of a natural gap exhibits less seed loss, an increase in the percent and rate of germination, and an increase in seedling survival for Darcryodes (a late-successional species), than the lower temperature environment under an intact canopy. Low soil water is also a good cue for Dacryodes germination as it is for Prestoea and Cecropia. Finally, the lower temperature environment found under the forest canopy (compared to the natural gap) leads to less seed loss and more germination for Guarea (a mid-successional species). Our results suggest that a good patch for regeneration of many species in this forest, early- as well as late-successional species, would have high light and a litter layer that moderates extremes.

Effects of light, moisture, temperature, and litter on the regeneration of five tree species in the tropical montane wet forest of Puerto Rico

To study regeneration on landslides, we counted the number of woody stems at least 1 m tall in 2 m × 5 m permanent plots on Puerto Rican landslides for 5 y and generated successional pathways for all plots in 16 different landslides using principal components analysis (PCA). PCA separated plots well, with early successional plant species (Cecropia schreberiana, Schefflera morolotoni, Alchornea latifolia, Gonzalagunia spicata) positively associated with axes I and II, and the negative portion of axis II associated with a mix of those species and mid-successional species (Inga vera, Piper glabrescens). Pathways generally showed (1) a positive association with axis I in early succession, (2) a negative association with both I and II later in succession, (3) little evidence of a decrease in the rate of succession, and (4) a lack of convergence of pathways over the 5 y of our study.

Plant successional pathways on Puerto Rican landslides

. Seed mortality (caused by predators and pathogens) and germination were compared between Puerto Rico and Costa Rica on landslides in lower montane wet forest. Seeds of six common species on five Puerto Rican landslides and four common species on two Costa Rican landslides were used with a Cecropia species and a Gonzalagunia species included at both sites. In the Puerto Rican experiments Cecropia schreberiana was the only species to show significant seed predation (which was due to insects), pathogens grew from all species and fewer seeds were lost to predators than pathogens. Also in Puerto Rico mean germination across all species was 57 % before dispersal (filled seeds collected while still on the tree) and 71 % after, with Phytolacca rivinoides seeds germinating most abundantly, followed in descending order by Ocotea leucoxylon, Cecropia spec, Miconia racemosa, Palicourea riparia and Gonzalagunia spicata. In the Costa Rican experiments three species had significant predation: Cecropia polyphlebia and Urera caracasana (both due to insects) and Witheringia coccoloboides (due to mammals); pathogenic disease caused more seed loss than predation, and germination was high (61 % pre-dispersal, 69 % post-dispersal).



Similarities between these island and mainland sites included (1) percentage of seeds lost to predation and percentage lost to pathogens (all in the 5–15 % range), (2) generalist pathogens which claimed more seeds than predators and (3) majority germination with a general increase after dispersal. Finally sites were dissimilar only in the number of species with significant predation loss and whether it was by insects or mammals, casting doubt on the traditional island/mainland dichotomy.

Randall W. Myster

Papers

Ecosystem development and plant succession on landslides in the caribbean

Tree invasion and establishment in old fields at Hutcheson Memorial Forest

Effects of light, moisture, temperature, and litter on the regeneration of five tree species in the tropical montane wet forest of Puerto Rico

Plant successional pathways on Puerto Rican landslides

Seed predation, disease and germination on landslides in Neotropical lower montane wet forest