The difference between commensalism and habituation, exemplified by Spilopelia chinensis in Australian suburbs

@gposs

Certain species of animals associate particularly with humans, usually for the food inadvertently supplied by an affluent and wasteful primate.

Commensalism (https://en.wikipedia.org/wiki/Commensalism) implies habituation. This is because the association implies proximity.

To see the house sparrow (https://www.inaturalist.org/taxa/13858-Passer-domesticus), pecking crumbs close to a table in an outdoor cafe, surprises nobody who understands that this species is exclusively commensal, never living in the wild (https://en.wikipedia.org/wiki/House_sparrow).

However, some commensal species remain unhabituated, instead remaining as shy of humans as fully wild animals would be. They live among us - indeed, in some situations exclusively among us - but they never get used to us enough to trust us any more than wild animals do.

And this vigilance, apprehensiveness, and distrust may continue, generation after generation, even where humans pay no attention to the animal in question, do nothing deliberately to harm it, and indeed hold it in kind regard.

Such a species is Spilopelia chinensis (https://www.inaturalist.org/taxa/1455918-Spilopelia-chinensis), at least in the urban/suburban environment of Perth, Western Australia, where I live.

This species is common in suburban streetscapes, where its peaceful cooing (https://www.youtube.com/watch?v=WEHe6W_D4lo and https://xeno-canto.org/species/Spilopelia-chinensis and https://avocet.integrativebiology.natsci.msu.edu/recordings/5987 and https://search.macaulaylibrary.org/catalog?taxonCode=spodov&mediaType=audio&sort=rating_rank_desc#_ga=2.93916062.2047640190.1660101380-673334876.1658548323 and https://ebird.org/species/spodov) is a familiar sound, disliked by nobody, and a reminder of the inoffensiveness associated with doves.

Most persons in Australia, including naturalists, take S. chinensis for granted in a neutral or positive way.

However, when one thinks about it, there is something odd in its relationship with us.

Spilopelia chinensis forages exclusively on the ground. It habitually flees from approach at the same distances that any wild, terrestrially foraging columbid would, if approached in some remote area of grassland or savanna. This behaviour seems stereotyped, instinctive, and unaffected by experience.

Australia has a rich indigenous fauna of Columbidae (https://www.inaturalist.org/observations?place_id=6744&taxon_id=2715&view=species).

However, the most successful spp. of columbids in Australian built-up areas are introduced deliberately from elsewhere.

These are:

Playing a minor role in the metropolitan area of Perth are two indigenous species.

Phaps chalcoptera (https://www.youtube.com/watch?v=v8x07ONuZb8 and https://www.inaturalist.org/taxa/3335-Phaps-chalcoptera) was indigenous to what is now the Perth Metropolitan area, before Europeans arrived about 1820.

It has been marginalised by settlement, remaining only in peripheral, densely wooded suburbs. Here it remains shy of humans, and tends to be detected more by its hooting call (https://ebird.org/species/combro1?siteLanguage=en_AU and https://xeno-canto.org/species/Phaps-chalcoptera) than by being actually seen.

Ocyphaps lophotes (https://www.inaturalist.org/taxa/3681-Ocyphaps-lophotes) is similar in body size and diet to S. chinensis. It was indigenous only to the dry interior when Europeans arrived. It has benefited from agriculture, becoming common in the farmed countryside where wheat is grown.

And, like several other ecologically similar indigenous birds, it has tended to expand its range, towards relatively rainy, formerly well-wooded regions, now altered by the human species (https://absa.asn.au/wp-content/uploads/2021/03/Cor-Vol-35-Pg73-81-Created-Pigeons.pdf and https://birdlife.org.au/bird-profile/crested-pigeon).

Ocyphaps lophotes has recently begun to encroach on the Perth metropolitan area, where it seems to be increasing in the southern suburbs. I have lived in this metropolitan area since 1977, but the first time I spotted O. lophotes in the suburbs was in 2019.

