Much of the literature about biological invasion has relied on the “tens rule”. First enunciated in the mid-1990s by Williamson and Fitter (1996), it was actually conceived a decade earlier by Williamson and Brown (1986).
The “tens rule” hypothesizes that about 10% of all species transported to a new environment will be released or escape and become introduced species. Subsequently, 10% of those introduced species establish viable populations in the wild. Finally, about 10% of the established species become highly damaging. That is, 1% of the number originally transported to the new environment is a highly damaging invader.
Is the “tens rule” supported by evidence?
Empirical support for the hypothesis has been mixed; the number of studies questioning it has increased over the decades (Jeschke and Pyšek 2018). So Jeschke and Pyšek (2018) decided to evaluate the basis for the hypothesis. First, they divided the hypothesis into two sub-hypotheses so they could separate the concept of impact from the process of introduction, establishment, and spread. They justified this separation by noting that novel species can have an impact at any stage. The two sub-hypotheses:
1st sub-hypothesis: At each of the three transitions between the invasion stages listed here the number of species completing the transition is reduced by 90% (invasion tens rule).
- transport to exotic range
transition
- introduction (release or escape into the environment)
transition
- establishment of a least one self-sustaining population
transition
- spread
2nd sub-hypothesis: about 10% of established non-indigenous species cause a significant detrimental impact. This sub-hypothesis applies to the transition from establishment (iii, above) to significant impact (iv). Stepping back to the earlier introduction, so as to consider the situation overall, about 1% of all introduced non-native species cause a significant detrimental impact; this sub-hypothesis thus relates to the transition from introduction (ii) to significant impact (iv).
Jeschke and Pyšek carried out a quantitative meta-analysis of 102 empirical tests of the tens rule drawn from 65 publications. They found no support for the “invasion tens rule”. Indeed, their analysis found that about 24% of non-native plant and 23% of non-native invertebrate species are successful in taking consecutive steps of the invasion process. Among non-native vertebrates, about 51% are successful in taking consecutive steps of the invasion process.
The “impact tens rule” is also not supported by currently available evidence. However, Jeschke and Pyšek decided that more data are needed before a reasonable alternative hypothesis can be formulated.
Findings
Jeschke and Pyšek state that the “tens rule” is not based on a model or other defensible concept. It is also hampered by confusion of terms. Thus, different authors define the invasion process differently. Particularly confounding is the mixing of “impact” with steps in the invasion process. At the same time, there have been few studies of the “impact tens rule” hypothesis.
Finally, the “tens rule’s” predictions are not adjusted to consider changes in temporal and spatial scales. That is, it does not recognize that more invaders will be detected in any given place during more recent times than in the past. Furthermore, more invaders will find suitable niches in large areas than small.
The note that analysis is hampered by the paucity of reliable data about establishment success – especially for taxa other than mammals and birds. They do not discuss how this lack might affect efforts to analyze proportions of entering species that succeed in becoming invasive, especially among the small and inconspicuous taxa such as insects and fungal organisms that concern thus of us that focus on threats to forests. This same data gap has limited other studies as well; see, for example, Aukema et al. (2010) – who restricted their discussion of pathogens to “high impact” species.
Although Jeschke and Pyšek (2018) do not specify which studies they relied on to determine the proportion of successful invaders among species belonging to particular taxa, it seems likely that they relied principally on Vila et al. (2010) in determining that on average 25% invertebrates that are introduced (that is, proceed to the second stage in the process given above) become invasive. Vila et al. analyze introductions to Europe. They found that 24.2% of terrestrial invertebrates caused recognized economic impacts.
Jeschke and Pyšek (2018) Results and Discussion
Considering the “invasion tens rule”, two-thirds of the empirical tests in the dataset focused on the “invasion tens rule”. The majority of these focused on the transition from introduction to establishment (the transition from (ii) to (iii). The observed average percentage of species making this transition is more than 40% – or greater than four times larger than the “tens rule’s” prediction.
At the next transition, from establishment to spread (from iii to iv), the observed percentage of species making the transition is greater than 30% – or greater than three times the predicted value under the “tens rule”.
