“No, Mr Baggins has gone away. Went this morning, and my Sam went with him: anyway, all his stuff went. Yes, sold out and gone, I teller. Why? Why’s none of my business, or yours. Where to? That ain’t no secret. He’s moved to Bucklebury or some such place, way done yonder. Yes it is – a tidy way. I’ve never been so far myself; they’re queer folks in Buckland. No, I can’t give no message. Good night to you!” JRR Tolkien – The Fellowship of the Ring

I’ve always liked this passage where old Gaffer Gamgee is talking, unbeknown, to a nazghul. It is an important story point but delivered in the type of conversation that you could hear all over the world. ‘Those people that live 20 – 30 km away are just so different and weird!‘ Are the people of Buckland really so different to the good, honest folk of the Shire? If so, how did this happen by simply crossing a river?

There is a question around invasive species whether the individuals that arrive in a new area are just a random selection of the individuals (and their traits) that live in their home area or whether they represent a group of individuals with consistent and particular traits that make them more likely to have successfully invaded the new area.

For example, all humans in Aotearoa/New Zealand have arrived from outside these shores over the last 1000 years. Were the people that made their way here more bold and explorative than the rest who stayed behind? Or were they no different than their neighbours who stayed at home? Maybe they just simply had the opportunity to go?

These ideas are important in thinking about why invasive species are successful at establishing or not. If any old random subset of the population can turn up then they are less often going to successful at establishing (they may not be fit-for-purpose!) compared to if they arrive with skills that allow them to survive better in a new environment (or even to survive the journey).

Being large might help give invasive individuals an advantage over native species. Likewise, producing more offspring, growing faster, being bold, exploring more, dispersing sooner, having a broader diet, could all help with invading and establishing.

What about our Bucklanders?

“Long ago Gorhendad Oldbuck, head of the Oldbuck family, one of the oldest in the Marish or indeed in the Shire [has had high evolutionary fitness over many generations], had crossed the river [successfully able to disperse relative to other hobbits and to explore more], which was the original boundary of the land eastwards. He built (and excavated) Brandy Hall, changed his name to Brandybuck, and settled down to become master of what was virtually a small independent country. His family grew and grew [high fecundity in offspring production], and after his days, continued to grow, until Brandy Hall occupied the whole of the low hill, and had three large front-doors, many side-doors, and about a hundred windows. The Brandybucks and their numerous dependants then began to burrow, and later to build, all round about … The people in the Marish were friendly with the Bucklanders … But most of the folk of the old Shire regarded the Bucklanders as peculiar, half foreigners as it were [suggests a slightly different distribution of traits compared to the parent population].Though, as a matter of fact, they were not very different from the other hobbits of the Four Farthings. Except in one point: they were fond of boats, and some of them could swim [bold and innovative behaviours].” JRR Tolkien- The Fellowship of the Ring

We are also told elsewhere that the Brandybucks and Tooks (another bold lineage of hobbits) are generally taller than average Shire hobbits. Tolkien, as I have said in many other places (taxonomy of orcs and hobbits, evolutionary biology ideas, burrow architecture, mammal pest management, fire and ecosystems), was rather accurate when it came to integrating biology into his writing. Did he get it right here?

To test this invasion idea you need a species that is well-studied in it’s native range as well as in its colonising range. You also need to be able to measure all of those traits. Spiders fit the bill nicely. They’re small and have short generations, are easy to fit into small experimental set ups, and some are venomous and, therefore, well studied. Enter the redback spider (Latrodectus hasselti), invasive in Japan and New Zealand and well studied in its Australian homeland.

Cor Vink, New Zealand’s leading arachnologist, joined a group based in Toronto, Canada headed by Monica Mowery, to look at individuals from these three areas. They measured the size of individuals (bigger is usually better in interactions with competitors), their egg sac production (producing more young may give you more opportunities for at least some surviving), and length of generation times in captive populations (shorter allows for faster replacement, longer allows for larger more long-lived individuals).

They measured the behaviour of the redbacks, such as frequency of cannibalism (you never know when a snack might come in handy!). Also, individual spiders were placed in a new environment and the speeds at which they started spinning webs (exploration) or moving after being exposed to a puff of wind (boldness) were measured. Spiders were also placed into a warm arena with a small simulated breeze to see whether they would balloon (effectively float away in the wind) or rappel (climb using their web silk) away from the start point (dispersal).

The outcomes from this work were published in Biological Invasions. Redbacks from the invasive populations showed more dispersal behaviour than the home populations. They also tended to be larger in size, more cannibalistic, and produced more offspring. Interestingly, the redbacks in Japan and New Zealand did not seem to be more bold or explorative than in Aussie. Overall though, the invasive populations looked and acted differently to the source population.

It appears that populations that successfully disperse and establish in new areas might do so because they are settled by individuals with useful traits that differ a little from the source population. This may help us to figure out which species potentially pose the most invasive threats.

What about those strange Bucklanders? The Gaffer was mostly right. They are a bit different. Bucklanders are a population that managed to successfully disperse to an isolated area. Bucklanders are larger and more fecund. Tolkien does not record whether the Bucklanders tended to be more cannibalistic than hobbits in the Shire, but that would be a prediction!

We can certainly sympathise with the Gaffer’s concerns about his Sam going to live among them.