However, the indigenous O. lophotes has yet to usurp most of the suburban habitat of the introduced S. chinensis. The former tends to occur in parks, rather than the streetscapes continually exploited by the latter.

And, significantly, O. lophotes already has achieved a level of habituation not seen in S. chinensis.

This is surprising, because S. chinensis

  • is indigenous to parts of Asia that have, for centuries, been heavily populated by humans,
  • was deliberately introduced to Australia as an 'instant feral', mainly for its aesthetic qualities, and
  • has failed to penetrate wild areas in Australia.

The anomaly, then, is that that S. chinensis, a species somewhat co-evolved with humans from the start, actively promoted by humans in the sense of being deliberately introduced, and universally tolerated - if not liked - by the current human population, shows no sign of treating its human benefactors as anything other than potential predators.

How can we explain this anomaly?

In contrast to the failure of S. chinensis to habituate, various birds, indigenous to Australia, have become so habituated that they can be approached closely in suburbs and streetscapes, and even voluntarily approach one to within 1 metre.

Prime examples are Rhipidura leucophrys (https://www.inaturalist.org/taxa/8143-Rhipidura-leucophrys), Gymnorhina tibicen (https://www.inaturalist.org/taxa/8575-Gymnorhina-tibicen), and Grallina cyanoleuca (https://www.inaturalist.org/taxa/8583-Grallina-cyanoleuca).

This lack of shyness is in line with faunas of islands, and is consistent with the relatively small incidence of indigenous predators in the fauna of Australia - the island continent.

So, here is a hypothesis:

Spilopelia chinensis originates in a part of Asia where there is a minimal ethic of conservation. Instead, the cultural norms are such that animals are a resource for consumption by humans. It has long been the custom, in the farmlands of China, to harvest any edible animal, from invertebrates through amphibians and fishes, to birds and rodents.

At the same time, it is the nature of columbids that they are among the less intelligent of birds. They rely not on an ability to learn, but mainly on a combination of fecundity and explosive flight, for their survival.

Because of risks of predation to S. chinensis in its original, anthropogenic habitat, natural selection has 'hardwired' into it an instinctive pattern of wariness and flightiness, in which proximity to people is avoided.

Because S. chinensis has minimal cognitive versatility, the instinctive pattern of anti-predator behaviour has tended to remain unmodified. And this has applied despite generation after generation of actual experience, in a situation free of predation by the human species, that should have taught the bird that it can relax around people.

What we thus find, in the Australian suburbs, is a species with an odd combination of characteristics. Spilopelia chinensis is one of the least approachable species of birds encountered in the metropolitan area: unhabituated even where it is the commonest 'feral' species present.

And this suggests that, eventually, we may be left with the following ironic situation in Perth.

If the indigenous, and not particularly shy, O. lophotes continues to adapt to the suburban environment, as it has been doing in recent decades, then it may eventually replace, and marginalise, S. chinensis on the basis of competition for food.

And, to the extent that this happens, it may transpire that S. chinensis will have come and gone from many Australian streets, its remarkable degree of caution proving to be ultimately futile - and perhaps self-sabotaging - in an adaptive sense.

הועלה ב-אוגוסט 10, 2022 12:16 לפנה"צ על ידי milewski milewski

תגובות

In contrast to Streptopelia chinensis, the Australian species, Ocyphaps lophotes, has shown little potential as an introduced species.

Ocyphaps lophotes was introduced to New Zealand, Europe, and the USA, including Hawaii (see page 231 of https://books.google.com.au/books/about/Introduced_Birds_of_the_World.html?id=fssQAQAAMAAJ&redir_esc=y and https://catalogue.nla.gov.au/Record/174442).

It failed in all of these locations.

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Streptopelia chinensis has been introduced to not only Australia, but also Hawaii, California, Mauritius, New Zealand, New Caledonia, New Britain, Sulawesi, the Moluccas, and Fiji.

It has succeeded in all these areas. This means that S. chinensis occurs in the suburbs of not only the various Australian cities, but also Los Angeles and Auckland.