Considering the “impact tens rule”, on average a quarter of established non-indigenous species have a significant detrimental impact, which is again significantly more than the 1 out of 10 species predicted by the rule. Specifically by taxon, 18% of established plants have shown detrimental impacts. Among invertebrates and vertebrates that estimate is greater than 30%. All these observations are higher than predicted by the rule. However, sample sizes are low so more studies are needed to test whether these values hold true.
Regarding the fullest possible extent of the invasion process, 16 out of 100 species that were introduced (stage ii) had a significant impact. This is 16 times greater than the 1% predicted by the “tens rule”. Considering specific taxa, 6% of established plants and 15% of established invertebrates had a significant impact. Data were too poor to support an evaluation for vertebrates.
I note that the alarmingly high “impact” estimates for invertebrates are probably biased by scientists’ and funding entities’ lack of interest in species that don’t cause noticeable impacts.
Poor data preclude an analysis of the transition from transport (i) to introduced (ii).
Strengthening The Estimates
Might these introduction and impact estimates be tightened by analysis of additional sources, such as the studies led Seebens, forest pest impact analyses by Potter et al. (2019) and Fei (2019) and reviews of pest introduction numbers by Haack and Rabaglia (2013)?
Is it worth pursuing efforts to refine the Jeschke and Pyšek (2018) estimates? I think it is. An underestimation of the risk of introduction might lead decision-makers to downplay the need for a response.
Some scientists have accepted the new “rule of 25” (Schulz, Lucardi, and Marsico. 2021. Full citation at end of blog; also cited by USFS report – Poland et al. 2021). Others have not. Venette and Hutchison (2021; full reference at end of blog) continue to cite the estimate of approximately one “invasion success” for every 1,000 attempts – that is, a low-probability, high-consequence event. This challenges those responsible for managing invasive species.
Or are there other conundrums of introduction, establishment, and predicting impacts that have more direct relationship to improving programs? I note that the recent Forest Service report on invasive species (Poland et al. 2021) does not address the “rule of tens”.
Other Reasons Why Bioinvasion Damage is Underestimated
Jaric´ and G. Cvijanovic´ (2012) note that scientists lack a full understanding of ecosystem functioning, so they probably often miss more subtle – but still important – impacts.
Jeschke and Pyšek (2018) note that the percentage of introduced or established species with a quantifiable impact is not always the most important information. A single introduced species can have devastating impact by itself. They cite the amphibian disease chytrid (Batrachochytrium dendrobatidis) and such mammals as rats and cats.
SOURCES
Aukema, J.E., D.G. McCullough, B. Von Holle, A.M. Liebhold, K. Britton, & S.J. Frankel. 2010. Historical Accumulation of Nonindigenous Forest Pests in the Continental United States. Bioscience. December 2010 / Vol. 60 No. 11
Brockerhoff, E.G. and A. M. Liebhold. 2017. Ecology of forest insect invasions. Biol Invasions (2017) 19:3141–3159
Fei, S., R.S. Morin, C.M. Oswalt, and A.M. Liebhold. 2019. Biomass losses resulting from insect and disease invasions in United States forests. Proceedings of the National Academy of Sciences of the United States of America, 12 Aug 2019, 116(35):17371-17376
Haack, R.A. and R.A. Rabaglia. 2013 Exotic Bark and Ambrosia Beetles in the USA: Potential and Current Invaders. CAB International. 2013. Potential Invasive Pests of Agricultural Crops (ed. J. Pena)
Jaric´, I. and G. Cvijanovic´. 2012. The Tens Rule in Invasion Biology: Measure of a True Impact or Our Lack of Knowledge and Understanding? Environmental Management (2012) 50:979–981 DOI 10.1007/s00267-012-9951-1
Jeschke J.M. and P. Pyšek. 2018. Tens Rule. Chapter 13 of book by CABI posted at http://www.ibot.cas.cz/personal/pysek/pdf/Jeschke,%20Pysek-Tens%20rule_CABI%202018.pdf
Poland, T.M., Patel-Weynand, T., Finch, D., Miniat, C. F., and Lopez, V. (Eds) (2019), Invasive Species in Forests and Grasslands of the United States: A Comprehensive Science Synthesis for the United States Forest Sector. Springer Verlag. (in press).
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