Reference: pages 220-222 in https://books.google.com.au/books/about/Introduced_Birds_of_the_World.html?id=fssQAQAAMAAJ&redir_esc=y and https://catalogue.nla.gov.au/Record/174442.

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@jeanshelton @paul_contreras @bbunny

Dear observers of Streptopelia chinensis in Los Angeles, do you - like me here in an Australian city - find that this species remains shy and wary of people, not allowing close approach?

פורסם על-ידי milewski לפני בערך 2 שנים

@jeremygilmore
In my suburb in Western Australia, the normal 'flight-threshold' of Streptopelia chinensis is about 10 m. What this means is that, as one walks about in the streets, or goes out from the house into one's garden, any encountered individual(s) of this species of columbid are so wary and lacking in trust that they generally fly off when one is still no closer than 10 m away.

I know of no other terrestrially-foraging bird in the metropolitan environment that shows such 'wildness', which seems fully in line with what it would be in a bird living in a predator-rich environment with no habituation. This is puzzling, because a) this species has a long history of association with humans, and b) there is categorically no persecution of it by humans in this metropolitan area.

My question to you is: in Cape Town, where Streptopelia capicola is indigenous, what would you estimate the 'flight-threshold' to be in its case?

פורסם על-ידי milewski לפני יותר משנה

Hmm, it's not something I've ever given much thought. It would have to be directed at a more bird-oriented person or @tonyrebelo

פורסם על-ידי jeremygilmore לפני יותר משנה

I hardly know the Cape Turtle Dove. We have Redeye and Laughing Doves in our area. Their flight distances vary if you have a dog or if you feed them. And if they are on the ground or in a tree or on a roof. But generally in the region of 5-10m if one approaches casually. But I have never measured it or tried to vary it. One usually encounters them closer in gardens, but they fly off as one arrives. One does not generally encounter them on the ground in urban parkland unless someone is feeding them.

פורסם על-ידי tonyrebelo לפני יותר משנה

@tonyrebelo

Many thanks for your valuable observations.

Streptopelia senegalensis, introduced to Western Australia, is fairly common in the Perth Metropolitan area. Here, it behaves similarly to what you have described, and I do not find it to be anomalously apprehensive.

Streptopelia semitorquata seems, over the last few decades, to have replaced S. capicola in the suburbs of the southwestern Cape of South Africa. I suspect that S. capicola was fairly common in the suburbs of Cape Town during my boyhood, in the 'sixties (see map in http://thebdi.org/2022/03/08/cape-turtle-dove-streptopelia-capicola/).

Streptopelia semitorquata is not really comparable with S. chinensis, because it is partly arboreal in its foraging.

So, I realise that the comparison I have requested is difficult to make.

In order to understand the context:

Several of the terrestrially-foraging birds in metropolitan Australia allow remarkably close approach by humans.

The most striking example in my garden is Grallina cyanoleuca (https://www.inaturalist.org/taxa/8583-Grallina-cyanoleuca), which is so trusting that I would find its behaviour surprising even for a deliberately fed bird. In reality it needs no particular encouragement to approach me as closely as one metre, in my garden.

Other good examples are Rhipidura leucophrys (https://www.inaturalist.org/taxa/8143-Rhipidura-leucophrys), Chenonetta jubata (https://www.inaturalist.org/taxa/7176-Chenonetta-jubata), Eolophus roseicapilla (https://www.inaturalist.org/taxa/144560-Eolophus-roseicapilla), and Cacatua tenuirostris (https://www.inaturalist.org/taxa/116839-Cacatua-tenuirostris).

However, the phenomenon is most striking in cases where there is a close relative in Africa. The local form of Porphyrio (https://www.inaturalist.org/taxa/418530-Porphyrio-melanotus) forages out in the open on lawns in a way seldom seen for its congener in South Africa (https://www.inaturalist.org/taxa/474333-Porphyrio-madagascariensis).

The following is a frequent scene in my local suburban park: https://www.inaturalist.org/observations/66240760.

Given this 'Galapagos-like' tameness on the part of certain birds in Perth, I find it surprising that a species so thoroughly associated with humanity as Streptopelia chinensis is more wild, and less trusting, than any comparable indigenous bird, in the suburbs.

פורסם על-ידי milewski לפני יותר משנה
פורסם על-ידי milewski לפני יותר משנה

In my suburb, both Spilopelia chinensis (which I notice every day) and Spilopelia senegalensis (which I notice every month) occur perennially.

The former is much larger-bodied (about 150 g) than the latter (about 100 g).

There is also a difference in visual 'flight-announcement', as follows.

On my normal approach, Spilopelia chinensis consistently bursts into flight from the ground at about 10 m from me. In the case of the more confiding species, S. senegalensis, the figure is more like 5 m.

When S. senegalensis bursts into flight, the tail is not particularly noticeable, despite the fact that the outer caudal feathers are whitish distally. This is because a) the tail is not fanned during this action, and b) there is no display of dark pigmentation on the tail feathers.

By contrast: when S. chinensis bursts into flight, there is accentuation by means of a) dark/pale contrast, b) fanning of the tail, and c) rocking of the tail. Whitish is restricted (as in S. senegalensis) to the outer (not medial) long feathers. However, what is displayed on these feathers is broadly dark subterminal pigmentation. The tail is simultaneously fanned and rocked as the bird bursts from the ground, the rocking action being rapid and about the longitudinal axis.

What this means is that the species with the more apprehensive reaction to human approach is also the species with the more conspicuous 'flight-announcement'.

However, my impression is that an auditory 'flight-announcement' goes the other way. When S. senegalensis bursts into flight on my approach, the wing feathers make a noticeably high-pitched chuckling sound, so much so that for years I thought that the name 'laughing dove' referred to this (non-vocal and non-visual) effect. When S. chinensis bursts into flight in similar circumstances, there is no such sound.

To hear the auditory 'flight-announcement' of S. senegalensis, listen to the end of the second-last audio recording on the first page in https://xeno-canto.org/species/Spilopelia-senegalensis?language=sv&dir=0&order=cnt, viz. the one by Andrew Spencer in Jemma Valley, Ethiopia.

Also see https://www.perplexity.ai/search/which-spp-of-columbidae-make-a-zP4x10WkQQiWAf5fGD2KwQ and https://www.youtube.com/watch?v=TdVofb6jStM and https://www.youtube.com/watch?v=WKEHUxxmqKA and https://www.youtube.com/watch?v=y9_Hll8sF98 and https://www.youtube.com/watch?v=1N9nMkZt1l0.

To be clear:
S. senegalensis is similar to S. chinensis in possessing conspicuous white on the tail feathers, and this pale signature is conspicuous when S. senegalensis flies about quietly, e.g. from one perch to another during its normal activities. However, when S. senegalensis bursts into flight from the ground on human approach, it seems to be true that the tail is kept folded and the sound of the wings is instead activated.

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@tonyrebelo @jeremygilmore @ludwig_muller @christiaan_viljoen @gposs

Testimony to the importance of search-images for naturalists:

Please see https://xeno-canto.org/species/Spilopelia-senegalensis?language=sv&dir=0&order=cnt.

This is a worthy repository of nearly 200 audio recordings of Streptopelia senegalensis, made throughout the range of this species. The dedication of naturalists is admirable.

However, what is most remarkable is that not one of these recordings targeted the adaptively significant, species-distinctive 'flight-announcement' made by the wing feathers as the bird bursts into flight in mild alarm.

While the vocalisations have been recorded 'ad nauseam', it seems that nobody has set out to record what is perhaps the most interesting sound made by S. senegalensis.

As it happens, there is actually one recording of this 'flight-announcement', at the end of the second-last item on page one. I refer to the recording by Andrew Spencer in Jemma Valley, Ethiopia. However, this seems to have been made merely by accident.

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